United States

Department of Proceedings of Symposium on

Agriculture

Forest Service
Effects of Air Pollutants
Pacific Southwest
Forest and Range
Experiment Station on Mediterranean and
Berkeley
California 94701
Temperate Forest Ecosystems

General Technical
Report PSW-43 June 22-27, 1980, Riverside, California
Acknowledgments:
We gratefully acknowledge major conference support from the following
organizations:
National Committee for Man and the Biosphere, Project 2-Temperate
and Mediterranean Forest Ecosystems, U S . Department of State
Environmental Research Laboratory, U.S. Environmental Protection
Agency, Corvallis
U.S. National Science Foundation
Forest Service, U.S. Department of Agriculture
Other organizations providing additional forms of cosponsorship include:
Statewide Air Pollution Research Center, University of California,
Riverside
Department of Forestry and Resource Management, University of Cali-
fornia, Berkeley
Lawrence Livermore Laboratory, University of California, Livermore
Southern Plains Range Research Station, Agricultural Research, Science
and Education Administration, U.S. Department of Agriculture
Oak Ridge National Laboratory, U.S. Department of Energy
International Union of Forestry Research Organizations
Stan Ranch Audubon Sanctuary, Division of Science and Sanctuaries,
National Audubon Society
Society of American Foresters
State of California Air Resources Board
Many individuals deserve recognition for making the Symposium a suc-
cess. At the University of California's Statewide Air Pollution Research
Center, 0. Clifton Taylor, Neva Friesen, and Merle Johnson deserve special
mention for managing the funding aspects of the conference. As symposium
administrator, Neva Friesen performed many difficult tasks with very effec-
tive results. Other staff who helped during the symposium are Marian Carpe-
Ian, Marjorie McDennont, Joe Lick, Mark Seminoff, Mary Cadman, Donna
Shaw, Dave Lick, Ronald Oshima, Patrick Temple, David Duncan, and Rob
Lennox. From the Pacific Southwest Forest and Range Experiment Station,
Forest Service,' U.S. Department of Agriculture, Berkeley, California, Marcia
Wood handled public information and mass-media contacts; John K.
Publisher:
Pacific Southwest Forest and Range Experiment Station
P.O. Box 245, Berkeley, California 94701
October 1980
McDonald, Vincent Y. Dong, and Audrey F. Hakim managed the details
necessary to the publicationof these proceedings. Anthony Gomez and Robert
Van Doren of the Station's Riverside staff handled the audiovisual require-
ments and assisted with the field trip.
Special credit is due Michael A. Fosberg and Lowell Smith who volun-
teered to substitute for scheduled speakers who canceled within 2 weeks of the
symposium. They also managed toprepare manuscripts on a tight schedule. In
a similar situation, William H. Smith agreed to present a review of the subject
listed as "Forests and Other vegetation as sinks for Air Pollution'' at theiast
minute, in addition to presenting his own scheduled paper.
We are also grateful for the contributions of the Symposium Advisory
Committee at all stages of organization. These individuals are Jay H. Blowers
and Russel M. Bums, both Man and the Biosphere Program (MAB), U.S.
Department of State; Robert Z. Callaham, C. Eugene Conrad, Jimmie L.
Hickman, and Charles W. Philpot, all of the Station staff, Peter F. Ffolliott,
University of Arizona and MAB 2; Jeffrey B. Froke, National Audubon
Society; Herb Hahn, Forest Service and society of American Foresters; James
N. Pitts, Jr., and 0. Clifton Taylor, University of California, Riverside; and
Lawrence C. Raniere, U.S. Environmental Protection Agency.
Technical Program Committee:
Paul R. ~ i i e rPacific
, Southwest Forest and Range Experiment Station,
Forest Service, U.S. Department of Agriculture, Riverside, California
(Chairman)
Patrick I. Coyne, Agricultural Research, Science and Education Adminis-
tration, U.S. Department of Agriculture, Woodward, Oklahoma
Leon S. Dochinger, Northeastern Forest Experiment Station, Forest Serv-
ices, U S . Department of Agriculture, Delaware, Ohio
Joe R. McBride, Department of Forestry and Resource Management,
University of California, Berkeley, California
Samuel B. McLaughlin, Jr., Oak Ridge National Laboratory, U S . De-
partment of Energy, Oak Ridge, Tennessee
David T. Tingey, Cornallis Environmental Research Laboratory, U.S.
Environmental Protection Agency, Corvallis, Oregon
Proceedings of Symposium on

Effects of Air Pollutants

Mediterranean and
Temperate Forest Ecosystems
June 22-27, 1980, Riverside, California

Paul R . Miller
Technical Coordinator

CONTENTS
Introduction
Welcoming Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Forest Genetics and Air Pollutant Stress . . . . . . . . . . . . . . . . . . . 100

James N . Pitts , Jr. Stefan Biafobok

Opening Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Interactions of Air Pollutants and Plant Disease. . . . . . . . . . . . . 103

Robert Z . Callaham Michael Treshow

Air Pollution in Forests: Social Costs, Predictive Models, Studies on Relationship Between Air Pollutants
and Public Policy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
and Microorganisms in Japan . . . . . . . . . . . . . . . . . . . . . . . . . 110

Charles F. Cooper Kiyoshi Tanaka

Historical Perspectives and International Concerns About Sensitivity of Lichens to Air Pollution with an Emphasis
Air Pollution Effects on Forests . . . . . . . . . . . . . . . . . . . . . . . . 10
on Oxidant Air Pollutants . . . . . . . . . . . . . . . . . . . . . . . . . . . .117

Edwin Donaubauer Thomas H. Nash III and Lorene L. Sigal

Influence of Air Pollution on Population Dynamics of Forest
Natural Influences of Forests on Local and Regional Insects and on Tree Mortality . . . . . . . . . . . . . . . . . . . . . . . . . 125

Air Quality Donald L. Dahlsten and David L. Rowney

Emissions and Air Resource Management Within Forests. . . . . . 13 Air Pollutants and Their Effects on Wildlife with Particular
Michael A. Fosberg and Hollis Record Reference to the House Wren (Delichon urbica) . . . . . . . . . . 131

Hydrocarbon Emissions from Vegetation . . . . . . . . . . . . . . . . . . . 24

James R. Newman
David T . Tingey and Walter F . Burns
Background Levels of Trace Elements in Forest Ecosystems . . . 31 Chronic Effects of Acidic Precipitation and Heavy Metals
G . Bruce Wiersma and Kenneth W . Brown on Forest Ecosystems
The Acidity Problem-Its Nature, Causes,
Effects of Chronic Exposures to Gaseous Pollutants and Possible Solutions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136

on Primary Production Processes Lowell Smith

Photochemical Oxidant Impact on Mediterranean and Temperate Acid Precipitation Impact on Terrestrial and Aquatic Systems
Forest Ecosystems: Real and Potential Effects . . . . . . . . . . . . . 38
in Norway . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145

John M. Skelly Lars N. Overrein

Primary Productivity, Sulfur Dioxide, and the Forest Ecosystem: The Impact of Acidic Precipitation and Heavy Metals
an Overview of a Case Study . . . . . . . . . . . . . . . . . . . . . . . . . .51
on Soils in Relation to Forest Ecosystems . . . . . . . . . . . . . . . 152

Allan H . Legge Stephen A. Norton, Denis W . Hanson, and Richard J . Campana

Effects of Airborne F on Forest Ecosystems . . . . . . . . . . . . . . . . 63
Impact of Heavy Metals on Terrestrial and
Robert G . Amundson and Leonard H. Weinstein Aquatic Ecosystems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158

Air Pollution-a 20th Century Allogenic Influence Tom C . Hutchinson

on Forest Ecosystems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
.9
Effects of Acidic Precipitation on Health and the Productivity
William H. Smith ofForests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165

Ellis B. Cowling and Leon S. Dochinger

Secondary and Interactive Effects of Chronic
Gaseous Pollutant Exposure of Producers, Simulation Modeling of the Effects of Chronic Pollutant Stress
Consumers, and Decomposers on Plant Processes and Plant Community Dynamics
Influence of Chronic Air Pollution on Mineral Cycling Modeling Pollutant Uptake and Effects on the Soil-Plant-Litter
inForests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .88
.
System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 174

Paul J . Zinke R. J . Luxmoore

 Data-Based Ecological Modeling of Ozone Air Pollution Effects Estimation of Adverse Effects of Air Pollution
in a Southern California Mixed Conifer Ecosystem . . . . . . . . 181
on Danish Forests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. 37

Ronald N. Kickert and Barbara Gemmill Ib Johnsen

Response of Plant Communities to Air Pollution . . . . . . . . . . . . 187
Changes in Southern Wisconsin White Pine Stands Related
R . Guderian and K. Kueppers to Air Pollution Sensitivity . . . . . . . . . . . . . . . . . . . . . . . . . . . 238

Forecasting Effects of SO, Pollution on Growth and Succession David F.. Karnosky
in a Western Conifer Forest . . . . . . . . . . . . . . . . . . . . . . . . . . 2. 00
+
Effect of 0, and 0, NO2 on Growth of Tree Seedlings . . . . 239

J. R. Kercher, M.C. Axelrod, and G.E. Bingham Lance W . Kress

Forest Models: Their development and Potential Applications Impact of Oxidant Air Pollution on Ponderosa and Jeffrey
for Air Pollution Effects Research. . . . . . . . . . . . . . . . . . . . . .203
Pine Cone Production . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 240

H. H. Shugart, S.B. McLaughlin, and D.C.West Robert F. Luck

Lichens as Air Quality Monitors . . . . . . . . . . . . . . . . . . . . . . . . . 241

P.D. Lulman, R.J. Fessenden, and S.A. McKinnon

Synthesis Session
Acid Precipitation in California and Some Ecological Effects . 242

Opening Remarks and Summary of Panel/Audience

John G . McColl and Mary K. Firestone
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .215

Leaf Litter Decomposition in the Vicinity of a Zinc Refinery . 243

Walter E. Westman
W.D. Mcllveen
Integration: a Role for Adaptive Environmental Assessment
Effects of Chronic Air Pollution Stress on Allocation
andManagement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. 21

of Photosynthate by White Pine . . . . . . . . . . . . . . . . . . . . . . . 244

Nicholas C . Sonntag, Robert R. Everitt. and Michael J. Staley

S.B. McLaughlin, R.K. McConathy, and D. Duvick
Effects of SO2 and Ozone on Photosynthesis and Leaf Growth
Poster Summaries in Hybrid Poplar.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245

Effects of Oxidant Air Pollutants on Pine Litter-fall Reginald D. Noble and Keith F. Jensen
and the Forest Floor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .225
Behavior of Airborne Flourides in Soils . . . . . . . . . . . . . . . . . . . 246

Rodney J. Arkley and Rudolph Glauser Janina Polomski, Hannes Fliihler, and Peter Blaser
Regional Air Pollution Impacts on Forest Growth . . . . . . . . . . . 226
Multiple Pollutant Fumigations Under
Thomas V . Armentano, Orie L. Loucks, and Wayne T. Williams Near Ambient Environmental Conditions
Canopy Analysis of Pollutant Injured Ponderosa Pine Using a Linear Gradient Technique. . . . . . . . . . . . . . . . . . . . . 247

in the San Bernardino National Forest . . . . . . . . . . . . . . . . . .227

P.B. Reich, R .G. Amundson, and J.P. Lassoie
M.C. Axelrod, P.I. Coyne, G,E. Bingham, J.R. Kercher, P.R. Changes in Plant Communities with Distance
Miller, and R.C. Hung from an SO2 Source . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248

Photosynthesis and Stomatal Behavior in Ponderosa Pine Paul R. Scale

Subjected to Oxidant Stress: Water Stress Response . . . . . . . 228
Lichens as Ecological Indicators of Photochemical Oxidant
Gail E. Bingham and Patrick I. Coyne AirPollution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249

Oxidant Impact on Ponderosa and Jeffrey Pine Foliage Lorene L. Sigal and Thomas H . Nash 111

Decomposition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .229
The Effects of Air Pollutants on Forest Ecosystems
J.N. Bruhn, J.R. Parmeter, Jr., and F. W . Cobb, Jr. inS.R.Slovenia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .250

Integrated Lake-Watershed Acidification Study . . . . . . . . . . . . .230

Marjan Solar
Carl W. Chen and Robert A. Goldstein Population Differences in Response to Sulfur Dioxide:
Photosynthesis and Stomatal Response to Light a Physiological Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 252

and Temperature in Ponderosa Pine Exposed to Long-Term G.E. Taylor, Jr. and D.T. Tingey
Oxidant Stress.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. 33
Ozone Injury to Pines in the Southern Sierra Nevada
Patrick I. Coyne and Gail E. Bingham of California . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 253

The Effect of Air Pollution on Western Larch as Detected Detlev R. Vogler and John Pronos
by Tree-Ring Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 234
Modification of Chemical Contents of Precipitation
Carl A. Fox and Thomas H.Nash 111
by Passage through Oak Forests . . . . . . . . . . . . . . . . . . . . . . . 254

Acid Rain: Threshold of Leaf Damage in Eight Species George T . Weaver and Jon D. Jones
from a Forest Succession . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. 35
Seasonal Variation of Inorganic and Organic Sulfur
B.L. Haines, M. Stefani, and F . Hendrix in Coniferous Needles Intensified by SO, Pollution. . . . . . . . 255

Prioritization of Research on Air Quality Related Resources Karl Friedrich Wentzel and Gunther Gasch
of the National Parks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 236
Sulfur Dioxide and Oxidant Effects on Californian
E.A. Howard, L.M. Boyd, W.M. Brock, Coastal Sage Scrub.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 256

D.W. Morse, S.S. Shin, and K.L. Steel-Sabo W.E. Westman and K.P. Preston
PREFACE
Sulfur dioxide from fuel combustion and ore smelt-
ing operations has caused significant damage to forest
communities throughout the industrialized world. In
the temperate regions, notably in Europe, the United
States, Canada, and Japan, examples of damage and
losses resulting from this pollutant are well-
documented. Hydrogen fluoride emissions from alu-
minum reduction plants, brick kilns, and phosphate
fertilizer plants also have caused significant damage in
many localities. In Mediterranean climates, the com-
bination of abundant sunshine and poorly controlled
emissions of nitrogen oxides and hydrocarbons has
resulted in extensive forested regions being exposed to
photochemical oxidant air pollution. Ozone is the most
damaging pollutant in this mixture. Acidic precipita-
tion, derived principally from sulfur oxide emissions,
recently has been shown to have severe effects on
aquatic ecosystems in northeastern United States,
Canada, and northern Europe. The projected increase
in the use of coal for energy generation and the continu-
ing growth of urban centers, accompanied by automo-
bile emissions, are two conditions that suggest a con-
tinuing and more pervasive influence of air pollution
on terrestrial and related aquatic ecosystems.
A large body of knowledge has been assembled that
describes pollutant effects on individual species as a
result of both field observations and controlled exper-
iments. Efforts are being made to use the tools of
systems analysis (modeling) to interpret and predict
pollutant effects on processes at both the individual
species and plant community levels. The ultimate goal
is to improve interpretation of pollutant effects on
ecological systems so that optimal protective and man-
agement measures can be taken to assure a more
healthy environment.
Experimenters and modelers can advance more
rapidly if a better exchange of ideas and essential data
can be stimulated. A symposium was planned to
encourage closer communication between experimen-
talists carrying out specialized studies of the effects of
major air pollutants on individual forest species and
researchers using computer simulation models to
interpret and predict long-term pollutant effects at the
plant community and ecosystem levels. This Sym-
posium, held in Riverside, California, June 22-27,
1980, was designed to report and discuss the state of
knowledge of single species-single pollutant relation-
ships, the interactions of producers, consumers, and
decomposers under pollutant stress, and the use of
ecological systems models for interpretation and pre-
diction of pollutant effects. In addition, the present
state of knowledge was examined in relation to an
overarching ecological concept: resilience of ecosys-
tems. Another important question was the search for
indicators of systems-level effects of air pollution on
ecosystems. For example, is an effect on nutrient cy-
cling a reliable indicator of system-level change in-
duced by pollution?
Twenty-eight papers were presented in the formal
sessions and 29 poster summaries were displayed con-
currently. Registered participants numbered 128. Most
participants attended a field trip to the San Bernardino
mountains for one-half day. Fifteen nations were rep-
resented including Austria, Canada, Czechoslovakia,
Denmark, Egypt, West Germany, Japan, Mexico,
Norway, Poland, Saudi Arabia, Sweden, Switzerland,
United States of America, and Yugoslavia.
To facilitate the publication of the Symposium Pro-
ceedings, we decided to have each author assume full
responsibility for submitting manuscripts in photo-
ready format by the time the conference convened. The
views expressed in each paper are those of the author
and not necessarily those of the sponsoring organi-
zations. Trade names are used solely for necessary
information and do not imply endorsement by the
sponsoring organizations.
Paul R. Miller
Forest Service, U.S. Department of
Agriculture
Technical Coordinator
Introduction
Welcoming ~emarks'
James N . P i t t s , Jr.
Good morning. On behalf of D r . David Saxon,
P r e s i d e n t of t h e U n i v e r s i t y of C a l i f o r n i a , D r .
om& Rivera, Chancellor of UCR, and we of t h e
Statewide A i r P o l l u t i o n Research Center, I would
l i k e t o welcome you t o t h i s i n t e r n a t i o n a l
symposium. We t r u s t you w i l l have a s c i e n t i f i -
c a l l y i n t e r e s t i n g and c h a l l e n g i n g experience
during t h i s week.
The s u b j e c t of t h i s meeting i s timely and
important. The a c c u r a t e assessment of
b i o l o g i c a l , economic, and a e s t h e t i c impacts of
a i r p o l l u t a n t s on f o r e s t ecosystems i s e s s e n t i a l
i f we a r e t o develop c o s t - e f f e c t i v e c o n t r o l
s t r a t e g i e s of a i r p o l l u t i o n . Overcontrol can
l e a d t o economic p e n a l t i e s i n t h e form of e x t r a
c o s t s f o r expensive technologies f o r p o l l u t a n t
removal. On t h e o t h e r hand, undercontrol can
l e a d t o economically unacceptable p l a n t damage
which impacts n o t o n l y o u r a g r i c u l t u r a l and
f o r e s t industry but a l s o our recreational
a c t i v i t i e s . We t r u s t t h i s symposium w i l l
e l u c i d a t e a r e a s of f u t u r e r e s e a r c h t h a t w i l l
provide a more e x t e n s i v e d a t a b a s e upon which
t o g e n e r a t e r e l i a b l e models t h a t can be used f o r
such c o s t - e f f e c t i v e c o n t r o l s t r a t e g i e s .
We i n C a l i f o r n i a a r e p a r t i c u l a r l y s e n s i t i v e
t o t h e t h r e a t , indeed t h e a c t u a l i t y , of s e r i o u s
a i r p o l l u t i o n damage t o c r o p s , f o r e s t s , e t c .
A g r i c u l t u r e remains o u r number one i n d u s t r y ,
with tourism and a s s o c i a t e d r e c r e a t i o n a l
a c t i v i t i e s a l s o making a major c o n t r i b u t i o n t o
t h e economic w e l l b e i n g of o u r s t a t e .
The f a c t t h a t t h e symposium i s being h e l d
here a t UCR seems a p p r o p r i a t e s i n c e i t was a
group of p l a n t s c i e n t i s t s headed by John Middle-
ton who, i n t h e l a t e 1940s, f i r s t showed t h a t
p l a n t damage seen in Los Angeles County was in
f a c t due t o a new type of a i r p o l l u t i o n .
p r e s e n t e d a t t h e Symposium on E f f e c t s of Air
P o l l u t a n t s on Mediterranean and Temperate
F o r e s t Ecosystems, June 22-27, 1980, R i v e r s i d e ,
C a l i f o r n i a , U.S.A.
2 ~ r o f e s s o rof Chemistry and D i r e c t o r , Statewide
A i r P o l l u t i o n Research Center, U n i v e r s i t y of
California, Riverside, California.
2
Through t h e i r e f f o r t s and t h e pioneering re-
s e a r c h of t h e l a t e A r i e Haagen-Smit and o t h e r s ,
i t became c l e a r t h a t we were d e a l i n g w i t h an
o x i d i z i n g atmospheric system formed by t h e
a c t i o n of s u n l i g h t on hydrocarbons and o x i d e s
of n i t r o g e n .
During t h e l a s t twenty y e a r s , much of i t
under t h e l e a d e r s h i p of C l i f Taylor, r e s e a r c h
has been conducted h e r e in two major a r e a s :
S t u d i e s of p o l l u t a n t e f f e c t s on p l a n t s , vegeta-
t i o n , and f o r e s t ecosystems, and t h e chemistry
of a i r p o l l u t i o n . This has been a p a r t i c u l a r l y
u s e f u l combination of i n t e r e s t s because we have
one group of s c i e n t i s t s working on one a x i s of
t h e c l a s s i c dose-response curve, t h a t i s , t h e
atmospheric chemists whose f u n c t i o n i s t o
d e s c r i b e t h e dose received by man, animals, o r
p l a n t s , and a n o t h e r group working on t h e response
axis, the plant scientists investigating the
i n t e r a c t i o n s of a i r p o l l u t a n t s w i t h v e g e t a t i o n .
Most of you a r e f a m i l i a r w i t h t h e work which
has been done h e r e in t h e p l a n t s c i e n c e s a r e a .
Let me j u s t mention t h a t one of t h e major r o l e s
of t h e atmospheric chemists a t t h e Center h a s
been t h e unequivocal s p e c t r o s c o p i c i d e n t i f i c a -
t i o n and measurement of s e v e r a l new gaseous
oxygenated and n i t r o g e n o u s s p e c i e s t h a t a r e
formed in photochemical a i r p o l l u t i o n . These
i n c l u d e formaldehyde, formic a c i d , n i t r i c a c i d ,
n i t r o u s a c i d , and t h e n i t r a t e r a d i c a l , NO3.
Whether o r n o t such gaseous s p e c i e s w i l l prove
t o be s i g n i f i c a n t phytotoxicants is a question
t h a t we l e a v e t o you "response" s p e c i a l i s t s .
I n c l o s i n g I want t o thank D r s . P a u l Miller
and C l i f .Taylor and M r s . Neva F r i e s e n f o r
t h e i r o u t s t a n d i n g e f f o r t s in o r g a n i z i n g t h i s
symposium. Many o t h e r people deserve a g r e a t
d e a l of c r e d i t a s w e l l , b u t t h e r e simply i s n o t
time t o acknowledge them i n d i v i d u a l l y . L e t me
j u s t s a y t h a t w e a r e p l e a s e d t o h o s t a meeting
of t h i s importance and we l o o k forward t o
l e a r n i n g of t h e s i g n i f i c a n t r e s u l t s t h a t w i l l
emerge from your g a t h e r i n g . I am c e r t a i n
t h e s e r e s u l t s w i l l be of l a s t i n g importance t o
t h i s c r i t i c a l a r e a of t h e a i r p o l l u t i o n problem.
 Opening ~emarks'
Robert Z. Callaham
My r e a s o n s f o r being h e r e a r e t o welcome you
on b e h a l f of t h e F o r e s t S e r v i c e , U.S. Department of
A g r i c u l t u r e , and t o i n t r o d u c e you t o t h i s confer-
ence. I want t o e x p l a i n t o you t h e o b j e c t i v e s of
t h e c o n f e r e n c e , why i t was o r g a n i z e d , and who made
i t possible. But I a l s o want t o i n t r o d u c e you t o
each o t h e r , s o t h a t you w i l l know which c o u n t r i e s
you r e p r e s e n t , and why you a r e h e r e .
I w i l l b e g i n by s t a t i n g t h e t h r e e objec-
t i v e s of t h i s c o n f e r e n c e . The f i r s t i s t o review
c u r r e n t i n f o r m a t i o n on s p e c i f i c gaseous and p a r t i c -
u l a t e p o l l u t a n t s and t h e i r e f f e c t s on f o r e s t eco-
systems. F o r e s t s a s s o u r c e s of p o l l u t a n t s and a s
s i n k s f o r p o l l u t a n t s a r e i n c l u d e d . The second
o b j e c t i v e i s t o a n a l y z e primary, secondary, and
i n t e r a c t i v e e f f e c t s of c h r o n i c p o l l u t a n t s on
ecosystems. Modeling w i l l b e explored a s a t o o l t o
s i m u l a t e observed and expected e f f e c t s . L a t e i n
t h e program, s t r a t e g i e s f o r a s s e s s i n g and managing
environmental impacts of a i r p o l l u t a n t s w i l l b e
d i s c u s s e d . The t h i r d o b j e c t i v e i s t o s t i m u l a t e
i n t e r n a t i o n a l communication t o a s s e s s t h e s t a t e of
knowledge and t o i d e n t i f y gaps i n our knowledge.
S e v e r a l y e a r s ago, t h e U.S. Environmental
P r o t e c t i o n Agency (EPA) gave a g r a n t f o r r e s e a r c h
t o D r . C l i f Taylor i n t h e Statewide A i r P o l l u t i o n
Center h e r e a t R i v e r s i d e . The F o r e s t S e r v i c e h a s
a c t i v e l y c o o p e r a t e d and p a r t i c i p a t e d i n t h e r e s u l t -
i n g r e s e a r c h up t o t h e p r e s e n t time. Research
under t h a t g r a n t i s drawing t o a c l o s e . T h i s con-
f e r e n c e was planned, t h e r e f o r e , t o sum up what h a s
been accomplished. S c i e n t i s t s involved i n t h i s
m u l t i m i l l i o n d o l l a r r e s e a r c h e f f o r t have t h i s
o p p o r t u n i t y t o t e l l o t h e r s what t h e y have l e a r n e d
1
P r e s e n t e d a t t h e Symposium on E f f e c t s of
A i r P o l l u t a n t s on Mediterranean and Temperate
F o r e s t Ecosystems, J u n e 22-27, 1980,
R i v e r s i d e , C a l i f o r n i a , U.S.A.
D i r e c t o r , P a c i f i c Southwest F o r e s t and Range
Experiment S t a t i o n , F o r e s t S e r v i c e , U.S.Department
of A g r i c u l t u r e , Berkeley, C a l i f o r n i a .
2
and t o i d e n t i f y problems t h a t remain t o b e s o l v e d .
Another purpose i n o r g a n i z i n g t h i s c o n f e r e n c e i s
t o broaden l o c a l p e r s p e c t i v e s by importing e x p e r t s
from around t h e world t o t a l k about o u r problems.
The f i n a l purpose i s t o t r a n s f e r technology gen-
e r a t e d by t h i s r e s e a r c h program t o t h e managers of
a i r , l a n d , and f o r e s t r e s o u r c e s .
T h i s c o n f e r e n c e was made p o s s i b l e through t h e
c o o p e r a t i v e e f f o r t s of s e v e r a l a g e n c i e s . The
F o r e s t S e r v i c e , U.S. Department of A g r i c u l t u r e ,
and t h e S t a t e w i d e A i r P o l l u t i o n Research C e n t e r ,
U n i v e r s i t y of C a l i f o r n i a , a t R i v e r s i d e , a r e spon-
s o r s . IUFRO, t h e I n t e r n a t i o n a l Union of F o r e s t r y
Research O r g a n i z a t i o n s , having about 380 member
o r g a n i z a t i o n s i n 86 c o u n t r i e s around t h e world and
a s u b j e c t group concerned w i t h a i r p o l l u t i o n , i s a
cosponsor. The U.S. Department of Energy and t h e
U.S. Environmental P r o t e c t i o n Agency have made
t h e i r p e o p l e and t h e i r r e s o u r c e s a v a i l a b l e . EPA
w i l l h e l p t h e F o r e s t S e r v i c e t o p u b l i s h t h e pro-
ceedings. The U.S. Department of S t a t e , p a r t i c u -
l a r l y t h e Man and t h e Biosphere Program through
i t s P r o j e c t 2 ~ M e d i t e r r a n e a nand Temperate F o r e s t
Ecosystems--has provided f i n a n c i a l s u p p o r t .
UNESCO, t h e i n t e r n a t i o n a l home of t h e Man and t h e
Biosphere Program, h a s p a i d t o b r i n g t h r e e i n t e r -
n a t i o n a l p a r t i c i p a n t s h e r e . For a l l of t h i s sup-
p o r t and c o o p e r a t i o n , t h e o r g a n i z e r s a r e most
gratef ul.
My f i n a l and, perhaps, unexpected r e a s o n f o r
being h e r e i s t o i n t r o d u c e you t o each o t h e r .
Although t h i s i s n o t o f t e n done a t c o n f e r e n c e s , I
have found i t t o b e a n e f f e c t i v e means of stimu-
l a t i n g communication. I am going t o c a l l t h e r o l l
of c o u n t r i e s , more o r l e s s i n a l p h a b e t i c a l o r d e r .
and a s k t h e i n d i v i d u a l s named t o s t a n d . (Intro-
d u c t i o n s followed.) About 20 p e r c e n t of t h e
people h e r e a r e from o u t s i d e North America. L e t
u s g i v e t h e s e v i s i t o r s a s p e c i a l welcome. Walk up
t o them. I n t r o d u c e y o u r s e l v e s . Ask t h e s e v i s i -
t o r s about programs and problems i n t h e i r
countries.
Now t h a t we know what c o u n t r i e s a r e r e p r e s e n t e d ,
l e t me c a l l f o r a show of hands t o f i n d o u t why
130 of you a r e h e r e . How many a r e p r i m a r i l y
t e a c h e r s o r p r o f e s s o r s ? About 7 p e r c e n t . How
many a r e managers of l a n d o r f o r e s t r e s o u r c e s ?
About 9 p e r c e n t . How many a r e managers of a i r re-
sources? Only a b o u t 2 p e r c e n t . That i s s u r p r i s -
ing. How many of you a r e s t u d e n t s , n o t y e t i n t o
p r o f e s s i o n a l a c t i v i t i e s ? About 2 p e r c e n t . The
remainder of y o u ~ a b o u t80 p e r c e n t ~ a r es c i e n t i s t s
and i n v e s t i g a t o r s . That i s a b o u t what I expected.
And now my r o l e i s f u l f i l l e d . I have
i n t r o d u c e d you t o t h e c o n f e r e n c e and t o each o t h e r .
I e x p e c t you t o b e n e f i t b o t h p r o f e s s i o n a l l y and
p e r s o n a l l y from t h i s c o n f e r e n c e . L a s t l y , I
e x p r e s s my deep a p p r e c i a t i o n t o a l l who have con-
t r i b u t e d t o t h e o r g a n i z a t i o n of t h i s c o n f e r e n c e
and p a r t i c u l a r l y t o Dr. P a u l M i l l e r .
On b e h a l f of t h e F o r e s t S e r v i c e , I t h a n k you
f o r coming. The u l t i m a t e s u c c e s s and meaning of
t h i s c o n f e r e n c e depend on you.
 Air Pollution in Forests: Social Costs,
Predictive Models, and Public ~ o l i c y '
C h a r l e s F. Cooper 2

A b s t r a c t : Long t i m e s c a l e s , s p a t i a l v a r i a t i o n i n ecosystems, and d i f f e r i n g v a l u e judgments make

models almost e s s e n t i a l f o r s o c i e t a l consensus about a i r p o l l u t i o n . Three c a t e g o r i e s of
p o l i c y - o r i e n t e d models a r e d e s c r i b e d . Empirical time s e r i e s models a r e good f o r immediate
d e c i s i o n s but a r e i n h e r e n t l y a short-term device. D e t a i l e d s t r u c t u r a l - f u n c t i o n a l models
emphasize r e l a t i o n s h i p s among components and demonstrate t h e s i g n i f i c a n c e of i n t e r c o n n e c t i o n s .
Small e r r o r s , however, can l e a d t o e r r o n e o u s q u a n t i t a t i v e r e s u l t s , l i m i t i n g t h e i r v a l u e f o r
direct policy decisions. Aggregated p o l i c y - o r i e n t e d models p r o v i d e b e t t e r compliance between
model o u t p u t and v a l i d a t i o n d a t a a t t h e c o s t of l o s s of r e s o l u t i o n . Good models should be
c l e a r l y documented, r e s u l t s s h o u l d be comprehensible, l i m i t s and probable e r r o r bands c l e a r l y
s t a t e d , t h e y should be f l e x i b l e enough t o d e a l with u n a n t i c i p a t e d problems without a t t e m p t i n g
t o t a l g e n e r a l i t y , and r e s u l t s should be c l e a r l y d i s p l a y e d . A model is a n a i d t o decisionmaking,
not a d e c i s i o n maker. For i t t o be e f f e c t i v e i n t h a t r o l e , t h e r e must be mutually s u p p o r t i v e
i n t e r a c t i o n among modelers, b i o l o g i c a l and s o c i a l s c i e n t i s t s , and d e c i s i o n makers. Perhaps t h e
most s i g n i f i c a n t r o l e of models i s i n h e l p i n g t o avoid s u b o p t i m i z a t i o n and i n f a c i l i t a t i n g
communication among d i s c i p l i n e s and p r a c t i t i o n e r s .

Atmospheric p o l l u t i o n i s a f f e c t i n g f o r e s t The e c o l o g i c a l and s o c i a l consequences of a i r

ecosystems i n much of t h e world. A major purpose p o l l u t i o n i n f o r e s t s a r e t h e r e s u l t of complex
of t h i s symposium is t o e s t a b l i s h a s c i e n t i f i c i n t e r a c t i o n s of p r o c e s s e s w i t h many t e m p o r a l ,
consensus about t h e n a t u r e , magnitude, and t i m e s p a t i a l , and v a l u e s c a l e s . Long-t ime s c a l e s ,
t r e n d of t h e s e e f f e c t s . A scientific resolution, v a r i a t i o n s among ecosystems, and d i f f e r e n c e s of
however, is n o t enough. A n a l y s i s of "The E f f e c t s o p i n i o n about v a l u e s make c l a s s i c a l l a b o r a t o r y
of A i r P o l l u t a n t s on Mediterranean and Temperate e x p e r i m e n t a t i o n almost u s e l e s s f o r d e f i n i n g t h e s e
Forest Ecosystems" must also take societal large scale consequences. Models are thus
o b j e c t i v e s and l i m i t a t i o n s i n t o a c c o u n t . e s s e n t i a l f o r helping t o a r r i v e a t a s o c i e t a l
consensus about how t o t r e a t a i r p o l l u t i o n .
Ecosystems used and enjoyed by man a r e
embedded i n a l a r g e r s o c i a l system; d e a l i n g w i t h COMPLEXITY OF THE FOREST AIR POLLUTION PROBLEM
e f f e c t s of a i r p o l l u t i o n on t h e s e ecosystems,
l o c a l l y , r e g i o n a l l y , o r g l o b a l l y , t h u s becomes a It i s a t r u i s m t h a t t h e f o r e s t a i r pollution
q u e s t i o n of p u b l i c p o l i c y . E s s e n t i a l t o sound problem is complex, but t h e n a t u r e of that
p u b l i c p o l i c y f o r m u l a t i o n is knowledge of t h e complexity must be understood i f we are t o
s o c i a l c o s t s of a i r p o l l u t i o n and i t s c o n t r o l . A v i s u a l i z e t h e r o l e of models i n d e a l i n g with i t .
m a j o r problem, of c o u r s e , i s how t o measure t h e s e The p o i n t s mentioned h e r e a r e e l a b o r a t e d by o t h e r
costs. T h i s symposium should l e a d u s some way a u t h o r s i n t h i s volume.
toward b e t t e r assessment of t h e r e a l c o s t s of
forest a i r pollution. Air pollution affects individual plants
d i r e c t l y , i n ways which change d u r i n g t h e c o u r s e of
the plant's l i f e history. It a f f e c t s p l a n t s
i n d i r e c t 1y, through impacts on s o i l s and on
p r e s e n t e d a t t h e Symposium on E f f e c t s of A i r consumers and decomposers. E f f e c t s on p l a n t s , i n
P o l l u t a n t s o n M e d i t e r r a n e a n and Temperate F o r e s t turn, a r e reflected i n other trophic levels.
Ecosystems, J u n e 22-27, 1980, R i v e r s i d e , C a l i f o r - C r i t i c a l l y important i s t h e dynamic r e a c t i o n of
n i a , U.S.A. ecosystems, which u s u a l l y cannot be p r e d i c t e d from
a simple summation of t h e r e s p o n s e s of i n d i v i d u a l
2 ~ r o f e s s o rof Biology and D i r e c t o r of C e n t e r f o r organisms .
Regional Environmental S t u d i e s , San Diego S t a t e
University, SanDiego, California 92182. Social consequences stem from loss of
p r o d u c t i v e r e s o u r c e s and amenity v a l u e s . Wood and
forage growth may d i m i n i s h , a l o s s t h a t w i l l be judgment, h o p e f u l l y good, of t h e people, hopeful 1 y
increasingly s i g n i f i c a n t i f f o r e s t s a r e i n greater experienced, i n charge. T h i s Is t h e common
demand f o r biomass a s a s o u r c e of energy o r procedure. There is, however, a more organized
s t r u c t u r e d chemicals, o r i f i n c r e a s i n g need f o r s c i e n t i f i c approach t o complex problems with many
g r a i n a s food l e a d s t o more p r e s s u r e on rangelands. temporal and s p a t i a l s c a l e s . T h i s process i n c l u d e s
A e s t h e t i c and r e c r e a t i o n a l v a l u e s a r e l o s t . The eight basic s t e p s .
degraded appearance of t h e smog-affected f o r e s t s of
t h e San Gabriel Mountains a r e apparent t o a l l who 1. Make a model of t h e p r o c e s s , based on
have seen them. W i l d l i f e , both game and non-game, e x i s t i n g knowledge and understanding of t h e
may s u f f e r . Lakes and s t r e a m s i n s e v e r a l p a r t s of system. The k i n d s of model which might be
North America and Europe have l o s t much of t h e i r undertaken i n t h i s s t e p a r e d i s c u s s e d i n more
c a p a c i t y f o r f i s h production (Loucks 1980). Of d e t a i l below.
course, there may be beneficial impacts a s
well--a1 l e v i a t i o n of l o c a l s u l f u r d e f i c i e n c i e s , f o r 2. F i t t h e parameters of t h e model t o d a t a ,
instance. Both b e n e f i c i a l and d e t r i m e n t a l e f f e c t s p r e f e r a b l y obtained from l a b o r a t o r y o r f i e l d
vary i n t i m e and s p a c e . experimentation; o t h e r w i s e from o b s e r v a t i o n a l

A i r p o l l u t i o n o p e r a t e s a t many time s c a l e s .
studies .
Hol l i n g (1973) h a s d i s t i n g u i s h e d between f a s t and 3 . V a l i d a t e t h e model. T h i s i n v o l v e s comparison
slow v a r i a b l e s . Fast variables a r e generally of model r e s u l t s with r e a l world outcomes i n
amenable to conventional laboratory systems o t h e r t h a n t h o s e used i n f i t t i n g t h e
e x p e r i m e n t a t i o n , and a r e t h e kind t h a t a r e u s u a l l y parameters. T h i s is a c r u c i a l but a l s o a
studied i n biological research. Slow v a r i a b l e s , most d i f f i c u l t s t e p , because t h e o b j e c t i v e of
however, t a k e long enough t o m a n i f e s t themselves t h e whole modeling e x e r c i s e i s o f t e n t o
that controlled experimentation is impractical i n p r e d i c t r e s p o n s e s of systems under s t r e s s e s
many r e a l world s i t u a t i o n s where a c t i o n cannot t h a t exceed t h e r a n g e of e x i s t i n g v a l i d a t i o n
wait. Decision makers must a l s o d e a l with a high data.
degree of s p a t i a l h e t e r o g e n e i t y i n both p o l l u t a n t s
and t h e i r t a r g e t ecosystems. Theory and r e s e a r c h 4. Test the sensitivity of t h e model to
i n ecology h a s n o t u n t i l now d e a l t very well with parameter changes. T h i s can h e l p t o l o c a t e
s p a t i a l processes. c r i t i c a l f e a t u r e s where b e t t e r understanding
o r more a c c u r a t e d a t a a r e needed. It can
F i n a l l y , a i r p o l l u t i o n i s o n l y one of many a l s o h e l p l o c a t e p a r t s of t h e system where
stresses a f f e c t i n g f o r e s t ecosystems. Multiple r e l a t i v e l y small changes may have l a r g e
s t r e s s e s may i n t e r a c t s y n e r g i s t i c a l l y o r they may effects. S e n s i t i v i t y a n a l y s i s is o f t e n s a i d
h e l p t o c o u n t e r a c t one a n o t h e r . We need t o know t o be one of t h e g r e a t v i r t u e s of a modeling
more, f o r i n s t a n c e , about t h e combined impact of approach, i n t h a t i t l e a d s t o d i r e c t i n g
a i r p o l l u t i o n and c l i m a t e change, whether due t o l i m i t e d r e s o u r c e s t o a r e a s where t h e y w i l l do
d e l i b e r a t e weather m o d i f i c a t i o n o r i n a d v e r t e n t t h e most good, o r c o n v e r s e l y a v o i d s t h e
c l i m a t i c change. Increased atmospheric carbon e x p e n d i t u r e of e f f o r t on measures u n l i k e l y t o
d i o x i d e from burning of f o s s i l f u e l seems l i k e l y t o have much effect. Points of special
warm t h e e a r t h ' s c l i m a t e and, perhaps, t o s t i m u l a t e s e n s i t i v i t y a r e of t e n hard to f i n d , however.
p l a n t growth d i r e c t l y . How w i l l t h e s e p r o c e s s e s Both c o n t r o l theory and p r a c t i c a l e x p e r i e n c e
i n t e r a c t with a i r p o l l u t a n t s ? F o r e s t h a r v e s t and a r e i n c r e a s i n g l y demonstrating t h a t many
r e g e n e r a t i o n is i t s e l f a s t r e s s on t h e ecosystem complex i n t e r 1 inked systems a r e r e l a t i v e 1 y
which w i l l i n t e r a c t with increased pollutant i n s e n s i t i v e t o small changes i n one o r two
l o a d i n g . A i r p o l l u t i o n i n f o r e s t s t h u s i s p a r t of variables. T h i s should come a s no s u r p r i s e
a complex network of biological and social t o t h o s e who have observed t h e e v i d e n t
i n t e r a c t i o n s whose i n t e g r a t e d impacts a r e almost resilience of ecosystems under stress
impossible to untangle through single-factor (Holling 1973). Sensitivity analysis
analysis. remains, nevertheless, an important
a p p l i c a t i o n of p o l i c y - o r i e n t e d models.

A SCIENTIFIC APPROACH TO COMPLEXITY 5. Use t h e model o u t s i d e t h e r a n g e s of s t r e s s e s

p r e v i o u s l y experienced. One of t h e p i t f a l l s
There are some four possible societal t h a t a l l of us have been warned t o avoid i n
responses to complex problems such as air s c i e n c e i s e x t r a p o l a t i o n , yet i t is j u s t
pollution. We can a t t e m p t t o t r e a t t h e symptoms because of t h e need f o r e x t r a p o l a t i o n t h a t
through such means a s f e r t i l i z a t i o n o r i r r i g a t i o n , models a r e c a l l e d f o r i n p r e d i c t i n g ecosystem
a l l e v i a t e t h e cause through emission c o n t r o l , consequences of a i r p o l l u t i o n . The response
accept t h e degradation a s gracefully a s possible, of t h e model system w i l l u s u a l l y need t o be
o r c o n v e r t t h e a f f e c t e d ecosystem t o one more e s t i m a t e d under p o l l u t i o n l o a d s g r e a t e r t h a n
r e s i s t a n t t o s t r e s s . Actual p o l i c y s o l u t i o n s will t h o s e a l r e a d y experienced by t h a t system.
probably i n c l u d e some combination of t h e s e . Even more important i s t h e time dimension. A
primary goal o f t e n i s assessment of t h e
How do we go about choosing the- a p p r o p r i a t e long-term consequences of c h r o n i c o r e p i s o d i c
p o l i c y response? One way i s simply t o r e l y on t h e a i r p o l l u t i o n . It is j u s t because of t h i s
 extended t i m e dimension t h a t models a r e KINDS OF POLICY-ORIENTED MODELS
needed, and yet t h i s is perhaps t h e most
d i f f i c u l t element i n t h e i r c o n s t r u c t i o n . Models may be merely conceptual and verbal--an
i n t u i t i v e , possibly q u i t e accurate, v i s u a l i z a t i o n
6. Array t h e output f o r p u b l i c d i s c u s s i o n . of how t h e world works. We a r e concerned h e r e ,
S i n c e t h e purpose of t h e models we a r e however, with mathematical models a b l e t o d e a l with
c o n s i d e r i n g h e r e is t o h e l p i n a r r i v i n g a t i n t e r a c t i o n s among more v a r i a b l e s t h a n t h e unaided
some s o r t of consensus about a p p r o p r i a t e human mind can r e a d i l y handle. These a r e of
societal response, p r e s e n t a t i o n must go s e v e r a l b a s i c kinds, which d i f f e r i n both t h e i r
beyond t h e immediate s c i e n t i f i c community. underlying structure and their range of
Seldom, if e v e r , w i l l t h e output of a application. (This section owes much to
realistic air p o l l u t i o n model lead to d i s c u s s i o n s with W. R. Emanuel , Environmental
deterministic predictions. Rather, t h e r e Sciences D i v i s i o n , Oak Ridge National L a b o r a t o r y . )
w i l l be a range of a l t e r n a t i v e outcomes, each
with a p r o b a b i l i t y l e v e l a t t a c h e d . It i s 1. Empirical time series a n a l y s i s . The emphasis
notoriously difficult to interpret risk h e r e i s on a n a l y s i s of t h e s e c u l a r t r e n d of
p r o b a b i l i t i e s i n terms of p u b l i c a t t i t u d e s . t h e v a r i a b l e s of i n t e r e s t . It i s assumed
There is a growing literature on t h a t t h e processes d u r i n g t h e period of
p r o b a b i l i s t i c r i s k assessment which i s h i g h l y record w i l l c o n t i n u e over t h e i n t e r v a l of
p e r t i n e n t t o t h e a i r p o l l u t i o n problem (e.g., extrapolation. A time s e r i e s model need not
Kates 1978, S t a r r and Whipple 1980). include e x p l i c i t casual relationships. It
must, however, i n c o r p o r a t e s u f f i c i e n t d a t a t o
7. Amalgamate with output of o t h e r r e l e v a n t e s t a b l i s h t h e s t a t i s t i c a l s i g n i f i c a n c e of t h e
models of s o c i e t a l i s s u e s f o r f i n a l p u b l i c observed p a t t e r n s . I n t h e words of Dennis
evaluation. A i r p o l l u t i o n i s only one of Meadows (1975), i t i s "data r i c h , t h e o r y
many problems f a c i n g s o c i e t y . Measures t a k e n poor. "
to alleviate the consequences of air
p o l l u t i o n a r e l i k e l y t o ramify i n t o many Such a time s e r i e s a n a l y s i s is o f t e n i d e a l
o t h e r a s p e c t s of s o c i e t y . I n my view, t h e f o r d e c i s i o n s which must be made immediately
most important single use of a but which can be revoked i n t h e l i g h t of new
policy-oriented model is a s an a i d i n information without lasting damage,
avoiding suboptimization. By subopt imiza- biological, or political. Time series
t i o n , of c o u r s e , i s meant choosing what is a n a l y s i s h a s t h e advantage t h a t i t i s e a s i l y
c l e a r l y and l o g i c a l l y t h e best s o l u t i o n t o a understood by d e c i s i o n makers who a r e not
s m a l l p a r t of a problem without adequately analytically inclined. It i s i n h e r e n t l y a
c o n s i d e r i n g t h e impact of t h a t s o l u t i o n on short-term t o o l , however. Lack of e x p l i c i t
the total system. An example of how c a s u a l r e l a t i o n s h i p s makes e x t r a p o l a t i o n even
s u b o p t i m i z a t i o n along a narrow path may t u r n more r i s k y t h a n with o t h e r models.
o u t t o be n o t j u s t s l i g h t l y wrong, but
e x a c t l y wrong i n a broader c o n t e x t is t h e 2. D e t a i l e d s t r u c t u r a l - f u n c t i o n a l models. These
d i s p o s a l of chemical wastes a t Love Canal, i n c o r p o r a t e t h e s t r u c t u r e and f u n c t i o n of t h e
N. Y. Out-of - s i g h t , out-of -mind b u r i a l was a system t o t h e e x t e n t t h a t it is known. There
good s o l u t i o n a t t h e time f o r t h e p o t e n t i a l i s a wide v a r i e t y of suggested procedures and
hazard t o workers and t h e p u b l i c of t h i s approaches f o r c o n s t r u c t i n g such models;
m a t e r i a l ; i t s consequences a r e now a f f e c t i n g s e v e r a l a r e discussed i n t h i s volume. The
a l l Americans, a s t a x p a y e r s , i f not a s emphasis throughout i s on u n d e r s t a n d i n g
r e c i p i e n t s of d i r e c t chemical i n s u l t . Less r e l a t i o n s h i p s and p r o c e s s e s , n o t t r e n d s . In
extreme c a s e s of s u b o p t i m i z a t i o n may be more Meadows' (1975) words, t h e y a r e "theory r i c h ,
d i f f i c u l t t o i d e n t i f y b e f o r e a c t i o n is taken. d a t a poor."
I f a p r o p e r l y designed model, by e x p l o r i n g a
w i d e r range of a l t e r n a t i v e s t h a n can t h e T h e i r p r i n c i p a l v a l u e i s a s an a i d t o
human mind alone, h e l p s t o avoid t h e understanding r e l a t i o n s h i p s among components
long-term costs of suboptimization, the and t h e s i g n i f i c a n c e of I n t e r c o n n e c t i o n s .
e f f o r t i n i t s c o n s t r u c t i o n w i l l be well They a r e o f t e n u s e f u l f o r p o i n t i n g out t o
rewarded. d e c i s i o n makers why c e r t a i n r e l a t i o n s h i p s
whose importance is not i n t u i t i v e l y obvious
8. Move toward a d e c i s i o n . There is no hope a r e a c t u a l l y more s i g n i f i c a n t t h a n they seem.
t h a t a d e c i s i o n , even i f based on t h e best
conceivable model, will satisfy all Proper1 y c o n s t r u c t e d f u n c t i o n a l models can
interested p a r t i e s i n a controversial issue. l e a d t o p r e d i c t i o n of ecosystem r e s p o n s e s t o
One c o u l d e x p e c t , however, t h a t t h e d e c i s i o n s t r e s s e s , which a r e l i k e l y t o d i f f e r markedly
would be more r a t i o n a l t h a n i f based on from t h o s e of i n d i v i d u a l organisms t e s t e d i n
emotion and maximization of each isolation. West and o t h e r s (1980) used a
participating individual's personal model of s u c c e s s i o n a l dynamics t o t e s t t h e
objectives. long-term impact of a i r p o l l u t i o n on e a s t e r n
deciduous f o r e s t s . T h e i r model p r e d i c t e d
 enhanced growth of some s p e c i e s d e s p i t e provides a q u a n t i t a t i v e framework f o r t h e
p o l l u t a n t stress, s i n c e they may g a i n a t a s k faced by a l l d e c i s i o n makers, t h a t of
c o m p e t i t i v e advantage because they a r e l e s s achieving an a c c e p t a b l e compromise among a
s e n s i t i v e t h a n o t h e r s p e c i e s with which t h e y s e t of competing o b j e c t i v e f u n c t i o n s . The
i n t e r a c t i n t h e s u c c e s s i o n a l process. v a r i o u s a n a l y t i c a l techniques used f o r t h i s
purpose a g r e e i n using t h e model t o o f f e r
Neither e c o l o g i c a l models, nor e c o l o g i c a l tradeoff f u n c t i o n s of some kind t o t h e
t h e o r y i n g e n e r a l , d e a l well with s p a t i a l d e c i s i o n maker. But h e , not t h e model o r t h e
dynamics i n f o r e s t s . Most emphasis h a s been modeler, e s t a b l i s h e s p r i o r i t i e s among t h e
on s u c c e s s i o n a l dynamics over time a t a p o i n t planning c r i t e r i a . The model is t h u s j u s t
o r i n a small a r e a . There have been one more t o o l f o r bringing p r e c i s i o n i n t o t h e
a t t e m p t s , a s by Shugart and o t h e r s (1973) t o planning process and f o r e v a l u a t i n g t h e
model t h e "flow" of one form of land use o r consequences of a l t e r n a t i v e c h o i c e s .
v e g e t a t i o n c o n d i t i o n t o a n o t h e r , but t h i s
approach is c h i e f l y u s a b l e f o r very l a r g e Ecological policy models would be
units. Because air pollution i s both s u b s t a n t i a l l y improved i f t h e y could d i r e c t l y
s p a t i a l l y extended and s p a t i a l l y v a r i a b l e , i n c o r p o r a t e human d e c i s i o n making i n response t o
t h e r e is a need t o i n c o r p o r a t e t h e s e f e a t u r e s ecosystem change. A c t i v e r e s e a r c h is i n p r o g r e s s
i n t o e c o l o g i c a l models of a i r p o l l u t i o n . A t o make t h i s p o s s i b l e . For i n s t a n c e , C. L. Smith,
promising approach seems t o be t h e l i n k i n g of J. M. Stander, and A. V. Tyler (personal
existing forest succession models with communication 1980), of Oregon S t a t e U n i v e r s i t y , a n
c a r t o g r a p h i c models developed by geographers anthropologist, an ecosystem modeler, and a
f o r dynamic map a n a l y s i s . E f f o r t s t o do t h i s fisheries biologist, have collaborated in
a r e now underway i n s e v e r a l r e s e a r c h u n i t s . developing an i n t e r a c t i v e model of a mythical human
The r e s u l t s should be v a l u a b l e f o r a i r h u n t i n g and g a t h e r i n g s o c i e t y and an e x p l o i t e d
pollution studies. fishery. Human participants are faced with
a l t e r n a t i v e s which f o r c e them t o make c h o i c e s . The
D e s p i t e t h e i r g r e a t v a l u e f o r many purposes, consequences of t h e c h o i c e s a r e t h e n e v a l u a t e d by
however, detailed structural-functional t h e models i n accordance with previous1 y developed
models a r e u s u a l l y u n s u i t a b l e f o r d e c i d i n g d e c i s i o n r u l e s . They found t h a t d e c i s i o n making i n
upon s p e c i f i c a c t i o n s o r p o l i c i e s . Small simulations did alter model outcomes.
e r r o r s , e i t h e r i n t h e o r e t i c a l underpinnings Decisionmaking i n t h e f i s h e r y s i m u l a t i o n l e d t o a
o r i n parameter e s t i m a t i o n , can l e a d t o s t a b l e e q u i l i b r i u m ; without i t , t h e r e was p e r i o d i c
q u a n t i t a t i v e p r e d i c t i o n s t h a t t u r n out t o be c y c l i n g of f i s h i n g v e s s e l s and f i s h biomass. Wrong
q u i t e wrong when t e s t e d a g a i n s t t h e l i m i t e d decisions, though, l e d t o extinction-economic
validation data usually available. We can e x t i n c t i o n of the fishery before biological
hope, though, that improvement i n both e x t i n c t i o n of the fish. Similar i n t e r a c t i v e
modeling t e c h n i q u e and i n b i o l o g i c a l and modeling i d e a s a r e being developed by H o l l i n g
s o c i a l knowledge w i l l l e a d t o f u n c t i o n a l (1978) and h i s a s s o c i a t e s and f o l l o w e r s under t h e
models t r u l y u s e f u l a s decision tools. r u b r i c of Adaptive Environment Assessment.

3. Aggregated pol icy-orient ed models. Here,

t h e r e i s a n attempt t o combine t h e many DESIRABLE CHARACTERISTICS OF MODELS
s t r u c t u r a l elements of t h e system i n t o a
r e l a t i v e 1 y few we1 1-understood components f o r To be e f f e c t i v e t o o l s f o r a s s i s t i n g i n p o l i c y
which good cause and e f f e c t d a t a e x i s t . The decisions, models should have several
major s t r u c t u r a l and f u n c t i o n a l r e l a t i o n s h i p s characteristics (Cooper 1976). It i s almost
a r e preserved, but a t a lower l e v e l of t o t a l l y i r r e l e v a n t i n t h i s c o n t e x t whether t h e
resolution with respect to their model uses d i f f e r e n t i a l o r d i f f e r e n c e e q u a t i o n s o r
interconnections. F u l l understanding of t h e whether i t i s w r i t t e n i n FORTRAN o r BASIC. There
complex system is traded for greater a r e more fundamental f e a t u r e s which determine
computational t r a c t a b i l i t y , and a g r e a t e r whether a model i s l i k e l y t o be accepted and used
p o s s i b i l i t y of showing time s e r i e s d a t a . i n decisionmaking.

With t h i s s o r t of model i n hand, s c i e n t i s t s 1. It should be extensively and clearly

c a n i n t e r a c t w i t h d e c i s i o n makers i n a documented. How was t h e model developed and
q u a n t i t a t i v e way t o prepare an a r r a y of what a r e i t s assumptions? T h i s should be
alternative actions and their probable e v i d e n t , i f not t o t h e u l t i m a t e u s e r , a t
consequences. Various t o o l s of o p t i m i z a t i o n , l e a s t t o t h o s e who might s e r v e a s t e c h n i c a l
a n a c t i v e a r e a of c u r r e n t r e s e a r c h , come i n t o consultants. Unfortunately, this is a
play h e r e . Particularly valuable i n a s e r i o u s weakness of most models. T h i s a r i s e s
problem with a s many c o n f l i c t i n g v a l u e p a r t l y from funding limitations--documents-
judgments a s e f f e c t s of a i r p o l l u t i o n may be t i o n i s an obvious t h i n g t o skimp i f t h e
the multiple objective optimization budget h a s t o be c u t . I suspect t h a t not
e x t e n s i v e l y used i n e v a l u a t i n g water r e s o u r c e i n f r e q u e n t l y , though, t h e r e i s a wish, maybe
development a l t e r n a t i v e s (Cohon and Marks subconscious, t o keep what t h e modeler did t o
1975). T h i s i s a planning concept which h i m s e l f . A f t e r a l l , knowledge i s power.
 Its results should be understandable-- A polIcy-oriented model is not a d e c i s i o n
surprising, perhaps, but not maker. It is an a i d t o informed decisionmaking.
incomprehensible. F o r r e s t e r (1971), i n a I f i t is t o function e f f e c t i v e l y i n t h a t r o l e ,
widely quoted article, discussed the t h e r e needs t o be mutually s u p p o r t i v e i n t e r a c t i o n s
c o u n t e r i n t u i t i v e n a t u r e of s o c i a l systems, among modelers, b i o l o g i c a l and s o c i a l s c i e n t i s t s ,
and p r e s c r i b e d computer modeling a s an and d e c i s i o n makers.
a n t i d o t e . But F o r r e s t e r was a b l e t o e x p l a i n
quite clearly how he obtained his A modeler d e a l i n g with e c o l o g i c a l p u b l i c
counterintuitive results (others disagree policy questions related t o a i r p o l l u t i o n i n
with h i s a n a l y s i s , but t h a t is i r r e l e v a n t f o r e s t s needs t h e knowledge of a b i o l o g i s t , t o
here). He would have had no credence understand t h e essence of t h e mechanisms by which
whatever i f h e had not been a b l e t o provide a i r pollution a f f e c t s biological processes. He
such e x p l a n a t i o n . needs t h e s k i l l s of a n a p p l i e d mathematician, t o
understand t h e s t r u c t u r e of t h e model and i t s
3. The l i m i t s and probable range of e r r o r s i m p l i c a t i o n s , and u s u a l l y t o d i r e c t t h e programmers
should be w e l l explained. Few computer preparing the actual computer code. Most
models y i e l d d e t e r m i n i s t i c r e s u l t s , and a l l important, h e needs t h e p a t i e n c e and i n t e r p e r s o n a l
a r e l i m i t e d i n t h e i r a c c e p t a b l e degree of a b i l i t i e s of a diplomat, t o persuade t h e b i o l o g i s t
extrapolation. T h i s i s o f t e n not well and t h e d e i c i s o n maker a l i k e t h a t h e is h e l p i n g
understood by those not analytically them t o do t h e i r job b e t t e r , and not usurping t h e i r
i n c l i n e d ; i t needs t o be made c l e a r . legitimate roles.

4. The model should be f l e x i b l e enough t o deal B i o l o g i c a l and s o c i a l s c i e n t i s t s d e a l i n g w i t h

with problems t h a t had not been f u l l y modelers need t o r e c o g n i z e t h a t modelers a r e t h e r e
a n t i c i p a t e d , but a g e n e r a l all-purpose model t o h e l p them, not v i c e v e r s a . A good model,
i s not a d e s i r a b l e g o a l . S e n a t o r S. I. properly presented, will usually provide t h e
Hayakawa's dictum, "The map is not t h e d e c i s i o n maker with a f u l l e r e x p l a n a t i o n of t h e
t e r r i t o r y ," although made i n q u i t e a n o t h e r consequences of a l t e r n a t i v e pol i c i e s t h a n w i l l
context i n h i s r o l e a s a semanticist, is unaided s c i e n t i f i c s t a t e m e n t s o r p o s i t i o n papers.
wholly a p p l i c a b l e t o modeling. A model i s a Thus, t h e s c i e n t i s t may be b e t t e r a b l e t o g e t h i s
map t h a t t e l l s us how t o g e t from one p l a c e points across through the medium of a
t o a n o t h e r , even t o some p l a c e s we had not we1 1-constructed and we1 1-presented model .
o r i g i n a l l y intended t o v i s i t . It cannot
reproduce every f e a t u r e of t h e system and Decision makers should r e a l i z e t h a t a good
s t i l l r e t a i n its u s e f u l n e s s a s a guide. model is p r i m a r i l y a means f o r e x p l o r i n g t h e
consequences of a l t e r n a t i v e p o l i c y c h o i c e s of
5. The r e s u l t s should be d i s p l a y e d i n an n e a r l y equal rank. I f t h e model shows one o r two
e f f e c t i v e and u n d e r s t a n d a b l e manner. TOO c h o i c e s t o be s o s u p e r i o r t o o t h e r s t h a t o n l y t h e y
o f t e n model r e s u l t s a r e presented a s a r c a n e should be c o n s i d e r e d , t h i s w i l l s u r e l y be obvious
and incomprehensible p r i n t o u t . Territorial t o competent a n a l y s t s i n t h e absence of a model.
defensiveness again? R e s u l t s need t o be The r e a l u t i l i t y of a p o l i c y - o r i e n t e d model i s t o
p r e s e n t e d i n a form comprehensible t o t h o s e e x p l o r e t h o s e s i t u a t i o n s where t h e r e are a l a r g e
who r e a c t t o g r a p h s r a t h e r t h a n t o columns of number of a1 t e r n a t i v e s which a r e " n e i t h e r a1 1 good
figures, to pictures rather than to n o r a l l bad. The broader t h e a r r a y of c h o i c e s s e t
equations. Visual p r e s e n t a t i o n almost never up f o r e x p l o r a t i o n , t h e g r e a t e r t h e f i n a l r a n g e of
r e c e i v e s enough a t t e n t i o n . o p p o r t u n i t i e s w i l l be.

6. The model should be portable--usable on o t h e r F i n a l l y , I s u g g e s t t h a t t h e g r e a t e s t v a l u e of

computers with a minimum of reprogramming. I a modeling approach t o e c o l o g i c a l problem s o l v i n g
have t h e f e e l i n g t h a t i n c o m p a t i b i l i t y h a s
become worse i n r e c e n t y e a r s . This w i l l
presumably c o r r e c t i t s e l f e v e n t u a l l y , but f o r
now i t i s a s e r i o u s problem.
CONCLUSIONS
The impact of a i r p o l l u t i o n on f o r e s t s ,
e s p e c i a l l y when combined with o t h e r s t r e s s e s , h a s
biological, s o c i a l , and p o l i t i c a l i m p l i c a t i o n s
which o p e r a t e a t s e v e r a l time s c a l e s extended over
space. The unaided human mind i s not well adapted
t o e x p l o r e t h e consequences of each of t h e l a r g e
number of p o s s i b l e combinations of v a r i a b l e s .
T h e r e f o r e , some s o r t of computer model i s v i r t u a l l y
essential if t h e most reasonable array
a l t e r n a t i v e s i s t o be presented f o r r a t i o n a l
choice.
may be i t s s t i m u l u s t o exchange of i n f o r m a t i o n
among d i s c i p l i n e s and among p r a c t i t i o n e r s . I
mentioned, e a r l i e r , t h e r e l a t i v e l y u n s a t i s f a c t o r y
i n c o r p o r a t i o n of s p a t i a l dynamics i n f o r e s t models.
U n t i l a few y e a r s ago, t h i s would have been t r u e of
environmental biology g e n e r a l 1 y. There have been
r e c e n t s i g n i f i c a n t advances, though, i n d e a l i n g
with s p a t i a l h e t e r o g e n e i t y i n i n s e c t d i s p e r s a l and
i n dynamics of marine plankton. I n taxonomically
o r i e n t e d s c i e n c e , such advances t a k e a long time t o
come t o t h e a t t e n t i o n of workers i n a d j a c e n t
f i e l d s . The f a c t t h a t modeling approaches tend t o
c r o s s taxonomic boundaries seems l i k e l y t o c u t t h i s
unnecessary time l a g . T h i s a l o n e would be adequate
j u s t i f i c a t i o n f o r wider u s e of modeling i n a
complex i n t e r d i s c i p l i n a r y f i e l d such a s t h e e f f e c t
of of a i r p o l l u t a n t s i n f o r e s t s .
 LITERATURE CITED Kates, Robert W.
1978. Risk assessment of environmental
hazard (SCOPE 8 ) . John Wiley and Sons,
Cohon, J. L., and D. H. Marks New York, 112 pp.
1975. A review and e v a l u a t i o n of m u l t i o b j e c t i v e
programming t e c h n i q u e s . Water Resour. Res. b u c k s , Orie
11:208-220. 1980. Acid r a i n : l i v i n g r e s o u r c e i m p l i c a t i o n s
and management needs. Trans. 45th North
Cooper, C h a r l e s F. Amer. W i l d l i f e Nat. Res. Conf. [In press]
1976. Ecosystem models and environmental
policy. Simulation 26:133-138. Meadows, Dennis
1975. A c r i t i q u e of short-term perspectives
F o r r e s t e r , J . W. i m p l i c i t i n most r e s o u r c e models. I n Mineral
1971. Counterintuitive behavior of social Materials Modeling: A state-of-the-art
systems. Tech. Review 73(3):53-68. Review. W. A. Vogely, ed., pp. 66-80.
H o l l i n g , C. S. Shugart, H. H., Jr., T. R. Crow, and J. M. Hett
1973. R e s i l i e n c e and s t a b i l i t y of e c o l o g i c a l 1973. F o r e s t s u c c e s s i o n models: a rationale
systems. Ann. Rev. Ecol. Systematics 4: 1-23. and methodology for modeling forest
H o l l i n g , C. S. [Ed.]
s u c c e s s i o n over l a r g e regions. F o r e s t Sci .
19 :203-2 12.
1978. Adaptive environmental assessment and
management. Wiley Inter-Science, Chichester , S t a r r , Chauncey, and C h r i s Whipple
U.K. 377 pp. 1980. Risks of r i s k decisions. Science
206: 1114-1119.

West, D. C., H. B. McLaughlin, and H. H. Shugart

1980. Simulated f o r e s t response t o c h r o n i c a i r
p o l l u t i o n s t r e s s . J. Env. Q u a l i t y 9:43-49.
 Historical Perspectives and
International Concerns About Air
Pollution Effects on Forests1

Edwin Donaubauer2

Abstract: Air pollution from man's activities

has a long history; real hazards for forest veg-

etation occurred more than a century ago which

marked the starting point of forest research

in the field. The objectives of research have

been subject to a steady metamorphosis from sim-

ple causal relationships to the task of invest-

igating the long-term influence of pollutant

mixtures on trees directly and on entire eco-

systems. The IUFRO Subject Group S2.09, Air

Pollution, encourages interdisciplinary work

in the field and offers an organizational home

for close cooperation on an international basis.

The consideration of air pollution effects on

forest ecosystems goes far beyond limited for-

est interests, the findings can provide fund-

amental data on the deterioration of the hu-

man environment.

Some forest insect pests develop outbreaks is still in high demand as an energy source, and

characterized by high insect population den- in fact is still the most important source for

sities that use up the food resource represented over 90 percent of the world's population. Col-

by the host species. The direct damage by in- lection of fuel wood and the harvesting of

sects and the predisposition of weakened trees timber for many uses have influenced the ex-

to certain diseases both result in a serious tension and quality of forests over long per-

degradation or even destruction of the local iods of prehistoric and historic time. All

forest environment. these activities had, at least in some major

To some extent man-made air pollution prob- regions of the globe, serious consequences for

lems show some similar aspects. In many re- the ecosystem and human environment. We are

gions of the world mankind cleared the forests reminded of the historical fact, that 2000 years

for agriculture, for settlements, for roads, ago forests existed around the Mediterranean

for mining, for outdoor recreation, etc. Wood Sea and protected large and prospering ag-

ricultural areas in North Africa, where de-

serts are now present. These were direct in-

fluences, but among others an additional in-

1 direct threat developed by the quick progress

Presented at the Symposium on Effects of Air in technology and by increasing quantities

Pollutants on Mediterranean and Temperate Forest

and numbers of toxic substances released into

Ecosystems, June 22-27, 1980, Riverside,

the air.

California, U.S.A. Certainly, man-made air pollution has been

mentioned long ago in historic times, but the

'~ead, Institute of Forest Protection, dimensions of the problem have changed in re-

Federal Forest Research Institute, lation to regional economic and technical de-

Vienna, Austria velopments. The effects of air pollution to

Deputy Coordinator, IUFRO, Division 2 forest ecosystems were and seem still to be
underestimated, maybe for these reasons:

- symptoms a r e o f t e n u n s p e c i f i c , o r develop
l a t e and slowly, o r a r e even i n v i s i b l e
( a s depression of growth increment) ;
- s e r i o u s changes i n f o r e s t ecosystems may be-
come obvious only a f t e r many years of ac-
cumulation of low concentrations o r amounts
of t o x i c substances (heavy metals, a l -
kaline dusts, acid rain).
- t r e e s a r e more s e n s i t i v e t o widely d i s t r i b -
uted p o l l u t i o n than humans and t h i s d i f -
ference i s not f u l l y known o r r e a l i z e d .
REMARKS ON HISTORY
Perhaps Pliny (65 A.D.) was t h e f i r s t t o
observe and d e s c r i b e apparent Sop-damage t o
vegetation surrounding a smelter. Later on
we find r e f e r e n c e s f r e q u e n t l y i n documents of
t h e Middle Ages concerning a i r p o l l u t i o n by
c o a l burning; i n general people of higher so-,
c i a 1 c l a s s e s f e l t inconvenienced. Especially
i n such r e p o r t s from England and Central
Europe persecutions happened even a t t h a t time
i f a i r p o l l u t i o n was caused i n a prohibited
a r e a o r time.
A i r p o l l u t i o n became more than a l o c a l and
occasional a f f a i r when i n d u s t r i a l i z a t i o n and
the use of f o s s i l energy resources increased
r a p i d l y i n Western and Central Europe during
t h e past century. By t h e time f o r e s t research
i n s t i t u t e s and f o r e s t f a c u l t i e s were estab-
lished - more than a hundred years ago - they
had s e v e r a l r e p o r t s o f p r i o r experience and
observations of b o t a n i s t s t o draw upon. There-
f o r e f o r e s t research was stimulated t o pay a t -
t e n t i o n t o a i r p o l l u t i o n problems from t h e be-
ginning. The present r e s e a r c h p r o j e c t s provide
numerous p u b l i c a t i o n s on a i r p o l l u t i o n e f f e c t s .
CHANGING AIMS AND TASKS
The o b j e c t i v e s of r e s e a r c h p r o j e c t s con-
c e n t r a t e d f i r s t on sulphur dioxide including
methods f o r d e t e c t i o n and e v a l u a t i o n of a i r
p o l l u t i o n damage. Among t h e s e e a r l y works we
find very forward-looking ones, a s Rusnov' s
(1919) study about t h e i n f l u e n c e of ' a c i d
r a i n ' ( t h i s term was introduced much l a t e r )
on f o r e s t s o i l s . Several a u t h o r s proved t h e
use of chemical a n a l y s i s of f o l i a g e f o r de-
t e c t i o n and e v a l u a t i o n of a i r p o l l u t i o n (SO2,
HF) and i t s e f f e c t s on f o r e s t stands. These
p u b l i c a t i o n s caused long l a s t i n g d i s c u s s i o n s
and s t i m u l a t e d f u r t h e r i n v e s t i g a t i o n s ; how-
ever, many years passed u n t i l t h e chemical
a n a l y s i s of f o l i a g e (needles) became a u s e f u l
t o o l f o r a g r e a t number of cases. Step by
s t e p t h e method was r e f i n e d , so t h a t today
s u f f i c i e n t knowledge i s a v a i l a b l e f o r proper
sampling, and t h e t h r e s h o l d s of n a t u r a l s u l f u r
and f l u o r i n e c o n t e n t s and t h e i r v a r i a t i o n a r e
b e t t e r defined.
C e r t a i n l y , s u l f u r dioxide i s s t i l l t h e pol-
l u t a n t occurring over l a r g e s t a r e a s , but t h e
l i s t of substances t h u s f a r i d e n t i f i e d a s r e -
sponsible f o r damage t o f o r e s t t r e e s becomes
longer and longer. More a t t e n t i o n i s now given
t o i n d i r e c t and chronic e f f e c t s of a c i d i c o r
a l k a l i n e p r e c i p i t a t i o n and heavy metals on
f o r e s t ecosystems.
Formerly, t h e i n t e r e s t o f f o r e s t r e s e a r c h
concentrated on a s i n g l e - p o l l u t a n t - s i n g l e - t r e e
species r e l a t i o n s h i p . Today, increased e f f o r t s
a r e invested i n e l u c i d a t i n g t h e complex i n -
fluences on f o r e s t ecosystems and studying
t h e i n t e r a c t i o n of p o l l u t a n t mixtures which
occur i n many cases.
Regional i n v e s t i g a t i o n s on a i r p o l l u t e d
f o r e s t zones use a l l a s s i s t a n c e of modern
technology, such a s remote sensing combined
with physiological, chemical and t r e e men-
s u r a t i o n methods f o r v e r i f i c a t i o n .
Forest a i r p o l l u t i o n problems a r e linked
with t h e economic standard i n general and with
c e r t a i n i n d u s t r i a l development.
But i t i s n o t n e c e s s a r i l y so, t h a t only
point sources of a i r p o l l u t i o n o r concentra-
t i o n s of i n d u s t r i e s cause s e r i o u s problems.
Automobile t r a f f i c o r even a s i n g l e source
a r e a can under s p e c i f i c o r o g r a f i c and meteor-
o l o g i c c o n d i t i o n s lead t o p o l l u t a n t accumu-
l a t i o n and perhaps t o heavy damage. F u r t h e r ,
we a r e l e a r n i n g more about long d i s t a n c e pol-
l u t a n t t r a n s p o r t over hundreds and thousands
of kilometers.
The experience i n n a t i o n s with highly de-
veloped i n d u s t r y makes i t a d v i s a b l e t o s t i m -
u l a t e more i n t e r e s t i n t h i s f i e l d of r e s e a r c h
i n many developing c o u n t r i e s i n t h e world.
F i r s t of a l l t h e experiences of o t h e r s should
be transmitted and applied i n a l l planning
f o r economic development t o avoid t h e same
h i s t o r i c a l lessons received by many developing
c o u n t r i e s . There a r e many f o r e s t r e s e a r c h
workers and i n s t i t u t i o n s i n North America,
Europe (including t h e A s i a t i c p a r t of USSR),
and Japan who a r e experienced with t h e v a r i o u s
a s p e c t s of a i r p o l l u t i o n e f f e c t s on f o r e s t
ecosystems and could provide s c i e n t i f i c i n -
formation and a s s i s t a n c e .
I U F R 0 AND AIR POLLUTION RESEARCH
The I n t e r n a t i o n a l Union of F o r e s t Research
Organizations (IUFRO) has s i x major organi-
z a t i o n a l d i v i s i o n s . Division 2 , 'Forest
P l a n t s and Forest P r o t e c t i o n ' i s home f o r t e n
Subject Groups, one of which i s S2.09 - ' A i r
Pollution.' The t i t l e s of t h e various Working
P a r t i e s e l u c i d a t e . t h e i n t e r d i s c i p l i n a r y scope:
Researchers a r e included from t h e f i e l d s of
f o r e s t mensuration, s o i l s c i e n c e , pathology,
remote sensing, t r e e physiology, e t c . They work
t o g e t h e r , and exchange research r e s u l t s t h a t
s t i m u l a t e t h e i r f u r t h e r research. This IUFRO
Subject Group has a long h i s t o r y of a c t i v i t y .
For more than a q u a r t e r of a century meetings
have brought t o g e t h e r an i n c r e a s i n g number of
p a r t i c i p a n t s . A few western and c e n t r a l European
c o u n t r i e s were represented a t f i r s t ; now more
c o u n t r i e s from e a s t e r n and southern Europe a r e
represented too, a s w e l l a s a slowly i n c r e a s i n g
number of p a r t i c i p a n t s from Canada and t h e U.S.A.
The eleventh meeting of a l l Working P a r t i e s w i l l
be held this year in Graz, Austria and previous
I feel that this international symposium

conferences were hosted by nine other European

sets the stage for new conceptualizations by

countries. It is somewhat disappointing that

placing the view of effects of air pollutants

the group met only once in another continent:

on forest ecosystems into the center of the de-

Gainesville, Florida, U.S.A. at the occasion

liberation. The discussions here should stim-

of the IUFRO Congress in 1971. We expect to

ulate further research and interdisciplinary,

have a meeting at the occasion of the 1981

international cooperation.

IUFRO Congress in Japan.

In addition I will emphasize that air pol-

The interdisciplinary scope of all Subject

lution effects on forests are not only a

Group meetings makes them attractive to a cer-

question for researchers or foresters, but

tain group, seldom represented at such scien-

of fundamental interest to mankind. That may

tific conferences. We could characterize this

seem overstated but forest ecosystems demon-

group as 'consumers' of research results in-

strate due to the long rotation periods the

cluding: forest managers, representatives from

dynamics and the consequences of long-term

industries, and sometimes politicians. There

influences on complex ecosystem-level processes.

are of course some difficulties in accommodating

Various pollutant effects that are dangerous

the needs of each group, but I consider the

not only to the forests themselves, as an im-

interest in such meetings as an expression for

portant resource, but also as an invaluable

a demand to receive the information from the

part of the human environment can be predicted

scientists directly and not via more tradition-

by ecosystem-level research. In this sense one

al channels, i.e., technical publications. I

can consider the forest ecosystems as sensitive

feel we should think of possibilities for such

warning systems for the vital, fundamental

direct information flow in specific fields like

interests of life.

air pollution where we find more and more vital

interest from the public.

Natural Influences of Forests on Local

and Regional Air Quality

Emissions and Air Resource

Management Within Forests '
2
Michael A. Fosberg and H o l l i s Record

Abstract: S u b s t a n t i a l p o r t i o n s of t h e emissions i n v e n t o r y
w i t h i n f o r e s t e d l a n d s a r e from d i s p e r s e d i n t e r m i t t e n t
sources. Chief s o u r c e s a r e smoke from w i l d f i r e and pre-
s c r i b e d f i r e , emissions a s s o c i a t e d with c o n c e n t r a t e d rec-
r e a t i o n and second-home developments , and f u g i t i v e d u s t
from unpaved r o a d s and e o l i a n s o i l s . E f f e c t s of smoke o n
f l o r a range from reduced p h o t o s y n t h e t i c e f f i c i e n c y a t low
dosages t o t i s s u e n e c r o s i s a t high dosages. E f f e c t s on
fauna a r e n o t c l e a r l y d e f i n e d . E f f e c t of smoke on s o c i a l
v a l u e s , p r i m a r i l y v i s i b i l i t y , is recognized but not under-
s t o o d . D i s p e r s i o n p r o c e s s i n complex t e r r a i n , t h e physio-
g r a p h i c s e t t i n g f o r most f o r e s t e d l a n d s , i s complicated by
topography and s p a t i a l l y varying wind f i e l d s , a h i g h e r
degree of a n i s o t r o p y of t u r b u l e n c e , and a wider range of
t u r b u l e n c e i n t e n s i t i e s t h a n found over l e v e l ground.
Management of a i r r e s o u r c e s w i t h i n f o r e s t e d a r e a s is
l i m i t e d t o land management planning a c t i v i t i e s because of
t h e complexi t y of emission c h a r a c t e r i s t i c s , d i s p e r s i o n
p r o c e s s e s , and e f f e c t s of p o l l u t a n t s from w i t h i n f o r e s t
sources.

Much of t h i s symposium t r e a t s e f f e c t s of pol- s i o n s from f i r e a r e Con, CO, p a r t i c u l a t e s , and

l u t a n t s from major s t a t i o n a r y s o u r c e s on eco- hydrocarbons. R e c r e a t i o n a l and second-home devel-
systems. Significant pollutants treated i n other opments emit C O , CO, p a r t i c u l a t e s , and hydro-
papers a r e n i t r o u s o x i d e s , o x i d e s of s u l f u r , reac- carbons from f i r e p l a c e s and campfires; SO2 and NOx
t i v e hydrocarbons, and t h e photochemical d e r i v e d from dispersed transportation systems--namely,
p o l l u t a n t , ozone. E f f e c t s of NOn, SO2, and O n a r e p r i v a t e automobiles. F u g i t i v e d u s t from unpaved
w e l l documented i n t h e companion papers i n t h i s r o a d s and e o l i a n s o i l s a l s o c o n t r i b u t e t o t h e
volume. p a r t i c u l a t e loading.

P o l l u t a n t s from w i t h i n f o r e s t s o u r c e s a l s o
i n c l u d e emission from w i l d f i r e , p r e s c r i b e d f i r e ,
unpaved r o a d s , e o l i a n s o i l s , c o n c e n t r a t e d recrea-
t i o n , and second-home developments, Chief e m i s -
p r e s e n t e d a t t h e Symposium on E f f e c t s of A i r
P o l l u t a n t s on Mediterranean and Temperate F o r e s t
Ecosystems, J u n e 22-27, 1980, R i v e r s i d e ,
C a l i f o r n i a , U.S.A.
2 ~ r o j e c tLeader, F o r e s t and Brushland Meteorol-
ogy, P a c i f i c Southwest F o r e s t and Range Experiment
S t a t i o n , F o r e s t S e r v i c e , U.S. Department of Agri-
c u l t u r e , Berkeley, C a l i f . , s t a t i o n e d a t R i v e r s i d e ,
C a l i f . ; and G e o l o g i s t , Los Padres N a t i o n a l , F o r e s t ,
F o r e s t S e r v i c e , U.S. Department of A g r i c u l t u r e ,
Goleta, C a l i f .
E f f e c t s of t h e above p o l l u t a n t s , p a r t i c u l a r l y
smoke, f l o r a , fauna, and s o c i a l v a l u e s a r e poorly
understood. Documented e f f e c t s range from reduced
p h o t o s y n t h e t i c a c t i v i t y through blockage of s o l a r
radiation t o t i s s u e necrosis. E f f e c t s on micro-
organisms range from i n h i b i t i o n of some s p o r e s and
fungi t o i n c r e a s e d germination of one fungus.
E f f e c t s of smoke on fauna a r e documented, but with-
out e x p l a n a t i o n .
V i s i b i l i t y i s both a physical and s o c i a l v a l u e .
V i s i b i l i t y can be q u a n t i f i e d i n terms of v i s u a l
range and a b i l i t y t o d e f i n e d e t a i l s a t s p e c i f i e d
distances. V i s i b i l i t y is a l s o a personal v a l u e
based on p a s t and expected experiences. A recent
popular c o u n t r y and western song (McCall and o t h e r s
1976) goes "..... One of t h e guys from New York
s a i d 'Hey, look a t t h e smog i n t h e sky, smog c l e a r
out h e r e i n t h e s t i c k s ' . Someone s a i d , 'Hey J o e ,
t h a t ' s not smog, t h a t ' s t h e Milky Way', J o e had
never s e e n t h e Milky Way ...." Contrast Joe's
r e a c t i o n t o t h a t of a r e s i d e n t of t h e Four Corners
a r e a of Utah v i s i t i n g any urban a r e a i n t h e world
on a c l e a r a i r day. Local r e s i d e n t s would comment
on t h e c l a r i t y of t h e a i r , but t h e v i s i t o r would
n o t i c e t h e impairment of v i s i b i l i t y . and o t h e r s 1971).
The r e l a t i o n s h i p between a s o u r c e of p o l l u t i o n
and t h e ef f e c t of t h a t p o l l u t a n t on Mediterranean
and temperate f o r e s t ecosystems is through dis-
p e r s i o n of t h e p o l l u t a n t between t h e s o u r c e and t h e
r e c e p t o r point. Nearly a l l d i s p e r s i o n c a l c u l a t i o n s
a r e based on t h e Gaussian model i n which t r a n s p o r t
is t r e a t e d through d e f i n i t i o n of a mean windspeed
and d i r e c t i o n and t u r b u l e n t d i f f u s i o n i s based on a
t r a n s f o r m a t i o n of t h e t u r b u l e n c e s t r u c t u r e t o a
Gaussian s t a t i s t i c a l d i s t r i b u t i o n (Turner 1969).
V a l i d i t y of t h e c o e f f i c i e n t s used i n t h e Gaussian
model a r e u n c e r t a i n i n s i t u a t i o n s where t e r r a i n
f e a t u r e s a r e complex. I n p a r t i c u l a r , winds a r e
known t o c o n t a i n a high degree of s p a t i a l and
temporal v a r i a b i l i t y (Fosberg and o t h e r s 1980) and
3
t h e turbulence i n t e n s i t i e s a r e highly anisotropic.
Because much of t h e Mediterranean and temperate
f o r e s t ecosystems a r e found i n complex t e r r a i n
throughout t h e world, t h e d i s p e r s i o n o r d e l i v e r y
system of p o l l u t a n t s from t h e s o u r c e t o t h e
r e c e p t o r must account f o r complex t e r r a i n atmos-
pheric processes.
An understanding of each p h y s i c a l and b i o l o g i c a l
process i s necessary but not s u f f i c i e n t t o develop
management p l a n s f o r a i r r e s o u r c e s w i t h i n f o r e s t s
and brushlands. A c t s , laws, r e g u l a t i o n s , and codes
e s t a b l i s h e d by Congress down through l o c a l county
r e g u l a t o r y agencies s p e c i f y g o a l s and o b j e c t i v e s
f o r a i r q u a l i t y and f r e q u e n t l y s p e c i f y t h e methods
i n which a i r q u a l i t y o b j e c t i v e s w i l l be met. As
example, o r g a n i c a c t s of most Federal a g e n c i e s i n
t h e United S t a t e s r e q u i r e t h a t t h e agency p r o t e c t
o r p r e s e r v e , o r meet a i r q u a l i t y o b j e c t i v e s . The
Clean A i r Act of 1977 (U.S. Congress 1977) s p e c i f i-
c a l l y r e q u i r e s t h a t a i r q u a l i t y o b j e c t i v e s be met
through emissions c o n t r o l . Because i t is not
f e a s i b l e t o i n s t a l l s c r u b b e r s on p r e s c r i b e d f i r e ,
emission c o n t r o l is achieved through emissions
d e n s i t y planning.
Each of t h e following s e c t i o n s a d d r e s s e s t h e
s p e c i f i c t o p i c s of emission c h a r a c t e r i z a t i o n , dis-
p e r s i o n i n complex t e r r a i n , e f f e c t s of smoke on
f l o r a , fauna, s o c i a l v a l u e s , and a i r r e s o u r c e
management . s c r i b e d f i r e s and w i l d f i r e s a r e shown i n f i g u r e 3.
EMISSIONS FROM WITHIN FOREST SOURCES
Major emissions from w i l d f i r e and p r e s c r i b e d
f i r e a r e CO CO, hydrocarbons, and p a r t i c u l a t e s .
2'
anh ham, Lucy M. 1980. Wintertime d i s p e r s i o n
p r o c e s s e s i n t h e Lake Tahoe Basins. Proc. 2nd
Conf. o n A p p l i c a t i o n of A i r P o l l u t i o n Meteorology.
Amer. Meteorol. Soc. [ i n press]
Although C02 i s n o t a p o l l u t a n t a s such, C02 i s of
c o n s i d e r a b l e i n t e r e s t i n a n a l y s i s of t h e g l o b a l
h e a t balance. Carbon d i o x i d e emissions range from
1000 kg per m e t r i c t o n of f u e l t o 1750 kg per
m e t r i c t o n of f u e l (Ryan and McMahon 1976), w i t h
extreme v a l u e s n e a r 1830 kg per m e t r i c t o n (Vines
Carbon monoxide emissions from f i r e a r e h i g h l y
dependent on can bust i o n e f f i c i e n c y . Values r a n g e
from 17 t o 98 kg per m e t r i c t o n (Sandberg and
Martin 1975, Darley and o t h e r s 1966, G e r s t l e and
Kemnitz 1967). The U.S. Environmental P r o t e c t i o n
Agency (1978) recommends a v a l u e of 4 5 kg per
m e t r i c t o n from hemlock, Douglas-f i r , and c e d a r and
98 kg per m e t r i c t o n from ponderosa pine. Inef f i-
d e n t combustion f o r smoldering damp f u e l s have
r e s u l t e d i n emissions a s high a s 250 kg per m e t r i c
t o n of f u e l (Ryan and McMahon 1976). Emissions a s
high a s 250 t o 400 kg per m e t r i c t o n of f u e l have
been reported when energy r e l e a s e from f i r e i s less
t h a n 750 w a t t s per s q u a r e meter (Sandberg and
Martin 1975).
Hydrocarbon emissions range from 2 t o 7 kg p e r
m e t r i c t o n of f u e l (U.S. Environmental P r o t e c t i o n
Agency 1978) although emissions a s high a s 20 kg
per m e t r i c t o n have been r e p o r t e d (Ryan and McMahon
1976, Darley and o t h e r s 1966). S p e c i a t i o n of
hydrocarbons ( f i g . 1) shows s a t u r a t e d hydrocarbons
(mostly methane) comprise about 30 p e r c e n t a t peak
f i r e i n t e n s i t y and about 1 5 percent a t low f i r e
intensities (Sandberg and o t h e r s 1979). Low
molecular weight o l o f i n e s make up about 17 percent
of t h e emission from flaming f i r e and 3 p e r c e n t
from smoldering f i r e (Sandberg and o t h e r s 1979).
Bnissions of SO and N O a r e n e g l i g i b l e . Most
x
f u e l c o n t a i n s l e s s t h a n 0.2 percent s u l f u r and com-
b u s t i o n temperatures a r e low, p r e v e n t i n g formation
of NOx.
P a r t i c u l a t e emissions from f i r e a r e g i v e n a s 2
t o 6 kg per m e t r i c t o n of f u e l (U.S. Environmental
P r o t e c t i o n Agency 1978). A range of emissions a r e
given i n t a b l e 1. P a r t i c u l a t e s i z e s a r e mainly i n
t h e submicron diameter c l a s s e s with o n l y a few
p a r t i c l e s l a r g e r t h a n a micron ( f i g . 2). Particu-
l a t e s i z e s a r e dependent on combustion e f f i c i e n c y
(Schaefer 1976) with t h e most e f f i c i e n t f i r e -
producing s i z e s i n t h e r e s p i r a b l e range.
National emissions of p a r t i c u l a t e s from pre-
Prescribed f i r e c o n s i s t s of less t h a n 20 percent of
a l l f i r e emissions n a t i o n a l l y . Emissions of t h e
major p o l l u t a n t s , p a r t i c u l a t e s , CO, and hydro-
carbons by S t a t e and r e g i o n s a r e given i n t a b l e 2
f o r p r e s c r i b e d burning.
I n a d d i t i o n t o w i l d f i r e and p r e s c r i b e d f i r e ,
c o n c e n t r a t e d r e c r e a t i o n and second-home develop-
ments c o n t r i b u t e smoke through f i r e p l a c e burning
and more i n c r e a s i n g l y through use of wood f o r home
heating .
 100-

IOOM GLASS SCOT OV-101

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iz
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TIME (MINUTES)

PEAK COMPOUND PEAK COMPOUND PEAK COMPOUND

2 homw 24A Wbotdihyd* ( t i l t ) 54 n-maw
26 2.5-dlmethylfuran 5 4 ~ 2- lsopro~>lfuran
28 1-pentun
zc furon 26A n-haptone 5s anlwlÃ
3 n-pentaw 27 a - 2 - htptanà 56 2-mathyl-5-IÇÈprÃ
3A isoprene 29 2-vinylfuran
2 3-dimçlhyl-2 m y If uron
30 ocetorr 31
4A iwpropanol Pcntona 57 cumene
48 cyclopç"t<"lhn 32 2.4-dlmçthylhtxo 58 n-decone
7 diacet~l 36 tOllWM 60
59 n-propyltfnzew
camphena (tçnl.
8 I-hexw 39 I-octene
8A methyl vinyl k à § t 40 2.3- dimçthylhçi 61 m-ethyltoluene
1.4-diem
9 2-methylfuran
41 n- octane 62 p- ethyltoluiw
10 n-hexone 2.3.5-trimtliylfuron 66 I-deceni
15 2,4- h e x a d h 42 67 benzofuron
16 1.3.5-hexatrlç 44 furfurol
49 ethyl benzene 71 m-diethylhzme
17 3-methylbutanal 72 .lne
20 benzene SO p-aybne
77 p-a-dlrnethylçtyrç
21 cyclohexom 50A 2-propionylfuranftfrtJ
22 4 - methylpentaw 78 n-undccane
styrene
23 214-dlmethylixf;t(~
52
53 Q-XYI~W
an n-dodecona
24 I-trans-2-dimethyl-
cyclopentone

F i g u r e IÑChromatogra of o r g a n i c vapors i n l o b l o l l y p i n e smoke (from Ryan and

McMahon 1976).

Table I--Summary of p a r t i c u l a t e emission y i e l d s r e p o r t e d from w i l d l a n d f u e l s

(from Sandberg and o t h e r s 1979)

P a r t i c u l a t e s (kg per m e t r i c t o n
of f u e l burned)

Lablfield Type of f i r e - ~

Fuel t y p e experiment Heading Backing Reference

Logging r e s i d u e s Field 14-53 Sand berg (1974)

(Western)
Laboratory Sandberg (1974)
Field Radke and o t h e r s (1978)
Laboratory F r i t s c h e n and o t h e r s (1970)

Landscape r e f u s e Laboratory 12 F e l d s t e i n and o t h e r s (1963)

Grass burning Field 8 Boubel and o t h e r s (1969)

Live u n d e r s t o r y Field Vines and o t h e r s (1971)

(Australia)
Laboratory Vines and o t h e r s (1971)
(Southern) Field 7-15 Ward and o t h e r s (1976)
Laboratory 12-49 Ryan (1974)

Pine l i t t e r Field 22-27 Ward and o t h e r s (1976)

(Southern)
Laboratory 3-1 4 Ryan and McMahon (1976)
Laboratory 11-63 Ryan and McMahon (1976)
 99.99 -
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PARTICLE DIAMETER, MICRONS
(C1
F i g u r e 2--Particle s i z e d i s t r i b u t i o n : (A) S i n g l e f i r e i n f o u r f u e l t y p e s ; (B)
Grand Average, a l l f u e l t y p e s ; and (C) Normalized d i s t r i b u t i o n , number, s u r f a c e
a r e a , and volume f o r a high and low c o n c e n t r a t i o n (from Ryan and McMahon 1976).

Some hydrocarbons and NO a r e a l s o e m i t t e d from fungi (Panneter and Uhrenholt 1975a, b). Effects
x of smoke on p h o t o s y n t h e s i s a t low dosages reduces
t r a n s p o r t a t i o n systems and p r i v a t e v e h i c l e s i n
t h e p h o t o s y n t h e t i c r a t e by d i r e c t blockage of s o l a r
r e c r e a t i o n a l and second-home developments.
radiation. Increased C O c o n c e n t r a t i o n s , however,
F u g i t i v e d u s t from unpaved roads and e o l i a n could i n c r e a s e carbon f i x a t i o n and p h o t o s y n t h e t i c
s o i l s occasionally contribute substantially t o the a c t i v i t y (Green and Wright 1977).
p a r t i c u l a t e l o c a t i o n of p o l l u t a n t s , but t h i s h a s
n o t been completely q u a n t i f i e d (Singer 1980). E f f e c t s of smoke on s o c i a l v a l u e s , p r i m a r i l y
v i s i b i l i t y , a r e not c l e a r l y defined. Although
p h y s i c a l a s p e c t s of v i s i b i l i t y ; t h a t i s , v i s u a l
EFFECTS OF SMOKE ON FORESTS range, maximum d i s t a n c e a n o b j e c t can be s e e n , and
d i s c r i m i n a t i o n of d e t a i l s on a d i s t a n t o b j e c t can
Few s t u d i e s e x i s t t h a t c l e a r l y d e f i n e t h e be defined q u a n t i t a t i v e l y (Malm 1979). perceived
e f f e c t s of smoke and f o r e s t b i o t a . E f f e c t s of psychological b e n e f i t s (Driver and o t h e r s 1979) of
smoke on micro-organisms s u g g e s t t h a t smoke reduced v i s i b i l i t y a r e i n t e r r e l a t e d with o t h e r demands on
growth of s p o r e germination of s e v e r a l fungal t h e sensory system. Paraphrasing D r i v e r and o t h e r s
pathogens, but i n c r e a s e d s p o r e germination on one (1979) a l i t t l e n o i s e p o l l u t i o n , a l i t t l e l i g h t
 LEGEND:
SEASON
1 JAN- - MAR
2 APR- - JUNE
-
3 JULY - SEPT
-
4 OCT - DEC
FIRE TYPE:
R -
WILDFIRE
- PRESCRIBED FIRE

REGIONS AND SEASONS

F i g u r e 3--Forest f i r e p a r t i c u l a t e production by r e g i o n and season (from

Ward and o t h e r s 1976).

p o l l u t i o n , a l i t t l e l o s s of open s p a c e , awareness pollutants. The most f r e q u e n t l y used method of

of water p o l l u t i o n , nonbiodegradable s u b s t a n c e s , q u a n t i f y i n g t h e d i s p e r s i o n process is through t h e
a e r o s o l cans and c a n c e r , change expected v a l u e s of so-called Gaussian d i s p e r s i o n model (Turner 1969) .
visibility. R e f e r r i n g t o t h e quote from a popular Downwind c o n c e n t r a t i o n s , X , a r e r e l a t e d t o emission
song used i n t h e i n t r o d u c t i o n of t h i s paper, we can Q by
a s k , d i d J o e ' s a t t i t u d e s toward v i s i b i l i t y change
o n h i s v i s i t t o Colorado, and how would t h o s e
changes i n f l u e n c e h i s view of t h e New York s k y l i n e ?
The p o i n t D r i v e r and o t h e r s (1979) a r e emphasizing where u i s t h e mean windspeed and
is t h a t t h e perceived v a l u e s a r e t h e r e a l v a l u e s
and t h a t t h e p h y s i c a l l y measurable v a l u e s of v i s i -
b i l i t y a r e i n d i c e s of t h e v a l u e s . In particular,
s c a t t e r i n g and a t t e n u a t i o n of l i g h t i s n o t a s o c i a l
value.
Here, o and a a r e t h e v a r i a n c e s i n t h e Gauss-
Y
i a n s t a t i s t i c a l d i s t r i b u t i o n ; y and z a r e t h e d i s -
DISPERSION PROCESSES I N
t a n c e of t h e p o l l u t a n t element from t h e plance
COMPLEX TERRAIN
c e n t e r l i n e . The v a r i a n c e o i s r e l a t e d t o t h e t u r -
bulence s t r u c t u r e through
Wind p a t t e r n s i n complex t e r r a i n a r e h i g h l y
v a r i a b l e i n t i m e and space. Local mountain and 2Kx
o2 = - (3)
v a l l e y c i r c u l a t i o n s f r e q u e n t l y mask t h e l a r g e - s c a l e u
p a t t e r n s such t h a t a mean t r a n s p o r t wind f o r pol-
l u t a n t movement i s d i f f i c u l t t o d e f i n e (Fosberg and
i n which K i s t h e eddy t u r b u l e n c e c o e f f i c i e n t and x
o t h e r s 1976a, b; Fosberg and Fox 1978). Spatial
i s t h e d i s t a n c e downwind from t h e s o u r c e . Tradi-
v a r i a b i l i t y of winds i s c l e a r l y i l l u s t r a t e d i n
t i o n a l i n t e r p r e t a t i o n s of atmospheric processes
f i g u r e 4 over t h e Oregon Coast Range and over t h e
t h a t were developed f o r l e v e l ground suggest t h a t
Cascade Mountains. C o n t r a s t t h i s v a r i a b i l i t y with
t h e K ' s be defined through a n a l y s i s of t h e i n e r t i a l
t h e u n i f o r m i t y of winds over t h e P a c i f i c Ocean and
subrange of t u r b u l e n c e and t h a t t h i s mean wind is
w i t h i n t h e Willamette Valley.
c o n s t a n t over s u b s t a n t i a l d i s t a n c e s , t h i s i s , 1 0 ' s
of km. Such assumptions a r e extremely d i f f i c u l t t o
I n a d d i t i o n t o mean t r a n s p o r t of p o l l u t a n t s ,
s a t i s f y i n complex t e r r a i n .
t u r b u l e n t d i f f u s i o n i s important i n d i s p e r s i n g a i r
Table 2 ~ S u m m a r yof p r e s c r i b e d f i r e a c r e s burned and t o n s of c r i t e r i a p o l l u t a n t e m i t t e d (by geographic
r e g i o n , annual b a s i s ) (from Sandberg and o t h e r s 1979)

Fuel consumed Particulates Carbon monoxide Hydrocarbons

S t a t e s by Area Metric Metric 8 kglmetric 10 k g l m e t r i c 5 kglmetric
Region hectares tonslha tons tons tons tons

California
Oregon
Washington
Total
ROCKY MTN .
Arizona
Colorado
Idaho
Montana
New Mexico
North Dakota
Total
N. CENTRAL
Michigan
Minnesota
Wisconsin
Total
EASTERN
Delaware
New J e r s e y
Total
SOUTHERN
A1a bama
Arkansas
Florida
Georgia
Louisiana
Mississippi
N. C a r o l i n a
S. C a r o l i n a
Texas
Virginia
Total
USA Tot a1

An a1 t e r n a t i v e method of d e f i n i n g t h e v a r i a n c e s
f o r complex t e r r a i n i s mathematically i d e n t i c a l but
does n o t r e q u i r e t h a t t h e t u r b u l e n c e l i e i n a
p a r t i c u l a r p o r t i o n of t h e energy spectrum (Fosberg
and o t h e r s 1976b, Fosberg and Fox 1978). I n par-
t i c u l a r , a mean wind i s d e f i n e d s t a t i s t i c a l l y over
t h e d i s p e r s i o n d i s t a n c e of i n t e r e s t . D e v i a t i o n of
wind about t h i s mean, whether i n t h e t u r b u l e n t
i n e r t i a l subrange o r produced by organized flows of
s c a l e s m a l l e r t h a n t h e averaging d i s t a n c e s , a r e
t r e a t e d mathematically as components of K. The
d e v i a t i o n s about t h e s t a t i s t i c a l l y d e f i n e d mean
wind a r e u'. The K ' s a r e t h e n defined by
d e f i n e t h e wind v a r i a b i l i t y a t s c a l e s s m a l l e r t h a n
t h o s e used t o d e f i n e t h e mean wind.
MANAGEMENT OF AIR RESOURCES
Emission c o n t r o l is r e q u i r e d t o meet a i r q u a l i t y
o b j e c t i v e s (U .S. Congress 1977). Because d i r e c t
l i m i t s on emission from open burning can be
achieved o n l y by l i m i t i n g t h e mass of f u e l burned
a t any given time, a model f o r a i r r e s o u r c e a l l o -
c a t i o n was developed. The A i r Resource A l l o c a t i o n
Model (ARAM) i s based on t h e Gaussian d i s p e r s i o n
model d e f i n e d i n e q u a t i o n 1. Because t h e i n t e n t i s
t o l i m i t emissions, e q u a t i o n 1 i s r e a r r a n g e d t o

i n which t h e l i n e over t h e s q u a r e of t h e d e v i a t i o n s Q = X-"L

G (5
i s t h e a v e r a g i n g o p e r a t o r . The time c o n s t a n t T i s
r e l a t e d t o t h e averaging t i m e and space. These K ' s The c o n c e n t r a t i o n X is i n t e r p r e t e d h e r e a s t h e
do n o t r e p r e s e n t t u r b u l e n c e . I n s t e a d , t h e K's increment of a i r q u a l i t y a v a i l a b l e f o r p r e s c r i b e d
 Table 3--Change i n annual burn ( i n h e c t a r e s by p o l l u t a n t )

Basin C.O. 1 T.S.P. 1 H.C.

Monterey Ranger D i s t r i c t
1. L i t t l e Sur
2. Big Sur
3. Carmel
4. Arroyo Seco
5. Ocean Front
6. San Antonio
7. Nacimiento

Santa Lucia Ranger D i s t r i c t

8. S a l i n a s (A)
9. S a l i n a s (B-)
10. Lopez Canyon
11. Cuyama (A)
12. Sisquoc

Mount Pinos Ranger D i s t r i c t

13. Cuyama (B)
14. an Joaquin Val l e y
15. P i r u

Ojai Ranger D i s t r i c t
16. Sespe 20,500 170 2,700
17. ~ a n t aPaula 6,200 -17 no d a t a
18. Ventura 25,500 -7 0 no d a t a

S a n t a Barbara Ranger D i s t r i c t
19. Santa Ynez
20. Santa Barbara Front

burning. ARAM i s based on e a r l i e r development on

emission l i m i t s f o r s i n g l e sources, s i n g l e receptor
r e l a t i o n s d e f i n e d i n t h e TAPAS model (Fosberg and
Fox 1976, Fox and Fosberg 1976). ARAM d i f f e r s from
TAPAS i n t h a t ARAM c o n s i d e r s m u l t i p l e s o u r c e s and The g e n e r a l i z e d form of ARAM expressed i n e q u a t i o n
m u l t i p l e r e c e p t o r s i t e s and c o n t a i n s improvements 5 is t h i s
i n c h a r a c t e r i z a t i o n of t h e d i s p e r s i o n p r o c e s s e s .
R e l a t i o n s between m u l t i p l e s o u r c e s and m u l t i p l e
r e c e p t o r s i t e s a r e d e f i n e d through m a t r i x a l g e b r a
as ARAM h a s been a p p l i e d on one National F o r e s t i n
C a l i f o r n i a , t h e Los Padres, i n support of use of
p r e s c r i b e d f i r e i n v e g e t a t i o n management. Current
a i r quality regulations i n California recognize
t h a t prescribed f i r e is an a l t e r n a t i v e t o w i l d f i r e
f o r v e g e t a t i o n management. In particular, the
r e g u l a t o r y a g e n c i e s accept t h e concept t h a t a t o n
of f u e l burned i n p r e s c r i b e d f i r e can be used t o
o f f s e t a t o n of f u e l burned i n w i l d f i r e .
Here x, , and s o on a r e t h e increments of
p o l l u t i o n allowed a t r e c e p t o r s i t e s 1, 2, and s o I n t h e following example, t h e increment f o r
on; Q,, Q2, ...a r e t h e a l l o w a b l e emissions a t p r e s c r i b e d burning is d e f i n e d a s t h e incremental
d e p a r t u r e from t h e e x i s t i n g emission from pre-
s o u r c e sites 1, 2, ... and Gll is the dispersion s c r i b e d f i r e and w i l d f i r e s . T h i s approach circum-
v e n t s t h e d i f f i c u l t i e s a s s o c i a t e d with development
relation between s o u r c e 1 and r e c e p t o r 1; G of a complete regionwide e m i s s i o n i n v e n t o r y base.
21 is The assumption h e r e i s t h a t t h e e x i s t i n g emission
t h e d i s p e r s i o n r e l a t i v e between source. 2 and from p r e s c r i b e d f i r e and w i l d f i r e a r e d e f i n e d
r e c e p t o r 1, and s o on. Expressing e q u a t i o n 6 i n w i t h i n t h e S t a t e Implementation Plan. The follow-
m a t r i x format i n g c a l c u l a t i o n s t h e n r e p r e s e n t a n a n a l y s i s of
where p r e s c r i b e d burning can be i n c r e a s e d and where
burning must be reduced. The a i r q u a l i t y database
i s t h e C a l i f o r n i a A i r Resources Board (1977) Three-
Year Summary of A i r Quality. Nearly a l l t h e a i r
q u a l i t y monitoring s t a t i o n s a r e i n urban a r e a s and,
t h e r e f o r e , do n o t n e c e s s a r i l y r e f l e c t c o n d i t i o n s i n
t h e wildlands. The following c a l c u l a t i o n s a r e
c o n s e r v a t i v e e s t i m a t e s because of t h e b i a s i n t h e
d a t a b a s e . The m e t e o r o l o g i c a l d a t a b a s e i s from t h e
N a t i o n a l F i r e Weather L i b r a r y (Furman and Brink
1975). T h i s d a t a b a s e i s t h e o n l y r e a d i l y acces-
s i b l e d a t a b a s e f o r wildlands. Plume r i s e c a l c u l a -
t i o n s were made through t h e e q u a t i o n s developed by
Craig and Wolf (1980) f o r p r e s c r i b e d burning.
T h r e e c r i t e r i a p o l l u t a n t s were e v a l u a t e d . These
p o l l u t a n t s were p a r t i c u l a t e s , hydrocarbons, and
carbon monoxide.
The p h y s i c a l s e t t i n g of t h e Los Padres National
F o r e s t i s along t h e C a l i f o r n i a Coast extending from
n e a r Monterey i n t h e n o r t h , around Point Concep-
t i o n , t o n e a r Santa Barbara i n t h e ~ a l i f o r n i a
F i g u r e 4a--Location of weather s t a t i o n s i n northwest
Oregon used t o c a l c u l a t e wind p a t t e r n s shown i n
f i g u r e 4b.
Bight. The Los Padres National F o r e s t l i e s w i t h i n
t h e C a l i f o r n i a Coast Range and, t h e r e f o r e , can be
broken up i n t o a s e r i e s of small a i r s h e d s . In
particular, 20 a i r s h e d s were defined. Several
C l a s s I w i l d e r n e s s a r e a s a r e included. Emissions
i n each of t h e a i r s h e d s were converted t o h e c t a r e s
through a f u e l s i n v e n t o r y of t o n s of f u e l per
h e c t a r e and t h e emission c h a r a c t e r i s t i c s defined i n
t h e second s e c t i o n of t h i s paper. Calculated
changes i n combined emissions of p r e s c r i b e d and
w i l d f i r e s a r e defined by t h i s l e a s t a c r e s f o r
i n c r e a s e o r l a r g e s t n e g a t i v e numbers f o r d e c r e a s e
from c u r r e n t emissions. A s a n example, a l l a i r -
sheds on t h e Monterey D i s t r i c t a r e l i m i t e d by
p a r t i c u l a t e p o l l u t a n t s ( t a b l e 3 ) . Most a i r s h e d s on
t h e Los Padres National Forest could s u s t a i n minor
i n c r e a s e s i n p r e s c r i b e d burning. Only t h r e e a i r -
sheds show a need t o decrease t h e combined pre-
s c r i b e d f i r e , w i l d f i r e emissions. A l l three air-
sheds a r e h e a v i l y populated, and t h e F o r e s t must
compete with numerous o t h e r p o l l u t a n t s o u r c e s f o r
t h e a i r resource.
WASHINGTON
F i g u r e 4 b à ‘ C a l c u l a t e wind p a t t e r n s . Mesh l e n g t h i s 4 km by 4 km. Wind-
speed is p r o p o r t i o n a l t o l e n g t h of arrows. Note t h e u n i f o r m i t y of wind
d i r e c t i o n and speed over t h e P a c i f i c Ocean and w i t h i n t h e Willamette
Valley. Winds i n complex t e r r a i n , t h e Coast Range, and t h e Cascade Range
show a h i g h d e g r e e of speed and d i r e c t i o n v a r i a b i l i t y .
 LITERATURE CITED Fox, Douglas G. , and Michael A. Fosberg .
1976. Estimating r e g i o n a l a i r p o l l u t i o n impact.
Boubel, R. W., E. F. Darley, and E. A. Schuck. -
I n Proc. 4 t h I n t . Clean A i r Congress. Int.
1969. Emissions from burning g r a s s s t u b b l e and Union of A i r P o l l u t . Control Assoc. p. 229-
straw. J. A i r P o l l u t . Control Assoc. 19:497- 301.
520.
F r i t s c h e n , Leo, Harley Bovee, Konrad B u e t t n e r ,
C a l i f o r n i a A i r Resources Board. Robert Charlson, Lee Monteith, Stewart P i c k f o r d ,
1977. Three year summary of California air James Murphy, and El 11s Darley.
q u a l i t y d a t a . 346 p. 1970. Slash f i r e atmospheric p o l l u t i o n . Res.
Paper PNW-97, 42 p. P a c i f i c Northwest F o r e s t
C r a i g , C. D . , and M. A. Wolf. and Range Exp. Stn., U. S. Dep. Agric.,
1980. F a c t o r s i n f l u e n c i n g p a r t i c u l a t e concen- .
P o r t l a n d , Oreg
t r a t i o n s r e s u l t i n g from open f i e l d burning.
Atmos. Environ. 14:433-443. Furman, R. William, and Glen F. Brink.
1975. The f i r e weather l i b r a r y : what i t is and
Darley, E. F., F. R. Burleson, E. H. Mateer, J. T. how t o use i t . Gen. Tech. Rep. m-19, 8 p.
Middleton, and V. P o s t e r l i . Rocky Mountain F o r e s t and Range Exp. Stn.,
1966. C o n t r i b u t i o n s of burning a g r i c u l t u r a l Forest Serv., U.S. Dep. Agric., F o r t C o l l i n s ,
waste t o photochemical a i r p o l l u t i o n . J. A i r Colo.
Pollut . Control Assoc. 16:685-690.
G e r s t l e , R. W., a n d D . A. Kemnitz.
D r i v e r , B. L., Donald Rosenthal , and Lynn Johnson. 1967. Atmospheric emissions from open burning.
1979. A suggested r e s e a r c h approach f o r quanti- J. A i r P o l l u t . Control Assoc. 17:324-327.
f ying t h e psychological b e n e f i t s of a i r v i s i -
bility. Proc. Workshop i n V i s i b i l i t y Values. Green, Kathleen, and Robert Wright.
USDA F o r e s t Serv. Gen. Tech. Rep. 18, p. 100- 1977. F i e l d response of p h o t o s y n t h e s i s t o CO 2
105. enhancement i n ponderosa pine. Ecology 58:
687-692.
F e l d s t e i n , M . , S. Duckworth, H. C. Wohlers, and B.
Lusky . Malm, William.
1979. V i s i b i l i t y : a physical p e r s p e c t i v e . Proc.
1963. The c o n t r i b u t i o n s of t h e open burning of
land c l e a r i n g d e b r i s t o a i r p o l l u t i o n . J. A i r Workshop i n v i s i b i l i t y Values. USDA F o r e s t
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Pol l u t Control Assoc. 13: 542-545, 564. Serv. Gen. Tech. Rep. 18, p. 56-68.

Fosberg, Michael A., and Douglas G. Fox. McCall. C. W.. B i l l F r i e s . and Chip Davis.
1976. An a i r q u a l i t y index t o a i d i n determining 1976. ~ u r o r a ~ o r e a l i s on
' record a1 bum Wilder-
mountain land use planning. I n Proc. 4th -
ness, PD-1-6069. Polydor Inc., New York.
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N a t i o n a l Conf on F i r e and F o r e s ~ ~ e t e o r o l o ~ ~ .
USDA F o r e s t Serv. Gen. Tech. Rep. RM-32, p. Parmeter, J. R. , and B. Uhrenhol t .
167-170. 1975a. E f f e c t of smoke on pathogens and o t h e r
fungi. Proc. T a l l Timbers F i r e Ecology Conf .
Fosberg, Michael A., W. E. M a r l a t t , and Lawrence No. 14:299-304.
Krupnak.
l976a. Estimating a i r f l o w p a t t e r n s over complex Panneter, J. R. , and B. Uhrenholt .
terrain. Res. Paper RM-162, 16 p. Rocky Moun- 1975b. Some e f f e c t s of p i n e n e e d l e s o r g r a s s
t a i n F o r e s t and Range Exp. S t n . , F o r e s t Serv., smoke on fungi. Fhytopathol 65:28-31. .
U.S. Dep. Agric., F o r t C o l l i n s , Colo.
Radke, L. F., J. L. Smith, D. A. Hess, a n d P . V.
Fosberg, Michael A., Douglas G. Fox, E. A. Howard, Hobbs .
and J a c k Cohen. 1978. Airborne s t u d i e s of p a r t i c u l a t e and g a s e s
1976b. Non-turbulent dispersion processes i n from f o r e s t f i r e s . J. A i r P o l l u t . Control
complex t e r r a i n . Atmos. Environ. 10:1053-1055. Assoc. 28: 30-34.

Fosberg, Michael A . , and Douglas G. Fox. Ryan, P. W.

1978. Reply: non-turbulent d i s p e r s i o n processes 1974. The q u a n t i t y and q u a l i t y of smoke produced
i n complex t e r r a i n . Atmos. Environ. 12:976. by s o u t h e r n f u e l s i n p r e s c r i b e d burning opera-
tions. B u l l . h e r . Meteor01 . Soc. 55: 70.
Fosberg, Michael A., C h a r l e s D. C r a i g , and Marshall
P. Waters, 111. Ryan, P. W., and C h a r l e s K . McMahon.
1980. A p p l i c a t i o n of i n f r a r e d d a t a from a geo- 1976. Some chemical and p h y s i c a l c h a r a c t e r i s t i c s
synchronous m e t e o r o l o g i c a l s a t e l l i t e i n s u r f a c e of emissions from f o r e s t f i r e s . Paper No.
wind modeling. Proc. 6 t h Conf .
on F i r e and 76-2.3 p r e s e n t e d a t t h e 6 9 t h Annual Meeting of
t h e A i r P o l l u t i o n Control A s s o c i a t i o n [June
F o r e s t Meteorology. Soc. Amer. For. p. 265-
275. 22-July 1, 1976, P o r t l a n d , Oregon], 15 p.
 Hydrocarbon Emissions from vegetation1
David T. Tingey

Walter F. ~ u r n s ~

Abstract: A wide range of volatile organic compounds may be

emitted by vegetation. The identified emittants, however,

are mainly terpenoid in nature. Their emission rates are

controlled primarily by the physical/chemical processes that

regulate hydrocarbon vapor pressure. Emission rates vary

between species and are influenced by environmental factors

such as light and temperature. Regional emission estimates

indicate that vegetation may emit as much as 30 kg of hydro-

carbons k w 2 day-l. The measured atmospheric concentrations

are in reasonable agreement with the estimated emission

rates. Within the atmosphere, these hydrocarbons may par-

ticipate in photochemical reactions leading to aerosol

production and the consumption or formation of ozone.

High levels of ozone have been measured in 1955; Rasmussen 1972; Zimmerman 1979a). Individ-

rural and remote locations far from significant ual species have relatively distinctive emission

anthropogenic sources of oxidant precursors. profiles. For some species, only one or a few

These elevated concentrations may have resulted compounds dominate the emission profile; however,

from long distance transport and/or the photo- other species have a diffuse emission profile with

oxidation of locally-produced biogenic hydro- no dominant compounds (Rasmussen 1972; Zimmerman

carbons. Robinson (1978) proposed that ambient 1979a). Despite the wide range of potentially

hydrocarbon concentrations were governed by both volatile compounds, only isoprene, monoterpenes,

long distance transport and local production. and a few aromatics have been conclusively identi-

Volatile organics, including monoterpenes and fied as emission products from vegetation

isoprene, have been detected in the atmosphere (Rasmussen 1972; Zimmerman 1979a), hence they form

(Rasmussen and Went 1965; Schjoldager and Watine the basis for further discussion.

1978; Whitby and Coffey 1977; Arnts and Meeks

1980; Lonneman and others 1977) and in laboratory

studies shown to produce ozone (Arnts and Gay METHODS FOR ESTIMATING EMISSION RATES

1979), suggesting that they may contribute to

ambient ozone concentrations. A variety of experimental methods have been

used to estimate emission rates. A tree branch or

Plants contain a number of potentially volatile a few small plants were enclosed in a large Teflon

organic compounds including monoterpenes, iso- bag to estimate biogenic hydrocarbon emission

prene, aldehydes, alcohols, and ketones (Meigh rates in the field (Zimmerman 1979a, l979b). The

bag was sealed, evacuated and refilled with hydro-

carbon-free air. A small gas-exchange rate was

maintained through the bag. After an accumulation

Presented at the Symposium on Effects of Air period, the head space was sampled to determine

Pollutants on Mediterranean and Temperate Forest the gas phase concentration. Vertical gradients

Ecosystems, June 22-27, 1980, Riverside, Califor- of temperature, water, and a-pinene, both within

nia, U.S.A. and above the canopy of a loblolly pine (Pinus

Plant Physiologist, Office of Research and taeda L.) plantation were measured and used to

Development, U.S. Environmental Protection Agency, calculate emission rates (Arnts and others 1978).

Corvallis, Ore.; and Chemist, Northrop Services,

Inc., Corvallis, Ore.

Sandberg, D. V. Turner, D. Bruce.

1974. Measurement of p a r t i c u l a t e emissions from 1969. Workbook of atmospheric d i s p e r s i o n e s t i -
f o r e s t r e s i d u e s i n open burning experiments. mates. U.S. Dep. Health, Education, and
Ph. D. t h e s i s , Univ. of Washington. 165 p. Welfare Public Health Service Publ. N.
999-AP-26. 84 p.
Sandberg, D. V., and R. E. Martin.
1975. P a r t i c l e s i z e s i n s l a s h f i r e smoke. Res. U.S. Congress.
Paper PNW-199, 7 p. P a c i f i c Northwest F o r e s t 1977. Clean A i r Act Amendment of 1977. 91 S t a t .
and Range Exp. S t n . , F o r e s t Serv., U.S. Dep. 685, 42 U.S.C. 7401 of seq.
A g r i c . , P o r t l a n d , Oreg.
U.S. Environmental P r o t e c t i o n Agency.
Sandberg, D. V., J. M. P i e r o v i c h , D. G . Fox, and E. 1978. Compilation of a i r p o l l u t a n t emission
W. Ross. f a c t o r s , 3d ed., Supplemental No. 8. AP 42.
1979. E f f e c t s of f i r e on a i r . Gen. Tech. Rep.
WO-9, 40 p. F o r e s t Serv., U.S. Dep. Agric., Vines, Robert G., L. Gibson, A. B. Hatch, M. K.
Washington, D.C. King, D. A. MacArthur, D. R. Packhan, and R. J.
Taylor.
Schaef e r , Vincent J. 1971. On t h e n a t u r e , p r o p e r t i e s , and behavior of
1976. The p r o d u c t i o n of optimum p a r t i c l e smokes brush-fire smoke. CSIRO, Div. of Appl. Chem.
in forest fires. Proc. A i r Q u a l i t y and Smoke Tech. Paper 1. 32 p.
from Urban and F o r e s t F i r e s . National Academy
of Science-Nat i o n a l Research Council. p. Ward, D. E., C. K. McMahon and R. W. Johansen.
27-29. 1976. An u p d a t e on p a r t i c u l a t e emissions from
forest fires. Paper No. 62-2.2. 6 t h Annual
S i n g e r , Michael J . Meeting of A i r P o l l u t . C o n t r o l Assoc. 1 4 p.
1980. Climate v a r i a b l e s i n s o i l e r o s i o n pro-
cesses. Proc. of t h e N a t i o n a l Weather S e r v i c e
.
A g r i c u l t u r a l Meteorology T r a i n i n g Conf , Univ.
of C a l i f ., Davis. Atmos. S c i . Paper 19, p.
75-87.
 Hydrocarbon Emissions from Vegetation
David T. Tingey

Walter F. ~ u r n s ~

Abstract: A wide range of volatile organic compounds may be

emitted by vegetation. The identified emittants, however,

are mainly terpenoid in nature. Their emission rates are

controlled primarily by the physical/chemical processes that

regulate hydrocarbon vapor pressure. Emission rates vary

between species and are influenced by environmental factors

such as light and temperature. Regional emission estimates

indicate that vegetation may emit as much as 30 kg of hydro-

carbons k w 2 day-l. The measured atmospheric concentrations

are in reasonable agreement with the estimated emission

rates. Within the atmosphere, these hydrocarbons may par-

ticipate in photochemical reactions leading to aerosol

production and the consumption or formation of ozone.

High levels of ozone have been measured in 1955; Rasmussen 1972; Zimmerman 1979a). Individ-

rural and remote locations far from significant ual species have relatively distinctive emission

anthropogenic sources of oxidant precursors. profiles. For some species, only one or a few

These elevated concentrations may have resulted compounds dominate the emission profile; however,

from long distance transport and/or the photo- other species have a diffuse emission profile with

oxidation of locally-produced biogenic hydro- no dominant compounds (Rasmussen 1972; Zimmerman

carbons. Robinson (1978) proposed that ambient 1979a). Despite the wide range of potentially

hydrocarbon concentrations were governed by both volatile compounds, only isoprene, monoterpenes,

long distance transport and local production. and a few aromatics have been conclusively identi-

Volatile organics, including monoterpenes and fied as emission products from vegetation

isoprene, have been detected in the atmosphere (Rasmussen 1972; Zimmerman 1979a), hence they form

(Rasmussen and Went 1965; Schjoldager and Watine the basis for further discussion.

1978; Whitby and Coffey 1977; Arnts and Meeks

1980; Lonneman and others 1977) and i n laboratory

studies shown to produce ozone (Arnts and Gay METHODS FOR ESTIMATING EMISSION RATES

1979), suggesting that they may contribute to

ambient ozone concentrations. A variety of experimental methods have been

used to estimate emission rates. A tree branch or

Plants contain a number of potentially volatile a few small plants were enclosed in a large Teflon

organic compounds including monoterpenes, iso- bag to estimate biogenic hydrocarbon emission

prene, aldehydes, alcohols, and ketones (Meigh rates in the field (Zimmerman 1979a, 1979b). The

bag was sealed, evacuated and refilled with hydro-

carbon-free air. A small gas-exchange rate was

maintained through the bag. After an accumulation

Presented at the Symposium on Effects of Air period, the head space was sampled to determine

Pollutants on Mediterranean and Temperate Forest the gas phase concentration. Vertical gradients

Ecosystems, June 22-27, 1980, Riverside, Califor- of temperature, water, and a-pinene, both within

nia, U.S.A. and above the canopy of a loblolly pine (Pinus

Plant Physiologist, Office of Research and taeda I,.) plantation were measured and used to

Development, U.S. Environmental Protection Agency, calculate emission rates (Arnts and others 1978).

Corvallis, Ore.; and Chemist, Northrop Services,

Inc., Corvallis, Ore.

 Semi-quantitative estimates of emission rates

tion gradient. The larger the concentration

were made using static gas-exchange chambers

gradient, the larger the hydrocarbon flux; con-

containing either detached leaves, twigs or whole

versely, the larger the resistance to mass trans-

plants (Rasmussen 1970; Rasmussen 1972; Sanadze

fer, the smaller the flux (Nobel 1974).

and Kalandadze 1966a). Within these chambers

carbon dioxide may be elevated or depleted depend-

Only hydrocarbons with appreciable vapor pres-

ing on light intensity. This may modify plant

sures at ambient temperatures will be emitted at

metabolism and the stomatal aperture; humidity

significant rates. The vapor phase concentration

will increase; and high concentrations of hydro-

of hydrocarbons within the leaf is controlled by

carbon gases will build up within the chamber

the liquid phase concentration, vapor pressure,

reducing diffusion gradients. These factors can

and solubility. The vapor pressure of terpenoid

lead to an underestimation of emission rates.

compounds increases exponentially with the temper-

Dynamic mass-balance gas-exchange chambers and

ature (Jordan 1954). Monoterpene emission rates

leaf cuvettes which simulate the gaseous environ-

from black sage (Salvia mellifera Greene) and

ments of plants in the field have also been used

slash pine (Pinus elliottii Engelm.) also exhibit

to estimate hydrocarbon emissions (Kamiyama and

an exponential increase with temperature (Dement

others 1978; Tingey and others 1979 and 1980;

and others 1975; Tingey and others 1980), indi-

Tyson and others 1974). These chambers may be

cating that vapor pressure is a significant factor

used to determine the influence of environmental

in controlling emissions. Emission rates from

factors on emission rates.

dead slash pine needles and black sage leaves are

similar to emission rates from live tissue (Tingey

and others 1980; Dement and others 1975), support-

MECHANISM OF HYDROCARBON VOLATILIZATION

ing the concept that the volatilization is primar-

ily a physical process. Hydrocarbons with chain

Terpenoid Biosynthesis
lengths greater than Clo generally have low vapor

pressures and will not have a large emission rate.

Knowledge of the mechanism and sites of terpen-

oid biosynthesis aids in understanding the factors

When monoterpenes occur in high concentrations

controlling emission rates. Terpenoid biosynthe-

in resin ducts, oil cells or glandular trichomes,

sis starts with the conversion of carbon dioxide

their emission rates are essentially independent

to sucrose with its subsequent metabolism to

of concentration. Large pools of isoprene, how-

acetyl-CoA and mevalonic acid to form isopentenyl

ever, have not been detected. Below 35OC (the

pyrophosphate. The hemiterpene, isoprene, (Cg) is

boiling point for isoprene), the emission rate is

formed from isopentenyl pyrophosphate; monoter-

closely linked to its synthesis rate. Above 35OC,

penes (Clo) are formed from a condensation of

the emission rate is diffusion-limited (Tingey and

dimethylallyl pyrophosphate and isopentenyl pyro-

others 1979).

phosphate. Subsequent additions of isopentenyl

pyrophosphate units form higher homologs in the

Monoterpenes have a low aqueous solubility

terpenoid series (Loomis and Croteau 1980).

(hydrophobic) and higher vapor pressures than

similar, more hydrophilic compounds. Therefore,

The conditions that promote isoprene biosynthe-

monoterpenes would be emitted at a higher rate

sis indicate that it is likely synthesized within

than similar oxygenated compounds at equal concen-

the chloroplast. Isoprene biosynthesis is

trations within the tissue. Similarly, if the

affected by metabolic inhibitors that regulate

concentration exceeds its aqueous solubility

photorespiration (Loomis and Croteau 1980).

limit, then vapor pressure and emissions are

independent of tissue concentrations.

Monoterpenes appear to be ubiquitous in higher

plants (Loomis and Croteau 1980). The accumula-

Resistance to mass transfer can occur along

tion or secretion of significant quantities of

either a stomatal or cuticular pathway (Nobel

monoterpenes is associated with the presence of

1974). Either one or both pathways may be signif-

secretory structures such as glandular hairs or

icant, depending on the species. Stomata are

trichromes, oil cells, resin ducts or glandular

apparently the main pathway for diffusion of

epidermises, and lysogenous spaces. It is gener-

monoterpenes (Hanover 1972), isoprene and other

ally assumed that monoterpenes are synthesized

compounds synthesized within the leaves. However,

within the secretory cells, although this point

for plants with glandular trichomes or glandular

has not yet been conclusively demonstrated (Loomis

cells in the epidermis, such as in the Labiatae

and Croteau 1980).

and Solanaceae, the cuticular pathway is the main

one for diffusion.

Hydrocarbon Diffusion from Plants

BIOGENIC EMISSIONS

Gaseous diffusion between the plant and its

environment is controlled by the chemical poten-

Emission Rates

tial gradient between the inside and the outside

of the leaf and the resistance to mass transfer

Emission rates for several plant species are

along the diffusion pathway. The chemical poten-

shown in table 1. Total non-methane hydrocarbon

tial gradient can be approximated by a concentra-

and monoterpene emission rates are similar among

 Hydrocarbon emission rates/unit tissue multi-

Table 1--Biogenic hydrocarbon emission rates estimated at 30°C

plied by biomass density yield emission factors.

Species TNMHC1 Isoprene Monoterpenes References

Emission factors for the Tampa-St.Petersburg,

fg [g dry weightl-I hr-l- Florida, area (Zimmerman 1979a), indicate that 92

Slash Pine 4.1 2.6 Zimerman 1979a

percent of the total non-methane hydrocarbon emis-

Longleaf Pine 7.3 5.6 Zitmerman 1979a

Sand Pine 13.6 11.0 Zimerman 1979a

sions occur in the following four land use types:

Cypress 14.2 8.1

6.4
Zimerman 1979a

Tingey and other? 1980

evergreen forests (35 percent); citrus groves (22

Slash Pine
Loblolly Pine 3.7 Arnts and others 1978
percent) ; pasture and rangeland (19 percent) ; and
Cryptomeria 3.0 Kamiyama and others 1978
residential areas (16 percent). The remaining 8

Laurel Oak 12.6 10.0 Zimerman l979a

Turkey Oak 26.5 23.4 Zimmerman 1979a

percent was distributed among crop lands, decid-

Bluejack Oak 56.4 43.9 Zimerman 1979a

Live Oak 10.8 9.1 Zimnennan 1979a

uous forests, mangroves, freshwater, marine, and

Live Oak 41.2 Tingey and others 1980

barren lands. Emission factors for trees were

Willow 22.1 12.4 Zimerman 1979a

Saw Palmetto 11.5 8.6 Zimmerman 1979a

approximately 6 mg m-2 hr-l; shrubs, 2.0 mg m-2

hr-l; pastures, mud flats and other land use types

Mean 7 Hardwood
Trees--Isoprene 20'0

Flyckt and others 1980

were less than 0.1 mg m-2 hr-I .
Wax Myrtle Zimennan 19798

Persimon Zimerman 1979a

Emission factors were developed to characterize

Orange Zimerman 1979a

Grapefruit Zimnerman 1979a

the various biomes in the United States:

Red Maple Zimerman, l979a

Hickory Zimnennan, 1979a

Emission Rate (mg w2hr-l)

Mean 10 Hardwood
Trees--Nm-Isoprene 1 7'3

Flyckt and others, 1980

Biome : Night
Total non-methane hydrocarbons

Grassland 0.2

Sclerophyll Scrub 1.4

the conifers and as much as 50 percent less than

Temperate Rain Forest 9.4

emission rates from hardwoods that emit isoprene.

Deciduous Forest 2.7

Monoterpenes account for 50 to 75 percent of the

Coniferous Forest 2.7

total non-methane hydrocarbon emissions in con-

Desert 0.7

ifers. Similarly, isoprene accounts for 60 to 90

Tundra, Alpine Fields 0.4

percent of total non-methane hydrocarbon emissions

from isoprene emitters. Plants whose emissions

Data from Zimmerman

were not dominated by either isoprene or a few

monoterpenes had emission rates roughly similar to

the conifers.
Daytime biome emission factors ranged from a low

of 0.3 mg m-2 hr-I for grasslands to a high of

Total non-methane emission rates were estimated

10.7 for temperate rain forests. Nighttime emis-

at several locations in the United States on

sions were 10 to 60 percent lower reflecting, in

similar vegetation types:

part, the absence of isoprene emissions.

Vegetation Type Emission Rate1

and Location: Environmental Influences on Emission Rates

(ye [g dry weight]-I hr-l)

Conifers
Isoprene production is light dependent, and

Washington
persists for only a few minutes when plants are

Florida
darkened (Rasmussen and Jones 1973; Sanadze and

Texas
Kalanadze 1966b). Emissions increase with

increasing light intensity until a maximum is

Oaks
reached and then remain constant (Sanadze and

California
Kalandadze 1966a; Tingey and others 1979); similar

Florida
to a light saturation curve for photosynthesis.

Texas
Isoprene emissions are light saturated at moderate

light intensities (Sanadze and Kalandadze 1966a;

Non-Conifers, Non-Isoprene Emitter

Tingey and others 1979). In contrast, monoterpene

Washington 7.8
emissions from slash pine, black sage, and several

California 4.1
other plant species, are similar in the dark and

Florida 4.7
light (Tingey and others 1980; Dement and others

Texas 0.2
1975; Rasmussen 1972).

Data from Zimmerman 1979b and c.

Isoprene emissions increase sigmoidally with

temperature; low emissions occur at 18-20° and

increase exponentially between approximately 20

Emission rates for the conifers, oaks and non-

and 35OC, then plateau. At higher temperatures

conifer, non-isoprene emitting vegetation are

(between 43 and 47OC), depending upon the species,

similar within each vegetation type. This indi-

there is the large, precipitous decline in iso-

cates an apparent high uniformity in emission

prene emissions (Sanadze and Kalandadze 1966a;

rates among locations using the same estimation

Rasmussen and Jones 1973; Tingey and others 1980).

technique.

The increase in isoprene emissions with temper-

Table 2"-Estimated emissiops for biogenic hydrocarbons.

ature is greater at high light intensities than

Emission

low (Tingey and others 1979). Isoprene emissions

Location Emissions Factor* References

from several hardwood trees and live oak (Quercus

---metric tons- kg km-x day-'

virginiana Mill.) increased at approximately 20

World 1.75 x 10'/year Went 1960

World 4.38 x lo8/year Rasmussen and Went 1965

and 16 percentI0C (20-35OC), respectively (Flyckt

World 8.30 x 108/year Zimwrnan 1979b

and others 1980; Tingey and others 1979).

United States 0.23-4.64 x lo7/year Rasmussen 1972

United States 6.5 x lo7/year Zimeman 1979b

Florida 157.0lday 32.3 Zimeman 1979a

(81 x 60 km)

Monoterpene emissions from conifers, black sage

Texas . .
32.4ldav 27.5 Zimeman 1979~
and hardwood trees increase exponentially with the

temperature (Arnts and others 1978; Kamiyama and

(38 x 31 to)

Pennsvlvania 1 3.580.0/day 30.7 Flyckt and others 1980

others 1978; Rasmussen 1972; Flyckt and others

1980; Dement and others 1975; Tingey and others

1980). The relative percent increase per degree

Regional Emissions

temperature varies between species and ranges from

approximately 6 to 20 percent/OC. In conifers,

Biogenic hydrocarbon emission rates for a

extensive genetic variations in monoterpene pools

variety of plant species and biomass estimates,

(Hanover 1972) may explain the lack of an exponen-

were used to estimate emissions for various areas

tial relationship between temperature and emission

(table 2). The emission rate estimates for vari-

rates in some field studies (Flyckt and others

ous estimation scales (world, United States, or

1980).
regional) were approximately similar. The close

agreement between the emission estimates from the

Typical diurnal emission patterns for isoprene

three regional studies may have occurred because

and monoterpenes and environmental conditions for

the same experimental approach was used. Emission

an average of summer days in Tampa, Florida, were

estimates for Pennsylvania and the Tampa-

used to illustrate the interaction of light and

St.Petersburg, Florida, area indicate that bio-

temperature on terpenoid emissions (Tingey and

genic emissions range from 12 percent greater to

others 1979, 1980). During early morning and late

20 percent less than anthropogenic emissions

afternoon, when the leaves are not light-saturated

(Flyckt and others 1980; Zimmennan 1979a; Wayne

and the temperature is moderate, light would be

and Kochis 1978).

the main factor controlling isoprene emissions

from live oak. However, during most of the day,

the leaves of the canopy are light-saturated; thus

Relationship Between Primary Productivity

varying air temperature would control emission

and Emission Rates

rates. More than 80 percent of the isoprene

emissions were expected to occur after mid-

A relationship between biogenic hydrocarbon

morning, ceasing in the evening. Monoterpene

emissions and primary productivity should exist

emission rates . from slash pine increase after because they are ultimately derived from photo-

sunrise, peaking during early afternoon, and

synthetically fixed carbon dioxide. Measurements

declining to a minimum shortly before sunrise.

of the ratio of carbon lost as volatile terpenoids

Approximately 55 percent of the total daily mono-

to primary productivity for several tree species

terpene emissions occurred during daylight hours

indicated loss rates of 0.2 to 2 percent for

(0600-1800) with an additional 25 percent emitted

isoprene and 0.06 to 0.4 percent for monoterpenes

between sunset (1800) and midnight (2400).

(Sanadze 1969; Tingey and others 1979, 1980; Tyson

and others 1974). The relationship between pri-

Seasonal emission patterns were estimated for

mary productivity and biogenic hydrocarbon emis-

individual ponderosa pine and red oak trees

sions could be used to delineate geographic areas

(Flyckt 1979). Monoterpene emissions from pon-

where emissions would tend to be high. Based on

derosa pine were sinusoidal, at a maximum during

the work of Leith (1975), primary productivity is

late spring (May and June), declining to a minimum

highest in the Southeast, followed by the

around November, and then gradually increasing.

Mississippi Valley area and the central part of

In contrast, isoprene emissions from red oak were

the United States and lowest in the Great Basin

maximum during July and August and decreased

and the Southwest. Zimmennan (1979b) estimated

during the fall. No isoprene emissions were

that 45 percent of the total national biogenic

detected during the winter; emissions reappeared

hydrocarbon emissions occurred in the South, an

in the spring with the initiation of new leaves.

area with the highest primary productivity in the

It is not clear whether seasonal emission changes

United States (Leith 1975).

were due solely to changes in environmental condi-

tions or were, in part, due to changes in terpen-

oid pools. In addition to changes in the emission

AMBIENT CONCENTRATIONS OF TERPENOIDS

rates, there were also qualitative changes in the

monoterpene emissions throughout the year (Flyckt

Biogenic hydrocarbons were measured in the

and others 1980).

atmosphere over several vegetation types. The

average isoprene concentrations varied from 10 ppb

carbon for an oak forest to 0.1 ppb carbon for a

pine forest. The average monoterpene concentra-

tions ranged from 24 ppb carbon in the coniferous

forest in Norway to 2.7 ppb carbon in the conifer-

ozone, PAN, formic acid, acetone, aldehydes, CO,

ous forest in Idaho (Schjoldager and Watine 1978;

and Coy formation when terpenes were irradiated in
Arnts and Meeks 1980; Coffey and Westberg 1978).
the presence of nitrogen oxides. The amount of

Measured ambient concentrations of biogenic hydro-

ozone formed depended on the C / N O ratio. At a

carbons and ambient concentrations predicted from

low C/NO ratio, 1 ppb C from terpenoids produced

biogenic emission rates are in reasonable agree-

2-4 ppb ozone. However, when the ratio was large,

ment (Zimmerman 1979c; Flyckt and others 1980;

1 ppb C produced 0.3 to 0.1 ppb ozone, suggesting

Scully, 1979; Coffey and Westberg, 1978).

that terpenoids also consume ozone. Eschenroeder

(1974) and Coffey and Westberg (1978) concluded

Atmospheric hydrocarbon concentrations are

that emissions of biogenic hydrocarbons did not

dependent on emission rates, mixing height, and

significantly alter ambient ozone concentrations

the reactivities (photolysis and ozonolysis) of

through scavenging reactions.

the individual components. Peterson and Tingey

(1980) used a box model to estimate ambient air

Zimmerman (1979~) suggested that photooxidation

concentrations of isoprene and monoterpenes. The

of isoprene from forests could contribute 22 ppb

predicted isoprene concentrations increased during

ozone to the ambient concentration. Similarly,

the daylight hours, reaching a maximum at mid-

Coffey and Westberg (1978) suggested that emis-

afternoon and then disappearing during the early

sions from coniferous forests could react to add 1

evening when isoprene emissions ceased. Predicted

to 5 ppb ozone to the ambient air.

ambient monoterpene concentrations were the lowest

during mid-day when atmospheric dilution, photo-

oxidation, and ozonolysis, were the highest,

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682.

 Background Levels of Trace Elements in

Forest Ecosystems1

G. Bruce Wiersma and Kenneth W. ~ r o w n ~

Abstract: This study was conducted as part of a project to

develop a pollutant monitoring system for biosphere

reserves. Sampling was carried out in the Great Smoky

Mountains National Park and Olympic National Park. Results

are reported for copper, lead, manganese, aluminum, calcium

and phosphorus. Olympic National Park had much lower levels

of lead and copper than Great Smoky Mountains National Park.

Moss appeared to be a good collector for lead and copper.

Results indicate that reference levels for trace elements

can be established for remote areas, although they cannot be

considered true background levels.

Introduction The reserves are areas that can be used to mon-

itor the behavior of pollutants that have long

range transport characteristics such as trace

Biosphere reserves are remote, pristine areas elements (Zoller and others 1974; Duce and others

set aside in perpetuity. A pollutant monitoring 1975; Thrane 1978; Weiss and others 1971;

system is being developed for implementation on Scheslinqer and others 1974; and Chow and Earl

the reserves (Wiersma and others 1978a; Wiersma 1970).

and others 1979). Purpose of monitoring

pollutants on these areas are: Since many trace elements have the potential for

long-term transport the question becomes, in a

1. to serve as locales for background monitoring program for background areas, what

reference levels of certain pollutants . elements should be of prime interest. Two

parameters can be used to estimate potential for

2. to provide a frame of reference against long-term transport. First, elements that have a

which changes in impacted areas can be measured high vapor pressure, and second, elements that

would have a significant small particle (1.0 p or

3. to reflect changes of a global nature less) association. There is evidence that these

before such changes are obvious in more impacted two phenomena may work in conjunction. Ondov and

areas. others (1977a) analyzed the relationships existing

between particle size and elemental composition in

power plant emissions. They stated that elements

with low vapor pressures tended to be associated

-presented at the Symposium on Effects of Air with larger particle sizes. In a subsequent paper

Pollutants on Mediterranean and Temperate Forest Ondov and others (1977b) listed several elements

Ecosystems, June 22-27, 1980, Riverside, as having significant small particle association

California, U.S.A. including manganese, lead and copper.

2~coloqist,U. S. Environmental Protection

Agency, Environmental Monitoring Systems Kyser and others (1978), using a variety of

Laboratory, U.S. Environmental Protection Agency, microprobe analytical techniques, found that

Las Vegas, Nevada; Botanist, U.S. Environmental arsenic, cadmium, cobalt, chromium, manganese,

Protection Agency, Environmental Monitoring nickel, lead, sulphur antimony, selenium,

Systems Laboratory, U.S. Environmental Protection thallium, vanadium and zinc were present on

Agency, Las Vegas, Nevada. particles primarily as surface material. This

lends further support to the hypothesis that

elements with high vapor pressure tend to condense

on smaller particles.

Methods

Analytical

This paper will present data for trace element

levels in vegetation and forest litter. The ana-

lytical procedure used was spark source emission

spectroscopy (SSES) which determines 26 elements

per sample. The analytical procedure has been

previously described by Alexander and others

( 1975).

Every tenth sample submitted was a quality

1 Air
assurance sample, alternating between known value
T Water
samples and replicated samples. Samples were sub-
0 Vegetation-Soil
mitted in a set order. The analytical laboratory

was required to analyze the samples in the order

Figure 2-Sampling locations in the Olympic

submitted.
National Park Biosphere Reserve.

Field Sampling
Results and Discussion

Two biosphere reserves, the Great Smoky

Mountains and Olympic National Park, have been

Trace Element Selection

sampled as part of a pilot research project to

develop a cost effective pollutant monitoring

system. Great Smoky Mountains National Park was

The 26 trace elements analyzed by SSES are

originally sampled in the fall of 1977 and again

given in table 1.

in 1978. The results of this effort have been

reported by Wiersma and others (1979). Olympic

National Park was sampled in the summer of 1979.

Table l~elementsand detection limits for spark

source emission spectroscopy.

Figures 1 and 2 show sample site locations for

the Great Smoky and Olympic National Parks

respectively. Details of the Olympic study design

Element PPm Element PPm

are presented by Brown and Wiersma (1979).

Na

ca

M9

Zn

cu

Fe

Mn

A1

Si

Ti

*Â Sampled Autumn, 1977

Sampled Spring, 1978

Figure 1--Sampling locations in the Great Sknoky

Using this listing and selection criteria

Mountains Biosphere Reserve.

previously described, we are limiting our

presentation in this paper to copper, lead and

manganese. Also included are two biologically

essential elements, calcium and phosphorus and one

non-essential element, aluminum, which also is not

one of the elements which has the potential for

long-term transport.

 Quality assurance results are shown in table 2.

samples, but the rest of the vegetation samples

Good agreement exists between the analytical

reflected a slight decline in lead levels. A
results and certified levels. In addition, the
similar relationship was shown for copper.

replicated samples, which were analyzed sequen-

tially, showed no drift and had acceptable

Studies using vegetation, particularly for

reproducibility.
those elements where root uptake is small, as

indicators of airborne pollution have ranged from

interception phenomenon of vegetative surfaces for

Table 2--Quality assurance results for elements

modeling purposes (Shreffler, 1978; Davidson and

from Great Smoky Mountains.

others 1976) to the use of individual species as

collectors of airborne pollutants. Smith (1977)

has reviewed the probability of urban vegetation

malytical

filtering out airborne particulates. He reported

results

that fine hairs on vegetative surfaces increase

for QA

particle trapping phenomenon. Carlson and others

samples

(1976) and Wedding and others (1975) found in

controlled studies that rough, pubescent leaves

entrap seven times more particles than smooth

Phosphorus nonpubescent leaves and the particle load

1,950 2,000 to Not significant

increases with leaf area sometimes by a factor of

2,200
10. Removal of particle from vegetation surface

appears to be through solubilization in rain and

Lead 44.7 44 Not significant

not physical impaction from droplets (Carlson and

others 1976). Compounding this phenomenon is data

Copper 12.0 11.0 to Not significant

reported by Harris and others (1976) which states

13.0
"...all elements are in a far more soluble phase

Manganese in the ambient aerosol (and are associated with

92.4 87 to 95 Not significant

particles retained by biological and inert

surfaces) than in fly ash collected from in stack

Calcium 2.1 2.1 3 Not significant deflector plates...".

~luminum~ - - Not significant From the above discussion, plants with large

leaf surface areas, or those with very rough

pubescent surfaces (ferns, witch hobble) should

'AS determined by 95 pet. correlation coeffi-

collect larger particulate loads, but because of

wash off, they probably cannot be expected to

cient of replicated samples and site numbers

increase the particle load throughout a growing

analyzed in order with site number

season unless another phenomena were at work.

2 ~ standard
o

This is shown in table 4 by a decrease for lead

3pct. by weight
and copper concentrations for ferns and witch

hobble when compared to the results in table 3.

Manganese, however, does not follow this pattern.

Table 3 shows elemental levels for seven

Tyler (1972) states that mosses, via passive

species of vegetation collected in the Great Smoky

ion exchange, can accumulate a variety of airborne

Mountains in the spring of 1978. Lead levels are

elements. If this is the case then solubilization

highest in moss samples. Copper levels tend to be

of surface material will not be an important

higher in wood fern and witch hobble, while

removal process and fall moss samples should have

manganese appears to be higher than previously re-

higher levels than spring, particularly for copper

ported for agricultural crops (Hemphill 1972).

and lead. Data in tables 3 and 4 tends to support

However, Romney and others (1977) report manganese

this hypothesis.

levels for desert vegetation ranging from approxi-

mately 20 ppm to 220 ppm. Van Hook and others

The forest floor of the Great Smoky Mountains

(undated) report manganese values in chestnut oak

was sampled in the spring of 1978. Two types of

and hickory on the walker Branch water shed

samples were collected, the first was the unin-

ranging up to 1,000 ppm. Grodzinska (1978)

corporated organic material and the second was the

reported manganese levels in moss from Poland

partially decomposed material of the fermenta-

ranged from 79 to 880 ppm. Therefore, the levels

of manganese reported appears reasonable when

tion layer. The results are shown in table 5.

Except for manganese, significant differences

compared to other studies, particularly from

existed between the litter and fermentation layer

forested areas. Calcium and phosphorus levels

for all trace elements. The fermentation layer

appear to be equal to or slightly below calcium

showed an increase in lead, copper, phosphorus,

levels reported for agricultural crops (Hemphill

aluminum, and a significant decrease in calcium.

1972).

Site to site correlations for each element,

Table 4 presents the results for the second

excluding aluminum, were significant between

unincorporated organic matter and the fermentation

sampling that occurred in the fall of 1978. There

layer.

was a large increase in lead levels in moss

Table 3--Average concentration of selected elements for samples collected in the Great Smoky Mountains

biosphere reserve, spring 1978.

Lead Cop&= r Manganese Calcium Phosphorus Aluminum

^9/9 ^9/9 ^9/9 pet. ~919 M9/9

Moss 42.3 13.4 368 0.32 1,430 1,410

Yellow birch 12.2 13.2 2,090 1.38 2,540

165

Betula

allegheniensis

Red maple

--
Acer rubrum

New York fern

Thelypteris

novaborecensis

Wood fern

Dryopteris

spinulosa

Witch hobble

Viburnum

alnifolium

Fraser fir

Abies fraseri

Table &-Average concentration of selected elements for samples collected in the Great Smoky Mountains

biosphere reserve, fall 1978.

Lead CoPPr Manganese Calcium Phosphorus Aluminum

~ / l ~ / 9 P3/9 pet. Id9 ~-9/<3

Moss

Yellow birch 10.5 2,370 1.50 2,360 154

Red maple 6.6 1,070 1.14 1,920 66

1
1
New York fern 9.9 3.9 1,000 0.68 854 2,040

Wood fern 5.3 11.5 1,350 0.50 2,200 424

Witch hobble

Fraser fir
I 13*3

0.2
9.2

3.8
2,800

915
1.50

0.30
1,700

2,680
426

375

Reiners and others (1975) found lead levels in Wiersma and others (1978b, 1980) indicate that the

New Hampshire in litter increase with altitude. probable source of this lead is from anthropogenic

At extreme elevations, a slight decrease was

activities.

noted. The levels of lead in litter ranged from

35 to 336 ppm. The fir forest sites had the Some data are available from samples collected

highest lead concentrations. Figure 3 shows a in the summer of 1979 in Olympic National Park.

similar relationship from our data for lead in the The forest floor was not sampled by the method

Smoky Mountains. This relationship did not.exist previously used in the 1978 Smoky Mountains study.

for any of the other elements. Previous work by With the modified technique, unincorporated

Table 5 ~ C o m p a r i s o nof elemental l e v e l s i n unincorporated organic m a t e r i a l and t h e fermentation l a y e r ,

Great Smoky Mountains biosphere r e s e r v e , s p r i n g 1978.

Unincorporated Fermentation Coefficient

organic m a t e r i a l layer correlation ( 8 df)2

Manganese yg/g

calcium p e t . 1 0.89 0.58 -5.22 0 .W4

Phosphorus p e t .

Aluminum M / g
I 1,980 5,420 Not Calculated

Paired "T" t e s t
'Element i n unincorporated organic m a t t e r v e r s u s element i n fermentation l a y e r .
'95 p e t . confidence
'99 p e t . conÂidence

Copper l e v e l s i n moss i n t h e G reat Smoky Mountains

averag e about 3 t i m e s g r e a t e r than i n t h e Olympic
National Park. Lead l e v e l s i n t h e Smoky Mountains
a r e 7 t o 19 times g r e a t e r than t h e Olympic
National Park.

A similar relationship exists for the forest

f l o o r samples:

Comparison of f o r e s t f l o o r samples from

Olympic National Park and Great Smoky
Mountains National Park. (yg/g)

01ympic
litter

30'00 35'00 40'00 4500 5000 55'00 6000

Smokys
Elevation (Feet) unincorporated
organic m a t t e r
Figure 3--Relationship between l e a d r e s i d u e s and s p r i n g 1978
a l t i t u d e i n t h e Great Smoky Mountains Biosphere
Reserve. Smokys
fermentation
s p r i n g 1978
organic m a t e r i a l s were sampled along with t h e
fermentation l a y e r .
True "background" l e v e l s f o r many t r a c e
Copper and l e a d l e v e l s i n moss and l i t t e r a r e elements a r e probably impossible t o determine.
compared f o r t h e two biosphere r e s e r v e s : Lead l e v e l s f o r moss i n t h e Smoky Mountains appear
high b u t i n l i n e with v a l u e s r e p o r t e d i n t h e
Comparison of moss samples from Olympic literature. For example Ruhling and Tyler (1968)
National Park and Great Stooky Mountains analyzed museum samples of moss. They found
National Park. (pg/g) samples c o l l e c t e d a f t e r 1950 had l e a d concentra-
t i o n s of 80 t o 90 ppm. Samples c o l l e c t e d around
Copper -

Lead 1860 contained average l e a d l e v e l s of 20 ppm.

These r e s e a r c h e r s believed t h a t t h e 20 ppm l e v e l
Moss-Olympic 4.4 5.7 did not represent "natural" lead levels. They
Moss-Smoky Spring 13.5 42.4 suspected " n a t u r a l " l e a d l e v e l s might be
Moss-Smoky F a l l 15.3 108.0 considerably lower. The moss samples from Olympic
National Park averaged 5.7 ppm lead with some
Duce, R. A., G. L. Hoffman, and W. H. Zoller.

remote sites having average lead levels of 0.4 ppm

1975. Atmospheric trace metals at remote

and 2.2 ppm. Hirao and Patterson (1974) estimated

northern and southern hemisphere sites -
for a remote site on the high sierra crest that 97
Pollution or natural? Science 187:59-61.

percent of the lead detected was from anthro-

pogenic sources. Therefore, even for sites as

Grodzinska, Krystyna.

remote as the high Dosewallips/High Quinault (over

1978. Mosses as bioindicators of heavy metal

13 miles from the nearest road) in a park that

pollution in Polish national parks. Water,

primarily receives wind off the Pacific Ocean, it

Air and Soil Pollution 9:83-97.

may not be appropriate to consider the lead levels

natural background levels.

Harris, W. F., B. S. Ausmus, G. J. Dodson, Sidney

Draggan, G. K. Eddlemon, Cyrus Feldman, J. M.

It is our opinion that reference levels can be
Giddings, J. W. Huckabee, D. R. Jackson, S. A.

established in vegetation in remote areas for a

Janzen, M. J. Levin, S. E. Lindberg, L. K. Mann,

variety of trace elements. Sites should be

E. G. OINeill, R. V. OINeill, Cheryl B. Phillips,

regionally representative and sampling should be

B. M. Ross, W. J. Selvidge, D. S. Shriner, R. R.

repetitive through time, at least once a year,

Turner, P. Van Voris, Martin Witkamp.

preferably twice a year. Biosphere reserves are

1976. Environmental behavior of trace

ideal places to use because of their protected

contaminants. The role of vegetation aerosol

nature and the fact that they are selected to be

scavenging. ORNL-5257. Environmental

representative of various biological systems.

Sciences Div. Annual Progress Report. Oak

Ridge National Laboratory, Oak Ridge,

Monitoring systems can and should be estab-

Tennessee.

lished in these reserves for the purposes listed.

Hemphill, D. D.

1972. Availability of trace elments to plants

Literature Cited
with respect to soil-plant interaction.

Annals New York Academy of Sciences

199:46-61.

Alexander, G. V., D. R. Young, D. J. McDermott,

M. J. Sherwood, A. J. Mearns, and 0. R. Lunt.

Hirao, Y., and C. C. Patterson.

1975. Marine organisms in the Southern

1974. Lead aerosol pollution in the high sierra

California bight as indicators of pollution.

over-rides natural mechanisms which exclude

-
In International Conference on Heavy Metals in
lead from a food chain. Science

the Environment. [1975. Toronto, Canada],

184:989-992.

p. 955-972.

Keyser, T. R., D. F. S. Natusch, C. A. Evans, Jr.

Brown, K. W. and G. B. Wiersma.
and R. W. Linton.

1979. Pollutant monitoring in Olympic National

1978. Characterizing the surfaces of

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Francisco, Calif.] 12 p.
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Fisher, D. Silberman and B. A. Prentice.

Carlson, R. W., F. A. Bazzaz and J. J. Stukel and

1977a. Interlaboratory comparison of neutron

J. B. Wedding.
activation and atomic absorption analyses of

1976. Physiological effects; wind reentrainment

size-classified stack fly ash. Preprint

and rainwash of Pb aerosol particulate

UCRL-78194. Lawrence Livermore Laboratory,

deposited on plant leaves. Environmental

Livermore, Calif.

Science and Technology 10(12):1139-1142.

Ondov, J. M., R. C. Ragaini, A. H. Bierman, C. E.

Chow, T. J., and J. L. Earl.

Choquette, G. E. Gordon and W. H. Zoller.

1970. Lead aerosols in the atmosphere:

1977b. Elemental emissions from a western coal

increasing concentrations. Science

fired power plant: preliminary report on

l69:577-580.
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Preprint UCKL-78825. Lawrence Livennore

Davidson, C. I., S. V. Herring and S. IS.

Laboratory, Livermore, Calif. 7 p.

Friedlander.

1976. The deposition of ~b-containing

Reiners, W. A., R. H., Marks and P. M. Vitousec.

particles from the Los Angeles atmosphere.

1975. Heavy metals in subalpine and alpine

-
In Proc. International Conference on
soils of New Hampshire. OIKOS 26(3):264-274.

Environmental Sensing and Assessment. [Sept.

14-19, Las Vegas, Nevada] pp. 6-3 to 6-4.

Romney, E. M., A. Wallace and G. V. Alexander.

1977. Boron in relationship to a coal-burning

power plant. Communications in Soil Science

and Plant Analysis 8(9):803-807.

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Wiersma, G. B., C. W. Frank, K. W. Brown and C. I.

1968. An ecological approach to the lead

problem. Bot. Notiser 121:338-341.

Davidson.
1980. Lead particles in the Great Smoky

~ountainsBiosphere Reserve. EPA-600/

Schlesinger, W. H., W. ,A. Reiners and D. S. 4-80-002. Environmental Monitoring

Knupman. Systems Laboratory, U.S. Environmental

1974. Heavy metal concentrations and deposition

Protection Agency, Las Vegas, Nevada.

in bulk precipitation in montane ecosystems of

New Hampshire, U.S.A. Environmental Pollution

Wiersma, G. B.

6:39-47.
1979. Kinetic and exposure commitment analyses

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Shreffler, J. H.
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1978. Factors affecting dry deposition of so2

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19 p.

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Van Hook, R. I., W. F. Harris, G. S. Henderson and

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1978b, Development of a pollutant monitoring

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Effects of Chronic Exposures to

Gaseous Pollutants on Primary
Production Processes

Photochemical Oxidant Impact on

Mediterranean and Temperate Forest
Ecosystems: Real and Potential
Effects
2
John M. S k e l l y

Abstract: Photochemical oxidants Cprimarily ozone) a s

a i r p o l l u t a n t s pose a more s e r i o u s problem t o f o r e s t s of
t h e United S t a t e s than any o t h e r s i n g l e a i r p o l l u t a n t .
Temperate and Mediterraean f o r e s t s elsewhere have most l i k e l y
been s i m i l a r l y impacted and c u r r e n t i n v e s t i g a t i o n s of such
e f f e c t s a r e being pursued. Ozone Land i t s photochemically
r e a c t i v e precursors) has been demonstrated t o occur a t con-
s i d e r a b l e d i s t a n c e s downwind of major urban sources. Ozone
has induced p e r t u b a t i o n s t o vegetation over l a r g e a r e a s and
has t h e r e f o r e impacted innumerable and d i v e r s e f o r e s t eco-
systems. Direct i n j u r y due t o ozone has been documented t o
occur on numerous i n d i v i d u a l f o r e s t vegetation s p e c i e s but
d i r e c t a l t e r a t i o n s of f o r e s t ecosystems a s r e l a t e d t o ozone
induced e f f e c t s have only been e x t e n s i v e l y documented i n t h e
San Bernardino Mountains of C a l i f o r n i a ; t o a l e s s e r degree
s i m i l a r s t u d i e s have been done i n t h e Blue Ridge Mountains
of V i r g i n i a . Due t o t h e c u r r e n t s u l f u r dioxide (SO?)
problems confronting European f o r e s t s and due t o t h e planned
increased u t i l i z a t i o n of f o s s i l f u e l s i n much of t h e North
American Continent, t h e influence of ozone i n combination
with SO2 must be f u l l y considered. The influence of o t h e r
photochemical oxidants such a s nitrogen oxides and peroxy-
a c e t y l n i t r a t e on f o r e s t vegetation has remained r e l a t i v e l y
unknown.

The d i f f i c u l t y of understanding t h e r e a l and f o r e s t ecosystems must be s e v e r a l o r d e r s of magni-

p o t e n t i a l impact of a i r p o l l u t a n t s such a s ozone
(03) on i n d i v i d u a l s p e c i e s within any given p l a n t
community a s compared t o r e l a t i n g d i r e c t o r
i n d i r e c t e f f e c t s of OT, on e n t i r e and complex
p r e s e n t e d a t t h e Symposium on E f f e c t s of A i r
P o l l u t a n t s on Mediterranean and Temperate Forest
Ecosystems, June 22-27, 1980, Riverside,
C a l i f o r n i a , U.S.A.
Professor, Plant Pathology and D i r e c t o r , Labo-
r a t o r y f o r A i r P o l l u t i o n Impact t o Agriculture and
Forestry, V i r g i n i a Polytechnic I n s t i t u t e and
S t a t e University, Blacksburg, VA 24060.
tude i n d i f f e r e n c e . Several s t u d i e s have r e l a t e d
dose e f f e c t s t o f u l l - s i b c r o s s e s of various f o r e s t
t r e e s p e c i e s but such s t u d i e s have l i s t e d a
s e r i e s of caveats f o r t h e r e a d e r t o consider
while i n t e r p r e t i n g even t h e most b a s i c r e s u l t s
obtained. Such caveats have taken i n t o account
p o s s i b l e unknown e f f e c t s due t o exposure chamber
systems, physical and b i o t i c f a c t o r i n t e r a c t i o n s ,
and/or r e l a t e d monitoring and d a t a a n a l y s i s
methodologies. A s r e s e a r c h advances from s t u d i e s
of f u l l - s i b crossed p l a n t m a t e r i a l s through h a l f -
s i b (open p o l l i n a t e d , maternal l i n e s ) through
s p e c i e s and f o r e s t types, t o f o r e s t communities
and complex f o r e s t ecosystems, t h e number of
influencing v a r i a b l e s t o be considered while
i n t e r p r e t i n g t h e r e s u l t s becomes d i s p r o p o r t i o n a l
and d i f f i c u l t t o comprehend.
 Most s t u d i e s t h a t have d e a l t d i r e c t l y with 03
e f f e c t s t o any given p l a n t s p e c i e s have not
considered t h e e f f e c t s of p o l l u t a n t combinations
even a t h i g h e r doses. Under n a t u r a l conditions,
f o r e s t ecosystems a r e impacted by a m u l t i p l i c i t y
of atmospheric d e p o s i t i o n s including anthropogenic
p o l l u t a n t s i n gaseous, d r y , and wet formulations.
In a d d i t i o n such d e p o s i t i o n s do n o t occur s i n g l y
a t doses s i m i l a r t o those reported f o r various
exposure chamber s t u d i e s , e . g . , 1-8 hours p e r day
Background Concentrations
Ozone is a n a t u r a l l y occurring component of
t h e E a r t h ' s atmosphere and concentrations of
0.03-0.05 ppm 03 a r e g e n e r a l l y considered t o be
normal due t o mixing v i a t h e s t r a t o s u h e r i c t r a n s -
p o r t e f f e c t (Corn and o t h e r s 1975) Cfig.1 ) .

/
f o r 5 days/week f o r 10 weeks, but r a t h e r t h e y
occur a t low c o n c e n t r a t i o n s , i n various and
Net decay, but effects
c o n s t a n t l y changing combinations and over from long term
ur, photochemistryand
extended exposure periods. Thus, ecosystem photochemistry regional input
Stratospheric
s t u d i e s t h a t have n o t considered even t h e s e few ,-transport effect

f a c t o r s must be considered of l i m i t e d value f o r

any i n t r i n s i c o r e x t r i n s i c i n t e r p r e t a t i o n .
Natural
photochemicalinput

\
Loss to ground
I t i s a well known concept t h a t a i r p o l l u t a n t and aerosols

i n c i t e d p e r t u b a t i o n s lead t o s i m p l i f i c a t i o n of -Rural (upwind)- -Urban- -Run1 (downwtnd)-

f o r e s t ecosystems. Continued exposures t o SO2
complexes have r e s u l t e d i n s i m p l i f i c a t i o n and Figure I--The tropospheric ozone c y c l e . (from
r e v e r s a l o f once f o r e s t e d land t o grassland Corn and o t h e r s 1975).
communities o r i n some severe c a s e s even t o
s t e r i l i t y of t h e s o i l and accompanying erosion. The emission of oxidant forming p r e c u r s o r s such
However, such severe impacts due t o 0 p o l l u t i o n a s oxides of n i t r o g e n and hydrocarbons i n t o t h e
have n o t been r e p o r t e d t o occur w i t h t h e p o s s i - lower atmosphere l e a d s t o t h e buildup of photo-
b l e exception of major f o r e s t v e g e t a t i o n s p e c i e s chemically produced ozone and r e l a t e d peroxyacyl-
d e c l i n e i n t h e San Bernardino Mountains of n i t r a t e s [fig. 2) (NAS 1977).
Southern C a l i f o r n i a . The complexity of such
s i m p l i f i c a t i o n s c e n a r i o s involving minor changes
i n r e c e p t o r p l a n t physiology and r e l a t e d long-
term e f f e c t s on competitive a b i l i t i e s such a s
reproduction, n u t r i e n t c y c l i n g , and r e l a t e d food
chain e v e n t s h a s n o t been well documented.

The main purpose of t h i s paper i s n o t t o p r e -

s e n t a review of l i t e r a t u r e p e r t a i n i n g t o ozone
impact t o f o r e s t s p e c i e s and r e l a t e d ecosystems.
Several e x c e l l e n t reviews of t h e s u b j e c t have
been p r e v i o u s l y published and due t o t h e s e compre-
hensive up-to-date a n a l y s i s of t h e t o p i c any
f u r t h e r review would r e s u l t i n a redudancy i n
t h e l i t e r a t u r e (Miller 1973; M i l l e r and McBride
1975; Brown and o t h e r s 1979 and Kozlowski 1980).
Rather t h i s symposium paper w i l l attempt t o r e l a t e
t h e s e review a r t i c l e s and accompanying new l i t e r -
a t u r e i n t o an a n a l y s i s of t h e r e a l and p o t e n t i a l Figure 2--The normal n i t r o g e n oxide p h y t o l y t i c
long-term e f f e c t s of ozone on f o r e s t ecosystems. cycle [A) and t h e c y c l e a s a l t e r e d by t h e
A d d i t i o n a l l y , an e f f o r t w i l l be made t o r e l a t e a d d i t i o n of hydrocarbons leading t o increased
e x i s t i n g l i t e r a t u r e i n t o suggestions f o r f u t u r e ozone concentrations (B) (from NAS 1977).
c o n s i d e r a t i o n of ecosystem impacts i n t o t h e
National Ambient A i r Q u a l i t y Standards. PAN has n o t been d e t e c t e d i n non-anthropogeni-
c a l l y p o l l u t e d atmospheres but oxides of n i t r o -
gen do occur n a t u r a l l y i n t h e atmosphere and
BACKGROUND AND ANTHROPOGENIC CONCENTRATIONS concentrations of 0.02-0.10 ppm NOx have been
OF OXIDANTS IN FOREST SITUATIONS reported QNAS 1977). Brennan (Personal
communication) has reported a high PAN concentra-
A comprehensive review of ozone and o t h e r t i o n i n t h e Eastern United S t a t e s of 10 ppb but
ph'o'tochemical oxidants has been published by t h e suggested o v e r a l l concentrations of PAN a r e
National Academy of Sciences (NAS 1977); a well below West Coast observations.
r e c e n t compilation of papers has a l s o been
published (USEPA 1977b). These c o l l e c t i v e s e t s Anthropogenic Concentrations i n
of papers have provided an e x c e l l e n t review of Forested Areas
t h e o r i g i n , chemistry, t r a n s p o r t , and atmospheric
modeling of t h e s e p o l l u t a n t s . The long d i s t a n c e t r a n s p o r t of oxidant pre-
c u r s o r s and ozone i n t o remote f o r e s t e d a r e a s has
 been well documented (Miller and o t h e r s 1972;
Husar and o t h e r s 1977; Hayes and S k e l l y 1977;
and Cleveland and Kleiner 1975). Concentrations
of ozone above t h e c u r r e n t National Ambient A i r CHAROLETTSVILLE I
Q u a l i t y Standard (NAAQS) of 0.12 ppm 03 one hour
average p e r 24-hour p e r i o d ; twice (2 days) p e r
year have r e c e n t l y been r e p o r t e d within t h e
f o r e s t e d a r e a s of Eastern (Skelly and o t h e r s 1979)
and mid-Western United S t a t e s (Wolff and o t h e r s
1977). Numerous r e p o r t s of high ozone concen-
t r a t i o n s have been i s s u e d from t h e extensive San
Bernardino Mountain S t u d i e s (SBM) i n southern
C a l i f o r n i a (USEPA 1977a) .
Major episodes of
ozone have developed s p o r a d i c a l l y over t h e summer
months of May through October i n e a s t e r n f o r e s t s
whereas a b e t t e r defined and s t a b l e oxidant season
e x i s t s i n t h e f o r e s t s of southern C a l i f o r n i a .
Galloway and S k e l l y (1978) defined a major a i r
p o l l u t i o n epidose i n J u l y 1977 t h a t involved
high ozone c o n c e n t r a t i o n s and t h e highest ever
recorded l a r g e and f i n e a e r o s o l SO4 c o n c e n t r a t i o n s Figure 4--Total s u l f a t e (mg/m3) a s monitored a t
( f i g . 3, 4, and 5 ) . C h a r l o t t e s v i l l e , VA during a i r p o l l u t i o n episode
of J u l y , 1977. Note buildup of J u l y 15-20 and
By comparing monitoring d a t a from t h e SBM s i m i l a r sharp drop a s i n f i g u r e 3 on J u l y . (From
s t u d i e s with t h o s e of t h e Blue Ridge Mountain Galloway and S k e l l y 1978).
S t u d i e s (BRM) a 3-4 f o l d g r e a t e r c o n c e n t r a t i o n
of ozone i s apparent i n t h e former over t h e
l a t t e r . Peaks of 0.20 t o 0.40 ppm O3 d a i l y one-
hour maximums have occurred i n t h e SBM f o r e s t
( f i g . 6) but t h e peak one-hour average ever
recorded i n t h e BRM a r e a has been 0.166 ppm O r .
Table 1 p r e s e n t s t h e monthly and peak one-hour
c o n c e n t r a t i o n s of ozone a s monitored a t v a r i o u s
s i t e s i n t h e Blue Ridge Mountains of V i r g i n i a .
PAN and NO have n o t been monitored i n e a s t e r n
f o r e s t s oi^the United S t a t e s and although d e t e c t e d
i n western f o r e s t s l i t t l e has been done t o d i s -
tinguish differences i n t h e i r respective effects
over t h o s e induced by ozone a l o n e ( f i g . 6 ) .

F OXIDANT CONCENTRATIONS JULY, 1977

-SALT POND MTN.

-- APPLE ORCHARD MTN.
ROCKY KNOB
........PINNACLES

I 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 192021 22232425262728293031
DATE
Figure 3--Oxidant c o n c e n t r a t i o n s a s monitored
a t s e v e r a l l o c a t i o n s i n t h e Blue Ridge and
Southern Appalachian Mountains of V i r g i n i a
d u r i n g J u l y , 1977. Daily averages (24 hours)
a r e i n d i c a t e d . Note peak period of J u l y 15-
20.
Figure 5(A)--A review of t h e Peaks of O t t e r a s
taken on a c l e a r day i n t h e Blue Ridge
Mountains. Photograph taken from Pine Tree
Overlook a t a d i s t a n c e of 11.6 KM from t h e
Peaks. (B) Photo taken on J u l y 20, 1977
during t h e worst a i r p o l l u t i o n s t a g n a t i o n
experienced i n V i r g i n i a . Same view a s
(A) .
 Norway; an a r e a s i t u a t e d c l o s e t o t h e c o a s t and
considered t o be normally unaffected by l o c a l
oxidant producing sources. The h i g h e s t 03 concen-
t r a t i o n observed a t Rorvik was 0.20 ppm (August
1975) and i n Gothenburg t h e h i g h e s t 03 concentra-
t i o n was 0.13 pprn ( t a b l e 2 ) . They suggested
clockwise a i r movement a s a s s o c i a t e d with high
p r e s s u r e systems and long d i s t a n c e t r a n s p o r t from
i 4

-
U
OZONEIpphml UV, OASIBI
TELEMETRY,ONCE HOURLY
Europe t o be r e l a t e d t o t h e ozone episodes.
Skarby (1979) reported t h a t ozone v a l u e s recorded
i n t h e summertime have been too high t o be con-
s i d e r e d a s normal background concentrations f o r
Swedish c o n d i t i o n s . During t h e summer of 1977
1\
l a PAN ( p p b ) 5.C. ELECTRON
CAPTURE
high O3 concentrations (0.20 ppm) were recorded
on 21 out of 92 days. She a l s o suggested long-
range t r a n s p o r t t o be involved.

DAYS From t h e s e few r e p o r t s it i s q u i t e obvious

Figure 6--Comparative d a i l y maximum hourly
averages f o r ozone, t o t a l oxidant, PAN, and NO2
a t Sky Forest August, 1974. (From USEPA 1977).
Ozone concentrations i n European f o r e s t s have
not been e x t e n s i v e l y i n v e s t i g a t e d but i n i t i a l
s t u d i e s i n d i c a t e t h a t ozone appears t o be episodal
i n i t s occurrence i n a manner s i m i l a r t o t h a t
experienced i n Eastern United S t a t e s . Grennfelt
(1979) r e p o r t e d on ozone concentrations a s moni-
t o r e d a t Rorvik about 40 KM south of Gothenburg,
t h a t ozone concentrations i n excess of t h e NAAQS
occur f r e q u e n t l y i n t h e temperate and Mediter-
ranean f o r e s t s of t h e world. Therefore, it i s
a l s o obvious t h a t t h e i n j u r y t h r e s h o l d s f o r
numerous p l a n t s p e c i e s have a l s o been f r e q u e n t l y
exceeded and t h e e f f e c t s of t h e s e exposure doses
a r e discussed below.
DIRECT EFFECTS TO FOREST SPECIES
The responses of any given f o r e s t ecosystem t o

Table I--Ozone concentrations (pprn) monitored a t Rocky Knob, Floyd Co., Va. (Blue Ridge Parkway)
and a t Pinnacles and Big Meadows, Madison Co., Va. (Shenandoah National Park).

Month

Rocky Knob Pinnacles Rocky Knob Pinnacles ' Rocky Knob Big Meadows
1

4onth Peak Month Peak Month Peak Month Peak Month Peak Month Peak
her. Hour Aver. Hour Aver. Hour Aver. Hour Aver. Hour Aver. Hour
Aver. Aver. Aver. Aver. Aver. Aver.

January
February
March
Apr i1
May
June
July
August
September
October
November
December

Average
f o r moni-
t o r e d mo.

% i n n a c l e s s i t e moved 14 KM t o Big Meadow CSNP) May 1979.

o n l y 8 days out of t h e month were used f o r t h i s d a t a .

 Table 2--The number of days with high ozone concentrations in Goth-

enburg and Rsrvik, Norway 1972-1978 expressed as days with one hour

mean of ozone exceeding indicated values (Grennfelt 1979).

Year -
>0.08 >0.10
- >0.12
- >0.15
- Maxhourly
PPm PPm PPm PPm mean
(PP~)

Gothenburg

18 11 5 0.20
17 5 0 0.13
6 0 0 0.11
12 8 1 0.15
I

l~ncludesdata only until June 30, 1978.

photochemical oxidants must be as a result of the Extensive research has been justifiably done

expression of direct effects to the individual on specific forest species to better define the

component species within that ecosystem. The injury phase of this scenario of increasingly

ability of scientific research to define those inclusive terminology. However, even within this

direct effects to individuals and to relate such type of problem definition research a larger

defined effects to the whole has been limited. endeavor has been made to define visible plant

Three terms have been used somewhat interchangea- responses over the less easily measured physiolo-

bly to define these effects and for purposes of gical responses. The latter responses may actual-

this paper they shall be defined as: ly be of more importance to understanding the

damage and impact phases of subsequent ecosystem

Injury - the result of one or more deleterious deteriorations. Thus, three additional terms
alterations of normal physiological processes as emerge :
manifested by the presence of chronic or acute

visible symptoms and/or growth reductions (growth acute injury - involves expresion of clinical
reduction may be the only manifestation),. symptoms leading to death of cells, tissues,

organs, or entire plants and/or plant communities.

Damage - injury that results in measurable Such injury is usually initiated by exposure to
economic loss to specific crops e.g. reduced high doses of pollutants (^concentration x <time)_

height or radial increment growth of forest trees but may result from exposure to lower concentra-

resulting in reduced value of the commerical tions of pollutant over extended periods of time.

forest.

chronic injury - involves non-lethal types of

Impact - the total influence Qdetrimental or clinical symptom expressions such as reduced
beneficial) of air pollutants on all aspects of chlorophyll production and related pigmentation

the forest ecosystem including even minor shifts changes, and reduced growth rates. Such injury

towards reduced diversity of species, indirect usually results from still lower dose exposures.

effects to watersheds and water quality, or

direct effects to recreational values due to functional injury - involves injury to the
reduced visibility at vistas or overlooks in functional efficiency of the plant as expressed

National Forests and Parks. by reduced growth or other expression of loss

without the development of any other clinical

indigenous to a mixed hardwood-conifer forest

*
symptoms, i.e. injury is only of a physiological
ecosystem of the Northeastern United States is
and pre-clinical nature. Further visible
taken into account our knowledge is very limited

symptoms do not develop. This form of injury is

even for the clinical expression of symptoms by

most difficult to define and measure and is the

most species. Interpretation of available

result of lowest dose exposures at predominantly

knowledge in the forest community and forest eco-

near background concentrations of pollutants over

system columns may be somewhat harsh but most

extended periods of time.

probably realistic. A similar table as con-

structed for sulfur dioxide related investigations

An attempt to define the current status of

would be much more optimistic thus attesting to

knowledge concerning these various forms of di-

the relative ease of working on predominant point

rect effects due to ozone has been presented in

sources of pollutants and their related effects.

table 3. A more specific assessment of recent

However, our knowledge of SO2 induced pertubations

documented studies of chronic pollution effects

to forest ecosystems located at distances from

as induced by all species of air pollutants at the

various sources is relatively poor, e.g. knowledge

ecosystem level has been presented by Kickert and

of the subtle influences of S-related compounds

Miller (1978).
to the productivity of Northeastern United States

forests is virtually non-existent.

It is evident from table 3 that our knowledge

of ozone effects following high dose exposures

using artificial exposure systems for species

Direct Effects to Forest Trees

level investigations is considered somewhat

superior to that of the other levels of activity.

Davis and Wilhour (1976) have provided the

However, when the abundance of plant species

most complete listing of woody plant sensitivities

to sulfur dioxide and photochemical oxidants as

Table 3--The current status of knowledge concerning ozone induced effects to Temperate

and Mediterranean forest tree species, forest communities, and forest ecosystems. Com-

parisons of the San Bernardino Mountain Studies (SBM) versus all other investigations

(01) have been noted.

Effect Study
Forest species2 Forest Communities
Forest Ecosystems

I
Injury

01
moderate
poor
poor

overall
SBM
moderate
moderate
moderate

01
abundant
moderate
poor

acute
SBM
abundant
moderate
moderate

01
moderate
poor
non-existent

chronic

SBM
moderate
moderate
poor

01
poor
non-existent
non-existent

functional
SBM
moderate
moderate
poor

01
moderate
poor
non-existent

Damage

SBM
abundant
moderate
poor

Impact
non-existent
non-existent

moderate
poor

In consultation by author with P. R. Miller Cpersonal communication).

2~stimatesin this column include responses obtained in fumigation chamber studies

If such information was to be detected poor to non-existent descriptions would be

appropriate for each category.

^ B definition
~
not applicable.

derived from t h e i r review of United S t a t e s ,
Canadian, and European l i t e r a t u r e .
Eastern White Pine
The predominant f o r e s t t r e e s p e c i e s s t u d i e d
i n Eastern United S t a t e s has been e a s t e r n white
Pinus s t r o b u s L.; e x t e n s i v e l i t e r a t u r e reviews
by Gerhold (1977) and Nicholson (1977) a r e
a v a i l a b l e . I n t e r e s t i n t h i s s p e c i e s has
remained h i g h s i n c e t h e f i r s t discovery of i t s
somewhat unique s e n s i t i v i t y t o ozone by Berry
(1961). The response of t h i s s p e c i e s t o ozone
c o n c e n t r a t i o n s a s monitored i n t h e Blue Ridge
Mountains of V i r g i n i a ( t a b l e 1) has been t h e
s u b j e c t of s e v e r a l c u r r e n t i n v e s t i g a t i o n s . S k e l l y
and o t h e r s (1979) r e p o r t e d t h a t of 315 white p i n e s
surveyed by u s i n g a modified e v a l u a t i o n scheme a s
adapted from M i l l e r (1973) 17, 80, and 3 percent
were considered t o be t o l e r a n t , i n t e r m e d i a t e and
s e n s i t i v e , r e s p e c t i v e l y , t o ozone. Of t h e 315
t r e e s tagged i n 1977, 10 were reported t o have
d i e d following r e p e a t e d t y p i c a l c l i n i c a l symptoms
of oxidant induced i n j u r y . Subsequent r o o t exca-
v a t i o n s and i s o l a t i o n of fungi has yielded
V e r t i c i c l a d i e l l a p r o c e r a from t h e dying t r e e s .
The growth r a t e of t r e e s i n each c l a s s was a l s o
examined by Benoit (1980) and r a d i a l increment
growth over t h e period 1955-1978 f o r t h e s e n s i -
t i v e c l a s s was s i g n i f i c a n t l y l e s s (p = . 0 l ) than
t h a t of t h e t o l e r a n t c l a s s ( f i g u r e 7 ) . A general
d e c l i n e i n growth f o r a l l c l a s s e s was noted.
9
1 -- - =TOLERANT;53 YRS
-=
AVG AGE
INTERMEDIATE.52 YRS AVG AGE
\---
YEAR 1955 '60 '70 1978
AVERAGE RADIAL INCREMENT GROWTH OF SENSITIVITY CLASSES
Figure 7--The average r a d i a l increment growth
of e a s t e r n white p i n e i n t h r e e ozone s e n s i t i v i t y
c l a s s e s a s found i n t h e Blue Ridge Mountains of
V i r g i n i a . Trees were l o c a t e d i n groups of 3
p e r s i t e with each s e n s i t i v i t y c l a s s represented
i n each of 10 p l o t s (10 t r e e s / c l a s s ) .
Other Eastern Species
F o r e s t t r e e s - - t h e r e l a t i v e 03 s e n s i t i v i t y of
s p e c i e s has been i n v e s t i g a t e d u s i n g high dose
exposures, e . g . 0.25 ppm O3 f o r 8 hour exposure
p e r i o d s . Using such doses Davis and Wood (1972)
exposed 1.8 c o n i f e r s p e c i e s and found Austrian
Pine (Pinus -- n i g r a , Arnold), jack pine (c. E-
s i a n a , Lamb.) and V i r g i n i a pine (P. - virginiana,
M i l l . ) t o be t h e most s e n s i t i v e . However, they
reported v a r i a b l e symptom response among t h e
d i f f e r e n t s p e c i e s , among p l a n t s within s p e c i e s ,
and between branches and needles of i n d i v i d u a l
p l a n t s . In a s e r i e s of exposures, numerous
deciduous t r e e s were a l s o exposed t o s i m i l a r
ozone doses and green ash ( ~ r a x i n u spennsylvanica
Marsh.), white ash (F. americana L.) and t u l i p
poplar ( ~ i r i o d e n d r o n t u l i ~ i f e rLa. ) were r e p o r t e d
t o e x h i b i t f o l i a r i n j u r y by Wood (1970). Jensen
(1973) determined t h e s e n s i t i v i t y of 9 decidu-
ous t r e e s p e c i e s on t h e b a s i s o f height
growth during a 5 month exposure t o 0.30 ppm
03 f o r 8 hours p e r day. He determined t h a t
s i l v e r maple &A( saccharinum, L.) , green ash,
and sycamore (Platanus o c c i d e n t a l i s , L . ) were t h e
only s p e c i e s determined t o be s e n s i t i v e using
both parameters. Numerous s i m i l a r s t u d i e s u s i n g
high dose exposures have been reviewed by S k e l l y
and Johnston (1979).
Typical oxidant symptoms have been noted by
S k e l l y (unpublished) on s e v e r a l major f o r e s t
t r e e s p e c i e s indigenous t o Shenandoah National
Park of V i r g i n i a [ f i g . 8) .
Figure 8--Typical oxidant induced s t i p p l i n g
on hickory G a r y a spp.) a s observed i n t h e
Shenandoah National Park of V i r g i n i a . Note
asymptomatic a r e a of covered over p o r t i o n of
lower l e a f i n c e n t e r of photograph (upper
l e a f p u l l e d back).
Very few low dose exposure s t u d i e s have been
conducted t o determine e f f e c t s due t o c l o s e r t o
ambient p o l l u t a n t c o n c e n t r a t i o n o r due t o ambient
exposure c o n d i t i o n s . However, s e v e r a l r e c e n t
s t u d i e s have attempted t o reproduce ambient
concentrations of 03, SO2, and NOx (and v a r i o u s
combinations t h e r e o f ) and t o study t h e i r e f f e c t s
on t h e growth of l o b l o l l y p i n e (P. - -t a e d a L.)
Kress and S k e l l y (1980a) American sycamore
Kress and S k e l l y (1980b), and s e v e r a l o t h e r
e a s t e r n t r e e s p e c i e s (Kress 1980). I n t h e com-
bined p o l l u t a n t s t u d i e s using 0.05 ppm 03, 0.10
ppm N02, and 0.14 ppm SO2 f o r 6 hours p e r day f o r
28 consecutive days s i g n i f i c a n t height r e d u c t i o n s
were r e p o r t e d a s induced by O x alone t r e a t m e n t s s a r y t o c l i p (to a 1 . 3 cm h e i g h t ) and remove com-
f o r each s p e c i e s without c l i n i c a l symptoms p r e - p e t i n g n a t u r a l v e g e t a t i o n which was t h e n c o l -
s e n t on sycamore and with <5 percent f o l i a r l e c t e d f o r d r y weight measurements ( t a b l e 4 ) .
i n j u r y on l o b l o l l y pine. Kress (1980) r e p o r t e d
h e i g h t growth i n c r e a s e s and/or decreases f o r
10 t r e e s p e c i e s following exposure t o 0.05,
0.10, and 0.15 ppm 03 f o r 6 hours/day f o r 28
consecutive days. Lowest dose exposure s i g n i f i -
c a n t l y reduced t h e h e i g h t of l o b l o l l y p i n e and
0.10 pprn O3 reduced h e i g h t growth i n l o b l o l l y p i n e ,
green a s h , sycamore, p i t c h p i n e (P. r i g i d a M i l l . )
and sweetgum (Liquidambar s t y r a c i f l u a L . ) .
S l i g h t h e i g h t growth s t i m u l a t i o n s were r e p o r t e d
f o r s e v e r a l s p e c i e s following 0.05 ppm 03 t r e a t -
ment and sugar maple (A. saccharum Marsh.)
responded p o s i t i v e l y even a t t h e 0.10 ppm 03
t r e a t m e n t (p = 0.05) .

Understory VegetationÑHarwar and Treshow

(1971) have conducted one of few s t u d i e s designed
t o determine ozone e f f e c t s t o u n d e r s t o r y p l a n t s .
They exposed 17 r e p r e s e n t a t i v e s p e c i e s of an
aspen community t o v a r i o u s high and low doses
of 0 5 days p e r week throughout t h e growing
3
season f o r 3 consecutive y e a r s . Several s p e c i e s
were found more s e n s i t i v e t h a n expected and
s e n s i t i v i t y was s o v a r i e d between s p e c i e s t h a t
t h e a u t h o r s suggested major s h i f t s i n community
composition would be probable following only a year
o r two of exposure.

Kohut and Krupa (1978) determined t h e s e n s i -

t i v i t y of s e v e r a l herbaceous p l a n t s of t h e North-
c e n t r a l U.S. f o r e s t s . They l i s t e d a group of
p l a n t s found i n t h e f o r e s t s o f t h e North C e n t r a l
r e g i o n t h a t were a l s o s e n s i t i v e t o 0.08 and 0.15
ppm 0, f o r o n l y 4 hours. These p l a n t s a r e a l s o
n a t i v e t o t h e f o r e s t e d a r e a s of t h e n o r t h and
s o u t h e a s t e r n p o r t i o n s of t h e United S t a t e s .
The s e n s i t i v e p l a n t s l i s t e d were wild buck-
wheat, c h i c o r y , d a i s y , mustard, and Ribes.
Other work by S k e l l y (1977, unpublished) has
i d e n t i f i e d symptoms t y p i c a l l y induced by 0 on
Clematis s p . i n t h e Shenandoah National Park
of V i r g i n i a .

Figure 9 has been p r e s e n t e d t o f u r t h e r

demonstrate t h e e f f e c t of ambient c o n c e n t r a t i o n s
of ozone on a s p e c i e s t h a t i s widely d i s t r i -
buted a c r o s s North America i . e . common milk-
weed (Asclepias spp.) . Duchelle and o t h e r s
(1980) observed s e v e r e , moderate, and
only s l i g h t i n j u r y t o t h i s s p e c i e s t h a t occurred
n a t u r a l l y i n open p l o t s and i n n o n - f i l t e r e d and
f i l t e r e d open t o p chambers, r e s p e c t i v e l y , a s
l o c a t e d i n t h e Shenandoah National Park i n
V i r g i n i a . This s p e c i e s i s being t e s t e d f u r t h e r
f o r i n j u r y t h r e s h o l d s and f o r u s e a s p a r t of a
p l a n t b i o i n d i c a t o r system.

A s a n o t h e r p a r t of t h e Blue Ridge Mountain F i g u r e 9--Oxidant i n j u r y t o milkweed

S t u d i e s , Duchelle and S k e l l y (1980) e s t a b l i s h e d
4 r e p l i c a t i o n s of open t o p chambers r e c e i v i n g
c h a r c o a l f i l t e r e d a i r o r non-charcoal f i l t e r e d
a i r i n o r d e r t o i n v e s t i g a t e h e i g h t growth of
s e l e c t e d f o r e s t t r e e s . Four open p l o t s were
a l s o e s t a b l i s h e d . By mid-summer it became neces-
~ ~ & l e ~ spp.)i a s observed i n t h e Shenan-
doah National Park, VA a s grown i n (A) open
p l o t s , (B) non-f i l t e r e d open t o p chamber and
CC) c h a r c o a l - f i l t e r e d open t o p chamber.
Purple s t i p p l e was only noted on upper
leaf surfaces.
Table 4--The dry weight of foliage of composited
fixation and stomatal conductance usually associ-

clippings as collected from 4 filtered air and 4

ated with needle aging. Needles of sensitive

non-filtered air open top chambers and 4 open

trees senesced and abscissed prematurely thus

plots established in the Big Meadows Area of the

contributing to a steady decline in tree vigor

Shenandoah National Park, VA.

and increased vulnerability to other sources of

stress.

Exposure Clipping Dates1

Other Forest Tree Species

Aug. 14, 1979 Oct. 9, 1979

Dry Weight (Grams)

Miller (1973) ranked the following species for

their decreasing sensitivity to ozone following

fumigation tests:
Total Total
wt2 Average wt Average Most sensitive Western white pine (P. monti-
cola Doug.)
Filtered 7263 1816 1599 400 Jeffrey x Coulter pine hybrid
Non-filtered 4937 1234 1323 331
Open 3128 782 845 211

Monterey x Knobcone pine

~ l species
l
composited as clipped to 1.3 cm
hybrid (P. radiata x -
P.

height, 10 foot diameter plots.

altenuata)

z Ponderosa pine [c. ponderosa

4 replications.
Laws.)
Intermediate Coulter pine (P.
- coulteri D.

Western Species
Don")

Douglas fir (Pseudotsuga

Ponderosa Pine
menziesii (Mirb.) Franco
Jeffrey pine (P. jeffreyi

Since the decline of ponderosa pine (P.

Grev. & ~alf.)-
ponderosa Laws .) during the 1950's (and hence- White fir (Abies concolor

forth into the 1980's) in the South Coast air

(Gord. 6 Glend.1 Lindl.
basin and San Bernardino Mountains of California,
~i~ cone Douglas fir (Pseudo-

this species has become the most intensively

tsuga macrocarpa (Vasey) Mayr.

investigated of all western forest vegetation

Knobcone pine (P.
- attenuata

species. Declining ponderosa pines have most

.
Lemm )
recently been reported in the southern Sierra-
Tolerant Incense cedar (Libocedrus

Nevada mountains by Miller and Millecan (1971)

decurrens Torr.)

and in the Sequoia National Forest and Sequoia-

Sugar pine (c.lambertiana

Kings Canyon National Parks by Williams and

Doug1 ) .
others (1977).
Giant sequoia (Sequoia

gigantea (Lindl. ) Decne.

The decline of this species was initially

termed "X-diseasev by Parameter and others (1962);

In one of few low O3 dose exposure studies,

further study by Miller and others (1963) eluci-

9 western conifer species have been evaluated

dated ozone to be the direct incitant. Inter-

for foliar injury and growth responses by

relationships of foliar symptoms with increased

Wilhour and Neely 0.977). They found signifi-

root disease and increased incidences of bark

cant growth reduction in juvenile seedlings of

beetle infestations in injured trees have been

P. ponderosa and P. monticola exposed to 0.10

determined along with significant growth decreases

ppm 03 for 6 hours/day consecutively for up to

and mortality (Stark and others, 1968; McBride and

22 weeks. They noted no constant association

others, 1975). Several reviews of the major

between growth response and foliar injury.

studies that have dealt with ponderosa pine have

recently been published by Brown and others

(19791, Miller (1973) and Miller and McBride

DIRECT EFFECTS TO FOREST ECOSYSTEMS

(1975).

A fair number of published reports assessing

Most recent investigations by Coyne and Bingham

air pollution induced injury to agricultural crops

(in press) identified characteristics of ecotypic

are available. Damage estimates of yield losses

variation in E. ponderosa which varied in their have also been published e.g. air pollution

clinical symptom response to ozone under field

injury to potatoes in the Atlantic Coastal States

conditions. Light responses, photosynthetic

[Heggestad, 1973). Such evaluations have been

rates, and stomatal conductances were observed

based upon extensive knowledge of crop manage-

and differences among injury classes were mani-

ment practices and abundant information exists

fest as acceleration of the normal decline of C02

concerning expected yields. Therefore, yield

losses such as potato tuber number, size, and

weight may be easily determined and subse-

quently correlated with the degree of foliar

injury as induced by photochemical oxidants

(Heggestad, 1973) .
Advancing from a relatively simplistic agro-

nomic monocultural system of crop management to

CHARACTER-

a mixed hardwood conifer forest ecosystem poses

ISTICS

considerably different problems in determining

an effect due to any given single stress factor.

As noted previously, an abundance of literature

COMMUNITY
COMMUNITY

describing symptoms has become available for STRUCTURE STRUCTURE

several major forest species that appear sensi-

tive to photochemical oxidants (table 3). How- 5 6 7 8 9

ever, the overall impact of the more subtle

changes in functional efficiency of the plant

expressed, for example, as slightly reduced photo-

synthetic capability of otherwise asymptomatic

foliage or noted trends of slightly shorter pollen

tube length in the presence of low doses of 03

remain relatively little understood.

There have been no major investigations of

oxidant impacts to forest communities or forest Figure lo--Community-succession interactions in

ecosystems with the single well known exception a mixed-conifer forest ecosystem. (From Taylor,

of the San Bernardino Mountain Studies. The 1974).

intensity of the various studies and their

related interactions within one of many possible

areas of investigation has been illustrated in

figures 10 and 11. As noted in figure 10,

oxidant air pollutants are only one of many

natural or anthropogenic stress factors important

to the forests of the San Bernardino Mountains

0
POLLUTANTS

and if this figure were to be modified for illus-

tration of an Eastern U.S. or European forest, /&-$.A

SYNTHESIS
1.
SO2 and other forms of atmospheric depositions

would necessarily be added for emphasis. Figure

11 illustrates only a few of the complex inter-

actions which may take place within an ecosystem

and even a very subtle change in bark charac-

teristics or carbohydrate pools as initially

induced by oxidant injury of the foliage may be

sufficient to encourage a bark beetle attack of

0;

MOISTURE

BARK
weakened trees. As related in table 3, an
TEXTURE
evaluation of knowledge concerning injury,
.......... .. - -
THICKMESS
damage, and impact to forest species of the San .......- .... -
Bernardino Mountains (primarily E. ponderosa) is
PHLOEM THICKNESS
.... ...- .- - -
PHLOEM MOISTURE
abundant and knowledge is considered to be . - - - .....- - -
moderate for the forest community. However, due

to the complexity of any given forest ecosystem, RESIN QUANTITY

.... . - -- -- - -
including that which has been intensively studied

in the San Bernardino Mountains, relatively poor

information exists as to the subtle influences

to the functioning of any given forest ecosystem.

FUNGI

FUTURE RESEARCH EMPHASIS AND CONSIDERATIONS

Why do we know so little about oxidant induced Figure 11-411example of tree-level interactions

injury, damage, or impact to forest communities in a mixed-conifer forest ecosystem. Data from

or forest ecosystems? Why have only a few tree these types of studies would be integrated into

species been intensively studied and relatively overall effect illustrated in figure 11. (From

few others evaluated for foliar symptom response? Taylor, 1974) .

How does the lack of such information influence

decisions made regarding the establishment of Intensive management of agronomic crops through

National Ambient Air Quality Standards? selection of tolerant varieties for planting in

high oxidant areas and appropriate changes in

be statistically significant but may have

cultural practices such as withholding irrigation

greater long-term biological significance to

water during oxidant pollution episodes have all

the ability of a species to survive over

served to reduce the immediate injury (and there-

their natural range.

fore damage and impact) to such important crops

as potatoes, tobacco, soybeans, snapbeans and

The natural forested ecosystems of the temper-

certain horticultural plants. Through the develop-

ate and Mediterranean regions of the world may

ment of such practices, the agricultural scientist

serve to provide the most invaluable bioindicator

has assisted the immediate grower (as should be

of long-term photochemical oxidant air pollution

the case) but concurrently the economic cost/bene-

induced effects. National air quality standards

fit justifications for pollution abatement enforce-

must be reasonably developed to protect these

ment have been lessened since overall impact has

natural resources from even minor change but

potentially been greatly reduced. The evaluation

initially adequate financial support and associ-

of forest tree species for sensitivity to various

ated quality research must be continued to ade-

pollutants has also been attempted and likewise

quately define the real and potential effect of

long term growth losses may be averted. Such

oxidants to plant species, to plant communities,

investigations must be approached with a certain

and to entire forest ecosystems.

degree of caution since long term subtle changes

most undoubtedly are occurring in natural forest

ecosystems but as yet most remain undetected.

LITERATURE CITED

Natural ecosystems usually have a system of

checks and balances but the system remains deli-

Benoit, L. F.

cate and trends towards simplification are easily

1980. Ozone effects on long term radial incre-

initiated. Changes in primary productivity,

ment growth and reproduction of eastern

energy resource flow patterns, biogeochemical

white pine. (Abstr.) Proc. Potomac Div.

cycles, and species successional patterns may all

her. Phytopath. SOC., Morgantown, W. Va.

be challenged by oxidant air pollution but have

remained virtually non-studied. Berry, C. R.

1961. White pine emergence tipburn, a physio-

The difficulties encountered in developing

genic disturbance. (Sta. Paper No. 130.,

research aimed at isolating, identifying, and

8 p.). Southeast For. Exp. Sta., USDA-

subsequently integrating the known effects of oxi-

For. Serv., Athens, Ga.

dant air pollutants on the forest ecosystem are

too numerous to completely cover in the remaining

Brown, H. D., P. R. Miller, J. M. Skelly, D. B.

space available. A partial listing however, must

Drummond, and C. E. Carlson.

include these important points:

1979. Air pollution effects on forest vege-

tation and the analysis of the role of

1) Photochemical oxidant is insidious over

Forest Insect and Disease Management.

extremely large areas of diverse forest

134 p. USDA-For. Serv., Washington, D.C.

land and the establishment of control (non-

pollution exposed) areas has become virtually

Cleveland, W. S. and B. Kleiner.

impossible.
1975. Transport of photochemical air pollution

from The Camden-Philadelphia urban complex.

2) The introduction of charcoal filtration sys-

Environ. Sci. Tech. 9:869-872.

tems into such areas to establish kontrolsw

is at best artificial and due to physical

Corn, M., R. W. Dunlap, L. A. Goldmuntz and

restrictions the ability of such systems to

L. H. Rogers.

define a true ecosystem pertubation is like-

1975. Photochemical oxidants: sources, sinks

wise limited.
and strategies. J. Air Poll. Contr. Assoc.

25 :16-18.
3) Long-term investigations must take into

account innumerable variables some of

Coyne, P. I. and G. E. Bingham.

which are very transient in their occurrence

1980. Comparative ozone dose response of gas

and others for which relatively little is

exchange in a ponderosa pine stand exposed

known even under natural conditions.

to long term fumigations. (Submitted for

publication to J. Air Poll. Contr. Assoc.).

4) Modeling and related statistical procedures

must take into account the diversity and

Coyne, P. I. and G. E. Bingham.

complexity of a forest ecosystem and

1980. Variation in photosynthesis and

statistically probabilities of p = .01, .05, stomata1 conductance in an ozone-stressed

or .10 must not be the only acceptable

ponderosa pine stand: Light response.

limits of "biological significance." The

(Submitted for publication to For. Sci.).

prediction of a biological event with 70

percent accuracy may be valid. Observation

Davis, D. D. and R. G. Wilhour.

of only a 1-5 percent decrease in annual

1976. Susceptibility of woody plants to

radial increment growth or similar decreases

sulfur dioxide and photochemical oxidants.

in pollen production and viability may not

72 p. USEPA Ecol. Res. Series EPA-600/3-

76-102.

Davis, D. D. and F. A. Wood.

Kohut, R. J. and S. V. Krupa.

1972. The relative susceptibility of eighteen

1978. Sensitivity of selected species of

coniferous species to ozone. Phytopathology

native vegetation to ozone. (Abstr.).

62:14-19. Proc. Am. Phytopathological Soc. 4:88.

Duchelle, S. F., J. M. Skelly and L. W. Kress.

Kozlowski, T. T.

1980. The impact of photochemical oxidant

1980. Impacts of air pollution on forest eco-

air pollution on biomass development of

systems. Bio. Sci. 30:88-93.

native vegetation and symptom expression

of Asclepias spp. (Abstr.) . Proc. Kress, L. W.

Potomac Div. her. Phytopath. SOC.

1980. Effects of O3 and 0 + NOn on growth
Morgantown, W. Va.
of tree seedlings. Proc. Internatll.

Symp. on Eff. of Air Pollutants on

Galloway, J. N. and J. M. Skelly.

Mediterranean and Temperate Forest Eco-

1978. A pollution episode in Virginia.

systems. June 22-27. Riverside, Ca.

Virginia Climate Advisory 2(3):4-6.

Kress, L. W. and J. M. Skelly.

Gerhold, H. D.
1980. Growth imuact of 07. NO?, and/or SO2

1977. Effect of air pollution on Pinus

--
on Pinus taeda.
(~ubmi&ed for publi-
strobus L. and genetic resistance. 45 p.
cation to Atmos. Environ.).

USEPA Rep. No. EPA-600/3-77-002. Cor-

vallis, Or.
Kress. L. W. and J. M. Skellv.

1980. Growth impact of o;, NO,, and/or SO2

Grennfelt, P.

1979. Ozone monitoring in Scandanavia. In

o Atmos. Environ.).

Report from the workshop "Ozone effects

on vegetation in Europe." M. Eastmond and

McBride, J. R., V. P. Semino, and P. R. Miller.

L. Skarby Eds. p. 5-8. Swedish Water and

1975. Impact of air pollution on the growth

Air Res. Inst., Stockholm.

of ponderosa pine. Calif. Ag. 29 (12) :8-9.

Harward, M. R. and M. Treshow.

Miller, P. R.

1971. The impact of ozone on understory

1973. Oxidant-induced community change in a

plants in the aspen zone. Proc. 64th Ann.

mixed conifer forest. -In Air Pollution

Mtg. Air Poll. Contr. Assoc. Atlantic

Damage to Vegetation. J. A. Naegela Ed.

City, N.J.
p. 101-117. Adv. in Chem. Series 122.

Amer. Chem. Soc., Washington, D.C.

Hayes, E. M. and J. M. Skelly.

1977. Transport of ozone from the northeast

Miller, P. R. and J. McBride.

U.S. into Virginia and its effect on

1975. Effects of air pollutants on forests.

eastern white pine. Plant Dis. Reptr.

-
In Response of Plants to Air Pollution.

51 :778-782. B. Mudd and T. T. Kozlowski, eds. p. 195-

235. Academic Press, New York.

Heggestad, H. E.

1973. Photochemical air pollution injury to

Miller, P. R. M. H. McCutchan and H. P. Millegan.

potatoes in the Atlantic Coastal States.

1972. Oxidant air pollution in the central

Amer. Pot. J. 50:315-328.

valley, Sierra Nevada foothills, and Mineral

King Valley of California. Atmos. Environ.

Husar, R. B., D. E. Patterson, C. C. Paley, and

6:623-633.

N. V. Gillani.

1977. Ozone in hazy air masses. Jn- Inter-

Miller, P. R. and A. A. Millecan.

national Conference on Photochemical Oxi-

1971. Extent of oxidant air pollution damage

dant and its Control. p. 275-282. USEPA

to some pines and other conifers in Cali-

Ecol. Res. Series EPA-600/3-77/001b.

fornia. Plant Dis. Reptr. 55:555-559.

Research Triangle Park, N.C.

Miller, P. R., J. R. Panneter, Jr., O.C. Taylor,

Jensen, K. F.
and E. A. Cardiff.

1973. Response of nine forest tree species.

1963. Ozone injury to the foliage of Pinus

to chronic ozone fumigation. Plant Dis.

ponderosa. Phytopathology 53:1072-1076.

Reptr. 57:914-917.
National Academy of Sciences.

Kickert, Ronald, N. and P. R. Miller.

1977. Ozone and other photochemical oxidants.

1978. Responses of ecological systems. In Comm. on Med. and Biol. Eff. of Environ.

Handbook of methodology for the assessment

Pollutants. 719 p. NAS, Washington, D.C.

of air pollution effects on vegetation.

W. W. Heck, S. V. Krupa, S. N. Linzon,

eds. p. 14.1-14.45 Air Poll. Contr. Assoc.

Pittsburgh, Pa.

Nicholson, C. R.
USEPA.

1977. The response of 12 clones of eastern

1977. Photochemical oxidant air pollution

white pine (Pinus strobus) to ozone and

effects on a mixed conifer ecosystem -
nitrogen dioxide. 141 p. M.S. Thesis,
A progress report. 339 p. USEPA Ecol.

Virginia Polytechnical Institute and State

Res. Series EPA-600/3-77-104. Corvallis,

University. Blacksburg, Va.

Or.

Parmeter, J. R., Jr., R. V. Bega, and T. Neff.

USEPA.

1962. A chlorotic decline of ponderosa pine

1977. International Conference on photo-

in southern California. Plant Dis. Reptr.

chemical oxidant pollution and its

46 :269-273. control. 1169 p. Vol. I and 1 1 . USEPA

Ecol. Res. Series EPA-600/3-77-OOla, OOlb.

~karby,L.
Res. Triangle Park, N.C.

1979. Elevated ozone levels at the Swedish

west coast and in southern Sweden (Skane)

Wilhour, R. G. and G. E. Neely.

using tobacco as an indicator plant. & 1977. Growth response of conifer seedlings

Report from the workshop "Ozone effects

to low ozone concentrations. In Inter-

on vegetation in Europe." M. Eastmond and

national Conference on ~hotochemical

L. skirby Eds. p. 9-12. Swedish Water and

Oxidant Pollution and Its Control. p .
Air Res. Inst., Stockholm.
635-645. USEPA Ecol . Res . Series
600/3-77/OOlb.

Skelly, J. M., S. F. Duchelle, and L. W. Kress.

1979. Impact of photochemical oxidant air

pollution on eastern white pine in The

Williams, W. T., M. Brady and S. C. Wilson.

Shenandoah, Blue Ridge Parkway and Great

1977. Air pollution damage to the forests

Smoky Mountains National Parks. Proc. I1

of the Sierra Nevada Mountains of Cali-

Conf. on Sci. Res. in Natll. Parks. San

fornia. J. Air Poll. Contr. Assoc. 27:

Francisco, CA.
230-234.

Skelly, J. M. and J. W. Johnston.

Wolff, G. T., P. J. Lioz, G. D. Wight, R. E.

1979. Oxidant air pollution impact to the

Meyers, and R. T. Cederwall.

forests of eastern United States - a 1977. An investigation of long-range trans-

literature review 30 p. USEPA Rep. No.

port of ozone across the midwestern and

EPA-600/3-79-045. Corvallis, OR.

eastern United States. Atmos. Env.

11:797-802.

Stark, R. W., P. R. Miller, F. W. Cobb, Jr.,

D. L. Wood, and J. R. Parmeter, Jr.

Wood, F. A.

1968. Photochemical oxidant injury and

1970. The relative sensitivity of sixteen

bark beetle (Co1eoptera:Scolytidae)

deciduous tree species to ozone. Phyto-

infestation of ponderosa pine. I. Inci-

pathology 50:579. (Abstr.) .
dence of bark beetle infestation in injured

trees. Hilgardia 39:121-126.

 Primary Productivity, Sulfur Dioxide,

and the Forest Ecosystem: an
Overview of a Case study1

Abstract: The objective of the West Whitecourt case study

was to determine the consequence of chronic long term ex-

posure of a forest ecosystem to low concentrations of

sulphur dioxide emissions originating from a "sour gas"

processing plant in west central Alberta, Canada. An inter-

disciplinary ecological approach was utilized. The

vegetation and atmospheric environment were characterized.

A concept of ecologically comparable sampling site selec-

tion was developed and applied in the West Whitecourt study

area. Laboratory and field measurements revealed a reduc-

tion in photosynthetic rate in lodgepole pine x jack pine

(Pinus contorta x Pinus banksiana) in the field. Reduction

of adenosine triphosphate (ATP) concentration in pine tissue

during SO2 fumigation in the field followed by complete re-

covery after termination of SO2 fumigation and the disruption

of mineral nutrient cycling in the forest ecosystem were

observed. Basal area increment measurements of 200 lodge-

pole x jack pine trees from 5 ecologically comparable

sampling sites revealed a decrease in wood production

directly related to the presence of sulphur dioxide emissions.

It is recommended that the concepts of the assimilatory

capacity of the environment for sulphur gas pollutants and

irreversible ecological modification be utilized as

measures of environmental quality.

Many review articles have been written' addres- Kickert and Miller, 19791, very little emphasis

sing the problem of air pollutants and forest has been placed upon integrated research programs

ecosystems (Tam and Aronsson, 1972; Smith, 1974; concerning the impact of chronic long term low

Miller and McBride, 1975; and Linzon, 1978). concentration air pollution stress on forest eco-

These reviews documented the extreme examples of systems. The objective of this paper is to pre-

acute high concentration long term air pollution sent an overview of a four-year integrated forest

stress on ecosystems and were essentially post- ecosystem case study designed to determine the

mortem studies. Environmental change due to air consequence of chronic long term exposure of a

pollution stress was clearly visible in these conifer forest ecosystem to low concentrations of

cases. With the exception of the San Bernardino sulphur dioxide emissions originating from a

Mountain study investigating the effects of ''sour gas" processing plant in Alberta, Canada

photochemical oxidants on a mixed conifer forest (Legge and otters 1978).

ecosystem in California (Miller and others, 1977;

BACKGROUND TO THE CASE STUDY

^presented at the Symposium on Effects of Air Potential detrimental environmental impact of

Pollutants on Mediterranean and Temperate Forest sulphur dioxide emissions from the sour gas proc-

Ecosystems, June 22-27, 1980, Riverside, cessing industry upon the environment in Alberta

California, U.S.A. was a major concern of this industry in the early

2~rofessionalAssociate, Kananaskis Centre for 1970's. This concern lead to the formation of

Environmental Research, University of Calgary, the Whitecourt Environmental Study Group (WESG)

Calgary, Alberta T2N 1N4 Canada in 1971, a consortium of eight companies involved

in the production of saleable natural gas from

research indicated the potential for environmental

sour gas (natural gas containing hydrogen sul-

change in the field due to sulphur dioxide ex-

phide) in the Whitecourt district of west-central

posure. To resolve these contradictory research

Alberta. It was clear at that time that the

results it became clear that a detailed field

assessment of the impact of sulphur dioxide on

case study of a forest ecosystem surrounding a

the forest ecosystem was not a simple cause and

sulphur dioxide source was required.

effect relationship. A five-year environmental

research program therefore was initiated in 1972

The AMOCO Petroleum Company Limited West

and was cooperatively funded by both industry and

Whitecourt (Windfall) sour-gas processing plant

the Alberta Government. The objective of the re-

was chosen for the case study. This gas plant

search program which was called the Whitecourt

had the longest operational history in the

Environmental Study was to determine the environ-

Whitecourt study area beginning operation in 1962.

mental consequences of the operation of sour-gas

processing facilities on the forest ecosystem in

the Whitecourt district occupied by 11 sour gas

SOURCE OF SULPHUR GAS EMISSIONS

processing plants and defined as the Whitecourt

study area (figure 1).

The following is a brief outline of the opera-

tion of the West Whitecourt sour-gas processing

and sulphur recovery plant to familiarize the

reader with the origin of sulphur gas air pollu-

tion from the sour-gas processing industry in

Alberta.

Hydrogen sulphide is the principle sulphur com-

pound present in sour natural gas and is removed

by a series of chemical processes in a sulphur

recovery gas plant as elemental sulphur. The H2S

not converted to elemental sulphur is incinerated

in a high temperature reaction furnace (580'

Celcius) in excess air and methane where it is

oxidized to sulphur dioxide (SO2) and vented to

the atmosphere from a tall (122 meter) "candy-

striped" incinerator stack. In addition to an

incinerator stack, smaller stacks called flare

stacks, generally less than 46 meters in height,

Figure 1. Map of western Canada showing in bold

outline the province of Alberta and the

--
-
TOTAL EMISSIONS/MONTH
MAIN STACK/MONTH
... ........,... FLARE STACK 1 MONTH
Whitecourt Study Area.

3200

The Whitecourt study began in 1972 as a remote

sensing airborne environmental survey accompanied

by assessment on the ground carried out by INTERA

Environmental Consultants Limited to determine if

there were any visible large scale environmental

disturbances. After two years of general re-

connaissance no large scale environmental dis-

turbance was found (Whitecourt Environmental

Study 1972 and 1973). Controlled SO2 fumigation

experiments carried out in the laboratory by the

Kananaskis Centre for Environmental Research,

University of Calgary, on young lodgepole pine

(Pinus contorta Loud.) seedlings, however, re-

vealed a direct effect of SO2 on vegetation.

These experiments showed that, although plants

have the ability to adjust physiologically with-

in certain environmental limits, plants were

adversely affected when these limits were ex-

ceeded (Legge and Harvey 1974). A conflict

therefore arose; the general environmental field

Figure 2. Monthly sulphur emission history of

survey indicated no large scale modification of

the West Whitecourt Gas Plant from 1970 through

the vegetation while the preliminary laboratory

1976.

are used to burn small waste quantities of

growth (Katz 1949; Linzon and others 1978). The

sulphur recovery gas plant process and compressor

distinction between the assimilation and the

gases. Except in the case of a gas plant opera-

accumulation of sulphur of atmospheric origin by

tional upset, when for short periods the flare

plantsmust be addressed in any ecosystem study.

stack may contribute more sulphur gas emissions

to the atmosphere on a daily basis than the in-

cinerator stack, the incinerator stack is the

CONCEPTUAL APPROACH TO CASE STUDY

main source of sulphur gas emissions from the

West Whitecourt Plant.

To carry out an ecosystem case study one must

have a basic understanding of what an ecosystem

The monthly sulphur emissions (in long tons)

Is. The ecosystem is the basic functional unit
of the West Whitecourt Gas Plant for 1970 through
of ecology since it includes both the living

1976 is shown in figure 2; simply double the

organism and the non-living environment in which

sulphur emissions to obtain the SO2 emissions.

these organisms live. Odum (1971) has defined

It is important to note that this gas plant has

an ecosystem as "any unit that includes all of

reduced its sulphur emission output per day an

the organisms in a given area interacting with

order of magnitude since start-up in 1962 from

the physical environment so that a flow of energy

150 long tons/day to 18 long tonslday in 1976.

leads to clearly defined trophic structure, bio-

This reduction was achieved by enhanced operating

tic diversity and material cycling within the

procedures and the addition of "tail-gas" re-

system". Due to the inseparable nature and

covery units.
interdependence of the components of ecosystems

upon one another, however, any change that occurs

in one component of the ecosystem potentially

SULPHUR: THE NUTRITIONAL CONTROVERSY

affects all the components of that ecosystem.

The higher the diversity of an ecosystem there-

Sulphur is an essential nutrient element for

fore the more numerous the interrelationships

normal plant growth and metabolism. It is re-

within the ecosystem (Jernelou and Rosenberg 1976).

quired in intermediary metabolism and is a con-

The stability of an ecosystem can be viewed as a

stituent of many organic compounds such as amino

function of the balances amongst the components

acids and proteins in plant tissue. Sulphur

of that ecosystem. An environmental stress such

normally enters the plant via the root system in

as air pollution can modifiy the stability of an

the form of sulphate, which is biochemically re-

ecosystem by disrupting the balance amongst the

duced and then converted into numerous organic

ecosystem components.

compounds. Plants, however, can also take-up

and utilize SO2 from the atmosphere via the

Prior to the initiation of the West Whitecourt

stomates and utilize it as sulphur source for

case study, it was recognized that an ecosystem

plant nutrition. Faller (1971), for example, has

study was an interdisciplinary undertaking. The

shown that tobacco plants can utilize SOy as a

term interdisciplinary in this context means the

source of sulphur in sulphur deficient soils.

amalgamation of a set of specific disciplinary

This type of information has led many government

talents to work together to address a complex

and industry departments to say that SO2 emitted

environmental problem. An interdisciplinary re-

from industrial sources is actually beneficial as

search team was assembled by the Kananaskis Centre

an aerial fertilizer for plants growing on sul-

for Environmental Research of the University of

phur deficient soils (Terman 1978; Noggle and

Calgary, The University of Alberta, The University

Jones 1979). The situation is not quite -so

of Washington, San Jose State University and the

simple, however. Though a small amount of atmos-

Southern Alberta Institute of Technology. The

pheric SOo can be nutritional to plants in the

scientific expertise of the research team was

short term, the large amount and high frequency

broadly based and ranged from remote sensing,

of uncontrolled application of sulphur such as

ecology, taxonomy, genetics, plant physiology,

SOy to an ecosystem by sulphur sources such as

analytical chemistry, biochemistry, stable iso-

smelters, pulp and paper mills, coal-fired power

tope physics and meteorology to statistics and

plants, oil sand and oil shale extraction plants

electrical engineering.

and sulphur recovery gas plants can be detri-

mental in the long term. Different plant species

A conceptual model was developed to illustrate

not only have different nutritional requirements

the dynamic relationship between the sulphur

for sulphur, but the rate at which plant species

dioxide "source" and the generalized ecosystem

assimilate sulphur is influenced by many other

"sink" and is shown in figure 3. For purposes of

variables, such as physiological status, age,

communication among disciplines, the ecosystem

time during the growing season, temperature,

was sub-divided into the following four compart-

soil nutrient availability, and light intensity

ments: air, vegetation, soil, and water. The two-

to name a few. When more sulphur is available

way arrows indicate the inter-relationship of the

than can be assimilated it is accumulated in the

four ecosystem compartments. The expertise of

tissue (Ulrich and others 1967; Legge and others

each member of the research team was thus focussed

1977; Cowling and Koziol 1978; Thompson and Kats

on more than one of the ecosystem compartments at

1978). This foliar sulphur accumulation can

all times. This lead to the formulation of co-

reach toxic levels and adversely affect plant

operatively designed experiments to evaluate not

 fir [Abies balsamea (L.) Mill.] (Legge and others

A physiognomic classification of the vegetation

communities in the West Whitecourt study area

identified 13 major cover types out of 24 com-

munity types. Ten climax vegetation associations

VEGETATION were recognized. The vegetation of the study

area was mapped using a combination of LANDSAT

imagery, LANDSAT digital data and conventional

false colour infrared aerial photography both of

which were accompanied by ground based verifica-

tion. With the aerial colour infrared

photography of the West Whitecourt study area as

a subsample, computer mapping utilizing LANDSAT

digital data was completed on 116,000 hectares to

place the study area in a regional perspective.

The computer mapping of the study area was eight

times faster than conventional photography alone

u
WATER
and had a comparative accuracy of 93 percent.

ECOLOGICALLY ANALAGOUS SITES

One of the principle difficulties encountered

by air pollution researchers, before collecting

samples in the field for analysis, is the selection

of sampling locations. Most sampling locations in

air pollution studies are chosen solely on the

Figure 3. Conceptual model of the forest eco-

basis of a gradient which is usually a function of

system in the West Whitecourt case study area.

the prevailing wind and the distance from a poll-

ution source. Not enough emphasis is placed upon

the structure and composition of the plant communi-

only the interfaces between ecosystem compartments

ties from which samples are taken. Since an eco-

but also the processes within the ecosystem com-

system is composed of many interrelated highly

partments. Scientists with different areas of

variable biological and physical components, the

expertise were brought into the case study, how-

response of these components to a chronic environ-

ever, as a function of the needs of the research

mental stress such as air pollution will also be

program. The selection and the timing of inter-

highly variable. If the range of variability of the

action of disciplinary participants were viewed

responses of ecosystem components to an environmen-

as critically important factors for the success

tal stress is not considered, the expression of

of the program so the case study grew in terms of

the effect of the environmental stress on the eco-

disciplinary participation from 7 in 1974 to

system components may not be detected. There must

10 in 1975 and finally to 12 in 1976 and.1977.

be a common basis for comparison of ecosystem com-

This evolutionary interdisciplinary approach

ponents therefore to determine both the gross and

added a dimension of insight into the fate of sul-

subtle effects of an environmental stress along a

phur gas emissions in the forest ecosystem that

gradient.

would not have been possible had the separate

disciplines of the research team been operating

The concept and criteria for ecologically

in isolation.
analogous sample site selection were developed

and applied during the West Whitecourt case study

in an attemot to minimize the variability of eco-

VEGETATION CHARACTERIZATION
system components and hence to minimize the varia-

bility of the response of the ecosystem components

The vegetation of the Whitecourt area is in-

to air pollution stress. The key to this concept

cluded in the predominately forest subregion of

of sample site selection is based upon comparable

the Boreal Forest Region of Canada (Halliday 1937)

ecological variability of the ecosystem components

and is characterized as a transition forest area

and comparable environmental variability

between the Boreal and Subalpine Forest Regions.

at the sampling locations chosen along a distance

The transitional nature of the common species of

gradient. The criteria for ecologically analogous

trees occurring in the area are actually repre-

sample site selection are summarized in Figure 4.

sented by populations of hybrid individuals be-

When the ecological and environmental variation

tween lodgepole pine (Pinus contorta Loud.) and

of ecosystem components at all the sam~lingloca-

jack pine (Pinus banksiana Lamb.), while the true

tions are as similar as possible, the sampling

fir in the area represents hybrids between alpine

sites are said to be ecologically analogous. The

fir [Abies lasiocarpa(Hook.) Nutt.] and balsam

major difference amongst the sampling locations

 downwind of the West Whitecourt Gas Plant. The

ECOLOGICAL ANALOGUE CONCEPT intensive experimental site therefore, was prin-
cipally exposed to sulphur gas emissions from the
flare stacks. Only under short-lived meteorolog-
ASSUMPTIONS 1 = 2 + 3 ical conditions such as would occur during the
break-up of an inversion would sulphur gas
1. Ecological Variables (sites A, +An ) emissions from the main incinerator stock reach
slope soil moisture the intensive experimental site.
aspect species density
soil type species diversity
It must be emphasized at this point that the
2. Environmental Variables (other than pollutants) air quality standards for SO2 of 0.2 p p m h hr. /
temperature solar radiation 24 hr. as set by the Alberta Department of the
wind precipitation Environment, were only exceeded on three occas-
ions at the intensive experimental site 1.5 km
3. Pollutant Variables
east of the West Whitecourt Gas Plant in over
composition
location
2500 hours of ambient air monitoring during the
distance 1975 and 1976 growing seasons in the West White-
concentration/conversion court study area.
frequency/duration
Five ecologically analogous lodgepole x jack
pine sampling locations were selected in the West
Whitecourt case study area. The five sample
sites were chosen in locations which were prog-
ressively downwind in the main path of sulphur
dioxide emission corridor. These sites were
chosen in this manner so that the sulphur dioxide
Figure 4. Summary of criteria utilized for the emission impact gradient would be very steep
selection of ecologically analogous sample sites. across the five sampling locations to maximize
the differences in the responses of the ecological
analogues to pollution stress. The conceptual
therefore is distance from the pollution source. ecological model presented in Figure 3 can be
The assumption is that the pollutant variables generalized to express the relationships amongst
such as concentration, frequency of fumigation the five ecological analogues and is shown in
and duration of fumigation will decrease in Figure 5. The ecologically analogous sampling
magnitude with increasing distance from the
pollution source. The magnitude of the air
pollution stress on ecosystem components at the
sampling locations will correspondingly decrease
with increasing distance from the pollution
source. This procedure ensures that the expres-
sion of the air pollution stress on ecosystem
components will be maximized.

Meteorological and air quality data were

essential in sample site selection. The prevail-
ing winds during the growing season in the study
area were shown to have the highest frequency
of occurrence from the WNW and the second
highest frequency from the ESE. Sulphur dioxide
emissions from the West Whitecourt Gas Plant
therefore occurred with greatest frequency in
an ~ S corridor.
E Although idealized, the IMPACT GRADIENT
corridor concept provided an essential point
SOURCE SINK
of reference for the areas chosen for the
selection of sampling locations in the West
Whitecourt study area.
Figure 5. Replicated conceptual ecological model
The range of distances at which maximum illustrating the five ecologically analogous
ground level concentrations would occur from lodgepole x jack pine sampling locations along
sulphur gas emissions from the main incinerator a sulphur dioxide concentration gradient.
stack and flare stacks was calculated using the
simple Gaussian plume model under mean wind con-
ditions and a wide range of stability classes. sites were located at distances of 1.2 km (A=),
The maximum ground level concentrations would 2.8 km (AII), 6.0 km (AII~), 7.5 km (AIv) and 9.6
occur between 1.6 and 34.0 km downwind of the km (Av) from the Gas Plant. Sampling locations in
incinerator stack and between 0.4 and 2.6 km the study area other than ecological analogues
were designated by the letter S.
 CASE STUDY DATA OVERVIEW
increasing foliar age. The foliar sulphate-sulphur

data strongly support the concept of an environ-

Foliar sulphate-sulphur concentration in

mental stress gradient presented earlier.

lodgepole x jack pine trees was found to be a

better measure of foliar sulphur accumulation

The stable sulphur isotopic composition ( s /

than the foliar total sulphur concentration.

s3' ratio) of sulphur dioxide emissions from sour

The method of Johnson and Nishita (1952) was

gas plants in Alberta have been shown to differ

used to determine foliar sulphate-sulphur

from the natural environmental background stable

concentration while a Leco Furnace was used

sulphur isotopic composition (Lowe and others,

to determine foliar total sulphur. Figure 6

1971; Krouse, 1977). This is referred to as the

shows a plot of foliar sulphate-sulphur concen-

6 3 4 value.
~
The more positive the 63^S value the
tration in age-classed lodgepole x jack pine
greater the enrichment in s ~ ~The . background

stable sulphur isotopic composition in the West

Whitecourt study area is near 0.

The stable sulphur isotopic composition of

sulphur dioxide emissions from the incinerator

stack at the West Whitecourt Gas Plant was shown

I to be +22.2'/00 (per thousand). This difference

I W x t Whilmcourt Study Area

-- -
Pinut contort0 a P.banktiam
provided an environmental tracer for sulphur of

industrial origin.

1 Birch Mountain Firm Tmw

Pinut bonkeiana K R contww
The mean 6 3 4 value
~
of 1974-1976 foliage from

the ecologically analogous sampling locations

AI through Av remained close to the mean 6 3 4 ~

value for the incinerator stack at +22.2' l o o

while the mean foliar sulphate-sulphur concentra-

tion in 1974-1976 foliage decreased from 422 ppm

to 185 ppm. These data clearly show that lodge-

pole x jack pine trees were obtaining some of

their sulphur directly from the atmosphere from

sulphur gas emissions originating from the West

Whitecourt Gas Plant. Needles in the upper crowns

of many lodgepole x jack pine trees in the West

Whitecourt study area, however, displayed foliar

@S values which were greater than those asso-

ciated with incinerator stack emissions. These

data when compared to the laboratory data of

Wilson and others (1978) was suggestive of iso-

topically selective metabolic processes function-

ing in the lodgepole x jack pine foliage under

field conditions .
100' , I , , , , , The 28-meter radio mast tower erected at the

6 4 2
KILOMETERS WEST KILOMETERS EAST
intensive experimental site provided a framework

4
for measuring and characterizing SO2 concentration

profiles of ambient air above, within and below

the lodgepole x jack pine forest canopy. The

vertical SO2 profiles revealed that the SO2

Figure 6. Plot of foliar sulphate-sulphur con-

concentration minimum measured in the upper crown

centration in age-classed lodgepole x jack pine

at 16 meters was orimarily due to an aerodynamic

foliage from eight sampling locations in the

effect and was not due to the trees acting as a

West Whitecourt study area as a function of

biological sink. This aerodynamic effect was

distance from the West Whitecourt Gas Plant.

described as a splitting of the air flow above

and below the crowns of the lodgepole x jack pine

trees. The measurements of foliar sulphate-

foliage as a function of distance from the West

sulphur, foliar total sulphur and foliar S^^S

Whitecourt Gas Plant. The background foliar

values revealed that the trees, however, were

sulphate-sulphur concentration was determined

also a biological sink for sulphur gas emissions

from age-classed foliage of jack pine x lodge-

but that the rate of atmospheric sulphur uptake

pole pine from the Birch Mountain Fire Tower

by the trees was so slow that it was beyond the

Ill km (69 mi) NNW of Fort McMurray, Alberta.

resolution of the two Thermo Electron Model 43

One can clearly see the decrease in foliar

Pulsed Fluorescent SO2 analyzers used.

sulphate-sulphur concentration with increasing

distance from the sulphur gas emission source.

Photosynthetic rates and leaf resistances of

The decrease in foliar sulphate-sulphur concen-

lodgepole x jack pine trees in the West Whitecourt

tration with distance was more pronounced with

study area were shown to be modified. The amount

of this ecological modification was a function of

Soil pH profiles were measured at the same nine

the distance from the West Whitecourt Gas Plant.

vegetation sampling locations where foliar mineral

For example, the seasonal photosynthetic rates of

nutrient concentrations were determined since soil

1976 lodgepole x jack pine foliage were lower and

pH is known to affect the availablility of mineral

the leaf resistances higher when foliage from

nutrients to plants. The general trend or grad-

S5 (1.5 km east) was compared with foliage from

ient in soil pH over all nine sampling locations

Sin (5.2 km east) (4.38 Â 1.99 mg C02/dry g/hr was an increase in soil pH with depth and with

versus 6.42 Â 1.28 mg C02/dry g/hr and 11.3 2 distance from the West Whitecourt Gas Plant and

6.6 s/cm versus 7.8 5 1.9 s/cm for sample sites distance from the ~ S sulphur
E
gas emission

S5 and Sin respectively). This relative reduc-

corridor. The soil pH gradient was most striking

tion in photosynthetic rates, however, was only

when only the ecologically analogous sampling lo-

partially attributable to increased leaf re-

cations were considered. A direct relationship was

sistance. Additional ecological factors there-

found between lowered soil pH and the elevated

fore, such as foliar mineral nutrient status

levels of foliar Mn in lodgepole x jack pine trees.

and soil pH were considered since these para-

The foliar Mn concentration data and the soil pH

meters were known to modify plant response.

data suggest that foliar Mn concentration in

lodgepole x jack pine trees could be used as a

A detailed analysis of foliar mineral nutrient

mineral nutrient indicator of modification of the

concentration of N, P, K, Ca, Mg, Mn, Al, Fe and

forest ecosystem by sulphur gas emissions.

Zn in lodgepole x jack pine foliage from nine

sampling locations (including the ecological

Soil total sulphur concentration in the soil

analogues) in the West Whitecourt study area

profiles at the nine soil sampling locations also

revealed that the mineral nutrient status of

decreased with soil depth, distance and direction

the lodgepole x jack pine trees had been altered.

from the West Whitecourt Gas Plant. It is impor-

It must be remembered at this point that normal

tant to note, however, that there was no correla-

plant growth requires a balance of all essential

tion between soil total sulphur concentration and

mineral nutrients within the plant. The foliar

soil pH. There was also no direct correlation

concentration of P, K, Fe, Mg, N and Zn tended

between a given soil pH value and the soil f i 3 %
to increase while the foliar concentration of
value. These data suggest that the soil S^S

Ca and A1 tended to decrease with distance from

value can be used as an indicator of the presence

the West Whitecourt Gas Plant and distance from

and penetration of sulphur gas emissions into the

the WN-SE sulphur dioxide emission corridor.

soil profile while soil pH and soil total sulphur

Site type was shown to be a critical factor

can be used as indicators of sulphur loading of

influencing the concentration of these eight

the soil.

mineral nutrients. Foliar Mn concentration was

found to decrease dramatically with distance from

In terms of plant biochemistry and sulphur gas

the West Whitecourt Gas Plant and distance from

emissions the most significant observation in the

the W N W S E sulphur gas emission corridor.

field was a transient metabolic effect; the ATP

Variability of site type, however, did not

(adenosine triphosphate) concentration of foliage

modify this relationship. A low Fe to Mn ratio

cells from lodgepole x jack pine trees was found

was found in foliage from sampling locations

to be directly decreased upon exposure to low-con-

within 4 km of the West Whitecourt Gas Plant.

centration short duration SO2 fumigation (75%

The low foliar Fe concentration may contribute

decrease upon fumigation with 0.14 ppm SO2 for

to the chlorotic appearance of the foliage at

15 minutes). It is important to note that when

these locations. Foliar mineral nutrient analysis

the foliage was no longer exposed to SO2 the

of foliage 82, 85, A1 and Slo revealed that foliar

foliar ATP concentration increased to the pre-

K and P were lower in concentration in foliage

SO2 fumigation ATP concentration (see Harvey and

from S2, S5 and AT than from Sm.

Legge, 1979, for details). This decrease and

increase in the foliar ATP concentration was also

Since foliar K concentration has been linked

observed with excised lodgepole x jack pine

with stomatal activity, the reduced foliar K

branches from the West Whitecourt study area

concentration may be inhibiting stomatal opening

which were fumigated under controlled conditions

thus increasing leaf resistance which would then

at the Kananaskis laboratory. When lodgepole x

limit photosynthetic rate. Reduced foliar P

jack pine trees, which had been grown in the

concentration may inhibit phosphorylation and

absence of sulphur gas emissions in the laboratory,

thereby also limit photosynthetic rate. Foliar

were fumigated with SO2 no fluctuation in ATP

nutrient deficiencies of either P and K alone or

concentration was observed. It is important to

in combination therefore may be partially respon-

note that the laboratory trees had a foliar ATP

sible for the reduced photosynthetic rates ob-

content which was over twice the foliar ATP content

served in lodgepole x jack pine foliage in the

of the lodgepole x jack pine foliage from the

West Whitecourt study area.

West Whitecourt study area (658 nmoles/dry g

versus 1460 nmoles/dry g). The lower foliar ATP

The alteration of foliar mineral nutrient

concentration of field grown trees compared to

status in lodgepole x jack pine trees in the

laboratory grown trees suggests a partial explan-

West Whitecourt study area therefore is an

ation for the lowered photosynthetic capacities

important ecological factor contributing to

reported for lodgepole x jack pine trees in the

the modification of plant response.

West Whitecourt study area.

 The photosynthetic rate of lodgepole x jack

pine trees grown under controlled conditions in

a non-SO2 environment in growth chambers at the

Kananaskis laboratory and lodgepole x jack pine

trees grown in the sulphur dioxide emission en-

vironment in the field at the intensive West

Whitecourt study site S5 was measured when both

sets of trees were exposed to similar low concen-

tration short-duration SO2 fumigations. The

photosynthetic rate of the field grown plant

material was not depressed by SO2 fumigation

while the photosynthetic rate of the laboratory

grown material was depressed by the SO2 fumiga-

tions. The photosynthetic rate of the laboratory

trees was much greater than the trees in the

field. There was also no evidence of a plant-

water deficit in lodgepole x jack pine trees

severe enough to effect photosynthetic rate.

Adenosine triphosphate is the major bio-

chemical intermediate of energy transfer in

biological systems. A decrease in foliar ATP

Figure 7. Comparison plot of the mean photosyn-

content in lodgepole x jack pine trees caused

thetic capacity of 1976 foliage on lodgepole x

by SO2 fumigation therefore would also be a

jack pine branches from sampling sites S2, S5 and

decrease in the amount of biochemical energy

Slo from early in June to mid-September.

available for normal metabolic functions. Al-

though the ATP content of the lodgepole x jack

pine foliage recovered to the pre-SO2 exposure

When the observed ecological modifications of

concentration after the SO2 stress was removed,

the forest ecosystem such as reduced needle bio-

during the SO2 fumigation there would have been

mass, reduced biochemical energy, reduced photo-

a net loss of biochemical energy. The fact that

synthetic rates, reduced soil pH, the disruption

foliar ATP content increased after SO2 fumigation,

of mineral nutrient cycling, foliar sulphur load-

indicated that the trees were coping with the

ing and the shortened growing season are combined,

sulphur gas emissions at the cost of a metabolic

and considered in the long term time sense the net

energy drain.
effect should be measurable as a reduction in

forest productivity. This decrease in forest pro-

In summary, the contrast in the biochemical

ductivity, however, would be expected to decrease

and physiological responses of the lodgepole x

with increasing distance from the sulphur gas

jack pine trees fumigated with SO2 in the field

emission source.

and the laboratory strongly indicates that en-

vironmental pre-history and acclimation of the

Annual basal area increment measurements were

trees to ecological modification of components

taken from 40 lodgepole x jack pine trees at each

of the forest ecosystem are the critical factors

of the five ecologically analogous sampling loca-

determining plant response to sulphur gas

tions AI through Av in 1976 to determine if mod-

emissions in the West Whitecourt study area.

ification of the forest ecosystem was significant

enough in the long term sense to be reflected in

A comparison of the mean photosynthetic capac-

reduced wood production since the initial start-

ities of 1976 lodgepole x jackpine foliage

up of the West Whitecourt Gas Plant in 1961-1962.

through the 1976 growing season at sampling loca-

Statistical analysis of the basal area increment

tions S2, S5 and Slo revealed that S10 had a

data shown in Figure 8 revealed that distance from

positive net C02 fixation balance two to three

the West Wh'itecourt Gas Plant, time in years, and

weeks prior to the foliage at S2 and S5 and is

their interaction had significant effects on the

shown in Figure 7. The mean photosynthetic

basal area increment of the lodgepole x jack pine

capacity of 1976 Sin foliage was also always

trees from the five ecological analogues. An

higher than the mean photosynthetic capacity

exponential growth curve model was determined for

measured for 1976 foliage from S2 or S5 . Addi- lodgepole x jack pine trees from Airand the growth

tionally in terms of photosynthetically active

curve of the lodgepole x jack pine trees fromA1,

needle biomass lodgepole x jack pine branches

A1l, AIII and AIv were statistically compared to it.

sampled at Sl, 82, AI and S5 were chlorotic in

The basic underlying assumption was that sulphur

appearance with premature abscission (needledrop)

gas emissions had not had a significant effect on

of the third year needles and poor leader growth

the growth of the trees at Av. This analysis

while branches sampled at S q and Sinwere compar-

statistically revealed that there has been a defi-

atively darker green in color with a needle re-

nite reduction in basal area increment in lodgepole

tention of from four to six years and good leader

x jack pine trees since 1962 in AI, A,.,, AIII and

growth. The photosynthetic potential of lodge-

AIv compared to the basal area increment model

pole x jack pine trees based upon needle biomass

for AV which was attributable to sulphur gas

alone therefore was much greater at S9 and Sin

emissions from the West Whitecourt Gas Plant.

compared to Sl, Sy, A1 or $5.


ll I I I I
1950 1955 1960 1965 1970
YEAR O F GROWTH
Figure 8. Comparative p l o t s of t h e mean b a s a l
a r e a increments from 40 lodgepole
X jack pine t r e e s a t each of t h e
f i v e e c o l o g i c a l l y analagous sampling
s i t e s i n t h e West Whitecourt study
area.
The maximum r e d u c t i o n i n b a s a l a r e a increment
occurred a t A 1 and A 1 1 and p r o g r e s s i v e l y de-
c r e a s e d t o z e r o a t Ay. The e f f e c t of sulphur
gas emissions on b a s a l a r e a increment growth
i n lodgepole x j a c k pine t r e e s s i n c e 1961 a t
AI, AII, AIII, and AIv r e l a t i v e t o Av i s thus
a gradient with the reduction i n basal area
increment r e s u l t i n g from sulphur gas emission
f a l l i n g t o z e r o a t Av o r 9.6 km. I f t h e t o t a l
b a s a l a r e a increment r e d u c t i o n of AT r e l a t i v e
t o Av i s averaged over t h e f o u r t e e n y e a r s s i n c e
t h e s t a r t - u p of t h e West Whitecourt Gas P l a n t ,
t h i s would correspond t o approximately a one
t o two p e r c e n t r e d u c t i o n i n b a s a l a r e a i n c r e -
ment growth of AI r e l a t i v e t o Av.
The a r e a l e x t e n t of p o s s i b l e m o d i f i c a t i o n of
components of t h e f o r e s t ecosystem by sulphur
g a s emission from t h e West Whitecourt Gas P l a n t
can be e s t i m a t e d u s i n g t h e following assump-
t ions :
1. sulphur gas emissions r e a c h t h e f o r e s t
ecosystem w i t h i n 17 km. (10.6 mi.) of
t h e source; and
2. t h e impact of sulphur g a s emissions i s
r e s t r i c t e d t o a r e a s NW and SE of t h e
West Whitecourt Gas P l a n t .
The a r e a a f f e c t e d by sulphur d i o x i d e e m i s s i n s ,
t h e r e f o r e , i s approximately 454 km 2 (175 m i ) 5
o r 45,373 h e c t a r e s (112,130 a c r e s ) . This a r e a l
e x t e n t e s t i m a t e of impact i s c o n s e r v a t i v e be-
cause t h e 17 km d i s t a n c e i s o n l y one-half t h e
d i s t a n c e range c a l c u l a t e d u s i n g t h e simple
Gaussian plume model under a l l s t a b i l i t y c l a s -
s e s f o r maximum ground l e v e l c o n c e n t r a t i o n of
sulphur d i o x i d e emissions from t h e main i n -
cinerator stack.
It i s important t o b e a r i n mind a t t h i s
p o i n t , however, t h a t t h i s p r o j e c t e d impact
a r e a h a s n o t been uniformly modified by s u l -
phur gas emissions b u t r a t h e r h a s been mod-
i f i e d i n terms of an impact g r a d i e n t extending
NW and SE from t h e West Whitecourt Gas P l a n t .
I n o t h e r words, t h e e x t e n t of ecosystem com-
ponent m o d i f i c a t i o n w i l l d e c r e a s e w i t h d i s -
t a n c e from t h e sulphur g a s emission source.
Another f a c t o r must be considered a t t h i s
p o i n t . Sulphur emissions from t h e West White-
c o u r t Gas P l a n t have been reduced almost a n
o r d e r of magnitude s i n c e 1970 ( r e f e r t o F i g -
u r e 2 ) . This s i g n i f i c a n t r e d u c t i o n i n emissions
w i l l n o t o n l y g e n e r a l l y d e c r e a s e t h e magnitude
of t h e i m p a c t , o f sulphur emissions on t h e f o r -
e s t ecosystem, i t w i l l a l s o d e c r e a s e t h e a r e a l
e x t e n t of t h e a r e a impacted by s u l p h u r emissions
i n t h e p a s t thus allowing a p o r t i o n of t h e f o r -
I
e s t ecosystem t o recover from t h e previous
sulphur gas emission s t r e s s .
1975
When one uses f o l i a r s u l p h a t e - s u l p h u r con-
c e n t r a t i o n i n lodgepole x j a c k p i n e t r e e s a s a
measure of sulphur accumulation from exposure
t o t h e c u r r e n t l e v e l of sulphur g a s emissions,
i t appears t h a t a t o l e r a b l e c o n c e n t r a t i o n i s
reached by 9-12 km (5.6-7.5 mi) of t h e West
Whitecourt Gas P l a n t . This i s i n d i c a t e d by a
decrease i n f o l i a r sulphate-sulphur concentration
w i t h n e e d l e age which i s w i t h i n t h e range o f
t h e background f o l i a r s u l p h a t e - s u l p h u r con-
c e n t r a t i o n . The f o l i a r &^s v a l u e s , however,
could be used t o provide a more e x a c t measure
of t h e d i s t a n c e a t which t h e presence of s u l -
phur gas emissions become n e g l i g i b l e t o com-
ponents of t h e f o r e s t ecosystem.
CONC LUS I O N
It i s c l e a r from t h i s c a s e study t h a t s u l -
phur gas emissions from t h e West Whitecourt
Gas P l a n t have modified t h e f o r e s t ecosystem
i n a number of ways. The main e c o l o g i c a l pro-
c e s s which has been d i r e c t l y and i n d i r e c t l y
a f f e c t e d by sulphur d i o x i d e emissions i s min-
e r a l n u t r i e n t c y c l i n g . By p r o g r e s s i v e l y a l t e r -
i n g t h e mineral n u t r i e n t b a l a n c e s of ecosystem
components f o r example, t h e b i o l o g i c a l r e -
l a t i o n s h i p s amongst t h e components and t h e
p h y s i o l o g i c a l and biochemical f u n c t i o n s of t h e
components a r e modified. It i s t h e s e ecosystem
component modifications which are the ex-
The conceptual interdisciplinary nature of

pressions of environmental deterioration re-

the West Whitecourt case study has proven to

sulting from chronic exposure to sulphur di-

be the basis for the success in unravelling

oxide over time. Despite this measurable

the very complex interrelated consequences of

deterioration of the forest ecosystem, however,

the chronic exposure of the forest ecosystem

it does not appear at this time that sulphur

to sulphur gas emissions from the West White-

dioxide emissions from the West Whitecourt Gas

court Gas Plant. It is suggested that fu-

Plant have caused irreversible ecological de-

ture air pollution research on forest eco-

gradation. With the significant reduction in

systems follow a similar experimental design

sulphur emissions from the West Whitecourt

if the environmental perturbations caused by

Gas Plant (See Figure 2) it is not antici-

air pollution stress are to be understood.

pated that there will be significant irre-

Direct extrapolation of the data summarized

versible ecological modification of the forest

in this paper to other areas would be mis-

ecosystem in the remaining 10 to 20 years of

leading unless local environmental factors,

operation of the West Whitecourt Gas Plant.

vegetation and pollutant parameters, are taken

One philosophical dilemna has resulted

into consideration prior to interpretation.

from the West Whitecourt case study. There

is no relationship between air quality stan-

ACKNOWLEDGMENTS

dards and the maintenance of environmental

quality since the term environmental quality

The West Whitecourt Case Study could not

excludes environmental modification. No effort

have been completed without the enthusiasm

to date has been made to address or to quantify

and cooperation of the following interdisci-

acceptable limits of environmental modification

plinary team members: D.R. Jaques, G.W.

resulting directly or indirectly from air

Harvey, H.R. Krouse, H.M. Brown, E.C. Rhodes,

pollution stress despite the fact that the

and M. Nosal of the University of Calgary;

presence of air pollutants in the atmosphere

H.U. Schellhase of the Southern Alberta In-

implies that a certain amount of environmental

stitute of Technology; J. Mayo and A.P.

modification is acceptable.
Hartgerink of the University of Alberta;

Since, at the present time it is techno-

P.F. Lester from San Jose State University;

logically and economically impossible to re-

and R.G. Amundson and R.B. Walker of the

move all air pollutants from industrial pro-

University of Washington.

cesses, it is suggested that irreversible

The majority of the financial support for

ecological modification of the environment

this research was in the form of a grant-in-

be used as an additional criteria for limiting

aid of research to the Kananaskis Center for

pollutant emissions to the atmosphere. The as-

Environmental Research of the University of

similatory capacity of the environment, in other

Calgary from the Whitecourt Environmental

words, must be taken into account by both

Study Group. Additional financial support

industry and regulatory agencies. The uni-

was received from the Research Secretariat

form application and enforcement of fixed air

of Alberta Environment, the Oil Sands En-

quality standards over a geographical area

vironmental Study Group (OSESG) , the Alberta
the size of the p ovince of Alberta (661,183
Oil Sands Environmental Research Program

5
km2 or 255,285 mi ) with its physiographically (AOSERP), and the University of Calgary Inter-

complex terrain, heterogeneous vegetation and

disciplinary Sulphur Research Group (UNISUL).
diverse climatology is clearly not enough
A special note of thanks is in order for

to maintain environmental quality. Future

Mr. Ron Findlay and Mr. E. Baraniuk of AMOCO

research will be required to determine the

Canada Petroleum Company Limited and the rest

assimilatory and accumulatory capacity of

of the Whitecourt study group members companies

the environment to pollutants and to provide

who had the foresight to initiate the White-

the biological monitoring techniques to assure

court Environmental Study and the patience to

that the assimilatory and accumulatory capacity

see it through.

of the environment is not exceeded.

Finally I wish to thank my air pollution

After the assimilatory and accumulatory

colleagues throughout North America for their

capacity of the environment have been con-

genuine interest and support during the re-

sidered, flexible air quality standards may

search program.

be possible. These standards could be ad-

justed regionally and seasonally in order

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Faller, N.

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1971. Plant nutrient sulphur- SO2 vs. So&.

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environmental quality.

Halliday, W. E. D.
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1937. A forest site classification for

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Katz, M.
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Kickert, R. N., and P. R. Miller.

R. J. Arkley, F. W. Cobb Jr., D. L. Dahlsten,

1979. Responses of ecological systems. 2 P. J. Gersper, R. F. Luck, J. R. McBride,

Handbook of methodology for the assessment

J. R. Parmeter Jr., J. M. Wenz, M. White, and

of air pollution effects on vegetation.

W. W. Wilcox.

W. W. Heck, S. V. Krupa, and S. N. Linzon,

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Krouse, H. R.
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1977. Sulphur isotope abundances elucidate

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Noggle, J. C., and H. C. Jones.

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D
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Legge, A. H., D. R. Jaques, H. R. Krouse, H. M.

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Thompson, R. C . , and G. Kats. Wilson, L. G . , R. A . Bressan, and P. F i l n e r .

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U l r i c h , A . , M. A . Tabatabai, K. Ohki, and C . M.

Johnson.
1967. S u l f u r c o n t e n t of a l f a l f a i n r e l a t i o n
t o growth i n f i l t e r e d and u n f i l t e r e d a i r .
P l a n t and S o i l 26(2):235-252.
 Effects of Airborne F on Forest
Ecosystems'
Robert G. Amundson and Leonard H. Weinstein

A b s t r a c t : Although t h e r e a r e many r e p o r t s of f l u o r i d e (F)

i n j u r y t o f o r e s t s , t h e r e have been no s y s t e m a t i c s t u d i e s on
t h e f o r e s t ecosystem. I n t h i s p a p e r , we have reviewed t h e
p r e s e n t s t a t e of o u r knowledge on F p o l l u t i o n and e f f e c t s
on p h y s i o l o g i c a l p r o c e s s e s , t r e e growth, F accumulation
and p l a n t i n j u r y , community s t r u c t u r e , i n t e r a c t i o n w i t h
pathogens and i n s e c t s , and d i s t r i b u t i o n of F i n t h e environ-
ment. The p r e p a r a t i o n of t h i s review i n d i c a t e d t h e many
a r e a s of t h e F-plant i n t e r a c t i o n on which t h e r e i s no i n f o r -
n a t i o n , where i t i s poorly understood, o r where a v a i l a b l e
i n f o r m a t i o n i s h i g h l y c o n t r o v e r s i a l . I n some c a s e s , we
have j o i n e d t h e c o n t r o v e r s y .

I n j u r y t o f o r e s t t r e e s p e c i e s by a i r b o r n e
f l u o r i d e (F) h a s been r e p o r t e d i n many p a r t s o f
t h e world ( e . g . , Adams and o t h e r s , 1952; Horntvedt
and Robak, 1975; Niklfeld.1975; EPA, 1973; F l i i i l e r
and o t h e r s , 1979), b u t many F-emitting s o u r c e s a r e
i n a g r i c u l t u r a l o r urban a r e a s and r e p o r t s of
i n j u r y t o agronomic c r o p s , ornamental and urban
t r e e s ( e . g . , Bolay and Bovay, 1965; Facteau and
M e l l e n t h i n , 1976; de Ong, 1946; Leonard and Graves,
1966) o r on f l u o r i d e accumulation and p r o d u c t i o n
of f l u o r o s i s i n l i v e s t o c k and o t h e r h e r b i v o r e s
( S u t t i e , 1977) a r e a l s o common.
The p r i n c i p a l i n d u s t r i a l s o u r c e s of a i r b o r n e
F a r e primary aluminum s m e l t i n g ; s t e e l manufacture;
conversion of f l u o r a p a t i t e t o phosphate and phos-
phorus; and g l a s s , ceramic and b r i c k p r o d u c t i o n .
N a t u r a l s o u r c e s of a i r b o r n e F a r e p r i n c i p a l l y
from s o i l p a r t i c l e s , fumaroles, and volcanoes.
The a s h from t h e r e c e n t e r u p t i o n of Mount S t .
Helens c o n t a i n e d 8 ppm s o l u b l e F ( S t o i b e r and
o t h e r s , 1980) and 400 ppm t o t a l F and i t s impact
on f o r e s t s i n t h e n o r t h w e s t e r n U.S. w i l l b e
p r e s e n t e d a t t h e Symposium on E f f e c t s of A i r
P o l l u t a n t s on Mediterranean and Temperate F o r e s t
Ecosystems, J u n e 22-27, 1980, R i v e r s i d e ,
C a l i f o r n i a , U.S.A.
z
Research A s s o c i a t e and Program D i r e c t o r ,
Environmental Biology, r e s p e c t i v e l y . Boyce
Thompson I n s t i t u t e a t C o r n e l l U n i v e r s i t y , I t h a c a ,
New York.
watched c a r e f u l l y .
R e s u l t s of o n l y a few f i e l d s t u d i e s made n e a r
F-emitting s o u r c e s a r e a v a i l a b l e i n t h e s c i e n t i -
f i c l i t e r a t u r e . One r e a s o n f o r t h e absence of
more r e p o r t s i s t h a t t h e s t u d i e s were o f t e n rou-
t i n e and n o t q u a n t i t a t i v e , making p u b l i c a t i o n i n
r e f e r e e d j o u r n a l s d i f f i c u l t . A second r e a s o n i s
t h a t r e s u l t s of a s t u d y performed f o r a n i n d u s t r y
may b e s e q u e s t e r e d from p u b l i c a t i o n o r o t h e r u s e
because of a c t i v e , pending, o r p o t e n t i a l l i t i g a -
t i o n . Often, f i e l d s t u d i e s t h a t have been d i s -
t r i b u t e d were i n a form t h a t was n o t s u b j e c t e d t o
p e e r review, was c a r e l e s s l y assembled, a n d / o r
r e f l e c t e d t h e p e r s o n a l b i a s e s of t h e a u t h o r s .
Because of t h e s e problems, we have n o t confined
t h i s review t o works p u b l i s h e d i n j o u r n a l s , b u t
we have t r i e d t o judge t h e r e p o r t s t h a t we have
c i t e d i n terms of t h e i r p e r t i n e n c e a n d / o r a v a i l -
a b i l i t y , and o u r p e r s o n a l views a r e o f t e n pre-
s e n t e d . Host i n t e r n a l r e p o r t s were avoided, b u t
t h e l a c k of p u b l i s h e d i n f o r m a t i o n o f t e n l e f t no
r e c o u r s e b u t t o c i t e them. We hope t h a t we have
s t a t e d o u r c r i t i c i s m s of some s t u d i e s a s f a i r l y
a s possible.
LABORATORY STUDIES ON PLANT PRODUCTIVITY
The p r o d u c t i v i t y of t h e p l a n t depends upon t h e
c o o r d i n a t i o n and r a t e o f CO-; a s s i m i l a t i o n , res-
p i r a t i o n , t r a n s p i r a t i o n , t r a n s l o c a t i o n of photo-
s y n t h a t e , m i n e r a l n u t r i t i o n , growth, and repro-
d u c t i o n . The amount of i n f o r m a t i o n a v a i l a b l e on
t h e impact of a i r b o r n e F on t h e s e p r o c e s s e s
ranges from v i r t u a l l y none, e . g . , on t r a n s l o c a t i o n
of p h o t o s y n t h a t e s , t o a moderate amount, e.g., on
C02 a s s i m i l a t i o n (apparent p h o t o s y n t h e s i s ) and
respiration.
There a r e few d a t a on t h e chemical composi-
t i o n , d i s t r i b u t i o n , and p a t t e r n s and frequency
of exposure of atmospheric F i n t h e f i e l d . One
reason f o r t h i s is t h a t a i r monitors with
s h o r t a v e r a g i n g times have n o t been g e n e r a l l y
a v a i l a b l e . Because F c o n c e n t r a t i o n s i n t h e am-
b i e n t a i r have n o t been c h a r a c t e r i z e d and f l u c -
tuating F concentrations a r e d i f f i c u l t t o control,
t h e d e s i g n of meaningful l a b o r a t o r y o r c o n t r o l l e d
f i e l d experiments i s formidable. The i n f o r m a t i o n
t h a t i s a v a i l a b l e i s g e n e r a l l y f o r a v e r a g i n g times
of 1 2 o r 24 h o u r s (McCune and o t h e r s , 1976) and
t h e peak F c o n c e n t r a t i o n s t h a t occur n e a r s o u r c e s
a r e n o t known. T h i s i n f o r m a t i o n would b e v e r y
u s e f u l s i n c e i t h a s been shown f o r o t h e r a i r
p o l l u t a n t s , such a s S02, t h a t s h o r t - t e r m peak
exposures a r e more i m p o r t a n t i n e x p l a i n i n g p l a n t
damage t h a n a v e r a g e c o n c e n t r a t i o n s (McLaughlin
and o t h e r s , 1979). U n f o r t u n a t e l y , most l a b o r a t o r y
s t u d i e s have employed continuous exposures a t
c o n s t a n t c o n c e n t r a t i o n s t h a t do n o t s i m u l a t e
f i e l d exposures. For t h e s e r e a s o n s t h e d a t a
a v a i l a b l e a r e of l i m i t e d v a l u e i n p r e d i c t i n g t h e
impact of a i r b o r n e F on f o r e s t ecosystems.
Gas Exchange
Apparent P h o t o s y n t h e s i s
Given t h e problems o u t l i n e d above, i t i s n o t
s u r p r i s i n g t h a t t h e r e have been s o few s t u d i e s on
t h e e f f e c t s o f F on a p p a r e n t p h o t o s y n t h e s i s (AP)
of f o r e s t tree s p e c i e s (Table 1 ) . Consequently,
we have i n c l u d e d i n Table 1 n o t o n l y s t u d i e s on
f o r e s t t r e e s b u t a l s o t h o s e on h o r t i c u l t u r a l
s p e c i e s exposed t o hydrogen f l u o r i d e (HF) o r
s u p p l i e d w i t h sodium f l u o r i d e (NaF). We have
a r b i t r a r i l y s e p a r a t e d experiments w i t h HF i n t o
a c u t e exposures (over 1 0 pg m 3 f o r a few 'days o r
l e s s ) and c h r o n i c exposures (ca. 5 \E HF m 3 o r
l e s s f o r a few days t o more t h a n a growing s e a s o n ) ,
a l t h o u g h we r e c o g n i z e t h a t many exposures c l a s s i -
f i e d a s c h r o n i c could more r e a l i s t i c a l l y b e
c l a s s i f i e d a s a c u t e . Exposures t o s o l u t i o n s con-
t a i n i n g NaF have v a r i e d from s e v e r a l h o u r s t o
months and w i l l b e d i s c u s s e d i n d i v i d u a l l y .
Acute exposures -- With t h e e x c e p t i o n of c o t t o n ,
where h i g h c o n c e n t r a t i o n s of HF had no e f f e c t
(Thomas, 1 9 5 8 ) , a c u t e exposures have c o n s i s t e n t l y
reduced AP (Thomas and Hendricks, 1956; Thomas,
1958; H i l l , 1969; Bennett and H i l l , 1973).
Bennett and H i l l (1973) exposed a l f a l f a t o HF f o r
2 h o u r s and found t h a t (1) approximately 120 yg m-3
HF were needed t o produce f o l i a r n e c r o s i s ; (2)
about 40 pg m-3 were n e c e s s a r y t o c l e a r l y i n h i b i t
AP; (3) t h e d e p r e s s i o n of AP and subsequent recov-
e r y a f t e r exposure were slower f o r HF t h a n f o r t h e
o t h e r major a i r p o l l u t a n t s t e s t e d (S02, 03, N02,
NO and Cl2). They a l s o noted t h a t of t h e p o l l u -
t a n t s t e s t e d , HF p r o d u c e d - t h e g r e a t e s t r e d u c t i o n
i n AP f o r an e q u i v a l e n t p o l l u t a n t dose, b u t s t a t e d
t h a t t h e occurrence i n the f i e l d of a concentra-
t i o n t h a t would produce a 1 0 p e r c e n t r e d u c t i o n i n
AP would b e r a r e . One can conclude t h a t a s i g n i -
f i c a n t i n f l u e n c e of a n a c u t e exposure on p l a n t
community p r o d u c t i v i t y would l i k e l y b e preceded
by l e s i o n s and a b s c i s s i o n o f f o l i a g e .
Chronic exposures -- S e v e r a l i n v e s t i g a t o r s
have r e p o r t e d t h a t c h r o n i c exposure t o HF had no
e f f e c t on AP i f t h e r e was no v i s i b l e i n j u r y ( H i l l ,
1969; H i l l and o t h e r s , 1958; Thompson and o t h e r s ,
1967), and when f o l i a r i n j u r y o c c u r r e d , t h e re-
d u c t i o n i n AP was p r o p o r t i o n a l t o ( H i l l , 1969;
Thomas and Hendricks, 1956; Thomas, 1958) o r
g r e a t e r t h a n t h e amount of f o l i a g e i n j u r e d (Thomas,
1958, f o r f r u i t t r e e s ; and Woltz and Leonard, 1964,
f o r c i t r u s ) . Thomas (1958) proposed t h a t t h e r e i s
a t h r e s h o l d of F c o n c e n t r a t i o n and d u r a t i o n o f ex-
posure f o r each s p e c i e s above which AP i s reduced
more t h a n can b e accounted f o r by c h l o r o s i s and
necrosis.
McCune and o t h e r s (1976) d e s c r i b e d experiments
i n which field-grown sorghum was exposed f o r 14
days t o t h r e e c o n c e n t r a t i o n s of HF (0.7, 1 . 7 and
3.5 t o > 5 yg m-3) and AP of t h e whole p l a n t
canopy was measured t h r e e t i m e s d a i l y b e f o r e ,
d u r i n g , and a f t e r t h e exposure p e r i o d s . The l o w e s t
HF c o n c e n t r a t i o n had no e f f e c t on AP; t h e i n t e r -
mediate c o n c e n t r a t i o n reduced AP d u r i n g t h e expo-
s u r e p e r i o d , b u t immediate recovery o c c u r r e d upon
c e s s a t i o n o f t h e exposures. P l a n t s s u b j e c t e d t o
t h e h i g h e s t c o n c e n t r a t i o n a l s o had reduced r a t e s
of AP f o r t h e f i r s t week. But when t h e HF concen-
t r a t i o n was r a i s e d t o g r e a t e r t h a n 5 m-3 on t h e
e i g h t h day, s e v e r e f o l i a r i n j u r y o c c u r r e d , t h e
r a t e s of AP dropped d r a s t i c a l l y , and t h e r e was no
recovery i n t h e post-exposure p e r i o d .
I n an e x t e n s i v e s e r i e s o f experiments, K e l l e r
(1977) p l a c e d 11 d i f f e r e n t t r e e s p e c i e s ( s e e
Table 1 ) a t v a r y i n g d i s t a n c e s from a s o u r c e of a i r -
borne F f o r s e v e r a l months and measured r a t e s of AP
on t h e whole p l a n t s r e t u r n e d t o t h e l a b o r a t o r y .
Exposure t o F produced f o l i a r i n j u r y and a b s c i s s i o n ,
and reduced t h e r a t e o f AP of t h e whole p l a n t . The
r e d u c t i o n i n AP o f t h e whole p l a n t was due p r i m a r i l y
t o t h e l o s s of f o l i a g e , because t h e r a t e of AP of
n e e d l e s remaining on t h e p l a n t s was a s h i g h a s t h o s e
on c o n t r o l p l a n t s .
Sodium f l u o r i d e -- Navara (1963) r e p o r t e d b o t h
d e p r e s s i o n and s t i m u l a t i o n of AP of beans grown i n
s o l u t i o n c u l t u r e f o r 16 days w i t h 0.03 o r 0.3 ppm
NaF, w h i l e t h o s e s u p p l i e d w i t h 3 ppm had d e p r e s s e d
r a t e s of AP. When P i c e a e x c e l s a Link. c u t t i n g s were
watered p e r i o d i c a l l y through t h e w i n t e r and s p r i n g
w i t h d e i o n i z e d w a t e r c o n t a i n i n g 100 ppm NaF, t h e
AP r a t e s were n o t o n l y reduced b u t n e c r o s i s was
produced on t h e newly f l u s h e d f o l i a g e . The F con-
c e n t r a t i o n s i n t h e new f o l i a g e t h a t e x h i b i t e d
i n j u r y contained o n l y 3.7 t o 8 ppm when i n j u r y
f i r s t o c c u r r e d . By t h e end of J u l y , t h o s e n e e d l e s
t h a t s u r v i v e d c o n t a i n e d from 31.5 t o 52.2 ppm F
( K e l l e r , 1980).
 McLaughlin and Barnes (.I9751 exposed c u t
b r a n c h l e t s of t h r e e p i n e s p e c i e s and l e a v e s of s i x
deciduous t r e e s t o 0 , 1 . 9 , 1 9 , and 190 pprn NaF f o r
24 hours and then measured t h e r a t e s of APT With
1 . 9 pprn NaF, t h e r a t e s of AP of o l d e r n e e d l e s of
Pinus t a e d a L. and
-- P. e c h i n a t a M i l l . were
reduced w h i l e t h e o t h e r s p e c i e s were u n a f f e c t e d
(s-ee Table 1 f o r s p e c i e s used). Needles
w i t h reduced r a t e s of AP contained l e s s than 1 0
pprn F. Although low c o n c e n t r a t i o n s of f o l i a r F
reduced AP and s t i m u l a t e d r e s p i r a t i o n , t h e a u t h o r s
warn of t h e l i m i t a t i o n s of e x t r a p o l a t i n g labora-
t o r y d a t a t o t h e f i e l d s i t u a t i o n . However, t h e i r
d a t a r a i s e d s e v e r a l q u e s t i o n s : (1) What concen-
t r a t i o n of HF would be necessary t o i n c r e a s e t h e
f o l i a r F c o n c e n t r a t i o n 4-8 pprn i n a 24-hour p e r i o d
a s d i d t h e comparable dose of NaF? (2) I f upon
exposure t o HF, a branch on a t r e e accumulated F
a t t h e same r a t e , would t h e r e d u c t i o n i n AP be
permanent o r would i t recover t o t h e pre-exposure
r a t e ' a f t e r t h e exposure? (3) Would t h e 4-8 pprn
increase i n f o l i a r F associated with the reduction
i n AP produce v i s i b l e i n j u r y ? One could view t h i s
kind of exposure a s a c u t e , because t h e comparable
dose of HF t o accumulate t h i s amount of F i n 24
hours could be 4-8 -ng m--, assuming an accumula-
t i o n c o e f f i c i e n t of 1 pprn p g l m3 daym1.
Respiration
R e s p i r a t i o n (measured a s oxygen uptake) was
s t i m u l a t e d i n i n t a c t p l a n t s (Applegate and Adams,
1960a; Applegate and o t h e r s , 1960) o r i n t i s s u e s
from i n t a c t p l a n t s fumigated w i t h HF, i n t h e
absence (Weinstein, 1961; Applegate and Adams,
1960b; Yu and M i l l e r , 1967; M i l l e r and M i l l e r ,
1974) o r presence of f o l i a r l e s i o n s ( H i l l and
o t h e r s , 1959). F l u o r i d e i n h i b i t i o n of oxygen
uptake h a s a l s o been r e p o r t e d and was dependent
on p l a n t t i s s u e age (Bejaoui and P i l e t , 1975)
d u r a t i o n of exposure (Applegate and Adams, 1960a),
n u t r i e n t s t a t u s (Applegate and Adams, 1960b), and
t i s s u e F c o n c e n t r a t i o n s (Applegate and o t h e r s ,
1960). However, t h e r a t e of r e s p i r a t i o n of some
t i s s u e s i s r e l a t i v e l y i n s e n s i t i v e t o F ( H i l l and
o t h e r s , 1959; Givan and Torrey, 1968). I n t h e i r
experiments w i t h c u t b r a n c h l e t s of p i n e s and hard-
woods s u p p l i e d w i t h 1 , 9 , 19 o r 190 pprn NaF i n
s o l u t i o n f o r 24 h o u r s , McLaughlin and Barnes (1975)
found t h a t t h e lower two c o n c e n t r a t i o n s g e n e r a l l y
s t i m u l a t e d r e s p i r a t i o n (measured a s C02 e v o l u t i o n )
while the highest concentration both stimulated
and i n h i b i t e d r e s p i r a t i o n , depending on t h e
species.
T r a n s p i r a t i o n and Water Use
There a r e few r e p o r t s on t h e e f f e c t s of F on
t r a n s p i r a t i o n . Navara (1963) grew beans i n
s o l u t i o n c u l t u r e s c o n t a i n i n g 0.03, 0.3, and 3.0
pprn F and found reduced r a t e s of t r a n s p i r a t i o n
a f t e r 12 and 16 days i n p l a n t s s u p p l i e d w i t h t h e
two h i g h e s t c o n c e n t r a t i o n s of F. Soybeans fumi-
g a t e d w i t h 12 -ng m-- HF had d r a m a t i c a l l y reduced
r a t e s of t r a n s p i r a t i o n w i t h i n 4 hours (Poovaiah
and Wiebe, 1973). This a g r e e s w i t h Navara and
Kozinda (1967) who found s i m i l a r r e s u l t s i n bean
and a p r i c o t exposed t o 70 pg mV3 HF. But Thompson
and o t h e r s (1967) d i d n o t f i n d s i g n i f i c a n t d i f f e r -
ences i n water use of c i t r u s exposed over a growing
season t o e i t h e r ambient l e v e l s of F o r f i l t e r e d
a i r w i t h added F (both < 0.5 v& m-3) compared t o
c o n t r o l p l a n t s . Amundson and o t h e r s ( i n review)
exposed corn t o 1 . 5 lie mF3 HF continuously f o r
one week and found an i n c r e a s e d r a t e of t r a n s p i r a -
t i o n over c o n t r o l s . The d a t a a v a i l a b l e on F
e f f e c t s on t r a n s p i r a t i o n a r e v a r i a b l e and i n s u f f i -
c i e n t t o p r e d i c t p o s s i b l e e f f e c t s on p l a n t comu-
n i t y water r e l a t i o n s . However, s i n c e F can e l i c i t
changes i n stomata1 a p e r t u r e , t h e s e e f f e c t s may b e
important i n a f o r e s t ecosystem where water
d e f i c i t s l i m i t AP a t c e r t a i n times of t h e day o r
y e a r (Larcher, 1975; Kramer and Kozlowski. 1979).
p l a n t ' Metabolism
F has long been used a s a metabolic i n h i b i t o r
and t h e l i s t of published r e p o r t s of F e f f e c t s on
enzyme systems and metabolic p r o c e s s e s i s e x t e n s i v e .
Many of t h e e f f e c t s of F on p l a n t metabolism have
been reviewed (McCune and Weinstein, 1971; Chang,
1975) and Horsman and Wellburn (1976) have com-
p i l e d a u s e f u l l i s t of F-induced metabolic re-
sponses.
Since F a l t e r s normal p l a n t metabolism,
e f f o r t s have been made t o i d e n t i f y m e t a b o l i t e s
t h a t could be used a s i n d i c a t o r s of i n c i p i e n t F
i n j u r y . Yee-Meiler (1975) found t h a t non-specific
e s t e r a s e a c t i v i t y i n young Norway s p r u c e (Picea
a b i e s [L.] K a r s t . ) and European w h i t e b i r c h
(Betula verrucosa Ehrh.) exposed t o a i r b o r n e F
was i n c r e a s e d l a t e i n t h e growing season without
t h e appearance of i n j u r y symptoms. Needles of
c o n i f e r s placed a t varying d i s t a n c e s from an i n -
d u s t r i a l F source had s i g n i f i c a n t i n c r e a s e s i n
phenols i f they came from t r e e s w i t h F i n j u r y
(Yee-Meiler, 1977). The r e s u l t s were v a r i a b l e
f o r deciduous t r e e s . K e l l e r and Schwager (1971)
found i n c r e a s e d peroxidase a c t i v i t y i n l e a v e s of
seven t r e e s p e c i e s exposed t o an i n d u s t r i a l source
of HF and noted t h a t t h e enzyme a c t i v i t y i n c r e a s e d
b e f o r e o r i n t h e absence of development of F
i n j u r y symptoms. Unfortunately, many environmental
s t r e s s e s and l a b o r a t o r y manipulations can i n c r e a s e
pei-oxidase a c t i v i t y , l i m i t i n g t h e u s e f u l n e s s of
t h i s a s s a y (Endress and o t h e r s , 1980).
Mineral N u t r i t i o n
Wide d i f f e r e n c e s i n t h e response of peaches
t o HF l e d t o t h e f i r s t s t u d y of t h e i n f l u e n c e of
mineral n u t r i t i o n on HF s u s c e p t i b i l i t y (Brennan
and o t h e r s , 1950). With low o r d e f i c i e n t amounts
of N , Ca, and P i n tomato f o l i a g e , t h e r e was re-
duced uptake of NaF by r o o t s o r HF by l e a v e s ;
s i m i l a r r e s u l t s were found w i t h e x c e s s i v e amounts
of N and Ca (Brennan and o t h e r s , 1950). Other
s t u d i e s have r e s u l t e d i n i n c r e a s e d f o l i a r F i n
P-, K-, o r Fe-deficient beans (Applegate and Adams,
1 9 6 0 ~ ) ;reduced f o l i a r F i n Mg-deficient tomato
p l a n t s (MacLean and o t h e r s , 1969); s m a l l e r f r u i t s
i n Ca-deficient tomato p l a n t s (Pack, 1966); and
i n c r e a s e d t o l e r a n c e t o HF exposure i n tomato p l a n t s
 Table 1. Reported e f f e c t s o f f l u o r i d e (HF and NaF) on a p p a r e n t p h o t o s y n t h e s i s (as measured by changes i n
C02 u p t a k e ) o f h i g h e r p l a n t s .

Genus o r S p e c i e s Concentration Duration Response Reference

p g m-3 HF

Gladiolus 0.8 - 8.0 7 days p e t . r e d u c t i o n i n AP = p e t . i n j u r y Thomas &

Hendricks 1956
Hordeum 32 4-8 h o u r s t o t a l i n t e r r u p t i o n i n AF' w i t h
Medicago 200 4-8 h o u r s r e c o v e r y i n few h o u r s t o days
Fruit trees 16-40 4-8 h o u r s

Hordeum 32 2 hours AP reduced d u r i n g exposure w i t h Bennett &

Medicago 32 2 hours r e c o v e r y a f t e r exposure Hill 1973

Lycopersicon 1.4 - 5.2 4 weeks no e f f e c t Hill &

0.9 -11.2 3 weeks no e f f e c t others 1959

F r u i t trees 2.1 av. 183 h o u r s 14 p e t . r e d u c t i o n i n AP 1 0 p e t . i n j u r y Thomas 1958

Gladiolus 3.1 - 5.2 30-205 days p e t . r e d u c t i o n i n AP = p e t . l e a f i n j u r y

Gossypium 13.6 138 h o u r s no e f f e c t

Citrus .32 -.77 growing s e a s o n no e f f e c t Thompson o t h e r s 1967

Gladiolus 0.8 39 days no effect Hill 1969

1.2 27 days 3 pet. reduction over i n j u r y
Fragaria 2.3 6 3 days no effect
38 1 day 50 pet. reduction
Lycopersicon 5.1112 1 7 / 2 1 days no effect
Prunus 1.6 42 days no effect
Zea 7.7 1 6 days no effect

Sorghum 0.7 1 4 days no e f f e c t McCune &

2.2 t h e n 1.7 1212 days reduced w i t h r e c o v e r y a f t e r exposure o t h e r s 1976
3.5 t h e n 5+ 717 days reduced d u r i n g 3.5 exposure t h e n
severely injured l i t t l e recovery

Pinus s y l v e s t r i s ambient Nov- A p r i l reduced AP of whole p l a n t due t o l o s s Keller 1977

P.
- nigra near o f f o l i a g e w i t h v A s i b l e i n j u r y on
P.
- strobus source remaining f o l i a g e
Larix l e p t o l e p i s
Quercus b o r e a l i s 1
Pseudotsuga m e n z i e s i i
Picea excelsa
Ainus -
- incana
Sorbus A r i a
--
I
Acer p s e u d o p l a t a n u s
--
L a r i x decidua.

Cornus f l o r i d a 1 1.9 ppm 24 h o u r s AP reduced i n o l d e r n e e d l e s of 2. McLaughlin &

Liquidambar S t y r a c i f l u a NaF t a e d a and P. e c h i n a t a
-- others 1975
Plantanus occidentalis
Acer rubrum
-- I 1 9 PPm 24 h o u r s AP reduced i n a l l s p e c i e s
Liriodendron t u l i p i f e r a NaF
O x y d e n d r s arboreurn
Pinus s t r o b u s 1 9 0 ppm 24 h o u r s AP reduced i n all s p e c i e s
P.
- taeda
P.
- echinata

Picea excelsa 100 ppm winter- spring AP reduced i n o l d f o l i a g e l n e w i n j u r e d K e l l e r 1980

grown with excess Mg (MacLean and others, 1976).
sources. Thirdly, the source strength is often

known and can be applied to dispersion modelling.

There is little information on the effects of

Fourthly, F is not very mobile in plants and

F on forest tree nutrition, but there is a con-

tends to accumulate along the margins and distal

siderable amount of information on mineral cycling

end of the leaf. Consequently, most of the F that

in forest ecosystems (Grier and Cole, 1972; Bormann

enters a leaf remains, except for that lost by

and Likens, 1979), and airborne F can influence

weathering and perhaps a small amount by trans-

this cycling in forest vegetation (see "Tree

location. But, as mentioned earlier, one major

Growth").
drawback is the difficulty of monitoring ambient

concentrations.

Growth and Production

Smith (1974) recognized three broad classes of

Effects of F on the physiology and metabolism

air pollutant-dose relationships with respect to

of plants are ultimately manifested as changes in

potential impacts on forest ecosystems. The

the height, diameter, dry weight, and reproduction

Class I relationship pertains to a very low dose

of the plant. But most of the available litera-

where the forest acts as a sink for the pollutant

ture describes studies with agronomic crops.

and the impact may be immeasurable or stimulatory.

A moderate dose relationship (Class 11) is ex-

Relatively low concentrations of F have been

pected to cause significant direct and indirect

reported to stimulate growth, but growth can be

physiological impairment to individuals resulting

inhibited by amounts of foliar F that do not pro-

in reduced growth, reproduction and/or increased

duce chlorosis or necrosis in the same species

morbidity. With a high dose (Class 111), there is

(Treshow and Harner, 1968).

acute morbidity resulting in ecosystem simplifica-

tion with drastic changes in primary productivity,

The effects of F on reproduction have been

mineral cycling, succession, etc.

demonstrated and its possible implications dis-

cussed by Pack and Sulzbach (1976). They hypoth-

All three pollutant-dose relationships have

esized that lowered seed production was a result

been described in one form or another around F

of inhibition of pollen germination or pollen tube

sources (Bunce, 1978; Treshow and others, 1967;

growth, inhibiting or preventing, fertilization.

Carlson and Dewey, 1971; Wheeler, 1972).

Growth of pollen tubes in apricot (Facteau and

Rowe, 1977) and sweet cherry (Facteau and others,

Fluoride Accumulation in Soils

1973) was reduced by HF fumigation during flower-

ing, but Dinh and others (1973) found no effect on

The amount of total F in soils that has been

sweet cherry pollen tube growth after exposure to

reported ranges up to 8300 ppm but is generally

97 pg F m-3.
from 20-500 ppm (Weinstein, 1979). In general,

plants are poor accumulators of soil F (Hansen,

Joint Action with Other Pollutants

1958; MacIntire and others, 1949; Merriman and

Hobbs, 1962; McClenahen, 1976), but there are

Experiments on the joint action of HF with

some exceptions, notably species of Theaceae,

other pollutants have emphasized effects on F

such as tea and camellia (Zimmerman and others,

accumulation (Matsushima and Brewer, 1972; Mandl

1957; Zimmerman and Hitchcock, 1956), hickories

and others, 1975, 1980), foliar lesions (Solberg

and flowering dogwood (McClenahen, 1976).

and Adams, 1956; Hitchcock and others, 1962;

Mandl and others, 1975, 1980); and growth and

The deposition of fluoride in soils near sources

yield (Hatsushima and Brewer, 1972; Mandl and

of emission has been the subject of several in-

others, 1980). Field studies that attempt to

vestigations. McClenahen (1976) examined the geo-

determine the response of plants or plant connnuni-

graphic distribution of total F in soils at two

ties to F emissions must consider not only environ-

seasons and at different distances from an alumina

mental and edaphic factors (Treshow and others,

reduction smelter. Of course, the highest accumu-

1967), but also the presence of other pollutants

lations occurred in the direction of the prevail-

(Bunce, 1978; McClenahen, 1978; Carlson, 1978)

ing winds and extended about 10 km. In areas

that complicate assessment of the impact of F

where F deposition was lowest, total F increased

alone. McCune (1980) has discussed published and

with depth of the soil profile, but the opposite

unpublished results of experiments with HF in

was true in areas where deposition was heaviest.

combination with S02, 03, and N02.

There was a lower concentration of F in the soil

profile in outlying areas than near the source.

The total F in the soil profile in low and high

FIELD STUDIES
impact areas over the two-year study period was

consistently different. No attempt was made to

F has many characteristics that make it an

correlate soil F with the amount of F accumulated

ideal toxicant to study in an ecosystem. Firstly,

by plants.

it is an apparently non-essential element that

normally occurs in foliar tissues at a concentra-

Relatively large amounts of F-containing

tion of <10 ppm; thus, the presence and amount of

amendments are necessary to increase the accumula-

airborne contamination can be measured. Secondly,

tion of 7 in plants (Weinstein, 1977), and Israel

F is easily identified with specific emission

(1974) has estimated that each 120 vg/g increment

i n s o i l F r e s u l t e d i n a gain i n f o r a g e F of 1 ?~g/g.
The accumulation of F i n c o n i f e r n e e d l e s and i n
l'soil-humus" samples n e a r a phosphorus p l a n t i n
Canada h a s been s t u d i e d (Thompson and o t h e r s , 1979;
Sidhu, 1977). S e v e r i t y of damage t o v e g e t a t i o n was
r e p o r t e d t o be c o r r e l a t e d w i t h F c o n c e n t r a t i o n of
f o l i a g e and of "soil-humus". Because t h e d i s t r i -
b u t i o n of a i r b o r n e F i n s o i l s would be expected t o
follow t h e same p a t t e r n a s i n v e g e t a t i o n , i t would
be d i f f i c u l t t o e s t i m a t e t h e p r o p o r t i o n of F pres-
e n t i n f o l i a g e t h a t was accumulated from t h e atmos-
phere and t h a t from t h e s o i l . Water-soluble F from
t h e "soil-humus" was p o s i t i v e l y c o r r e l a t e d w i t h
f o l i a r F and because t h e s o i l s were h i g h l y a c i d i c ,
r o o t uptake could have been an important pathway
i n t o t h e vegetation.
The s o i l a s a s o u r c e of F t o p l a n t s has n o t been
adequately i n v e s t i g a t e d and t h e long-term e f f e c t
of a c i d i c p r e c i p i t a t i o n i n making s o i l F a v a i l a b l e
t o p l a n t s , especially i n a c i d i c , non-agricultural
s o i l s should be i n v e s t i g a t e d . There a r e many
o t h e r gaps i n our understanding of t h e c y c l i n g
of F i n f o r e s t ecosystems, such a s t h e e f f e c t s of
F accumulation on l i t t e r decomposition,on changes
i n n u t r i e n t a v a i l a b i l i t y , and on s o i l s t r u c t u r e .
F Accumulation and Occurrence of I n j u r y
I n many r e p o r t s , t h e a u t h o r s have p r e s e n t e d
v a l u e s f o r t h e F c o n t e n t of v e g e t a t i o n a t d i f f e r -
e n t d i s t a n c e s (and sometimes, d i r e c t i o n s ) from a
s o u r c e , b u t o f t e n they d i d n o t provide information
on t h e s o u r c e s t r e n g t h , ambient a i r c o n c e n t r a t i o n ,
o r t h e forms of a i r b o r n e F t h a t were p r e s e n t .
Often, q u a l i t a t i v e o r s e m i - q u a n t i t a t i v e e s t i m a t e s
of i n j u r y a r e given and t h e r e i s l i t t l e o r no con-
s i d e r a t i o n t o o t h e r p o s s i b l e causes of i n j u r y ,
such a s i n s e c t s , pathogens, environmental s t r e s s e s ,
o r even t h e presence of o t h e r p o l l u t a n t s . These
d a t a a r e most u s e f u l i n e v a l u a t i n g t h e r e l a t i v e
s e n s i t i v i t y of d i f f e r e n t s p e c i e s , t h e i n t e r s p e c i f i c
d i f f e r e n c e s i n s e n s i t i v i t y , t h e most s e n s i t i v e
s t a g e s of p l a n t o r f o l i a r development, t h e compo-
n e n t s of t h e f o r e s t ecosystem most v u l n e r a b l e t o
an e f f e c t , and, i f e v a l u a t e d c a r e f u l l y , t h e dose
of atmospheric F o r t h e amount of t i s s u e F
accumulated t o produce a measurable e f f e c t , whether
i t i s r e g i s t e r e d a s a metabolic o r p h y s i o l o g i c
change o r a s a c h l o r o t i c o r n e c r o t i c l e s i o n .
As one would e x p e c t , t h e atmospheric concen-
t r a t i o n of F and t h e amount accumulated i n vegeta-
t i o n d e c r e a s e s w i t h d i s t a n c e from t h e s o u r c e
(Treshow and o t h e r s , 1967; Sidhu, 1977, 1978;
Thompson and o t h e r s , 1979; Roberts and o t h e r s ,
1979; Bunce, 1978, 1979; Wheeler, 1972; Carlson
and Dewey, 1971). The amount of F accumulated i n
f o l i a g e , however, w i l l depend upon many f a c t o r s
i n c l u d i n g t h e dose and form of F, t h e s p e c i e s ,
a c c e s s i b i l i t y of t h e p o l l u t a n t t o t h e p l a n t (e.g.,
s c r e e n i n g of u n d e r s t o r y by o v e r s t o r y s p e c i e s ) ,
plant-to-plant v a r i a b i l i t y , e t c . I n g e n e r a l ,
b r o a d l e a f s p e c i e s w i l l accumulate more F than
c o n i f e r s when they occur t o g e t h e r (Sidhu, 1977,
1978); and g r e a t d i f f e r e n c e s can occur between
c o n i f e r s p e c i e s , c e t e r u s parabus, w i t h t h e most
t o l e r a n t ones accumulating t h e most F (Weinstein,
1 9 7 7 ) . A l i k e l y e x p l a n a t i o n f o r t h i s is t h a t when
t h e most s e n s i t i v e s p e c i e s a r e i n j u r e d (metaboli-
c a l l y o r p h y s i o l o g i c a l l y ) by a given dose of F,
continued a b s o r p t i o n and accumulation a r e reduced.
Not only a r e t h e most t o l e r a n t s ~ e c i e st h e most
e f f i c i e n t accumulators, b u t t h e amount of accumula-
t i o n and t h e t h r e s h o l d f o r i n j u r y w i t h i n a genus
( o r even s p e c i e s ) may b e v a s t l y d i f f e r e n t i n
d i f f e r e n t f o r e s t ecosystems. For example, Treshow
and o t h e r s (1967) d i d n o t f i n d n e e d l e i n j u r y i n
Douglas-fir (Pseudotsuga m e n z i e s i i [Mirb.] Franco)
i n Idaho a t F c o n c e n t r a t i o n s i n n e e d l e s t h a t
averaged 150 ppm (composite v a l u e f o r t h e c u r r e n t
y e a r , 2-year-old, and 3-year-old n e e d l e s ) , w h i l e
Carlson and o t h e r s (1979) r e p o r t e d t h a t "mottling
o r c h l o r o s i s was p r e s e n t a t 6-8 ppm" ( n e e d l e s of
u n c e r t a i n a g e ) . Obviously, t h i s d i f f e r e n c e is
r e f l e c t e d i n p a r t by t h e environmental d i f f e r e n c e s
between t h e s t u d y a r e a s (Idaho and Montana) b u t
o t h e r d i f f e r e n c e s between t h e r e s u l t s of t h e two
s t u d i e s should be considered. F i r s t l y , Carlson
and o t h e r s (1979) analyzed a number of n e e d l e
c h a r a c t e r i s t i c s a t d i f f e r e n t d i s t a n c e s from t h e
source of f l u o r i d e e m i s s i o n s i n Montana. The most
common n e e d l e i n j u r y observed was m o t t l i n g (pre-
sumably c h l o r o t i c m o t t l i n g ) and, although s i g n i f i -
c a n t i t was a weak a s s o c i a t i o n t h a t d i d n o t corre-
l a t e very c l o s e l y w i t h F c o n t e n t of n e e d l e s . The
~2 v a l u e s f o r F c o n t e n t of n e e d l e s and n e e d l e
m o t t l e i n Douglas-fir, lodgepole p i n e (Pinus -- -
c o n t o r t a v a r . l a t i f o l i a Engelm.) and w h i t e p i n e
(Pinus monticola Lamb.) were 0.0266, 0.0445, and
0.1604, r e s p e c t i v e l y (Carlson, 1980). But one
might conclude from t h e o r i g i n a l r e p o r t t h a t t h e
occurrence of m o t t l i n g on t h e s e c o n i f e r n e e d l e s
was i n c r e a s e d g r e a t l y by F. The p o s s i b i l i t y t h a t
o t h e r p o l l u t a n t s a s s o c i a t e d w i t h t h i s symptom, such
a s ozone o r another o x i d a n t , would be d i s t r i b u t e d
i n t h e same a i r mass a s F was n o t considered.
Carlson and o t h e r s (1979) a l s o concluded t h a t t h e r e
i s no t h r e s h o l d c o n t e n t of F i n n e e d l e s above
which i n j u r y w i l l occur, and t h a t any d e t e c t a b l e
amount of atmospheric F i s d e t r i m e n t a l t o c o n i f e r s .
This i s a s i m p l i s t i c e x p l a n a t i o n and i t i g n o r e s
s e v e r a l f a c t s . The f i r s t i s t h a t a l l c o n i f e r
needles a r e not equally s e n s i t i v e t o F injury, a s
i s noted i n many compilations ( e . g . , Weinstein,
1977; F l u o r i d e s , 1971; Thomas and A l t h e r , 1966).
Secondly, s e n s i t i v i t y t o F i s r e l a t e d t o t h e age
of t h e n e e d l e a t t h e time of exposure. It would
b e absurd t o a s s e r t t h a t "adverse e f f e c t s were
v i s i b l e o n n e e d l e s when t h e i r f l u o r i d e c o n c e n t r a t i o n
reached 8-10 ppm" (Carlson and o t h e r s , 1979) of t h e
F accumulated a f t e r t h e c o n i f e r n e e d l e s had com-
p l e t e d t h e i r e l o n g a t i o n . T h i r d l y , t h e form of F
t o which n e e d l e s a r e exposed and whether i t i s
i n t e r n a l o r s u p e r f i c i a l would a l s o determine t h e
kind of e f f e c t produced. F i n a l l y , i f t h e r e i s no
t h r e s h o l d f o r i n j u r y then t h e r e a r e no mechanisms
of d e t o x i f i c a t i o n i n p l a n t s and p h y s i o l o g i c o r
metabolic p r o c e s s e s , such a s p h o t o s y n t h e s i s o r
enzyme a c t i v i t y , t h a t have been a l t e r e d by I? should
evidence no recovery. There i s ample proof t h a t
once a fumigation c e a s e s , o r i f t h e periodsbetween
successive fumigations are sufficiently separated,
other objective criteria related to the intended

recovery processes (repair mechanisms) are active

use of the tree.

(see Dinman, 1972; McCune and Weinstein, 1971;

Thomas and Alther, 1966). Carlson and others

There is insufficient information to develop

(1979) appear to have equated injury from F with

these kinds of lists because existing compilations

such destructive agents as ionizing radiation.

are based primarily on field and laboratory observa-

tions of foliar injury. Sensitivity lists based on

Another interesting contrast in the sensitivity

foliar injury (Weinstein 1977, 1979) are only a

of conifers to F is exemplified by the conditions

guide iind do not provide evidence of relative

near a phosphorus plant at Long Harbour, Newfound-

effects on plant processes.

land and an alumina reduction smelter at Kitimat,

B.C. The Long Harbour area is classified as

Tree Growth

belonging to the Boreal Forest. Its productive

forests are 8-12 m tall and are composed princi-

Many studies have identified F as the cause of

pally of dense stands of balsam fir (Abies

tree mortality around industrial sources (Adams

balsamea [L.] Mill.) and black spruce (,Picea

and others, 1952; Scurfield, 1960; Jung, 1968;

mariana [Mill.] B.S.P.). Non-productive scrub

Robak, 1969). The determination of F as the causal

forests are less than 5 m tall and are composed

agent usually entailed determination of foliar F

of larch (Larix laricina IDuRoi] K. Koch), black

concentrations and, occasionally, air quality moni-

spruce, and balsam fir. Rock-barrens and peat-

toring. In these class 111 relationships (Smith,

lands are common (Thompson and others, 1979).

1974), determination of the area of impact is

Soils are generally shallow, precipitation is

normally easy to identify. This is not true for

heavy, and the forests are exposed to high winds

class I1 relationships for several reasons: (1)

containing saline aerosols.

environmental factors (mainly weather patterns)

change from year to year and not only distribute

Kitimat is in the Pacific Coastal Rain Forest

the pollutants randomly but also produce more or

area. The forest is an uneven-aged, overmature,

less favorable growing conditions for the impacted

decadent, and stable climax forest. Logging is

vegetation; (2) normal biotic factors (insects and

an important commercial activity. The forest

pathogens) and abiotic factors (soils) also

consists of about 60% western hemlock (Tsuga

account for variability in growth; (3) stage of

heterophylla [Raf.] Sarg.), 25% balsam fir (Abies

development of the stands of trees also dictate

amabilis [Dougl.] Forb.), 7% western red cedar

growth rates and the degree of competition between

(Thuja plicata Lamb.), 6% yellow cedar (Chamaecy-

individuals; and (4) all of these factors combined

paris nootkatensis [Lamb.] Spach.), and 2% sitka

with pollutant exposure produce a given effect.

spruce (Picea sitchensis [Bong.] Carr.). The

Therefore, to quantify the reduction in growth

average age of fir and hemlock is more than 300

caused solely by the pollutant, the variability

years, and it is not unusual to observe trees of

due to the other parameters must be accounted for.

1 or 2 m dbh. Total annual precipitation is about

115 inches and occurs on 53% of the days. The

Treshow and others (1967a) measured radial

site lies in a wide trough that runs north and

growthyneedlelength, needle dry weight, and foliar

south, and bisects the Coast Mountains (Reid,

fluoride concentrations in Douglas-fir (Pseudot-

Collins, 1976).
menziesii [Mirb.] Franco) located at differ-

ent distances from a fluoride source (also see

Emissions at the Long Harbour phosphorus plant

'F Accumulation and Occurrence of Injury"). They

are not known, but are certainly lower than those

classified the study plots into three groups based

at Kitimat, which have ranged from 2.5 to 6.6 tons

on foliar F concentrations of composite samples

of gaseous F/day between 1955 and 1977 (Alcan

o f four age-classes of needles. The groups were
Surveillance Committee, 1979). F concentrations
cpntrol (average 24 ppm F), intermediate fluoride

in conifers were frequently higher than 100 ppm

(average 150 ppm F), and high fluoride (average

in late summer without evidence of any foliar

225 ppm and with some needle necrosis). Signifi-

lesions. This can be contrasted to the published

cant reductions in radial growth were found in

threshold value for needle injury in balsam fir

both groups subjected to elevated F concentration

at Long Harbour of as low as 14 ppm (Sidhu, 1978).

There was a significant negative correlation of

Although the same species do not occur in the two

needle length with radial growth, but there were

areas, the different responses of conifers are so

no significant effects on needle dry weights.

striking that one must conclude that (1) it is

Thus, they found that (1) radial growth reduction

not possible to generalize from one site to another,

can occur without foliar lesions and (2) Douglas-

(2) foliar F contents alone may be a poor deter-

fir needles can average as much as 150 ppm F

minant of injury, and (3) environmental stresses

without foliar injury.

(such as wind, salt, nutrient, or water) are as

important predictors of an effect as is F content..

A study of the impact of F on nutrient cycling

in stands of loblolly (Pinus taeda L.) and slash

It is difficult to classify conifers and other

pines (Pinus elliotti Engelm.) and the impact on

tree species into groups based upon their relative

tree growth was made by Wheeler (1972). No

tolerance to airborne F because most compilations

injury symptoms attributable to F exposure were

are based upon foliar injury, and not according

noted in the sample plots but trees at the edge

to effects on timber volume, fruit production, or

of some stands did show some "burning of needles".

Increased foliar F concentrations (from 13 to 49

assumption, which implies that'^thereg no thres-

ppm in pooled samples) were correlated with in-

hold concentration of foliar F below which injury

creased return rates of Ca and K from greater

does not occur, cannot be substantiated. This

leaf leaching and of Ca, K, and Mg by greater

is not meant to imply that F is not phytotoxic,

litterfall. This altered nutrient cycling pre-

because it is the most toxic of the common atmos-

sumably should alter productivity but no relation-

pheric pollutants. But an understanding of its

ship was found between needle concentrations of

effects in the ecosystem, requires much research

F, Ca, Mg, and K and productivity as measured by

and the synthesis of an enormous amount of

amount of bole wood. Wheeler (1972) concluded

information. ~och'sPostulates were not written
that either these fluctuations in needle status
frivolously.
did not affect growth or that the sampling was

insufficient to detect differences that were

Community Structure
present.

Large areas of the Eastern United States are

Extensive studies on growth and F accumulation

subjected to a complex mixture of air pollutants

have been made at Kitimat, B.C. (Bunce, 1978) and

from urban centers and industrial sources. Most

Columbia Falls, MT (Carlson, 1978). Both areas

of the Eastern Deciduous Forests are subjected

were subjected to F for many years before any

to at least moderate air pollution (Class 11).

scientific assessment of growth reduction due to

McClenahen (1978) studied the effects of a mix-

F were made and each area was subjected to insect

ture of pollutants derived from industrial sources

infestation (see section "Insects"),

(containing F, SO?, NOx, chloride, and oxidant)

on changes in structure and composition of a mixed

Bunce (1978) used "foliage analysis, observa-

deciduous forest in the Ohio River Valley. The

tions of lichens, air flow patterns and topographic

study sites were arbitrarily divided into over-

features" to estimate the area of impact and to

story, subcanopy, shrub and herb layers and the

establish the distribution of his sample plots.

stands were measured for diversity (Pielou, 1975),

Tree ring cores were taken from western hemlock,

evenness (Williams, 1977), and species composition.

the dominant species, from all sample locations,

and were used to determine the amount of growth

In genera1,the average total stand densities

reduction due to F emissions. After variability

of the overstory and herb layers were found to

in growth rates due to weather, insect infesta-

decrease in proximity to the F source while the

tion, and another pollutant (SO2) were accounted

subcanopy and,shrub layer increased with the shrub

for, Bunce (1978) reported the annual loss of

layer being the only layer thatshoweda significant

wood production to be 950 cunits (95,000 cu. ft.)

correlation to F exposures. Chloride from another

per year compared to the 800,000 cunits attributed

source had a greater influence in the other layers.

to insect damage. Obviously the cause of the

insect outbreak is fundamental to the assessment

Murray (1979) conducted a study of plant comun-

of the magnitude of the F-related effects on

ity structure around an aluminum smelter in

growth and is discussed elsewhere. Although the

Australia. Although a number of study sites were

primary and secondary (bark beetles) insect out-

lost by fire, he was able to ordinate species

breaks ended by 1968, F emissions have continued

associations with F stress. More of these kinds

at a lower rate since 1975 and trees in the insect

of studies are needed to provide data to predict

damaged zone are regenerating satisfactorily. The

the risk of an effect when an ecosystem is exposed

question ofwhether there is a cause-and-effect

to airborne F.

relationship between F emissions and insect in-

festation has not yet been answered for reasons

Incidence and Severity of Diseases and Insects

discussed elsewhere.

There is evidence, from laboratory and field

F from an aluminum reduction plant in Columbia

experiments or observations, that airborne F may

Falls, MT caused growth reductions in Douglas-fir

alter the plant-pathogen and plant-insect rela-

fir, lodgepole pine, and western pine (Carlson,

tionships. The exact relationships between F and

1978, 1979). However due to questionable assump-

these biotic stresses and their underlying mech-

tions and miscalculations, an overestimate of the

anisms are only beginning to be understood.

loss of usable timber due to F pollution was made.

Statistically, the data (Carlson, 1978) showed

Plant pathogens-- Although there are many in-
only a weak correlation between foliar F concen-
dustrial sources of F, we are not aware of any

trations and reduced radial growth. The area of

field or laboratory reports that link airborne F

growth reduction was substantially smaller than

with incidence or severity of forest tree diseases.

reported previously (Carlson, 1980). It has also

It is necessary, therefore, to discuss some labora-

been stated (Carlson, 1978) that any increase in

tory research on the effects of HF on diseases of

foliar F above background concentrations is

crop plants in order to evaluate possible forestry

detrimental to tree growth. This assumption was

effects and to establish research needs.

generated by the implied growth reduction of trees

located in areas designated as having reduced

The plant-pollutant-pathogen interaction was

growth; but upon closer examination of the data,

reviewed by Heagle in 1973 and Laurence in 1978.

no growth reduction could be demonstrated

For the kinds of effects that have been found,

(Carlson, 1980). Consequently, the original

McCune and others (1973) provided three possible

e x p l a n a t i o n s : (1) t h e r e could be a d i r e c t e f f e c t
of t h e p o l l u t a n t on growth and development of t h e
organism; (2) t h e p o l l u t a n t could a f f e c t t h e sus-
c e p t i b i l i t y of t h e p l a n t t o t h e pathogen; and (3)
t h e p o l l u t a n t could a f f e c t t h e microbiota o r micro-
environment of p l a n t s u r f a c e s and thereby a f f e c t
t h e pathogen.
Tobacco l e a v e s i n f e c t e d with tobacco mosaic
v i r u s and c o n t a i n i n g 200-300 ppm F had a higher
t i t e r of v i r u s than c o n t r o l l e a v e s when a l o c a l
l e s i o n assay was used. The t i t e r was lower a t 500
ppm F (Dean and Treshow, 1966; Treshow and o t h e r s ,
1967b). But perhaps t h e b e s t evidence f o r a
d i r e c t e f f e c t of a i r b o r n e F on growth and develop-
ment of a pathogen was t h e c o n s i s t e n t reduction
i n bean powdery mildew (Erysiphe polygoni DC.)
found a s a r e s u l t of HF fumigation, i n d i c a t i n g
t h a t HF was a f f e c t i n g t h e i n f e c t i v i t y of t h e patho-
gen i t s e l f , because reduction i n d i s e a s e was pro-
p o r t i o n a l t o t h e l e n g t h of t h e exposure p e r i o d ,
i n f e c t i o n was continuous throughout t h e exposure
p e r i o d , and t h e pathogen i t s e l f i s e p i p h y t i c .
The most l i k e l y mechanism f o r an e f f e c t of F on
p l a n t pathogenic d i s e a s e s would be an a l t e r a t i o n
i n t h e s u s c e p t i b i l i t y of t h e h o s t p l a n t t o t h e
pathogen. The r e d u c t i o n i n t h e numbers of bean
r u s t (Uromyces p h a s e o l i [ P e r s . ] Wint.) u r e d i a by
pre- and post-inoculation exposures t o HF may have
been due t o a change i n h o s t metabolism by t h e
accumulation of F (McCune and o t h e r s , 1973). The
b e s t evidence a v a i l a b l e t h a t s u g g e s t s an i n d i r e c t
e f f e c t of F was found i n halo-blight of bean
(Pseudomonas p h a s e o l i c o l u s [Burkh. ] Dows ) where .
stem c o l l a p s e was a f f e c t e d , b u t f o l i a r symptoms
were n o t . Thus, t h e s i t e a f f e c t e d was s p a t i a l l y
removed from t h e s i t e of F accumulation, t h e l e a f
(McCune and o t h e r s , 1973).
There is no reason t o b e l i e v e t h a t crop p l a n t s
should respond d i f f e r e n t l y than f o r e s t s p e c i e s t o
a i r b o r n e F and p l a n t pathogens, and l a b o r a t o r y and
f i e l d s t u d i e s a r e needed t o determine and e v a l u a t e
e f f e c t s on t h e incidence of d i s e a s e and p o s s i b l e
epidemiological consequences.
Insects -- The c o n t r o v e r s i e s a s s o c i a t e d w i t h
t h e e f f e c t s of F on p l a n t s i n g e n e r a l , and eco-
systems i n p a r t i c u l a r , a l s o extend t o t h e p o s s i b i l -
i t y t h a t F a l t e r s t h e r e l a t i o n s h i p between p l a n t s
and d e s t r u c t i v e i n s e c t s , t h a t F k i l l s b e n e f i c i a l
i n s e c t s , o r t h a t accumulation of F i n i n s e c t s makes
them a v e h i c l e f o r t h e t r a n s f e r of F i n ecosystems.
There i s ample evidence t h a t an a s s o c i a t i o n can
e x i s t betwen F-contaminated y e g e t a t i o n and i n s e c t s ,
but t h e r e l a t i o n s h i p i s n o t understood and i t does
n o t occur under a l l c o n d i t i o n s o r with a l l i n s e c t s .
P f e f f e r (1962-1963) r e p o r t e d t h a t a t t a c k by
bark b e e t l e s , snout b e e t l e s , and f i r l e a f r o l l e r s
were a s s o c i a t e d w i t h F emissions i n a f i r f o r e s t
i n Czechoslovakia. Carlson and Dewey (1971) and
Carlson and o t h e r s (1974) have r e p o r t e d t h a t F
accumulation i n c o n i f e r f o l i a g e i s c l o s e l y r e l a t e d
t o i n f e s t a t i o n s by s e v e r a l d e s t r u c t i v e i n s e c t s :
p i n e needle s c a l e (Phenacapsia p i n i f o l i a e F i t c h ) ,
pine needle s h e a t h miner ( Z e l l a r i a haimbachi
Busck), needle miner (Ocnerostyma strobivorum
[ Z e l l e r ] ) , and sugar p i n e t o r t r i x (Choristoneura
lambertiana [Busck] ) t h a t ranged from no s i g n i -
f i c a n c e ( l a r c h casebearer) t o a non-significant
trend (pine needle s c a l e ) t o s t r o n g evidence of a
weak c o r r e l a t i o n (needle miners). Only about 6%
of t h e v a r i a t i o n i n needle damage by needle miners
was a s s o c i a t e d with f o l i a r F concentration. There
was an even more remote a s s o c i a t i o n between needle
miner population and f o l i a r F concentration.
Edmunds and Allen (1956) and Compton and o t h e r s
(1961) found no a s s o c i a t i o n between ~ i n eneedle
& a l e (Nuculopsis c a l i f o r n i c a [ ~ o l e m k ] )and t h e
-
e x t e n t of F i n j u r y o r F content of needles of
7
ponderosa p i n e (Pinus ponderosa Laws) and Edmunds
(1973) questioned t h e r e s u l t s of Carlson and Dewey
(1971). Thalenhorst (1974) and Wentzel (1965)
found p o s i t i v e r e l a t i o n s h i p s between spruce g a l l s
induced by Adelges a b i e t e s (L.) and F. But Temple
(personal communication) could f i n d no c o r r e l a t i o n
between t h e F content of washed s i l v e r maple f o l i -
age and g a l l s induced by t h e bladder-gall mite
(Vasates quadripes [deshimer]).
One of t h e most i n t e r e s t i n g examples of a possi-
b l e F-plant-insect a s s o c i a t i o n was observed n e a r
an alumina reduction s m e l t e r i n Kitimat, B.C. ( s e e
s e c t i o n on "F Accumulation i n P l a n t s and Occurrence
of Injury" and "Tree Growth"). Between 1960 and
1963, an e p i z o o t i c of saddleback l o o p e r s ( E c t r o p i s
c r e p u s c u l a r i a [Denis & S c h i f f . ] ) and spruce bud-
worms ( C h o r i s t m e u r a 9Free.) occurred t h a t
k i l l e d many t r e e s over a l a r g e a r e a t h a t coincided
w e l l w i t h t h e p a t t e r n o f f u m e d i s p e r s i o n . I n 1961,
balsam bark b e e t l e s (Pseudohylesinus g r a n d i s
[Swaine] and P. nebulosus [Lee.]) appeared a s
secondary p e s t s throughout t h e a r e a a t t a c k e d by
t h e looper and t h e budworm. We used t h e word
' p o s s i b l e " above i n r e f e r r i n g t o F as t h e c a u s a l
agent i n t h i s outbreak because (1) t h e r e i s no way
now t o e s t a b l i s h a cause-and-effect r e l a t i o n s h i p ;
(2) t h e emissions were a l s o high i n p a r t i c u l a t e
m a t e r i a l s , s u l f u r compounds, p i t c h v o l a t i l e s , and
even CO2; (3) t h e problem was n o t s t u d i e d a t t h e
time t h a t t h e outbreaks occurred; and (4) o t h e r
p o s s i b l e e t i o l o g i e s have been suggested by entomol-
o g i s t s from t h e Canadian F o r e s t r y Service. Several
of t h e t h e o r i e s t h a t might e x p l a i n t h e i n s e c t
a t t a c k s a t Kitimat a r e : (1) F absorbed by t h e
f o l i a g e of t h e t r e e a l t e r s i t s metabolism and in-
c r e a s e s i t s a t t r a c t i v e n e s s t o i n s e c t s ; (2) F weakens
t h e t r e e , rendering i t l e s s a b l e t o r e s i s t i n s e c t
a t t a c k ; (3) gaseous o r p a r t i c u l a r emissions a r e
t o x i c t o p a r a s i t i c and/or predaceous i n s e c t s t h a t
provide important c o n t r o l s of t h e population of
d e s t r u c t i v e i n s e c t s ; (4) t h e emissions have a
"blanket" e f f e c t t h a t r e s u l t s i n a s l i g h t tempera-
t u r e a l t e r a t i o n and gives t h e l a r v a e of t h e l o o p e r s
and budworm a competitive advantage over p a r a s i t e s
and p r e d a t o r s ; (5) loopers and budworm moths were
c a r r i e d on winds i n t o t h e Kitimat a r e a and dis-
persed i n t h e same p a t t e r n a s s m e l t e r emissions;
and (6) t h e r e were an unusually l a r g e number of
l i g h t s i n t h e v a l l e y above t h e smelter i n t h e
e a r l y 1960's and they provided l i g h t of wavelengths
t h a t a t t r a c t e d moths (Alcan S u r v e i l l a n c e Committee,
 Regardless of the approaches used, the path to

useful information can be a difficult and confusing

Although the primary and secondary insect

one, as is well-known for other atmospheric pollu-

attacks were extremely destructive, amounting to

tants. Some of these problems have been discussed

a total net loss of mature timber estimated at

by Weinstein and McCune (1970).

800,000 cunits (80,OOQOOO cu. ft.) (Reid, Collins,

1976) a considerable number of trees were not

There are a number of indigenous plant species

damaged and regrowth has been extensive. In some

that are sensitive to atmospheric F, including goat-

areas near the smelter, F-induced injury was a

weed (Hypericum perforaturn L.), common barberry

prominent feature on young hemlock, Sitka spruce,

(Berberis vulgaris L.), Oregon grape (Mahonia repens

black cottonwood, and even western red cedar

[Lindl.] Don. and g. nervosa [Pursh.] Nutt.), blue-

(Weinstein, unpublished field reports for 1971

berry (Vaccinium spp.), and young needles of many

and 1974 cited in Alcan Surveillance Committee,

conifers (see Table 2 for a list of F-sensitive

1979). In the intervening years, especially since

higher plants). One general conclusion can be made:

1975, there has been a substantial reduction in

field observations of plants can be a good qualita-

total emissions from the smelter (more than 50%

tive but is usually a poor quantitative indicator

between 1975 and 1977), accompanied by greatly

of effects.

reduced foliar injury. Nevertheless, vegetation

exhibiting no foliar symptoms often contains 100

In many cases of F pollution, there has been a

ppm F or more, and the incidence of insect attack

severe depletion of lichen populations (reviewed

is no-greater than in nearby areas not exposed

by Gilbert, 1973). In areas nearest the F source,

to the smelter emissions. Because any reduction

a lichen desert may exist, but they appear and in-

in emissions would include gaseous F, particles,

crease in frequency and diversity with increasing

and other components of the fumes, no cause- and

distance from the source (LeBlanc and others, 1972).

effect-relationship can be made. From subjective

Nash (1971) and LeBlanc and others (1971) trans-

observations, however, particulate emissions have

planted several species of lichens into the field

been reduced strikingly, at least since 1971, and

in areas of F-emitting industries and found that

we feel that this fraction of the emissions was

the species used were injured near the source (but

perhaps of great significance in the original out-

sometimes up to 10 km away) and were effective F

breaks. Certainly, the indirect relationship be-

accumulators. Corticolous lichens accumulate F

tween an increase in insect colonization and

more rapidly than saxicolous species, and consequent-

particles has been known for many years (Bartlett,

ly, demonstrate accelerated damage and reduced

1951). Even before Carlson and his colleagues

abundance (Perkins and others, 1980), but much

were attempting to demonstrate a relationship

research remains to characterize and classify the

between F and insects on U.S. Forest Service land

sensitivity of the different lichen types growing

and in Glacier National Park, an enormous outbreak

on their many kinds of habitats.

of mountain pine bark beetle (Dendroctonus

ponderosae [Hopkins]) was beginning on the Canadian

Problems associated with the measurement of

border many km to the north. It extended through-

biomass have been discussed in many treatises on

out the entire Flathead National Forest and the

forest mensuration, and Bunce (1978, 1979) and

western part of Glacier National Park, and has

Parker and others (1974) have discussed the problems

destroyed many thousands of lodgepole, ponderosa,

associated with discriminating between effects of F,

and white pines.

insects, and environmental stresses in evaluating

effects on tree growth. The relationship between

Although there is little doubt that an associa-

F accumulation and production of foliar lesions or

tion exists between airborne contaminants and

other effects, is discussed elsewhere, and fluorosis

insect outbreaks, the evidence for a cause-and-

in indigenous herbivores is beyond the scope of

effect relationship with F in unconvincing, and

this review.

at times it appears that some investigators have

forced a relationship. Insect outbreaks occur in

F Distribution in the Environment

unpolluted as well as polluted areas. The question

to be resolved is not whether there is or isn't a

In its most elementary form, the transfer of

relationship between airborne substances and in-

F to and from the atmosphere, waters, soils and

sects, but to determine the nature of this rela-

rocks, and living organisms, due to natural or anthro-

tionship and the features it has in common with

pogenic causes, has been described (Fluoride, 1971;

other stresses.
Weinstein, 1977). F that is accumulated in plants

enters the food chain through herbivores and passes

Evaluation of F Injury in the Field

into the soil in their wastes. The transfer from

one animal to another is possible in the case of

The most common measures of F injury to forests

insects that have accumulated F on or in their

include (1) assessment of the presence and amount

bodies and are eaten by birds or other carnivores,

of foliar injury, especially of indicator species;

but this has not beenstudied. It is also not known

(2) loss or depletion of sensitive receptors and

if increased levels of F associated with a variety

community changes; (3) measurement of biomass pro-

of insects was due to accumulation by ingestion or

duction; and (4) the accumulation of F in plant

by surface contamination. Hughes and others (in

tissues that might produce foliar injury or render

preparation) cultured cabbage loopers (Trichoplusia

the plant unsuitable for indigenous herbivores.

-
ni [Hubner]) on two kinds of diets. One contained

i n c r e a s i n g amounts of F a s NaF o r KF and equiva-
l e n t amounts of c o n t r o l cabbage. The o t h e r con-
t a i n e d F from HF-fumigated l e a v e s and was com-
bined w i t h c o n t r o l l e a v e s t o g i v e t h e same dose
curve. Analyses of prepupae and pupae showed
t h a t F accumulated i n t h e b o d i e s of t h e l o o p e r s
s u p p l i e d w i t h F s a l t (G. 1 0 p e t . of t h e concen-
t r a t i o n of t h e d i e t on a dry weight b a s i s ) ; no
F accumulated i n l o o p e r s grown w i t h t h e fumigated
cabbage d i e t . There was evidence t h a t l o o p e r s
grown w i t h t h e l a t t e r d i e t developed f a s t e r and
grew l a r g e r than t h o s e on t h e c o n t r o l cabbage
NaF o r KF d i e t s . These r e s u l t s s u g g e s t t h a t t h e
F r e p o r t e d a s having accumulated i n i n s e c t s prob-
a b l y was due t o s u r f a c e contamination of p a r t i c l e s
(Carlson and Dewey, 1971; Dewey, 1973). Inges-
t i o n of surface-contaminated i n s e c t s by c a r n i v o r e s ,
however, s t i l l t r a n s f e r s t h e F t o t h e n e x t t r o p h i c
l e v e l , b u t t h e amount of accumulation t h a t might
occur a t t h a t l e v e l i s n o t known.
There a r e g r e a t d i f f e r e n c e s i n t h e c a p a c i t i e s
of p l a n t s growing i n t h e same s o i l s t o accumulate
F. Most p l a n t s accumulate low c o n c e n t r a t i o n s of
F (0 - 1 0 ppm), b u t some s p e c i e s can accumulate
hundreds of ppm from t h e same s o i l . I n any
s p e c i e s , s o l u b l e F i n t h e s o i l s o l u t i o n can b e
r e a d i l y absorbed by p l a n t s (Romney and o t h e r s ,
1969). When d e p o s i t e d upon p l a n t s u r f a c e s , t h e
r e l a t i v e l y i n s o l u b l e forms of F have low phyto-
t o x i c i t y (McCune and o t h e r s , 1977), b u t t h e i r
i n g e s t i o n can b e harmful t o h e r b i v o r e s . Conse-
q u e n t l y , t h e main s o u r c e of p h y t o t o x i c a i r b o r n e
F i s HF.
Once i t e n t e r s a l e a f , F moves i n t h e t r a n s p i r -
a t i o n s t r e a m t o t h e t i p s o r margins of t h e l e a f
and s t a y s i n a form t h a t can be leached from many
l e a v e s (Leone and o t h e r s , 1956; L e d b e t t e r and
o t h e r s , 1960; Jacobson and o t h e r s , 1966). Con-
s e q u e n t l y , f o l i a r c o n c e n t r a t i o n s of F need n o t
s t e a d i l y i n c r e a s e (Knabe, 1970). Twigs of decid-
uous s p e c i e s can accumulate F i n t h e w i n t e r ,
presumably through t h e l e n t i c e l s , and e l e v a t e d
c o n c e n t r a t i o n s have been found i n young f o l i a g e
i n t h e s p r i n g ( K e l l e r , 1974, 1978). A s m a l l pro-
p o r t i o n of F e n t e r i n g a l e a f can a l s o be t r a n s -
l o c a t e d t o o t h e r p a r t s of t h e p l a n t (Kronberger
and o t h e r s , 1978).
F a l r e a d y p r e s e n t i n most s o i l s i s i n an i n s o l -
u b l e form and h a s l i t t l e i n f l u e n c e on v e g e t a t i o n ,
The f a t e of F leached from f o l i a g e and of F de-
p o s i t e d d i r e c t l y from t h e atmosphere h a s n o t been
s t u d i e d e x t e n s i v e l y . Any e f f e c t w i l l depend upon
t h e n a t u r e and chemistry of t h e s o i l . F l i i h l e r
and o t h e r s (1979) have shown t h a t l e a c h i n g of
p a r t i c u l a t e F depends upon i t s w a t e r s o l u b i l i t y
(NaF > powdered p i n e n e e d l e s c o n t a i n i n g F >
c r y o l i t e > CaFy). The F i n powdered p i n e n e e d l e
l i t t e r leached n e a r l y a s r a p i d l y a s d i d NaF.
When NaF s o l u t i o n s were a p p l i e d t o s o i l columns,
o r g a n i c m a t t e r , aluminum, and i r o n were l o s t ,
b u t t h e amount depended upon t h e s o i l t y p e ,
E f f e c t s of F on s o i l composition and s t r u c t u r e ,
m i n e r a l c y c l i n g , and l i t t e r decomposition a r e
important a s p e c t s of t h e impact of F on f o r e s t
ecosystems and r e s e a r c h i n t h e s e a r e a s should
have a h i g h p r i o r i t y .
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 Air Pollution-a 20th Century Allogenic

Influence on Forest ~cosystems'

W i l l i a m H. smith2

Abstract: Chronic doses of ozone, s u l f u r d i o x i d e ,

nitrogen oxides, hydrogen f l u o r i d e and other primary or
secondary gaseous a i r contaminants may cause subtle effects on
forest ecosystems. Air pollutants may influence reproduction,
n u t r i e n t cycling, photosynthesis, predisposition t o
entomological or pathological s t r e s s or q u a n t i t y of healthy
f o l i a r tissue. Forest ecosystem response t o chronic a i r
p o l l u t i o n may i n c l u d e a l t e r a t i o n s i n growth r a t e s and
successional patterns. The establishment of comprehensive
f i e l d and laboratory investigations t o systematically examine
chronic a i r p o l l u t i o n s t r e s s on f o r e s t ecosystems i n those
p a r t s of t h e world s u b j e c t t o atmospheric contamination is
concluded t o be of top priority. In the United States, forest
ecosystems judged t o be a t particular risk and i n need of more
intensive investigation include the Northern Hardwood f o r e s t ,
C e n t r a l Hardwood f o r e s t and Western Montane f o r e s t .

The interactions between a i r contaminants t o high dose - Class I11 r e l a t i o n s h i p - may

and f o r e s t ecosystems a r e extremely complex,
but can be conveniently divided i n t o - t h r e e
major c l a s s e s (Smith 1980). Under conditions
of low dose - Class I r e l a t i o n s h i p - t h e
vegetation and s o i l s of f o r e s t ecosystems
function a s important sources and sinks for a i r
pollutants. When exposed t o intermediate dose
- Class I1 r e l a t i o n s h i p - i n d i v i d u a l t r e e
s p e c i e s o r individual members of a given
species may be subtly and adversely affected by
n u t r i e n t s t r e s s , i m p a i r e d metabolism,
predisposition t o entomological or pathological
s t r e s s , or d i r e c t disease induction. Exposure
%?resented a t the Symposium on Effects of Air
Pollutants on Mediterranean and Temperate Forest
Ecosystems, June 22-27, 1980, Riverside, Cal-
ifornia, U.S .A.
p r o f e s s o r of Forest Pathology, School of
Forestry and Environmental S t u d i e s , Yale
University, New Haven, Conn., U.S.A.
induce acute morbidity or mortality of specific
trees. A t t h e ecosystem l e v e l t h e impact of
these various i n t e r a c t i o n s would be v e r y
variable. In t h e Class I relationship,
pollutants would be exchanged between the atmo-
s p h e r i c compartment, a v a i l a b l e n u t r i e n t
compartment, other s o i l compartments and
various elements of the biota. Depending on the
nature of the pollutant, the ecosystem impact
of this t r a n s f e r could be undetectable (inno-
cuous e f f e c t ) or s t i m u l a t o r y ( f e r t i l i z i n g
e f f e c t ) . I f t h e e f f e c t of a i r p o l l u t i o n dose
on some component of the biota is inimical then
a Class I1 r e l a t i o n s h i p is established. The
ecosystem impact i n t h i s c a s e could include
reduced productivity or biomass, alterations i n
species composition or community s t r u c t u r e ,
increased insect outbreaks or microbial disease
epidemics and increased morbidity. Under
conditions of high dose and Class I11 rela-
tionship, ecosystem impacts may include gross
simplification, impaired energy flew and bio-
geochemical cycling, changes i n hydrology and
erosion, climate alteration and major impacts
on associated ecosystems.
This paper is s p e c i f i c a l l y concerned with
Class I1 i n t e r a c t i o n s r e s u l t i n g from f o r e s t
ecosystem exposure t o chronic doses of ozone,
s u l f u r dioxide, nitrogen oxides, hydrogen
fluoride or other primary or secondary gaseous
a i r contaminants. It s p e c i f i c a l l y addresses
the relationship between these gases, and t h e i r
mixtures, on f o r e s t r e p r o d u c t i o n , f o r e s t
metabolism and direct forest s t r e s s as detailed
by previous c o n t r i b u t o r s t o t h i s s e c t i o n and
attempts t o examine resulting perturbations i n
ecosystem structure and function.
FOREST REPRODUCTION
Sexual reproduction of f o r e s t t r e e s is
c r i t i c a l l y important for maintenance of genetic
f l e x i b i l i t y and the persistence of most species
i n n a t u r a l f o r e s t communities. Reproductive
s t r a t e g i e s , however, a r e t y p i c a l l y b e s e t by a
v a r i e t y of "weak points" and reproductive
growth of many f o r e s t t r e e s is, a t best,
irregular and q u i t e unpredictable. Generally
t h e r e i s a very good c o r r e l a t i o n between t r e e
vigor and t h e capacity f o r flowering and
f r u i t i n g (Kramer and Kozlowski 1979). A
v a r i e t y of environmental c o n s t r a i n t s impose
r e s t r i c t i o n s on t r e e reproductive processes.
Because a i r contaminants may reduce t r e e vigor
and i n view of the f a c t t h a t numerous potential
p o i n t s of i n t e r a c t i o n have been i d e n t i f i e d
between a i r p o l l u t a n t s and r e p r o d u c t i v e
elements (Smith 19801, it has been hypothesized
t h a t a i r c o n t a m i n a n t s may i m p a c t f o r e s t
e c o s y s t e m s by i n f l u e n c i n g r e p r o d u c t i v e
processes.
Considerable evidence has been presented
i n d i c a t i n g a p o t e n t i a l a d v e r s e i m p a c t of
numerous g a s e o u s p o l l u t a n t s on p o l l e n
metabolism. Other p a p e r s have i n d i c a t e d
reduced cone and f r u i t production under f i e l d
conditions (Smith 1980). I f one o r more of
these various reproductive stress mechanisms is
o p e r a t i v e i n n a t u r a l f o r e s t ecosystems, it is
p o s s i b l e t h a t changes i n s p e c i e s composition
may ultimately occur. In their study of ozone
impact on t h e understory vegetation of an aspen
ecosystem, Harvard and Treshow (1975) concluded
t h a t only one or two y e a r s of ozone exposure
m i g h t be s u f f i c i e n t t o c a u s e s h i f t s i n
community c o m p o s i t i o n because of s e e d
production responses t o ozone exposure.
FOREST METABOLISM
Photosynthesis i s t h e most fundamental
metabolic process of forest ecosystems and is
the primary determinant of growth and biomass
accumulation. The r a t e of net photosynthesis
of mature t r e e s frequently is within the range
of 10-200 mg of carbon dioxide taken up per
gram of dry weight per day. The r a t e is
extremely variable, however, and influenced by
genetic, c l o n a l and provenance differences,
season of t h e year, time of day, p o s i t i o n
w i t h i n t h e crown of t h e t r e e , age of f o l i a g e ,
climate and edaphic factors.
Studies with a wide variety of agricultural
and herbaceous s p e c i e s , under c o n t r o l l e d
environmental conditions, have i n d i c a t e d t h a t
a i r contaminants must be added t o t h e list of
environmental v a r i a b l e s t h a t can p o t e n t i a l l y
a l t e r the r a t e of photosynthesis.
B e c a u s e of e a s e of h a n d l i n g a n d
experimental design, investigators studying the
r e l a t i o n s h i p between a i r p o l l u t a n t s and t r e e
photosynthesis have p r i m a r i l y employed t r e e
seedlings f o r research material and controlled
environmental f a c i l i t i e s for growth. Evidence
has been provided, under t h e above
circumstances, f o r photosynthetic suppression
caused by s u l f u r dioxide, ozone, f l u o r i d e ,
heavy metals and coal dust. The thresholds of
photosynthetic toxicity for tree seedlings vary
w i t h i n d i v i d u a l s p e c i e s and i n d i v i d u a l
pollutants. For s e v e r a l s e e d l i n g s t h e
threshold of s u l f u r dioxide photosynthet' c
influence may approximate 1 ppm (2620 pg m-3
o r l e s s f o r an exposure of s e v e r a l hours. For
ozone, the threshold of photosyn e t i c response
may approximate 0.5 ppm (980 g m or l e s s f o r9
an exposure of several days (Smith 1980).
Considerable r i s k i s associated with
extrapolation of seedling photosynthetic d a t a
accumulated i n c o n t r o l l e d e n v i r o n m e n t a l
f a c i l i t i e s t o older t r e e s i n n a t u r a l f o r e s t s .
Excised l e a f and s m a l l chamber techniques,
theref ore, have been employed t o assess the a i r
pollutant influence on photosynthetic rates of
t r e e s f ive-years-old and older. The use of
sapling-age experimental material avoids t h e
unique characteristics of seedling metabolism.
Evidence for f o r e s t t r e e sapling photosynthetic
suppression has been presented f o r s u l f u r
dioxide, ozone and cadmium. For sulfur dioxide
and ozone exposure, t h e s a p l i n g e v i d e n c e
suggests t h a t t h e threshold of photosynthetic
reduction may approximate 0.5 t o one ppm f o r 5-
10 hours for one or two days (Smith 1980).
Much of the seedling and sapling evidence
suggests t h a t t h e photosynthetic i n h i b i t i o n
caused by s u l f u r d i o x i d e and ozone i s
reversible i f the pollutant s t r e s s is removed.
Under t h e circumstance of v a r i a b l e p o l l u t a n t
concentration i n ambient atmospheres, photo-
synthetic recovery might be common. Synergism,
o r g r e a t e r s t r e s s resulting from simultaneous
pollutant exposure relative t o either pollutant
alone, appears f r e q u e n t l y i n t h e seedling and
sapling literature. Evidence f o r s y n e r g i s t i c
photosynthetic suppression by s u l f u r dioxide
and ozone and f l u o r i d e and cadmium has been
presented. Almost a l l of t h e s t u d i e s r e p o r t
photosynthetic depression in the absence, or a t
l e a s t p r i o r to, t h e appearance of v i s i b l e
f o l i a r symptoms.
The evidence f o r a i r pollution induced
photosynthetic suppression i n l a r g e t r e e s i n
n a t u r a l environmentsis extremely meager and
fragile. The seedling -
however, demonstrates a threshold of e f f e c t
sapling evidence
t h a t approaches ambient concentrations i n
numerous temperate environments. Because of
the profound importance of t h e photosynthetic
process and t h e p o t e n t i a l f o r suppression by
widespread a i r contaminants, appropriate f i e l d
s t u d i e s must be conducted i n s p i t e of t h e i r
d i f f i c u l t y and cost. The o p p o r t u n i t y t o
examine t h e i m p a c t of c o n t a m i n a n t s on
r e s p i r a t i o n and t r a n s p i r a t i o n should a l s o be
included i n experimental designs. Inclusion of
one or both of these physiologic processes i n
seedling - sapling s t u d i e s has revealed some
indication for significant alteration.
Increased r e s p i r a t i o n coupled with reduced
p h o t o s y n t h e s i s c o u l d e x a c e r b a t e growth
consequences.
FOREST FOLIAGE
Under conditions of s u f f i c i e n t dose, a i r
p o l l u t a n t s d i r e c t l y cause v i s i b l e i n j u r y t o
forest trees. The accumulation of particulate
contaminants on leaf surfaces or the continued
uptake of gaseous p o l l u t a n t s through l e a f
stomata w i l l eventually r e s u l t i n c e l l and
tissue damage that w i l l be manifest i n f o l i a r
symptoms obvious t o t h e trained, but unaided
eye. This direct induction of disease i n trees
by a i r p o l l u t a n t s i s t h e most dramatic and
obvious individual t r e e response of a l l Class
I1 interactions. I t i s t h e only Class I1
i n t e r a c t i o n t h a t can be detected i n t h e f i e l d
by c a s u a l o b s e r v a t i o n . Unlike a l t e r e d
reproductive strategy, n u t r i e n t cycling, t r e e
metabolism or insect and disease relationships;
t h e degree of f o l i a r symptoms induced by a i r
p o l l u t a n t s can be r e l a t i v e l y easily observed,
inventoried and quantified. In the presence of
s u f f i c i e n t dose, t r e e damage may be of
sufficient severity t o cause mortality.
Acute f o l i a r disease may be caused i n
f o r e s t v e g e t a t i o n by w i d e s p r e a d a i r
contaminants including; s u l f u r dioxide,
nitrogen oxides, ozone, peroxyacetyl-nitrates,
f l u o r i d e and s e v e r a l t r a c e m e t a l s , and
localized a i r contaminants including acid rain,
ammonia, chlorine, hydrocarbons and hydrogen
sulfide. The response of woody plants t o these
atmospheric p o l l u t a n t s is extremely variable
and dramatically controlled by genetic factors,
p l a n t age and h e a l t h and e n v i r o n m e n t a l
conditions. F i e l d symptoms of a i r p o l l u t i o n
injury a r e not highly specific, are mimicked by
a wide variety of other t r e e s t r e s s factors and
a r e u s e f u l o n l y t o experienced observers
f a m i l i a r w i t h t h e r a n g e of e d a p h i c ,
entomological and pathological s t r e s s factors
c h a r a c t e r i s t i c of a given f l o r a i n a given
location. The dose required t o produce acute
i n j u r y v a r i e s w i d e l y w i t h p o l l u t a n t and
vegetative type. There has been s u f f i c i e n t
work done t o enable a generalized ranking of
r e l a t i v e f o r e s t t r e e s e n s i t i v i t y t o t h e most
important a i r p o l l u t a n t s (Davis and Wilhour
1976). A summary treatment of general symptoms
and i n j u r y t h r e s h o l d s f o r t h e gaseous
contaminants included i n t h i s section is
contained i n Smith (1980).
FOREST ECOSYSTEM RESPONSE
The primary response of a forest ecosystem
t o sustained intermediate dose and Class I1
i n t e r a c t i o n would be reduced growth and con-
sequently reduced biomass. Reduced essential
element availability, decreased photosynthesis,
increased respiration, increased i n s e c t and
disease s t r e s s and decreased f o l i a r t i s s u e
would a l l contribute t o a reduction i n t r e e
growth rates and ultimately t o lessened forest
biomass. Alterations i n t h e reproductive
s t r a t e g i e s of i n d i v i d u a l t r e e s p e c i e s o r
d i f f e r e n t i a l response of these species t o
reduced nutrition, altered metabolism and pest
s t r e s s and t o d i r e c t f o l i a r i n j u r y may cause
changes i n competitive a b i l i t y and u l t i m a t e l y
lead t o a l t e r a t i o n s i n t r e e succession and
species composition. Recent reviews of Class
I1 vegetative responses t o a i r p o l l u t a n t s
include Heck and o t h e r s (19772, Jensen and
others (1976) and Weinstein and McCune (1979).
Forest Growth
Forest growth is complex i n concept and
measurement. Addition of woody tissue is the
dominant f e a t u r e of f o r e s t growth. The
accumulation of woody biomass (1iving weight)
represents gross photosynthetic production
l e s s respiratory losses. The most fundamental
characteristic of an ecosystem is its produc-
t i v i t y . Forest productivity is high r e l a t i v e
t o other ecos stems nd n e t productivity of
1200 dry g m'- year-' f o r t r e e s and shrubs
together is quite typical for temperate forests
(Whittaker 1975). Productivity i s strongly
controlled, however, by a variety of variables
i n c l u d i n g system a g e and e n v i r o n m e n t a l
parameters. The most important of the l a t t e r
include n u t r i e n t a v a i l a b i l i t y , water availa-
b i l i t y and temperature. Because of the variety
of Class I1 i n t e r a c t i o n s i d e n t i f i e d , a i r
quality also influences forest productivity i n
certain environments.
Productive f o r e s t s are critically
important, not only f o r t h e obvious relation-
s h i p between wood volume and commercial
products i n managed f o r e s t s , but a l s o f o r t h e
regulation and maintenance of q u a l i t y f o r
associated ecosystems, amenity functions and
general climatic and t e r r e s t r i a l stability. It
is disconcerting t o r e a l i z e , theref ore, t h a t
there is substantial and impressive evidence t o
indicate that two widespread a i r contaminants,
s u l f u r dioxide and ozone, a r e capable of
reducing f o r e s t growth. The more l o c a l i z e d
release of fluoride can also reduce the amount
of f o r e s t biomass (Smith 1980).
Evidence from a v a r i e t y of s t u d i e s
examining f o r e s t growth i n t h e v i c i n i t y of
l a r g e p o i n t sources of s u l f u r dioxide has
i n d i c a t e d s i g n i f i c a n t l y reduced growth.
Generally the correlation of growth impact w i t h
degree of f o l i a r i n j u r y c a u s e d by s u l f u r
dioxide is not high. Growth retardation occurs
i n t h e absence of any v i s i b l e i n d i c a t i o n of
stress. Most s u l f u r dioxide s t u d i e s have
accounted for precipitation influence on forest
growth over t h e study periods. Evidence f o r
ozone suppression of f o r e s t growth has been
provided by t h e comprehensive oxidant impact
study of the Western Montane forest ecosystem
i n California. Localized reduction of f o r e s t
growth may also occur i n environments subject
t o elevated l e v e l s of fluoride.
There a r e two s e r i o u s d e f i c i e n c i e s of
f o r e s t growth - a i r pollution s t r e s s research.
The f i r s t r e l a t e s t o the paucity of ambient a i r
quality determinations i n growth studies. This
makes establishment of dose thresholds or
c o r r e l a t i o n s of dose with growth influence
nearly impossible. The second serious l i m i t a -
tion relates t o the inability t o partition
reduced g r o w t h t o t h e v a r i o u s C l a s s I1
i n t e r a c t i o n s t h a t may actually be responsible
f o r it. For example, what percentage of
reduced growth may be due t o reduced nutrition,
reduced photosynthesis, increased i n s e c t or
disease a c t i v i t y or increased f o l i a r damage?
Future investigations of forest growth, a s
impacted by a i r q u a l i t y , must a l s o include
better accounts of growth inÂluencing f a c t o r s
other than p r e c i p i t a t i o n and a i r pollutants.
B e t t e r awareness of a d d i t i o n a l c l i m a t i c fac-
tors, impacts of insect and disease influence,
and management strategies must be indicated.
Forest Succession
A s a r e s u l t of t h e considerable v a r i e t a l
and s p e c i e s v a r i a t i o n i n r e l a t i v e
s u s c e p t i b i l i t y t o t h e v a r i o u s C l a s s I1
interactions, it is reasonable t o suppose t h a t
d i f f e r e n t i a l tolerance t o a i r pollution
influence a t the species level may be reflected
i n a l t e r e d p a t t e r n s of succession and s p e c i e s
composition a t the ecosystem level.
Ecologists recognize two major types of
processes t h a t influence ecosystem succession.
Autogenic processes a r e those r e s u l t i n g from
b i o l o g i c a l f a c t o r s w i t h i n t h e system. In
f o r e s t ecosystems autogenic processes would
i n c l u d e s i t e a l t e r a t i o n s c a u s e d by t h e
vegetation, i n f l u e n c e of one p l a n t s p e c i e s on
another and impact of native insect or disease
microorganisms. Allogenic processes, on t h e
other hand, a r e abiotic factors t h a t influence
s u c c e s s i o n from w i t h o u t t h e system.
Geochemical and climatic forces a r e especially
important examples of a l l o g e n i c f a c t o r s t h a t
inÂluence f o r e s t ecosystems. I d e a l i z e d eco-
system development c h a r a c t e r i s t i c a l l y i s
portrayed a s an o r d e r l y change of b i o l o g i c a l
p r o g r e s s i o n o c c u r r i n g i n a more o r l e s s
constant environment (Odum 1969, Woodwell
1974). It has been g e n e r a l l y assumed t h a t
autogenic processes dominate a l l o g e n i c
processes in t e r r e s t r i a l ecosystem succession.
This g e n e r a l i z a t i o n , however, is q u i t e incon-
s i s t e n t w i t h d a t a g e n e r a t e d by r e c e n t
imaginative s t u d i e s with f o r e s t ecosystems.
The importance of f i r e (an allogenic force) i n
influencing pre-settlement f o r e s t ecosystems i n
t h e North Central s t a t e s of t h e United S t a t e s
has been s u b s t a n t i a l (Loucks 1970, F r i s s e l l
1973, Heinselman 1973). The s i g n i f i c a n c e of
wind s t r e s s (an a l l o g e n i c f o r c e ) has been
suggested t o e x e r t s u b s t a n t i a l c o n t r o l over
successional development of f o r e s t ecosystems
i n New England (Stephens 1955, 1956, Raup 1957,
Henry and Swan 1974). F o r e s t management
p r a c t i c e s imposed by man, f o r example c l e a r -
cutting, may simulate the influence of natural
a l l o g e n i c f o r c e s on f o r e s t development and
i n t e r r u p t p r o g r e s s t o w a r d a steady s t a t e
c o n d i t i o n (Bormann a n d L i k e n s 1 9 7 9 ) .
Conversely other forest management procedures,
f o r example f i r e control, may e l i m i n a t e a
c o n t r o l l i n g a l l o g e n i c f o r c e and p e r m i t
succession t o proceed toward an unnatural
steady s t a t e condition. Class I1 s t r e s s e s
imposed on forest ecosystems by a i r pollutants
may be considered a 20th Century a l l o g e n i c
process of p o t e n t i a l importance t o f o r e s t
ecosystem development. A s i n t h e case of
clearcutting, t h i s human related force might be
expected t o a l t e r t h e a t t a i n m e n t of steady
s t a t e conditions. Air p o l l u t i o n s t r e s s would
appear t o have c e r t a i n unique q u a l i t i e s t h a t
may make it a n a l l o g e n i c i n f l u e n c e of
p a r t i c u l a r importance. Length of exposure t o
t h i s f o r c e precludes evolutionary adjustment
and i t s influence, i n c e r t a i n a r e a s , may be
q u i t e continuous r a t h e r than c y c l i c a s a r e
windstorms and f i r e s . What i s t h e evidence
a v a i l a b l e t o support t h e importance of a i r
pollution a s an allogenic force of significance
in forest ecosystem development?
I n 1968, p r i o r t o s o p h i s t i c a t e d
understanding of most Class I1 i n t e r a c t i o n s ,
Treshow (1968) provided an excellent review of
t h e i m p a c t of a i r c o n t a m i n a n t s on p l a n t
populations. Treshow's review, along w i t h a
v a r i e t y of a d d i t i o n a l l a t e 1960's papers, f o r
example Niklfeld (19671, ~ a j d f i kand ~ u s i 6 k a
(1968) and Trautmann and o t h e r s (19701, have
indicated a l t e r a t i o n s i n successional pattern
or s p e c i e s composition i n f o r e s t ecosystems
subject t o a i r pollution exposure.
The f o r e s t s of t h e San B e r n a r d i n o
Mountains i n southern C a l i f o r n i a have been
subject t o oxidant stress from the Los Angeles
m e t r o p o l i t a n complex f o r t h i r t y y e a r s .
Intensive investigations conducted i n t h e San
Bernardino National Forest over the years have
provided valuable insight and perspective on a
variety of forest a i r pollution relationships.
I n 1970, Cobb and Stark concluded t h a t i f a i r
pollution from the Los Angeles basin continued
t o increase, t h e r e w i l l be a conversion from
w e l l stocked f o r e s t s dominated by ponderosa
pine Qirui Doug ex. Laws) t o poorly
stocked stands of l e s s susceptible tree species
i n the San Bernardino Mountains. Miller (1973)
has provided a thorough discussion of this
oxidant induced f o r e s t community change.
Ponderosa pine is one of f i v e major species of
the "mixed conifer type" that covers wide areas
of the western S i e r r a Nevada and t h e mountain
ranges, including the San Bernardino Mountains,
i n southern C a l i f o r n i a from 1000 t o 2000 m
(3000-6000 f e e t ) elevation. Other s p e c i e s
r e p r e s e n t e d i n c l u d e s u g a r p i n e (W
l a m k r t h u Douql.) , white f i r Wigs
[Gord. & Glend.] L i n d l . ) , i n c e n s e - c e d a r
(Woce- Torr.) and C a l i f o r n i a
b l a c k oak ( Q u e r c u Newb.). The
response of t h e s e f i v e major t r e e s p e c i e s t o
oxidant a i r contaminants i n the San Bernardino
National F o r e s t has been variable. Ponderosa
pine exhibits the most severe f o l i a r response
t o elevated ambient ozone. A 1969 a e r i a l survey
c o n d u c t e d by t h e U.S.D.A. Forest Service
i n d i c a t e d 1.3 m i l l i o n ponderosa (or J e f f r e y ,
Pinus j e f f r e i Grev. & Balf.) p i n e s on more
2
than 405 km (100,000 acres) were s t r e s s e d t o
some degree. M o r t a l i t y of ponderosa pine has
been extensive. Actual death i s t y p i c a l l y
a t t r i b u t e d t o bark b e e t l e i n f e s t a t i o n of a i r
pollution stressed trees. White f i r h a s
s u f f e r e d s l i g h t damage, but s c a t t e r e d t r e e s
have e x h i b i t e d severe symptoms. Sugar pine,
incense cedar and black oak have exhibited only
s l i g h t f o l i a r damage from oxidant exposure. A
233 ha (575 acre) study block w a s delineated i n
t h e northwest s e c t i o n of t h e San Bernardino
N a t i o n a l F o r e s t i n o r d e r t o conduct a n
i n t e n s i v e inventory of vegetation present i n
v a r i o u s s i z e c l a s s e s and t o e v a l u a t e the
h e a l t h f u l n e s s of t h e f o r e s t . Ponderosa p i n e s
i n the 30 can (12 inch) diameter class or larger
were more numerous than any other s p e c i e s of
comparable s i z e i n the study area. These pines
were most abundant on t h e more exposed r i d g e
c r e s t sites of the sample area. Mortality of
ponderosa pine ranged from 8-10 percent during
1968-1972. The l o s s of a dominant species i n a
f o r e s t ecosystem clearly exerts profound change
i n t h a t system. Miller concluded from h i s
investigation that the lower two-thirds of the
study a r e a w i l l probably s h i f t t o a g r e a t e r
proportion of white f i r . I t was judged t h a t
incense cedar w i l l probably remain secondary t o
w h i t e f i r . Sugar p i n e was presumed t o be
r e s t r i c t e d by l e s s e r competitive a b i l i t y and
dwarf m i s t l e t o e i n f e c t i o n . The r a t e of
composition change w a s deemed dependent on t h e
r a t e of ponderosa pine mortality. The upper
one-third of t h e study area, c h a r a c t e r i z e d a s
more environmentally severe due t o climatic and
edaphic s t r e s s , supports less vigorous white
f i r growth. Following l o s s of ponderosa pine
i n this area, sugar pine and incense cedar may
assume g r e a t e r importance. M i l l e r judged,
however, t h a t n a t u r a l regeneration of t h e
l a t t e r s p e c i e s may be r e s t r i c t e d i n t h e more
barren, dry sites c h a r a c t e r i s t i c of t h e upper
ridge area. C a l i f o r n i a black oak and shrub
s p e c i e s may become more abundant i n t h e s e
disturbed areas. Additional and i n t e n s i v e
research on f o r e s t composition i n t h e San
Bernardino National Forest has been reported
(Miller 1977). Tree population dynamics were
examined on 18 permanent p l o t s established i n
1972 and 1973 and on 83 t e m p o r a r y p l o t s
e s t a b l i s h e d i n 1974 t o i n v e s t i g a t e f o r e s t
development a s a f u n c t i o n of t i m e s i n c e t h e
most recent f i r e . Generally the d a t a s t i l l
support the hypothesis t h a t f o r e s t succession
toward more tolerant species such as white f i r
and incense cedar occurs i n t h e absence of
f i r e . I n t h e presence of f i r e , pine may be
favored by seedbed preparation and elimination
of competing s p e c i e s . These more r e c e n t
studies suggest a larger number of f o r e s t sub-
types may e x i s t w i t h i n the f o r e s t ecosystem
than i n i t i a l l y realized.
The changes i n f o r e s t composition caused
by oxidants in this southern California f o r e s t
have c r e a t e d a management concern, a s w e l l a s
e c o l o g i c a l change, b e c a u s e t h e f o r e s t i s
intensively used as a recreational resource and
the l o s s of ponderosa pine is judged t o reduce
aesthetic q u a l i t i e s of the forest.
Other examples, not as dramatic a s the San
Bernardino example, can be found. Hayes and
Skelly (1977) have monitored t o t a l oxidants and
associated oxidant injury t o eastern white pine
i n t h r e e r u r a l V i r g i n i a sites between A p r i l
1975 and March 1976. V a r i e t i e s of p i n e
categorized a s s e n s i t i v e and i n t e r m e d i a t e t o
oxidant s t r e s s were judged t o be under stress.
The authors speculated that susceptible eastern
white pine LJ i n the Blue Ridge
and Southern Appalachian Mountains may be
rendered less competitive by a i r p o l l u t i o n
s t r e s s . S h i f t s i n s p e c i e s composition away
from white pine importance along with o t h e r
changes i n tree distributions may be occurring
i n certain eastern regions. Brandt and Rhoades
(1973) in t h e i r investigation of limestone dust
impact i n deciduous f o r e s t s i n southwestern
V i r g i n i a p r e d i c t e d changes i n s p e c i e s
composition r e s u l t i n g from d u s t influence.
Dusty s i t e s had reduced seedling and s a p l i n g
density of red maple Q & ~ L LJ, chestnut
oak (Quercw L.1 and red oak -(
Michx. £.I This observation along
w i t h documentation of reduced mean basal area
and l a t e r a l growth of t h e s e t r e e s , l e d t h e
a u t h o r s t o s u g. g.e s t t h a t y e l l o w - p o p l a r
L), more resistant t o
s t r e s s caused by d u s t accumulation, would
increase i n importance i n t h e s e hardwood
stands.
Treshow and Stewart (1973) have conducted
one of the few studies truly concerned with a i r
p o l l u t i o n i m p a c t on a n e n t i r e vegetative
community. Portable fumigation chambers were
placed over r e p r e s e n t a t i v e p l a n t s i n
intermountain grassland, oak, aspen and conifer
communities. Ozone fumigations were conducted
t o e s t a b l i s h i n j u r y thresholds f o r 70 common
plant species indigenous t o these communities.
G e n e r a l l y i n j u r y was e v i d e n t a t v a r yjJig
concentrations above 15 pphm (294 p g m 1.
Species that were found t o be most sensitive t o
ozone i n t h e grassland and aspen communities
investigated included some dominants which were
considered key t o community i n t e g r i t y . The
most dramatic example was aspen (m
h a n u l o i d e s Michx.) i t s e l f . Single two-hour
e x p o s u r e t o 1 5 pphm ozone caused s e v e r e
symptoms on 30 percent of the foliage exposed.
White f i r seedlings r e q u i r e aspen shade f o r
optimal juvenile growth. The authors judged
t h a t s i g n i f i c a n t aspen l o s s might r e s t r i c t
w h i t e f i r development and a l t e r f o r e s t
succession. In a companion study, Harward and
Treshow (1975) pursued t h e i r i n t e r e s t i n
evaluating ozone impact on aspen communities by
evaluating the growth and reproductive response
of 14 understory species t o ozone. Plants were
fumigated i n greenhouse chambers throughout
their growing seasons. It w a s concluded from
these fumigations t h a t p l a n t s e n s i t i v i t i e s
varied s u f f i c i e n t l y t o make probable major
s h i f t s i n composition i n aspen communities
following only a year or two of exposure t o
ozone above concentrations of 7-15
294 (tg m 3 ) . The a u t h o r s observed t h a t
(137-
comparable doses a r e widespread i n the vicinity
of urban a r e a s and t h a t widespread impacts on
p l a n t community s t a b i l i t i e s may be common i n
nature. The e f f o r t s of Michael Treshow and
c o l l e a g u e s h i g h l i g h t s t h e i m p o r t a n c e of
examining shrub and herb s t r a t a when assessing
a i r pollution impact on forest ecosystems.
McClenahen (1978) has provided a most
interesting study with quantitative data on the
impact of polluted a i r on the various s t r a t a of
a f o r e s t ecosystem. F o r e s t vegetation was
measured i n seven stands on similar s i t e s in a
50 km area of the upper Ohio River Valley. The
stands were s i t u a t e d along a g r a d i e n t of
polluted a i r containing elevated concentrations
of chloride, s u l f u r dioxide, f l u o r i d e and
perhaps other contaminants. Species richness
(number of d i f f e r e n t s p e c i e s ) evenness
(dominance index - low values i n d i c a t e domi-
nance by one or a few species) and Shannon
d i v e r s i t y index were typically reduced within
t h e overstory, subcanopy and herb s t r a t a near
i n d u s t r i a l sources of a i r contaminants. In-
creasing a i r pollutant exposure reduced canopy
stem density, but abundance of vegetation i n
other s t r a t a tended t o increase along the same
gradient. The r e l a t i v e importance of sugar
maple (AQX s a c c Marsh.) ~ was g r e a t t y
reduced i n all s t r a t a with increasing pollutant.
dose, while yellow buckeye ( A e s c w octandra
Marsh.) appeared tolerant of poor air quality.
In the shrub layer the importance of spicebush
(m b e n z a 1L.I B1.1 increased w i t h
increasing pollutant exposure.
I n southern C a l i f o r n i a t h e predominant
n a t i v e s h r u b l a n d v e g e t a t i o n c o n s i s t s of
chaparral and c o a s t a l sage scrub. The former
occupies upper e l e v a t i o n s of t h e c o a s t a l
mountains, extending i n t o t h e North Coast
ranges, e a s t t o c e n t r a l Arizona, and south t o
Ba j a California; while t h e former occupies
lower e l e v a t i o n s on t h e c o a s t a l and i n t e r i o r
sides of the coast ranges from San Francisco t o
Baja C a l i f o r n i a . Westman (1979) a p p l i e d
standard p l a n t ordination techniques t o these
shrub communities t o examine the influence of
a i r pollution. The reduced cover of n a t i v e
species of c o a s t a l sage scrub documented on
some s i t e s was s t a t i s t i c a l l y indicated t o be
caused by elevated atmospheric oxidants. S i t e s
of h i g h a m b i e n t o x i d a n t s w e r e a l s o
characterized by declining species richness.
Influence of a i r p o l l u t i o n s t r e s s on
succession and ecosystem species composition
probably varies with the age and successional
s t a t u s of t h e f o r e s t . Harkov and Brennan
(1979) have observed t h a t most woody p l a n t s
susceptible t o ozone injury are generally early
s u c c e s s i o n a l p l a n t s p e c i e s . Most t r e e s
intermediate or t o l e r a n t of ozone s t r e s s a r e
typically mid- or l a t e successional types. It
i s not unreasonable t o propose, a s Harkov and
Brennan did, t h a t l a t e successional f o r e s t
communities may be t h e most r e s i s t a n t t o
compositional change as a result of chronic a i r
pollution exposure. Mature ecosystems are a l s o
t y p i f i e d by other c h a r a c t e r i s t i c s t h a t may
increase t h e i r r e s i s t a n c e t o a i r p o l l u t i o n
stress. Low n e t p r o d u c t i o n may reduce
potential importance of restrictions imposed by
a i r contaminants on photosynthesis. Closed and
slow nutrient cycling may make nutrient capital
l e s s l i a b l e t o l o s s by a i r pollutant influence.
There is increasing appreciation of t h e
i m p o r t a n c e of a l l o g e n i c f o r c e s on f o r e s t
ecosystem succession. The significance of f i r e
and wind s t r e s s on f o r e s t development is
s u b s t a n t i a l i n c e r t a i n environments. I t is
concluded t h a t a i r p o l l u t a n t impact may a l s o
exert c r i t i c a l l y important control over forest
succession and species composition. Long-term,
continual s t r e s s tends t o decrease t h e t o t a l
f o l i a r cover of v e g e t a t i o n , decrease t h e
s p e c i e s r i c h n e s s , and t o i n c r e a s e t h e
concentration of dominance by favoring a few,
tolerant species.
 CONCLUSIONS
Large a r e a s of t h e t e m p e r a t e f o r e s t
ecosystem a r e c u r r e n t l y experiencing major
perturbation from a i r pollution. The influence
of a v a r i e t y of a i r c o n t a m i n a n t s on
biogeochemical cycling, patterns of succession
and competition and i n d i v i d u a l t r e e health,
designated Class I1 interactions (Smith 19801,
a r e causing s i g n i f i c a n t f o r e s t change i n t h e
temperate zone. A t t h e ecosystem l e v e l t h e
major p e r t u r b a t i o n s include decreased
productivity, biomass and d i v e r s i t y ; a t t h e
community l e v e l reduced growth; and a t t h e
population level a l t e r e d s p e c i e s composition.
E a r l y and m i d - s u c c e s s i o n a l f o r e s t s a r e
concluded t o be a t particular risk. Temperate
f o r e s t s have h i s t o r i c a l l y been subjected t o
major change resulting from the a c t i v i t i e s of
human beings. For c e n t u r i e s t h e major
i n f l u e n c e was g r o s s d e s t r u c t i o n f o r
a g r i c u l t u r a l , f u e l o r o t h e r wood-product
purposes. In the present Century reduced need
f o r a g r i c u l t u r a l land and increased f o r e s t
management has reduced t h e adverse impact on
f o r e s t s i n temperate latitudes. Human
a c t i v i t i e s of primary contemporary importance
t o f o r e s t structure and function have included
t h e i n t r o d u c t i o n of e x o t i c a r t h r o p o d and
m i c r o b i a l t r e e p e s t s i n t o f o r e s t systems
l a c k i n g e v o l u t i o n a r y exposure t o t h e s e
d e s t r u c t i v e agents, enhancement of native and
natural stresses by cultural practices, and the
creation of a r t i f i c i a l forests of one or a few
commercially important species. I n t h e p a s t
several decades, however, w e have accumulated
s u f f i c i e n t evidence t o i n d i c a t e t h a t an
a d d i t i o n a l major anthropogenic modifier of
temperate f o r e s t ecosystem development is a i r
pollution.
Research
During the l a s t decade f o r e s t researchers
have outlined numerous Class I1 interactions by
l a r g e l y u t i l i z i n g r e l a t i v e l y young f o r e s t
p l a n t s grown i n c o n t r o l l e d e n v i r o n m e n t
f a c i l i t i e s . During t h e next decade we must
make a n e f f o r t t o perform experiments i n
n a t u r a l f o r e s t e c o s y s t e m s t o confirm our
hypotheses t h a t a m b i e n t a i r p o l l u t i o n i s
reducing f o r e s t p r o d u c t i v i t y and a l t e r i n g
species composition.
The very highest research p r i o r i t y is
reserved f o r the establishment of comprehensive
investigations t o systematically examine Class
I1 interactions i n f o r e s t ecosystems located i n
t h o s e p o r t i o n s o f t h e t e m p e r a t e zone
particularly subject t o a i r pollution stress.
These investigations should include analysis of
a i r contaminant i n f l u e n c e on s o i l metabolism
and s t r u c t u r e f n u t r i e n t c y c l i n g , t r e e
reproduction, photosynthesis and r e s p i r a t i o n ,
important arthropod and microbial pathogens,
f o l i a r symptoms of important vegetation i n a l l
f o r e s t s t r a t a and a c a r e f u l examination of
f o r e s t p r o d u c t i v i t y and a l t e r a t i o n s i n
successional t r e n d s and s p e c i e s dominance.
These s t u d i e s w i l l be of extended term. They
w i l l r e q u i r e the p a r t i c i p a t i o n of numerous
s c i e n t i f i c d i s c i p l i n e s , minimally including
pathology, entomology meteorology, s o i l
science, s o i l microbiology, ecology and systems
analysis. Continuous meteorological and a i r
q u a l i t y monitoring w i l l be required. Air
p o l l u t a n t s measured should include s u l f u r
dioxide, nitrogen oxides, hydrocarbons, ozone
and p a r t i c u l a t e s , t h e l a t t e r t o i n c l u d e
determination of s u l f a t e s , n i t r a t e s and t r a c e
metals. P r e c i p i t a t i o n a c i d i t y w i l l be
routinely determined. The o b j e c t i v e of t h e s e
comprehensive s t u d i e s w i l l be t o c l a r i f y and
q u a n t i f y various Class I and I1 i n t e r a c t i o n s .
The ecosystems w i l l be evaluated f o r their
a b i l i t y t o r e s i s t ( i n e r t i a ) and respond
(resilience) t o disturbance from a i r pollution
s t r e s s . Model development f o r t h e various
i n t e r a c t i o n s w i l l h o p e f u l l y allow f u t u r e
projectionsr given various a i r q u a l i t y
sceneries, and allow extrapolation of findings
t o other ecosystems.
I n t h e United S t a t e s t h e only research
program presently addressing this need is the
oxidant study i n progress on the San Bernardino
National Forest i n California. I t is
imperative t h a t a d d i t i o n a l i n v e s t i g a t i o n s be
i n i t i a t e d a s soon a s possible. The s t u d i e s
should be established i n those areas judged t o
be under the greatest stress and they should be
initiated, where possible, i n association with
integrated and comprehensive f o r e s t ecosystem
studies currently in progress. Priority f o r e s t
ecosystems i n t h e United S t a t e s include: 1)
Northern Hardwood f o r e s t , 2) Central Hardwood
f o r e s t and 3) Western Montane f o r e s t (San
Bernardino p r o j e c t i n progress). Appropriate
l o c a t i o n s , i n t e r m s of e x i s t i n g r e s e a r c h
f a c i l i t i e s or abundant ancillary information,
f o r t h e Northern f o r e s t a r e t h e Hubbard Brook
Experimental Forest i n New Hampshire, the Isle
Royale National Park, Michigan and t h e I t a s c a
Forest, Minnesota. I n t h e Central f o r e s t the
CampBranchForestwatershed i n e a s t - c e n t r a l
Tennessee and the Coweeta Hydrologic Laboratory
i n western North Carolina would be appropriate.
With regard t o l o c a t i o n , t h e Wayne National
Forest i n Ohio would appear t o represent an in-
teresting research opportunity. In addition t o
t h e San Bernardino Forest study, t h e Andrews
Experimental Forest, Oregon and the Bitterroot
N a t i o n a l F o r e s t , I d a h o would be o t h e r
s t r a t e g i c a l l y located sites f o r the Western
Montane forest.
Policy
It is recognized t h a t a i r pollution is one
of t h e most s i g n i f i c a n t c o n t e m p o r a r y
anthropogenic s t r e s s e s imposed on temperate
forest ecosystems. Gradual and s u b t l e change
in forest m e t a b o l i s m and composition over wide
areas of the temperate zone over extended time, New d i r e c t i o n s i n c e n t u r y t h r e e :
rather than dramatic destruction of forests i n Strategies for land and water use, p. 193-
t h e immediate v i c i n i t y of p o i n t sources over 202, Proc. 32nd Annual meeting, S o i l
s h o r t periods, m u s t be recognized a s t h e Conserva. Soc. Amer., Aug. 7-10, 1977,
primary consequence of a i r p o l l u t i o n stress. Richmond, Vir.
This realization means t h a t forest interactions
with a i r contaminants must be given con- Heinselman, M.L.
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Henry, J.D. and J.H.A. Swan.
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Secondary and Interactive Effects of
Chronic Gaseous Pollutant Exposure
of Producers, Consumers, and
Decomposers
Influence of Chronic Air Pollution
on Mineral Cycling in Forests1

Paul J. Zinke 2

Abstract: This paper reviews the literature

concerning the impact of'chronic air pollution on

mineral element cycling in forests. The concept

involves the forest trees taking up essential and

other elements from the soil and surrounding en-

vironment eventually to return them to the soil

upon mortality and decay. Chronic pollutants are

considered in the context of this cycling as another

form of elemental addition to the site subject to

cycling in the same manner. The review is organized

to assess the addition of the major elements, carbon,

nitrogen, and sulfur, and the trace metallic elements.

In addition, pollutant effects upon the ecosystem

living components in terms of producers and decompos-

ers are considered along with potential changes in the

redox and pH state of the different portions of the

forest. Some oriuinal data on foliar comnosition of

Big Cone Spruce (Pseudotsuga macrocarpa (vasey) Mayr)

in relation to pollution exposure, and the evaluation

of a soil subject to increments o$ hydrogen in simu-

lated acid rain leaching are presented.

INTRODUCTION
T h i s paper w i l l p r e s e n t a review of
various aspects of mineral element
c y c l i n g i n f o r e s t s a s a f f e c t e d by c h r o n i c
atmospheric p o l l u t i o n .
L e t c . Return t o t h e s o i l s t o r a g e occurs,
Presented at the Symposium on Effects
of Air Pollutants on Mediterranean and
Temperate Forest Ecosystems, June 22-27,
1980, Riverside, California, U.S.A.
Associate Professor of Forestry, Dep.
of Forestry and Resource Manage., Univer-
sity of California, Berkeley, Calif.
The f o r e s t w i l l be c o n s i d e r e d t o b e
t h e s t a n d of t r e e s and t h a t p o r t i o n o f
t h e atmosphere encompassed by canopy and
t r u n k s p a c e , and t h e p o r t i o n o f t h e s o i l
encompassed by t h e r o o t s p a c e . Mineral
c y c l i n g i s t h e p r o c e s s of c y c l i n g of e l e -
ments from t h e s o i l t h r o u g h u p t a k e by
r o o t s o r by f o l i a g e , t r a n s p o r t w i t h i n t h e
t r e e s and t h e e v e n t u a l r e t u r n t o t h e
f o r e s t s o i l i n t h e processes of f o l i a r
leaching, root exudation, f o l i a g e drop,
and t h e c y c l e i s c o n t i n u e d by u p t a k e
a g a i n , o r i t may be broken by l o s s from
t h e s o i l , o r t i e u p as i n s o l u b l e p r e c i p i -
t a t e s o r compounds. A i r p o l l u t i o n expo-
s u r e w i l l be c o n s i d e r e d t o b e t h e a d d i -
t i o n a l a t m o s p h e r i c i n p u t s which e n t e r t h e
nutritional and elemental cycle of
f o r e s t s due t o man made e f f e c t s on atmos-
p h e r i c composition.
 ELEMENTAL CYCLIdG
The i d e a o f t h e c y c l i n g of e l e m e n t s
from v e g e t a t i o n t o s o i l , f o l l o w i n g u p t a k e
from s o i l probably outdates written
r e c o r d s as i s a p p a r e n t from t h e p r i m i t i v e
a g r i c l t u r a l p r a c t i c e of s h i f t i n g cultiva-
t i o n wherein a f o r e s t i s c u t and burned
f o r s o i l enrichment by t h e a s h . This i s
f o l l o w e d by t h e regrowth o f a f a l l o w
f o r e s t c r o p , which i n t u r n i s burned
again. The f o r e s t v e g e t a t i o n c y c l e s e l e -
ments e s s e n t i a l f o r v e g e t a t i v e growth as
w e l l as u n n c e s s a r y e l e m e n t s a c c i d e n t a l t o
t h e p a r t i c u l a r environment. The 16 o r 1 7
e s s e n t i a l e l e m e n t s ( c , d, 0, P, K, N, S,
C a , Fe, Mg, Mn, Cu, Zn, Mo, B, C l , and
p e r h a p s S e ) , a r e c y c l e d by n e c e s s i t y f o r
w i t h o u t them t h e r e would be no f o r e s t .
I n a d d i t i o n a l m o s t e v e r y o t h e r element i s
t r a n s l o c a t e d by p l a n t s . T h i s concept i s
used i n geochemical p r o s p e c t i n g f o r many
o f t h e more v a l u a b l e e l e m e n t s , u s i n g t h e
p l a n t as a chemical p r o s p e c t o r (NASA
1 9 6 8 ) . These e l e m e n t s which a r e c y c l e d
may be e n r i c h e d i n c o n c e n t r a t i o n i n v a r i -
ous p o r t i o n s of t h e c y c l e , as i n t h e
foliage, the plant d e t r i t u s on t h e
ground, o r i n t h e s u r f a c e o f t h e s o i l
beneath t h e p l a n t s . Such enrichment i s
o f t e n t h e i n d i c a t i o n of an extraneous
element. S i m i l a r l y t h e input of extrane-
ous e l e m e n t s i n t h e p r o c e s s o f p o l l u t i o n
may b e evidenced by enrichment o v e r t h e
normal background amounts.
The s t a t e o f t h e v e g e t a t i o n and i t s
capacity f o r storage of elements w i l l
d e t e r m i n e t h e r e l a t i v e change induced by
t h e a d d i t i o n o f a p o l l u t a n t element. The
s t a t e o f t h e v e g e t a t i o n is p a r t l y d e t e r -
mined by t h e f a c t t h a t v e g e t a t i o n i s a
g i a n t r e d u c t i o n r e a c t i o n i n which pho-
t o s y n t h e s i s r e d u c e s c a r b o n from carbon
d i o x i d e t o c a r b o n i n reduced carbon com-
ounds as d e s c r i b e d by Stumm and -Morgan
71 970) . The f o r e s t t r e e s c a r r y o u t t h i s
r e d u c t i o n r e a c t i o n , f o l l o w e d by t h e i r
d e a t h and t h e s u b s e q u e n t o x i d a t i o n of t h e
carbon compounds in the soil. The
c y c l i n g of t h e e l e m e n t s on t h e s i t e
o c c u r s i n t h e c o n t e x t o f t h i s g i a n t redox
r e a c t i o n ( ~ a r r e l s ,e t a l . , 1 9 7 5 ) , and i t s
s t a t e and magnitude d e t e r m i n e t h e r a t e
and amount a t which t h e o t h e r e l e m e n t s
w i l l be c y c l e d .
The c y c l i n g of most o f t h e e l e m e n t s
which undergo redox changes i n t h e f o r e s t
n u t r i e n t c y c l e (C H 0 N S Mn ~ e w ) ill
tend to be reduced i n the living
v e g e t a t e d p o r t i o n o-f t h e c y c l e and com-
p l e t e t h e c y c l e t o an o x i d i z e d form i n
the soil. However, l o c a l s o i l f a c t o r s
f a v o r i n g a n a e r o b i c c o n d i t i o n s such as
w a t e r i n e x c e s s , may c a u s e t h e s o i l t o
become a s i n k f o r some e l e m e n t s i n t h e
reduced form, o r f o r o t h e r e l e m e n t s i n
t h e o x i d i z e d form.
Thus p o l l u t a n t s may e n t e r t h e e l e -
mental c y c l e s i n f o r e s t s and v a r y i n
m o b i l i t y depending upon t h e redox poten-
t i a l a t v a r i o u s p o r t i o n s of t h e e l e m e n t a l
cycle. Some p o l l u t a n t s may a l s o b e oxi-
dants i n r e l a t i o n t o the portion of t h e
ecosystem t h a t i s a b s o r b i n g them and t h u s
a f f e c t t h e redox p o t e n t i a l a t t h a t p o i n t
a c c o r d i n g t o Haagen-Smit ( 1 9 5 8 ) .
STORAGE POINTS AND SINKS
The soil and vegetation of the
forest have several points of long term
s t o r a g e o f m a t e r i a l s added t o t h e elemen-
t a l c y c l i n g system. It h a s been l e a r n e d
from p a s t f e r t i l i z a t i o n e x p e r i e n c e t h a t
e l e m e n t s added t o t h e f o r e s t , e i t h e r t o
s o i l o r d i r e c t t o t r e e s , may b e s t o r e d
f o r v a r y i n g p e r i o d s of t i m e i n d i f f e r e n t
p o r t i o n s of t h e f o r e s t . Obviously, t h e
t r e e trunks provide storage of t h e
m a t e r i a l s c o n t a i n e d i n t h e wood f o r t h e
l e n g t h of l i f e o f t h e t r e e and i t s subse-
q.uent decomposition t i m e . Deciduous por-
t i o n s of t h e t r e e r e t a i n m a t e r i a l s f o r
l e n g t h s of t i m e p r o p o r t i o n a t e t o t h e i r
r e s i d e n c e t i m e on t h e t r e e ; b a r k f o r
l o n g e r p e r i o d s , l e a v e s and t w i g s f o r
l e s s e r p e r i o d s . The s t o r a g e t i m e i n t h e
decomposing o r o x i d i z i n g p o r t i o n s o f t h e
d e t r i t u s dropped t o t h e s o i l depends upon
l o c a l environmental c o n d i t i o n s of tem-
p e r a t u r e , redox s t a t e , and a v a i l a b i l i t y
o r t o x i c i t y of e l e m e n t s t o decomposing
( oxidizing) organisms. For example,
decomposition t i m e s f o r s u r f a c e d e t r i t u s
on t h e s o i l s u r f a c e i n a f o r e s t may v a r y
from a f r a c t i o n o f a y e a r t o many y e a r s .
The s o i l b e n e a t h t h e f o r e s t i s a
g i a n t fixed-bed i o n exchanger, and once
elements e n t e r t h e s o i l f o l l o w i n g r e l e a s e
from s t o r a g e i n t h e v e g e t a t i v e p o r t i o n o f
t h e f o r e s t and i t s d e t r i t u s t h e y may b e
s t o r e d on t h i s exchange complex.
c a p a c i t y of t h i s i o n exchange bed as w e l l
The
as t h e n a t u r e o f a s s o c i a t e d c a t i o n s w i l l
determine t h e s t o r a g e p r o b a b i l i t y of ele-
ments added t o t h e f o r e s t whether as
additives (pollutants, f e r t i l i z e r s ) , or
i n t h e normal c o u r s e of m i n e r a l weather-
ing or r a i n f a l l additions. The t o t a l
c a p a c i t y o f t h i s bed f o r c a t i o n exchange
i n c o n i f e r o u s f o r e s t s o i l s r a n g e s from 40
t o 300 gm e q u i v a l e n t s per meter s q u a r e d
t o a meter d e p t h ( Z i n k e , & g . , 1 9 7 9 ) .
Whether t h e p o l l u t a n t added h a s a s i g n i -
f i c a n t e f f e c t depends upon t h e n a t u r a l
base l i n e composition o f o t h e r c t i o n s o n
t h e s o i l column ( u s u a l l y H , Ca¥f, Mg ,
K', N a ) maintained. by t h e f o r e s t and
the lyotropic series ( l e a c h i n g pre-
cedence) of t h e s e e l e m e n t s . The f o r e s t
s o i l a l s o h a s a n i o n exchange c a p a c i t y
a l t h o u g h u s u a l l y of l e s s e r amount. Thus,
depending upon t h e n a t u r e of t h e p o l l u -
t a n t a d d i t i v e , and whether i t a t t a i n s
i o n i c form, i t may be s u b j e c t t o scrub-
b i n g out a t t h e s o i l exchange complex
stage of t h e elemental cycle. T h i s would
a p p l y t o t h e hydrogen and s u l f a t e o f a c i d
r a i n , o r t h e c a t i o n i c forms o f some
m e t a l l i c p o l l u t a n t s such as l e a d o r z i n c .
L a b o r a t o r y s i m u l a t i o n through l e a c h i n g
w i t h s u c c e s s i v e i n c r e m e n t s of t h e p o l l u -
t a n t such as a c i d r a i n can be made
r e a d i l y t o d e t e r m i n e t h i s c a p a c i t y , and
an example o f t h i s f o l l o w s l a t e r .
The s o i l h a s o t h e r c a p a c i t i e s f o r
storage of a d d i t i v e s i n addition t o t h a t
of t h e i o n exchange c a p a c i t i e s . Either
f o l l o w i n g b r e a k t h r o u g h o r s a t u r a t i o n of
t h e exchange c a p a c i t y , an a d d i t i v e e l e -
ment may be s t o r e d on t h e m e t a l l i c o x i d e
complex of t h e s o i l , o r as i n s o l u b l e pre-
c i p i t a t e s or oxides. The r e l e a s e and
u p t a k e o f e l e m e n t s i n t o s t o r a g e i n asso-
c i a t i o n with t h e m e t a l l i c oxides ( u s u a l l y
i r o n and manganese) w i l l depend upon t h e
pH and redox p o t e n t i a l of t h e s o i l as
reviewed by Jenne ( 1 9 6 8 ) . R e t e n t i o n w i l l
be l e a s t under t h e r e d u c i n g c o n d i t i o n s
brought about by l a r g e amounts of o r g a n i c
m a t t e r and p o o r l y d r a i n e d - wet condi-
tions.
Thus, t h e f o r e s t s o i l i s a major
determinant i n the f a t e of t h e chronic
p o l l u t a n t element added t o t h e f o r e s t .
The b r e a k t h r o u g h c a p a c i t y of t h e s o i l
f o r t h e added m a t e r i a l w i l l depend upon
t h e c u r r e n t i n p u t o f s i m i l a r e l e m e n t s by
t h e n a t u r a l c y c l i n g of c a t i o n s from t h e
t r e e c o v e r , t h e i n p u t s by c u r r e n t m i n e r a l
weathering, t h e current additions of
hydrogen from r a i n f a l l and o r g a n i c com-
pounds produced by t h e f o r e s t , t h e deple-
t i o n o f b a s i c m e t a l l i c c a t i o n s t a k e n up
by t h e t r e e growth as t h e t r e e s a g e on
the s i t e . The g r e a t e r t h e s t o r a g e capa-
c i t y of t h e s o i l e i t h e r as i o n exchange
o r m e t a l l i c oxide (hydrous) c a p a c i t y t h e
g r e a t e r the buffering capacity of the
f o r e s t on t h e i n p u t o f p o l l u t a n t s .
It w i l l be o f i n t e r e s t now t o review
some o f t h e e x p e r i e n c e o f how t h e v a r i o u s
p o l l u t a n t s behave i n r e l a t i o n t o t h e s e
processes of elemental cycles in a
forest.
POLLUTANT ADDITIVES TO FOREST CYCLES
Any of t h e e l e m e n t s added t o t h e
f o r e s t as p o l l u t a n t s can i n f l u e n c e e l e -
mental c y c l i n g , e i t h e r by a c t i n g as
n u t r i e n t e l e m e n t s r e q u i r e d f o r t h e growth
o f t r e e s , by changing t h e redox s t a t e o f
any p a r t of t h e f o r e s t , o r by changing
t h e pH a t some p o i n t i n t h e system. Some
o f t h i s i n t e r a c t i o n may b e p h y s i o l o g i c a l
i n t h e v e g e t a t i o n , o r i n t h e s o i l micro-
f l o r a , o r may be a m a t t e r o f s o i l chemis-
try. Growth r a t e s of t h e v e g e t a t i o n may
b e changed, t h u s a f f e c t i n g t h e r a t e s of
e l e m e n t a l c y c l i n g . Each p o l l u t a n t may
have s e p a r a t e e f f e c t s depending upon i t s
chemical and p h y s i o l o g i c a l n a t u r e , and
t h e r e may b e s y n e r g i s t i c e f f e c t s w i t h
combinations of a d d i t i v e s . Some o f t h e
major c h r o n i c p o l l u t a n t s w i l l be con-
s i d e r ed s e p a r a t e l y .
Carbon Compounds &i Accompanying Oxi-
dan-fcs
Carbon a d d i t i o n s t o t h e f o r e s t may
b e i m p o r t a n t because o f t h e r o l e t h e y
p l a y i n p l a n t growth and s u b s e q u e n t
mineral cycling. F i n l a y s o n and P i t t s
(1976) have r e p o r t e d t h a t c a r b o n may
account f o r 45$ of t h e mass o f smog aero-
s o l . Much o f t h i s may b e t h e r e s u l t o f
secondary r e a c t i o n s c r e a t i n g c a r b o x y l i c
acids, esters, carbonyl compounds,
a l c o h o l s , p e r o x i d i c polymers, l o n g c h a i n
a l k a n e s and a l k e n e s , and f a t t y a c i d s . In
a d d i t i o n , c a r b o n i n t h e forms of v a r i o u s
compounds i s a world-wide atmospheric
p o l l u t a n t due t o human a c t i v i t i e s o x i d i z -
i n g reduced carbon f o r e n e r g y , and t o
i n d i r e c t e f f e c t s such as enhanced oxida-
t i o n of s o i l o r g a n i c m a t t e r due t o c l e a r -
ing forestland for agriculture. Thus
carbon d i o x i d e as a c h r o n i c p o l l u t a n t h a s
been measured by o b s e r v a t i o n s a t Mauna
Loa (Hawaii) t o b e r i s i n g a t t h e r a t e o f
from .35 t o 1 .79 ppm per y e a r t o a l e v e l
of 325 ppm i n 1 974 (Hobbs, g &.
P e t e r s o n (1969) i n a review s t a t e d t h a t
, 1974).
carbon d i o x i d e a t i t s p r e s e n t l e v e l i n
t h e atmosphere i s s t i l l l i m i t i n g t o p l a n t
growth and t h a t t h e p r o d u c t i v i t y of
p l a n t s should i n c r e a s e as t h e C o n c o n t e n t
increases. A r e s u l t i n g increase of plant
and f o r e s t growth of 5$ by t h e y e a r 2000
is a ' p r e d i c t e d , and t h i s would i n c r e a s e
t h e r a t e s of u p t a k e o f n e c e s s a r y growth
e l e m e n t s from t h e s o i l , t h u s i n c r e a s i n g
r a t e s of elemental cycling. This a l l
assumes o t h e r e l e m e n t s o r r e q u i r e m e n t s
such as water a r e n o t l i m i t i n g t o p l a n t
growth. On t h e o t h e r hand, r e d u c t i o n i n
p l a n t growth may occur where t o x i c c a r b o n
compounds from p o l l u t i o n c a u s e damage t o
t r e e f o l i a g e , thus reducing r a t e s of
mineral cycling. Ozone produced as a
secondary p r o d u c t from u l t r a v i o l e t radia-
t i o n on hydrocarbons i n t h e atmosphere
may r e d u c e p l a n t growth. Evans, g &.
( 1 974) found ozone c o n t e n t s of 0.1 ppm a t
t h e t o p of a smoke plume from a f o r e s t
fire. However p l a n t s s y n t h e s i z e methyl
c h l o r i d e a c c o r d i n g t o Lovelock (1 9 7 5 ) ,
and t h i s may d e s t r o y s u c h added ozone.
The o x i d a n t s formed from t h e o r g a n i c pol-
l u t a n t s i n t r o d u c e d t o t h e atmosphere w i l l
have a d v e r s e e f f e c t s upon v a r i o u s amino
a c i d s i n p l a n t s r e s u l t i n g i n damage
(Haagen-Smit , 1 9 5 8 ) . F a t t y a c i d s on
p l a n t s u r f a c e s may be a l t e r e d by t h e
s i n g l e t oxygen r e s u l t i n g from NO2, ben-
z a l d e h y d e , and p o l y n u c l e a r hydrocarbons,
increasing saturated acid content of t h e
f o l i a g e a c c o r d i n g t o Dowty &. (1 9 7 3 ) .
Other oxidants such as p e r o x y a c e t y l
n i t r a t e ( P A N ) , hydroxyl OH, hydroperoxyl
HO , a l s o have similar e f f e c t s on vegeta-
ti$n. T h e i r e f f e c t s as o x i d a n t s a r e
d e f i n e d by t h e i r c a p a b i l i t y t o o x i d i z e
i o d i d e i o n i n aqueous s o l u t i o n of potas-
sium i o d i d e a c c o r d i n g t o Kuntz, e t 9.
(1 973). Also, t h e i r a c t i o n may b e s y n -
e r g i s t i c , s o t h a t a l t h o u g h one o r a n o t h e r
i s i n low a t m o s p h e r i c c o n c e n t r a t i o n t h e
sum of t h e o x i d a n t s must be c o n s i d e r e d i n
e f f e c t on t h e v e g e t a t i o n . Also, t h e
v o l a t i l e t e r p e n e s produced by t h e f o r e s t
i t s e l f must b e t a k e n i n t o a c c o u n t , and
a c c o r d i n g t o Rasmussen (1 9 7 0 ) , as many as
1008 of t h e s p e c i e s i n some w e s t e r n coni-
f e r o u s f o r e s t s may produce such v o l a t i l e
hydrocarbons.
Nitrogen
N i t r o g e n as i t c y c l e s i n t h e f o r e s t
i s i n reduced form i n t h e p l a n t and f o l -
lowing m i n e r a l i z a t i o n to ammonia in
decomposing d e t r i t u s i n t h e s o i l it is
g r a d u a l l y o x i d i z e d t o n i t r a t e where i t i s
a g a i n t a k e n up by p l a n t s t o b e reduced i n
v a r i o u s n i t r o g e n compounds. Nitrogen is
added as a c h r o n i c a i r p o l l u t a n t as
ammonium, and n i t r a t e n i t r o g e n which a l s o
i s t h e end p r o d u c t o f o x i d a t i o n of 80 by
ozone. Where n i t r o g e n i s l i m i t i n g i n t h e
f o r e s t as an e s s e n t i a l element t h i s
s h o u l d s t i m u l a t e f o r e s t growth and decom-
p o s i t i o n p r o c e s s e s provided o t h e r e l e -
ments a r e n o t l i m i t i n g . However, i f
o t h e r e l e m e n t s a r e l i m i t i n g t h i s would
t e n d t o emphasize t h e o t h e r d e f i c i e n c i e s .
A d d i t i o n s as a c h r o n i c p o l l u t a n t should
show similar symptoms t o t h o s e o f t h e
a d d i t i o n o f n i t r o g e n as a f e r t i l i z e r .
Thus f o l i a r n i t r o g e n c o n t e n t s should be
h i g h e r . A r e s u l t similar t o t h i s encoun-
t e r e d i n Big cone s p r u c e f o l i a g e i n t h e
San B e r n a r d i n o mountains i s r e p o r t e d
l a t e r i n t h i s paper.
-
S-u l f u r
The s u l f u r c y c l e i n f o r e s t s i s such
t h a t t h e s u l f u r i s i n reduced form i n
a s s o c i a t i o n w i t h o r g a n i c compounds i n t h e
p l a n t s , b u t i s o x i d i z e d upon decomposi-
t i o n i n the organic d e t r i t u s returned t o
the soil. The decomposing s o i l micro-
f l o r a c o n v e r t t h e s u l f u r t o s u l f a t e . The
d e g r e e t o which t h i s t a k e s p l a c e depends
upon t h e redox p o t e n t i a l of t h e s o i l . For
example, i f i t i s v e r y low, t h e s u l f u r
w i l l be r e t a i n e d i n s u l f i d e form, f r e -
q u e n t l y t y i n g up heavy m e t a l s .
The forms o f s u l f u r added i n c h r o n i c
atmospheric p o l l u t i o n have been SO?,
SO ", w i t h a s s o c i a t e d c a t i o n s H , NH
a n t HSO as determined i n t h e p o l l u t h ~
plume trim S t . L o u i s M i s s o u r i by Charl-
s o n , g 9. ( 1 9 7 3 ) . They found t h a t
t r o p i c a l a i r masses were dominated by t h e
more a c i d i c NH HSO w h i l e n o r t h e r n a i r
masses were l e s s a c i d i c and dominated by
(NH ) SO (1 9 7 5 ) . U s u a l l y t h e s u l f u r i n
the4a?mo%-phere w i l l be o x i d i z e d t o s u l -
f a t e and t h i s w i l l o c c u r e i t h e r as
ammonium o r c a l c i u m s u l h a t e d u s t a c c o r d -
i n g t o daagen-Snit (1959), and s i n c e
t r a p p i n g s u r f a c e s o f v e g e t a t i o n a r e aero-
b i c , any reduced s u l f u r w i l l be o x i d i z e d
t o sulfate in the foliage.
S i n c e s u l f u r is a n e s s e n t i a l element
f o r p l a n t growth, t h e a d d i t i o n s may o r
may n o t b e d e l e t e r i o u s . A c t u a l d e l e t e r i-
ous e f f e c t s would occur t o t r e e s i f s u l -
f u r d i o x i d e were t h e main a d d i t i v e . This
would occur o n l y c l o s e t o t h e e m i s s i o n
s o u r c e as h a s happened a t K e n n e t t , C a l i -
f o r n i a ; Copper B a s i n , Tennessee; and Sud-
bury, Ontario. A t g r e a t e r d i s t a n c e t h i s
would be o x i d i z e d . Presumably i f t h e
p o l l u t a n t were added i n extreme amounts,
t h e t i t r a b l e a c i d i t y produced would b r e a k
t h r o u g h t h e s o i l exchange c a p a c i t y a f t e r
depleting other cations present. An
example o f a s i m u l a t i o n o f t h i s i s
offered i n t a b l e 2 explained l a t e r .
T r a c e Elements
' .
Washout, P a r t i c u l a t e s ,
lusts
Most o f t h e e l e m e n t s o t h e r t h a n C ,
N, and S w i l l be added i n d u s t and r a i n
as c h r o n i c p o l l u t a n t s . The p a r t i c u l a t e s
i n atmospheric p o l l u t i o n c o n t a i n elements
such as lead, sodium, magnesium,
aluminium, vanadium, and z i n c i n Los
Angeles t y p e smog a c c o r d i n g t o F i n l a y s o n
and P i t t s ( 1 976). Sedimentary c o r e s
t a k e n o f f s h o r e i n t h e P a c i f i c Ocean by
Bruland, e t S., ( 1 9 7 4 ) , i n d i c a t e d t h a t
l e a d , s i l v e r , c o p p e r , z i n c , chromium,
nickel, molybdenum, and cobalt are
c h r o n i c p o l l u t a n t s i n t h e Los Angeles
area. Lead, i r o n , manganese, n i c k e l ,
copper and z i n c were analyzed i n p r e c i p i -
t a t i o n as common t r a c e element p o l l u t a n t s
throughout t h e U.S. a c c o r d i n g t o Lazarus
-
e t a l . (1 9 7 0 ) . Beryllium is common i n
i n d u s t r i a l d u s t s n e a r s o u r c e s . The amount
of t r a c e element f a l l o u t l e s s e n s as t h e
s q u a r e o f t h e d i s t a n c e from s o u r c e s
a c c o r d i n g t o B e r t i n e & Goldberg ( 1 9 7 4 ) .
Cannon and B o l e s (1962) found t h a t high-
ways r e p r e s e n t e d l i n e a r s o u r c e s f o r l e a d ,
the amount in vegetation decreasing
r a p i d l y w i t h d i s t a n c e from t h e s o u r c e .
Dedolph e t ( 1 970) found t h a t t h i s
d i m u n i t i o n was l o g a r i t h m i c w i t h d i s t a n c e
f o r p a r t i c u l a t e l e a d . However, some e l e -
ments such as cadmium, n i c k e l , l e a d , and
z i n c may b e v a p o r i z e d a t t h e s o u r c e and
carried longer distances according t o
L a g e r w e r f f & S p e c h t (1 9 7 0 ) .
The t r a c i n g o f i n p u t o f t r a c e e l e -
ments i n t h e e l e m e n t a l c y c l e s i n vegeta-
t i o n h a s b e e n a t t e m p t e d by many i n v e s t i -
gators. C h e s t e r and S t o n e r (1 973) used
a n e n r i c h m e n t f a c t o r i n which a r a t i o o f
t h e element t o iron. i n t h e p a r t i c u l a t e
b e i n g added i s d i v i d e d by t h e a v e r a g e
r a t i o of t h e element t o i r o n i n t h e
earth's crust. They f o u n d that tin,
l e a d , and z i n c most o f t e n were t h e e l e -
ments enriched by p o l l u t i o n . Peirson, e t
-
a l . (1 9 7 4 ) employed a n e n r i c h m e n t f a c t o r
i n t h e form o f t h e r a t i o o f t h e e l e m e n t
t o Scandium c o n t e n t , compared t o t h e same
for the local soils. They f o u n d t h e
s o i l s e n r i c h e d f o r V, Co, N i , Zn, A s , S e ,
Sb, and Pb t h i s way. S i m i l a r enrichment
r a t i o s f o r o t h e r p o l l u t a n t s should iden-
t i f y augmentations of t h e elements i n
elemental cycles i n f o r e s t s .
The c o n t e n t s o f p o l l u t a n t s i n t h e
plants should be identifiable by
anomalies i n composition. Thus, Schack-
l e t t e and Connor (1 9 7 3 ) u s e d t h e r e g i o n a l
v a r i a t i o n of vanadium i n S p a n i s h moss
( ~ i l l a n d s i a )along t h e gulf coast of t h e
U.S. t o i d e n t i f y a r e a s where c h r o n i c p o l -
l u t i o n b y a i r b o r n e vanadium o c c u r s . Con-
t e n t s as h i g h as 560ppm V were found i n
c e r t a i n a r e a s , and t h e s e were assumed t o
r e p r e s e n t t h e o u t p u t from o i l r e f i n e r i e s
u s i n g V e n e z u e l a n c r u d e o i l which i s h i g h
i n vanadium. In soil-vegetation cycling,
vanadium i s f r e q u e n t l y h e l d a t t h e s o i l -
r o o t i n t e r f a c e due t o i m m o b i l i z a t i o n i n
t h e oxidized s t a t e (pentavalent form).
Thus, t h e r e d o x s t a t e o f the soil-
v e g e t a t i o n s y s t e m and i t s components i n
t h e f o r e s t may d e t e r m i n e where a c h r o n i c
p o l l u t i n g e l e m e n t may b e i m m o b i l i z e d .
Elements undergoing changes i n s o l u b i l i t y
w i t h v a r i o u s o x i d a t i o n s t a t e s may b e
i m m o b i l i z e d a t v a r i o u s i n t e r f a c e s where
pH o r Redox c h a n g e . Chromium, vanadium,
manganese, and i r o n a r e s u s c e p t i b l e t o
this.
Some o f t h e c h r o n i c p o l l u t a n t s added
t o t h e e l e m e n t a l c y c l e o f f o r e s t s may
l a t e r b e r e l e a s e d from t h e f o r e s t by
vaporization. F o r example, C u r t i n e t a l .
( 1 974) found t h a t t i n i s t r a n s p i r e d i n
v a p o r s from c o n i f e r s t o t h e amount o f
23-80 ppm i n t h e r e s i d u e o f t h e v a p o r .
They f o u n d t h a t some c o n i f e r o u s t w i g s
c o n t a i n e d up t o 6-40 ppm t i n i n t h e a s h .
The t i n w a s added as a t m o s p h e r i c p o l l u -
t a n t i n d u s t from i n d u s t r i a l a r e a s . How-
e v e r , n o m e n t i o n w a s made o f t h i s as
being deleterious.
The inputs of dry particulate
m a t e r i a l s as d u s t s may b e washed o f f b y
rain. T h i s w a s h o f f may b e a m a j o r pro-
c e s s of c y c l i n g o f t h e added e l e m e n t t o
soils. Carlson - et - a l . ( 1 976) found t h a t
a s i m u l a t e d r a i n removed 45% o f a n
a p p l i e d a e r o s o l o f PbC12, and t h a t l i g h t
m i s t y r a i n i s most e f f e c t i v e . H e i c h e l &
Hankin ( 1 9 7 2 ) found t h a t t h e p a r t i c u l a t e s
i n which l e a d a d h e r e d t o t r e e s a v e r a g e d 7
micrometers i n diameter. C h l o r i n e and
Bromine were t h e main a s s o c i a t e d n e g a t i v e
elements associated with them. The
assessment o f t h e i n t e n s i t y o f i n p u t o f
c h r o n i c p o l l u t a n t s s u c h as t h e s e h a v e
b e e n made b y a n a l y z i n g c o n c e n t r a t i o n s o n
t r e e b a r k b y L o t s c h e r t ( l 9 7 7 ) , and Grod-
z i n s k a (1 9 7 7 ) . Some o f t h e s e accumula-
t i o n s a r e washed down t h e t r e e t r u n k b y
s t e m f l o w and may b e a c c u m u l a t e d i n t h e
s o i l at t h e t r e e base.
POLLUTANT INPUT I N PRECIPITATION
Washout as w e l l as i n p u t o f e l e m e n t s
i n s o l u t i o n i n p r e c i p i t a t i o n is a major
path of input t o t h e elemental cycling
t h a t o c c u r s i n f o r e s t s . F o r example, t h e
n i t r o g e n c o n t e n t s t o r e d i n a 1000 y e a r
o l d redwood f o r e s t i s a b o u t t h e amount
c o n t r i b u t e d by r a i n f a l l d u r i n g t h a t t i m e
p e r i o d ( ~ i n k e ,5 g . , 1 9 7 9 ) . The geo-
c h e m i s t r y o f p r e c i p i t a t i o n w a s reviewed
t h o r o u g h l y b y C a r r o l l (1 9 6 2 ) . She found
t h a t t h e c a t i o n s which b a l a n c e t h e i o n s
i n r a i n w a t e r were mostly b a s i c m e t a l l i c
elements near coast lines or arid
regions, but t h a t along the storm v e c t o r
from t h e s e s o u r c e s a n d o v e r r e g i o n s w i t h
more v e g e t a t i o n , t h e r e would t e n d t o b e
a c i d i f i c a t i o n due t o l e s s d u s t o r o c e a n
aerosols t o provide t h e b a s i c m e t a l l i c
cations. The c o n t r o l o f wind e r o s i o n
o v e r t h e G r e a t P l a i n s t h u s would t e n d t o
add t o t h e a c i d f i c a t i o n of rainfall
downwind from t h e a r e a . However t h e
c u r r e n t e m p h a s i s o n p o l l u t i o n h a s added
the consideration of sources of anions i n
p r e c i p i t a t i o n t h a t come from b u r n i n g f o s -
sil fuels. A s C a r r o l l (1962) mentioned,
a pH below 5.7 i n d i c a t e s t h a t h y d r o g e n i s
i o n i z i n g t o meet t h e n e c e s s a r y b a l a n c e
with anions.
The e f f e c t o f p r e c i p i t a t i o n i n p u t
upon n u t r i e n t e l e m e n t c y c l i n g i n a f o r e s t
w i l l t e n d t o b e s p e c i f i c t o t h e s i t e con-
d i t i o n s . T h i s would b e d e t e r m i n e d by t h e
following l o c a l factors: 1. background
n a t u r a l composition of t h e s o i l s o l u t i o n .
2. t h e c a t i o n exchange c a p a c i t y o f t h e
s o i l and t h e c o m p o s i t i o n o f c a t i o n s occu-
p y i n g t h i s c a p a c i t y . 3. t h e volume o f
w a t e r e n t e r i n g as r a i n f a l l , 4. t h e a c i d
b a s e b a l a n c e o f t h e r a i n f a l l , 5. t h e
n a t u r e of t h e v e g e t a t i o n s p e c i e s w i t h
 r e g a r d t o i n t e n s i t y of c y c l i n g b a s i c
m e t a l l i c e l e m e n t s , and perhaps o t h e r s
unique t o t h e s i t e . The e f f e c t s of i n p u t
i n p r e c i p i t a t i o n w i l l most obvious on
i n p r e c i p i t a t i o n w i l l b e most o b v i o u s on
s i t e s w i t h s o i l s h a v i n g low exchange
m a t t e r c o n t e n t s o i l s ) and w i t h s p a r s e
slow growing v e g e t a t i o n ( p i n e s , s p r u c e s ,
h e a t h l a n d ) , and i n c l i m a t e s w i t h l a r g e
amounts o f p r e c i p i t a t i o n .
FOREST ECOSYSTEM EFFECTS
The m i n e r a l c y c l i n g in a forest
t a k e s p l a c e i n t h e c o n t e x t o f an ecosys-
tem of p r o c e s s e s l i n k i n g t h e v a r i o u s l i v -
i n g organisms and t h e components o f t h e
environment. Chronic a t m o s p h e r i c p o l l u -
t i o n i s one o f t h e s e components. The
e l e m e n t s added w i l l be u t i l i z e d by t h e
Producers i n t h e f o r e s t . These a r e t h e
organisms which c a r r y o u t t h e r e d u c t i o n
of carbon t o c a r b o n compounds which
comprise t h e f o r e s t . T h i s f i x e d carbon
t h e n s e r v e s as an energy s o u r c e f o r t h e
Decomposers i n t h e f o r e s t which a l s o have
requirements f o r t h e n u t r i e n t elements
e i t h e r i n t h e biomass o f t h e p r o d u c e r s o r
f u r n i s h e d d i r e c t l y by t h e s o i l . Thus
p o l l u t a n t elements w i l l i n t e r a c t i n the
c y c l e between P r o d u c e r s and Decomposers,
w i t h s t o r a g e o f e l e m e n t s and p o s s i b l e
s i n k s o f u n a v a i l a b l e m a t e r i a l accumulat-
ing i n t h e s o i l portion of t h e cycle.
Effects -
of Chronic P o l l u t i o n on Producers
These effects can range from
enhancement o f p r o d u c t i v i t y t o a marked
d e c r e a s e depending upon whether t h e addi-
. t i v e is a n e l e m e n t c u r r e n t l y l i m i t i n g
growth o r i s a t o x i c element. If toxic,
t h e e f f e c t s w i l l show up i n f o l i a r dam-
a g e , o r i n t e r f e r e n c e w i t h v a r i o u s physio-
l o g i c a l processes. The syposium proceed-
i n g s e d i t e d by Naegele (1973) c o n t a i n s
many d e s c r i p t i o n s o f t h e s e t o x i c e f f e c t s .
Mc Cune i n t h i s p u b l i c a t i o n f e l t t h a t
such t o x i c e f f e c t s should be s p e c i f i c
with regard t o t h e Receptor ( l e a f , t r e e ,
f o r e s t ) , t h e P o l l u t a n t , t h e E v e n t , and
t h e Environment. Acute e f f e c t s a r e r e l a -
t i v e l y e a s y t o d e t e r m i n e , as i n t h e d e a t h
o f l e a v e s n e a r a s m e l t e r r e l e a s i n g SO2.
Obviously t h i s would d e c r e a s e t h e r a t e o f
elemental cycling at the s i t e along with
a b r i e f s u r g e o f added m a t e r i a l s t o t h e
s o i l as t h e d e f o l i a t i o n t o o k p l a c e and
l e a v e s dropped t o t h e s o i l t o b e decom-
posed. Whether t h i s is a l o s s t o t h e
s i t e depends upon t h e c a p a c i t y o f t h e
u n d e r l y i n g s o i l as d i s c u s s e d i n r e l a t i o n
t o d e f o l i a t i o n by h e r b i c i d e s by Zinke
(1974). The e f f e c t s a r e more d i f f i c u l t
t o a s s e s s i n t h e c a s e o f c h r o n i c low
l e v e l p o l l u t i o n as d i s c u s s e d by 'Feder
(1 9 7 3 ) . The changes i n p r o d u c t i v i t y
would need t o be s t a t i s t i c a l l y s i g n i f i -
c a n t b e f o r e and a f t e r t h e i n i t i a t i o n o f
t h e p o l l u t i o n . M i l l e r (1 973) h a s made an
approach t o i n d i c e s of l e v e l of c h r o n i c
p o l l u t i o n i n terms o f v i s u a l c h a r a c t e r i s -
t i c s on t h e t r e e s ; n e e d l e r e t e n t i o n , nee-
dle length, needle chlorosis, branch
d e a t h i n t h e c a s e of ponderosa p i n e .
F o l i a g e composition should i n d i c a t e such
i n p u t s , and a paper by Arkley and Glauser
i n t h i s symposium c o v e r s t h i s . Also an
example i n t h e c a s e o f Big Cone Spruce
foliage is presented in table 1.
McBride, e t a l . (1975) made measurements
of a c t u a l g r o w t h r a t e r e t a r d a t i o n o f 26%
f o r h e i g h t and volume growth o f ponderosa
In a g r i c u l t u r a l plants, White
1974) found a d e c r e a s e i n C02 u p t a k e due
t o s y n e r g i s t i c e f f e c t s of NO + SO2 when
each were p r e s e n t a t l e v e f s n o t con-
s i d e r e d i n h i b i t i n g t o growth. S t i m u l a t i o n
of primary p r o d u c e r s by p o l l u t i o n i s n o t
o f t e n r e p o r t e d due t o t h e n e g a t i v e conno-
t a t i o n of t h e t e r m . However, Schnappinger
(1975) found a n i n c r e a s e i n growth d u e t o
response t o z i n c contained i n f l y ash.
There a r e numerous v a r i a b l e s which
may a f f e c t t h e i n t e n s i t y w i t h which a
g i v e n l e v e l of p o l l u t a n t w i l l a f f e c t t h e
p r o d u c e r s i n an ecosystem. Thus t h e
t r a p p i n g e f f i c i e n c y of t h e f o l i a g e s u r -
f a c e i s d i f f e r e n t f o r v a r i o u s s p e c i e s as
d i s c u s s e d by Zinke ( 1 9 6 6 ) , t h e v e l o c i t y
of t h e a i r p a s t t h e t r a p p i n g s u r f a c e as
h a s been e v a l u a t e d by Hori (1 953) and
S l i n n ( 1 976). Following t r a p p i n g t h e
m a t e r i a l may n o t be absorbed by t h e s u r -
f a c e as found by Motto ( 1 9 7 0 ) . T h i s i s
partially attributable t o the particle
s i z e a c c o r d i n g t o Natusch & Wallace
( 1 9 7 4 ) . There may be s y n e r g i s t i c e f f e c t s
a t t h e f o l i a g e s u r f a c e . iiovelock (1974)
has alluded t o t h e possible c r e a t i o n of
PAN on f o l i a g e s u r f a c e by t h e combination
of 0 and hydrocarbons as o c c u r s on dry-
ing linen.
Once absorbed by t h e producer t h e
p o l l u t i n g element w i l l e n t e r t h e elemen-
t a l c y c l e on t h e s i t e . The d i s p o s i t i o n
from t h e p l a n t may be n o t o n l y t h r o u g h
l e a f d r o p , b u t may b e by d i r e c t r o o t exu-
d a t i o n as Olson e t a1 (1 962) found w i t h
a d d i t i o n s of r a d i o a c t i v e Cesium added t o
a t r e e t r u n k . However upon b e i n g dropped
t o t h e s o i l as d e t r i t u s o r e n t e r i n g i n
o t h e r ways, t h e p o l l u t i n g e l e m e n t w i l l
t h e n b e a f f e c t i n g t h e decomposers i n t h e
forest.
on
Effects
nosers
-
of Chronic P o l l u t i o n Decom-
The e f f e c t s upon decomposers, t h e
s o i l m i c r o f l o r a and f a u n a t h a t complete
t h e e l e m e n t a l c y c l e s i n t h e s o i l , w i l l be
variable. A s w i t h t h e p r o d u c e r s which
f i x carbon d i o x i d e , t h e a d d i t i v e s t o t h e
s i t e t h r o u g h a i r p o l l u t i o n may i n c r e a s e
or decrease their activities. These
organisms r e q u i r e t h e same n u t r i e n t e l e -
ments as t h e p r o d u c e r s as w e l l as t h e
a d d i t i o n a l ones o f sodium, i o d i n e , and
vanadium f o r t h e s o i l f a u n a . If t h e pol-
l u t a n t s a r e added i n t o x i c q u a n t i t i e s
t h e y would presumably r e t a r d t h e decompo-
s i t i o n of o r g a n i c m a t t e r i n t h e s o i l and
t h u s slow o r b l o c k e l e m e n t a l c c l i n g .
They may b e s u b l e t h a l o r even mutagenic
t o t h e s e organisms a c c o r d i n g t o S t o t z k y
1974).
The d e t r i t u s o r l i t t e r l a y e r on t h e
f o r e s t f l o o r is t h e f i r s t major a r e a o f
a c t i v i t o f t h e decomposers. Wittkamp &
Frank (1 969) found t h a t l i t t e r samples
impregnated w i t h i n t r o d u c e d e l e m e n t s such
as C o b a l t 60 and Cesium 137 had loga-
rithmic r e l a t i o n s h i p with time i n the
f r a c t i o n r e m a i n i n g , and t h a t l o s s c o e f f i -
c i e n t s were d i f f e r e n t f o r d i f f e r e n t e l e -
ments. These v a r i e d w i t h t h e s p e c i e s of
t r e e yielding the detritus. The s t o r a g e
p e r i o d of t h e a d d i t i v e element depended
upon r e l a t i v e chemical m o b i l i t y of t h e
e l e m e n t , c o m p o s i t i o n of o t h e r e l e m e n t s
p r e s e n t , and t h e m o i s t u r e and t e m p e r a t u r e
conditions of the s i t e .
If a p o l l u t a n t i s an element which
i s l i m i t i n g r a t e o f decompostion, t h e
e f f e c t on t h e decomposers may b e p o s i -
tive. For example l e a f l i t t e r w i t h t o o
h i g h a ~ a r b o n / ~ i t r o g e nr a t i o ( n i t r o g e n
l i m i t i n g ) decomposes v e r y s l o w l y . Adding
n i t r o g e n as a c h r o n i c p o l l u t a n t would
a c c e l e r a t e t h i s decomposition and r e l e a s e
s t o r e d e l e m e n t s more r a p i d l y .
The s o i l f a u n a s e r v e a major t a s k i n
e l e m e n t a l c y c l i n g i n a f o r e s t ecosystem
by physically decomposing organic
detritus, reducing it in size and
i n c r e a s i n g s u r f a c e a r e a t o b r i n g about
f a s t e r r a t e s o f f u n g a l and b a c t e r i a l
decomposition. Earthworms of various
s p e c i e s were found by Gish & C h r i s t e n s e n
( 1 9 7 3 ) t o accumulate t r a c e e l e m e n t s from
particulate pollution in order of
i n c r e a s i n g a t o m i c w e i g h t : N i < Zn < Cd <
Pb. The s o i l f a u n a may s e r v e as a l i n k
f o r t r a n s f e r of n u t r i e n t e l e m e n t s t o
v a r i o u s p r e d a t o r s i n t h e f o r e s t ecosys-
tem.
S t o t z k y ( 1 974) characterizes the
s o i l as b o t h an a b i o t i c as w e l l as b i o t i c
s i n k f o r p o l l u t a n t s . E t h y l e n e and C O a r e
degraded by s o i l micro organisms. Fungi
i n s o i l m e t a b o l i z e v o l a t i l e s from p l a n t
f o l i a g e s u c h as e t h a n e , b u t a n e , , e t c .
Thus s o i l by v i r t u e o f t h e v a r i o u s organ-
i s m s present has a strong capability t o
detoxify pollutants. A t low c o n c e n t r a -
t i o n s p o l l u t a n t s may b r i n g about an
enrichment o f t h e micro organisms which
can u t i l i z e them.
The u l t i m a t e r o l e o f t h e decomposers
i s t h u s t o c y c l e e l e m e n t s from t h e vege-
t a t i o n t o the s o i l , usually oxidizing
them as e n e r g y s o u r c e s , and u l t i m a t e l y
r e l e a s i n g them t o t h e chemical and physi-
c a l p r o c e s s e s of t h e s o i l o r renewed
uptake by t h e r o o t s of t h e p r o d u c e r s .
Chronic Pollutants into Soil
The f i n a l f a t e of a d d i t i v e s t o t h e
ecosystem may b e t o remain i n t h e s o i l i n
i n s o l u b l e form. Thus once p r o c e s s e d by
t h e decomposers t h e e l e m e n t s may become
m i n e r a l p r e c i p i t a t e s which may a c t as
s i n k s ( N a t . Res. Council 1 9 7 7 ) . Thus
f l u o r i n e added t h r o u g h p o l l u t i o n may b e
precipitated i n the s o i l a s insoluble
CaF2. Page and Ganje (1970) showed t h a t
t h e t o p 2.5 cm. of s o i l w a s most a c t i v e
in trapping lead entering the soil-
v e g e t a t i o n system i n s o u t h e r n C a l i f o r n i a .
S o i l s sampled i n 1967 were compared w i t h
p r e v i o u s samples o b t a i n e d i n 191 9. An
average increase of 15-36 ppm lead
accounted f o r 1 / 5 of t h e l e a d c o n t a i n e d
i n t h e 47 b i l l i o n g a l l o n s of g a s o l i n e
which had been burned d u r i n g t h e p e r i o d
encompassed. Keaton ( 1937) added l a r g e
amounts of l e a d n i t r a t e t o s o i l (2784
ppm) and found t h a t v e r y l i t t l e ( 1 7 ppm)
remained i n s o l u b l e form. Lee & T a l l i s
( 1 9 7 3 ) found i t p o s s i b l e t o d a t e p e r i o d s
of l e a d p o l l u t i o n i n a p e a t bog i n G r e a t
B r i t a i n , f i n d i n g a peak i n 5 0 0 A.D. w i t h
t h e o p e r a t i o n of Roman l e a d m i n e s , and a
subsequent industrial peak beginning
about 1 460 AD. Similar b u t recent his-
t o r i c a l evidence of l e a d c o l l e c t i o n i n
ecosystem s i n k s w a s found i n wood c o n t e n t
o f l e a d d a t e d by a n n u a l r i n g s by Sheppard
& Funk ( 1 9 7 5 ) .
Soil organic matter generally
enhances t h e r o l e of s o i l as a s i n k f o r
added t r a c e e l e m e n t s . P u r r e t a1 ( 1 9 7 1 )
have noted t h a t pH w i l l d e t e r m i n e whether
s o i l is a source o r sink. Thus, a t neu-
t r a l t o s l i g h t l y a l k a l i n e pH; A s , B, C d ,
Go, Cu, F, Fe, Pb, Mn, N i , Sn, and Zn
w i l l t e n d t o b e immobilized, b u t Ca, Cr,
K Mg, Mo, Rb, Se, and S r may become more
available t o plants. Thus a n y t h i n g caus-
i n g a change i n s o i l pH may r e l e a s e pre-
v i o u s p o l l u t a n t s t h a t had t e m p o r a r i l y
become t r a p p e d i n t h e s o i l as a s i n k .
Various p r e c i p i t a t e s such as s o i l c a r -
b o n a t e s may form, a l s o depending upon pH.
The s o i l i o n exchange c a p a c i t y w i l l
r e t a i n e l e m e n t s i n a form r e a d y t o b e
Table 1: Big cone s p r u c e f o l i a g e a n a l y s e s across gradients of decreasing
chronic pollution i n t e n s i t y .

WESTERN SAN GABRIEL MTNS. Angeles N a t i o n a l F o r e s t

Location Weldon Canyon Woodwardia Canyon Cloudburst Can yon T i e Canyon Summit
NITROGEN
percent 1.80
rank 93

PHOSPHORUS
PPm 601
rank 7

N /P
ratio 20.2
rank 92

EASTERN SAN GABRIEL MTNS. Angeles N a t i o n a l F o r e s t

Location Lower San Antonio Camp Baldy Snowcrest Camp Big P i n e s
NITROGEN
pet.
rank

PHOSPHORUS
Ppm 1093
rank 25

N /P

ratio
rank

SAN BERNARDINO-SAN JACINTO MTNS. San Bernardino N.F.

Location Lower Waterman Upper Waterman
Cleveland N.F.
Crestline
-
Camp Angeles Keene Palomar
(young) (old) (young) (old)
NITROGEN
pot. 2.12 1.80 2.02 2.04 1.22 .99 1.06 1.13
rank 97 92 96 96 52 13 26 38

PHOSPHORUS
PPm 6 48 838 1016 1685 1658 1552 1544 1580
rank 1 7 18 . 81 79 70 69 72

N /P
ratio 32.7 21.4 19.8 12.2 7.3 6.4 6.8 7.1
rank 99 94 92 70 36 26 31 34

CHRONIC
POLLUTION HEAVIEST HEAVY MODERATE-LIGHT LOW NIL
INTENSITY
 T a b l e 2: S i m u l a t i o n of a c i d r a i n f a l l a d d i t i o n by l e a c h i n g a s u r -
f a c e s o i l developed under t h e i n f l u e n c e of a mature pon-
d e r o s a p i n e a t t h e a r i d l i m i t o f t h e o c c u r r e n c e of t h e
species.
( i n m i l l i e q u i v a l e n t s p e r 100 m s . f i n e e a r t h p o r t i o n ex-
c e p t l a s t column as i n d i c a t e d )

Exchangeable C a t i o n s
S o i l Treatment Exchange Remaining on S o i l
Capacity %

Leached s o i l s :
L e a c h a t e Composition-3 I
500 m l H20
+5.71 meq. H+
+11 . 4 0 meq. H+
+17.09 meq. H+
+22.9 meq. H+

1
' F i n e e a r t h p o r t i o n , 0-2.5 i n c h sampling h o r i z o n .
D i f f e r e n c e between t h e sum ( ~ a " + M~++
Exchange C a p a c i t y .
^-IThe 'H w a s added as 0.1N H C 1 t o the.500
s i n c e 1 m l of 0.1N H C 1 = 0.1 meq. H .
r e c y c l e d back t o t h e l i v i n g organisms on
t h e s i t e , and a l s o k e e p them from l e a c h -
i n g away from t h e s i t e . Depending upon
t h i s c a p a c i t y , e l e m e n t s added t h r o u g h
c h r o n i c p o l l u t i o n may b e r e t a i n e d i n t h e
e l e m e n t a l c y c l e o f a f o r e s t provided t h e y
a r e not trapped i n a s i n k o r leached
away.
FOLIAGE ANALYSES I N D I C A T I N G
FOLLrnION
The e f f e c t o f c h r o n i c a i r p o l l u t i o n
o f t h e t y p e t y p i c a l of t h e Los Angeles - SIMULATED A C I D RAINFALL LEACHING
San B e r n a r d i n o b a s i n on f o l i a r a n a l y s e s
of the Cone Spruce ( P s e u d o t s u g a
macrocarpaBi. Vasey) Mayr) w a s d e t e r m i n e d .
Samples o f t h e f o l i a g e were g a t h e r e d
t h r o u g h o u t t h e r a n g e o f t h e s p e c i e s , and
a l s o on g r a d i e n t s o f e l e v a t i o n o r d i s -
t a n c e from t h e a r e a s t y p i f i e d by "smog"
t o a r e a s r e l a t i v e l y f r e e o f "smog". Ana-
l y s e s were made o f 9 major e l e m e n t s i n
t h e v a r i o u s a g e c l a s s e s of t h e f o l i a g e .
The e l e m e n t s most o b v i o u s l y r e l a t e d t o
t h e i n t e n s i t y o f t h e smog and t h e i r rat-
ing i n a cumulative probability d i s t r i b u -
+ K' + ~a') and the
m l of d i s t i l l e d w a t e r ,
t i o n from l o w e s t to highest value are
shown i n t a b l e 1.
N i t r o g e n c o n t e n t of t h e f o l i a g e and
t h e r a t i o of n i t r o g e n t o phosphorus con-
t e n t were t h e most obvious i n d i c a t o r s o f
t h e e f f e c t of chronic air p o l l u t i o n o f
t h e Los Angeles t y p e . Where t h e r e w a s
a m b i g u i t y i n t h e n i t r o g e n v a l u e s as i n
San Antonio Canyon, t h e n i t r o g e n t o phos-
phorus r a t i o was t h e b e s t i n d i c a t o r i n
t h i s species.
A l a b o r a t o r y experiment w a s con-
ducted i n which samples o f a n e a r l y
c a t i o n s a t u r a t e d s o i l under t h e i n f l u e n c e
of ponderosa p i n e were l e a c h e d w i t h s o l u -
t i o n s c o n t a i n i n g s u c c e s s i v e i n c r e m e n t s of
H The s o i l samples were o b t a i n e d a t t h e
lower r a i n f a l l l i m i t o f ponderosa p i n e
where t h e s o i l was most l i k e l y t o b e
base saturated. F i v e s e p e r a t e t e n gram
samples of t h i s s o i l were l e a c h e d by
s o l u t i o n s which were r e s p e c t i v e l y 500 m l
of d i s t i l l e d w a t e r , and s o l u t i o n s t o
w h i c h i n c r e m e n t s o f 'H as H C 1 e q u i v a l e n t
t o 20$, 40$, 808, of t h e exchange capa-
c i t y o f t h e sample were added. The
r e s u l t s p r e s e n t e d i n t a b l e 2 show t h e
l e a c h i n g of t h e exchange complex of s o i l s
t h a t t a k e s p l a c e w i t h added i n c r e m e n t s o f
a cation. Presumably t h e a d d i t i o n s o f
i n c r e m e n t s of H i n a c i d r a i n f a l l would
have similar e f f e c t s which would have t o
be considered i n t h e context of other
c a t i o n s b e i n g c y c l e d by t h e f o r e s t , and
by t h e c o m p o s i t i o n o f c a t i o n s a l r e a d y on
t h e exchange complex of t h e s o i l . Follow-
i n g t h i s l e a c h i n g , each s o i l p o r t i o n w a s
analyzed for remaining exchangeable
metallic cations.
These d a t a show t h a t g i v e n t h e s t o -
chiometry o f t h e l e a c h i n g of c a t i o n s from
t h e s o i l exchange complex by H the
e f f e c t s of increments of acid r a i n f a l l t o
a g i v e n s o i l may b e s i m u l a t e d w i t h s o l u -
t i o n s having compositions s i m i l a r t o t h a t
of t h e r a i n f a l l . Also, as t h e increment
of H i n the precipitation increased,
e i t h e r b e c a u s e o f c o n c e n t r a t i o n o r of
volume o f r a i n f a l l l e a c h i n g t h e s o i l
would t e n d t o become hydrogen s a t u r a t e d ,
d i s p l a c i n g b a s i c m e t a l l i c c a t i o n s i n pro-
portion. In the f o r e s t s i t u a t i o n these
would be l e a c h e d o u t o f t h e s o i l p r o f i l e
i n a b a s e s a t u r a t e d s o i l , o r t a k e n up by
the vegetation.
CONCLUSIONS
T h i s r e v i e w of t h e l i t e r a t u r e and
t h e p r e s e n t a t i o n o f some o r i g i n a l work i s
applicable t o the e f f e c t of chronic a i r
p o l l u t i o n on m i n e r a l element c y c l i n g of
forests. The l i t e r a t u r e i s g e n e r a l ,
i l l u s t r a t i n g processes, but i s not s i t e
s p e c i f i c except f o r t h e l o c a t i o n of t h e
studies. S i n c e e f f e c t s a t any s p e c i f i c
s i t e may b e i n small i n c r e m e n t s each
y e a r , but severe i n cumulative e f f e c t s
o v e r a number o f y e a r s , a major remaining
problem i s t o f i n d ways o f d e t e r m i n i n g
s i t e s p e c i f i c indications of detrimental
e f f e c t s of s u c h c h r o n i c p o l l u t i o n . Some
p e r t i n e n t c o n c l u s i o n s from t h e l i t e r a t u r e
reviewed i n t h i s paper a r e :
1 . The f o r e s t v e g e t a t i o n as i s t r u e o f
any v e g e t a t i o n r e q u i r e s many e s s e n t i a l
growth e l e m e n t s which i t w i l l c y c l e i n
a d d i t i o n t o o t h e r s mobilized by l o c a l
m i n e r a l w e a t h e r i n g o r i n p u t t h r o u g h pre-
cipitation.
2. To t h e e x t e n t t h a t c h r o n i c p o l l u t i o n
adds elements normally required by
plants, o r c h e m i c a l l y mobile in the
p l a n t , t h e s e w i l l be c y c l e d much as t h e
o t h e r s on t h e s i t e .
3. The f o r e s t i s c r e a t e d i n a v a s t reduc-
t i o n r e a c t i o n wherein CO i s reduced t o
o r g a n i c compounds by Producers i n t h e
ecosystem, and s u b s e q u e n t l y o x i d i z e d i n
energy r e l e a s i n g s t e p s by t h e s e p l a n t s o r
by t h e Decomposers i n t h a t ecosystem.
4. The c y c l i n g of e l e m e n t s t a k e s p l a c e i n
t h e c o n t e x t o f t h e s e redox r e a c t i o n s , and
t h o s e e l e m e n t s which change s t a t e w i t h
redox p o t e n t i a l change a r e u s u a l l y i n t h e
reduced form i n t h e producer v e g e t a t i o n
and subsequently oxidized in energy
r e l e a s i n g s t e p s by t h e s e p l a n t s o r by t h e
decomposers on t h e s i t e .
5. Chronic a i r p o l l u t i o n a d d i t i v e s t o t h e
f o r e s t a f f e c t t h e c y c l i n g of e l e m e n t s n o t
o n l y by d i r e c t a d d i t i o n s of t h e p o l l u t i n g
e l e m e n t s t o t h e c y c l e s , b u t a l s o by
a f f e c t i n g t h e redox s t a t u s o f t h e absorb-
i n g p l a n t t i s s u e , and o t h e r p o r t i o n s o f
t h e s o i l - v e g e t a t i o n system.
6 . C o n c e n t r a t i o n p o i n t s and s i n k s f o r t h e
e l e m e n t s added i n c h r o n i c p o l l u t i o n e x i s t
i n t h e various p a r t s of t h e vegetation,
t h e s u r f a c e d e t r i t u s on t h e s o i l , and i n
t h e s o i l c a p a c i t i e s f o r a b s o r p t i o n on i o n
exchange; hydrous o x i d e complexes, and as
insoluble precipitates.
7. The e f f e c t s o f c h r o n i c p o l l u t a n t s
added t o a g i v e n f o r e s t and i t s e l e m e n t a l
c y c l i n g p r o c e s s e s w i l l be h i g h l y s i t e
s p e c i f i c ; depending upon c l i m a t e , d e g r e e
o f w e a t h e r i n g development of t h e s o i l ,
t h e t y p e of m i n e r a l s from which t h e s o i l
i s forming, t h e c h a r a c t e r i s t i c s o f t h e
f o r e s t s p e c i e s and t h e i r s t o c k i n g densi-
ties, t h e topographic s i t u a t i o n with
r e g a r d t o a i r f l o w , and t h e n a t u r e o f t h e
pollutant.
8. Local s i t e s p e c i f i c i n d i c a t o r s o f
i n t e n s i t y of p o l l u t i o n e f f e c t may be
v i s u a l as i n appearance and s i z e o f f o l i -
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ments and t h e i r r a t i o s t o background s i t e
e l e m e n t s , and s o i l assessment o f c a p a c i -
t i e s t o a b s o r b t h e added elements.
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 Forest Genetics and Air Pollutant stress'
2
S t e f a n Bialobok

A b s t r a c t : The breeding of t r e e s , p a r t i c u l a r l y c o n i f e r s ,
f o r c u l t i v a t i o n in r e g i o n s under t h e i n f l u e n c e of i n d u s t r y
i s of c o n s i d e r a b l e economic importance. The r o l e of g e n e t i c
polymorphism in t h e p r o c e s s of a d a p t a t i o n t o such environ-
ments i s d i s c u s s e d . R e s u l t s a r e p r e s e n t e d of r e s e a r c h on
t h e g e n e t i c v a r i a b i l i t y in S c o t s p i n e p o p u l a t i o n s growing
under t h e i n f l u e n c e of i n d u s t r y and i n r e g i o n s f r e e of
gaseous p o l l u t i o n . Methods of s e l e c t i n g t r e e p o p u l a t i o n s
t o l e r a n t t o i n d u s t r i a l emissions a r e discussed aimed a t
t h e i d e n t i f i c a t i o n of g e n e t i c markers of t o l e r a n c e .

I n o r d e r t o reduce l o s s e s in f o r e s t ecosystems a s w e l l a s i n some s p e c i e s of broadleaf trees

caused by i n d u s t r i a l emissions a t t e m p t s were made
t o determine t h e s e n s i t i v i t y of trees (depending
on t h e i r g e n e t i c p r o p e r t i e s ) t o a c u t e i n j u r y of
t h e a s s i m i l a t i o n a p p a r a t u s by t h e most common
gaseous p o l l u t a n t s . The degree of p l a n t i n j u r y
by g a s e s i s dependent a l s o on t h e system of
e c o l o g i c a l f a c t o r s o p e r a t i n g in t h e environment.
GENETIC VARIATION OF TOLERANCE BETWEEN AND WITH-
I N SPECIES
So f a r a wide s c a l e of g e n e t i c v a r i a b i l i t y was
observed w i t h i n v a r i o u s s p e c i e s of t r e e s and
shrubs i n t h e i r s e n s i t i v i t y t o airborne pollution.
The r e s u l t s l i s t e d i n reviews (Davis e t a l . , 1976,
Bialobok 1979) a r e only approximate. It appears
t h a t most t o l e r a n t a r e some s p e c i e s of broadleaf
trees. Coniferous t r e e s which a r e of g r e a t e r
economic importance a r e g e n e r a l l y c h a r a c t e r i s e d
by low t o l e r a n c e .
It was a l s o found t h a t t h e r e is g e n e t i c
v a r i a b i l i t y in response t o SO^, O3 and HF
w i t h i n one p o p u l a t i o n of Pinus s y l v e s t r i s ,
P. s t r o b u s , P. ponderosa and w i t h i n P i c e a a b i e s
p r e s e n t e d a t t h e Symposium on E f f e c t s of A i r
P o l l u t a n t s on Mediterranean and Temperate F o r e s t
Ecosystems, June 22-27, 1980, R i v e r s i d e , C a l i f o r -
n i a , U.S.A.
2 ~ r e eB i o l o g i s t , I n s t i t u t e of Dendrology, P o l i s h
Academy of S c i e n c e s , 63-120 Kornik, Poland.
(Gerhold e t al,., 1977). The g r e a t e r t o l e r a n c e of
S c o t s p i n e and Norway spruce t o t h e a i r p o l l u t i o n
w i t h i n a population i s sometimes a s s o c i a t e d w i t h
t h e g r e a t e r r e s i s t a n c e t o low temperatures d i s e a s e s
a i d a xerothermic a d a p t a t i o n (Bialobok, 1979). A
suggestion a l s o e x i s t s t h a t t h e cause of t o l e r -
ance t o some gases may depend on mechanisms t h a t
reduce t h e a c c e s s of g a s e s t o t h e p l a n t organism
(Gerhold e t a l . , 1977).
However, i t h a s been experimentally e s t a b l i s h e d
o r deduced from r e l i a b l e evidence t h a t t h e r e
e x i s t g e n e t i c mechanisms c o n d i t i o n i n g h i g h e r
t o l e r a n c e t o i n d u s t r i a l emissions (Niemtur, 1979).
The l a t t e r i n v e s t i g a t o r s t u d i e d 58 progenies of
S c o t s p i n e from 4 provenances o b t a i n e d from s e e d s
c o l l e c t e d on remnant l i v i n g t r e e s in t h e h i g h l y
p o l l u t e d c o n d i t i o n s of t h e Upper S i l e s i a n indus-
t r i a l region. He h a s shown t h a t t h e s e p r o g e n i e s
have a g r e a t e r t o l e r a n c e than t h o s e c o l l e c t e d
from t r e e s growing beyond t h e range of p o l l u t i o n .
POLYMORPHISM AND ITS ROLE
S t u d i e s on t h e g e n e t i c v a r i a b i l i t y of enzymes
in f o r e s t t r e e s developed s t r o n g l y a f t e r S t e m ,
(1974) p o i n t e d o u t t h e g r e a t importance of gen-
e t i c polymorphism f o r t h e development of ecolog-
i c a l g e n e t i c s , f o r e s t g e n e t i c s and t r e e breeding.
Thanks t o t h i s method i t i s p o s s i b l e t o
recognize t h a t g e n e t i c s t r u c t u r e of a p o p u l a t i o n
( a l l e l e f r e q u e n c i e s and degree of h e t e r o z y g o s i t y )
in space and time, t o conduct s t u d i e s on i n h e r i t -
ance and l i n k a g e , t o determine t h e g e n e t i c d i s t a n c e
between i n d i v i d u a l s and p o p u l a t i o n s , t o i d e n t i f y
mechanisms of a d a p t a t i o n t o an environment and
t o employ i t in p r a c t i c a l f o r e s t r y .
Polymorphism of f o r e s t trees which a r e long-
l i v i n g organisms r e p r e s e n t s a d a p t i v e s t r a t e g i e s
of p o p u l a t i o n s t o a changing environment ( S t e r n
e t a l . , 1974). I n t h i s g e n e t i c system hetero-
zygotes a r e f a v o r e d and t h u s t h e population is
capable of e x i s t i n g i n a h e t e r o g e n i c environment
s u r v i v i n g numerous and major changes.
, Tree p o p u l a t i o n s in i n d u s t r i a l r e g i o n s grow
under s i m i l a r c o n d i t i o n s a s t h e herbaceous
p l a n t s t h a t c o l o n i z e m i n e s p i l l s having h i g h
c o n c e n t r a t i o n s of heavy elements (Pb, Zn, Cu).
This phenomenon S t e m (1974) a l s o c o n s i d e r s a s
being a good example of t h e developing poly-
morphism.
I n c o n d i t i o n s where a t l e a s t a p a r t of t h e
t r e e s can s u r v i v e i n an i n d u s t r i a l environment
t h a t has n o t been t o o s t r o n g l y p o l l u t e d , t h e
p o p u l a t i o n a d o p t s t h e optimal s t r a t e g y i n reac-
t i o n t o t h e changes t a k i n g p l a c e . This, of
c o u r s e , i s t h e r e s u l t of t h e e v o l u t i o n a r y
a d a p t a t i o n p r o c e s s t o t h e new environmental
c o n d i t i o n s where t h e g e n e t i c system i s under a
c o n s t a n t s e l e c t i o n p r e s s u r e . T o l e r a n t geno-
t y p e s appear t o have a c e r t a i n l e v e l of f i t n e s s
t o t h e i n d u s t r i a l environment. What i s most
important t o us i s t h a t t h e d r a s t i c changes in
t h e i n d u s t r i a l environment should n o t exceed
t h e l e v e l of g e n e t i c f l e x i b i l i t y of t h e popula-
t i o n s of t h e most important f o r e s t forming
s p e c i e s . I n t h e acute" c o n d i t i o n s of change i n
an i n d u s t r i a l environment, t h e coded g e n e t i c
system w i t h i n a p o p u l a t i o n , t h a t c o n d i t i o n s i t s
adaptive capacity, gradually ceases t o operate
s i n c e t h e p o t e n t i a l of i n d i v i d u a l genotypes f o r
a d a p t a t i o n h a s been surpassed.
GENETIC VARIABILITY OF ENZYMES I N SCOTS PINE
POPULATIONS
The u s e of e l e c t r o p h o r e t i c a n a l y s i s of
enzymes f o r t h e i d e n t i f i c a t i o n of v a r i a b i l i t y
determining g e n e t i c t o l e r a n c e t o i n d u s t r i a l
p o l l u t i o n i s s o f a r o n l y i n an i n t r o d u c t o r y phase.
The r e s u l t s o b t a i n e d s o f a r a r e encouraging.
The s t u d i e s conducted on S c o t s p i n e a r e t r e a t e d
o n l y a s a model.
Mejnartowicz (1978) h a s s t u d i e d t h e v a r i a -
b i l i t y of izozymes of leucyloaminopeptidase (LAP)
and a c i d phosphotase (APH) in 19 p o p u l a t i o n s of
S c o t s p i n e from t h e whole range of t h e s p e c i e s
in E u r a s i a . The Poland p o p u l a t i o n s were chosen
fromregions under i n d u s t r i a l i n f l u e n c e and from
r e g i o n s beyond p o l l u t i o n . I n a l l t h e p o p u l a t i o n s
APH was coded in l o c u s B w i t h 1 5 a l l e l e s , and
t h e r e was c o n s i d e r a b l e polymorphism in t h e
s t u d i e d p o p u l a t i o n s , and LAP was coded in 2
l o c i , LAP-A having 5 a l l e l e s and LAP-B 6
alleles.
It i s i n t e r e s t i n g t h a t t h e population of S c o t s
p i n e from Babki ( c e n t r a l Poland--strong indus-
t r i a l p o l l u t i o n ) h a s had t h e lowest degree of
h e t e r o z y g o s i t y on l o c u s LAP-A, ha0.1124 and t h e
h i g h e s t v a l u e i n LAP-B, h=0.3970. Among i n d i v i d -
u a l s s e n s i t i v e t o SO2 t h e genotype APH-B4/B6
dominated and among i n d i v i d u a l s t o l e r a n t t o t h i s
gas t h e a l l e l e APH-B5 appeared commonly
(Mejnartowicz, 1977-78). Szmidt (1978) when
studying t h e g e n e t i c polymorphism of c a t a l a s e i n
t h r e e p o p u l a t i o n s of S c o t s p i n e from i n d u s t r i a l
r e g i o n s and 5 p o p u l a t i o n s from r e g i o n s f r e e of
p o l l u t i o n , h a s found 7 phenotypes of t h e
c a t a l a s e . When comparing t h e p o p u l a t i o n s on t h e
b a s i s of h e t e r o z y g o s i t y he has shown t h a t popula-
t i o n s from r e g i o n s f r e e of p o l l u t i o n were charac-
t e r i z e d by g r e a t e r p a r t i c i p a t i o n of homomorphic
i n d i v i d u a l s of t h e type C-l/C-1 compared t o
p o p u l a t i o n s under t h e i n f l u e n c e of emissions.
Mejnartowicz, e t a l . , (1978) having taken
m a t e r i a l from p l u s t r e e s in a S c o t s p i n e seed
orchard e s t a b l i s h e d a l a c k of c o r r e l a t i o n between
t h e degree of h e t e r o z y g o s i t y of t r e e s i n LAP
l o c i and t h e degree of t h e i r s e n s i t i v i t y t o t h e
a c t i o n of SO2 in a l a b o r a t o r y t e s t .
These i n t r o d u c t o r y r e s u l t s i n d i c a t e t h a t t h e r e
i s d i f f e r e n t i a t i o n in t h e frequency of some
a l l e l e s in a p o p u l a t i o n depending on t h e i n c l u -
ence of i n d u s t r i a l emissions. The important
t h i n g i s t h a t an enzymatic marker b e found f o r
t o l e r a n c e which would b e independent of environ-
mental i n Âl u e n c e s .
INHERITANCE OF TOLERANCE TO INDUSTRIAL EMISSIONS
Not many i n v e s t i g a t o r s have attempted determin-
i n g in g e n e r a t i v e progeny t h e combining a b i l i t y ,
h e r i t a b i l i t y and r e p e a t a b i l i t y of t o l e r a n c e t o
i n d u s t r i a l emissions. I n t h e c a s e of S c o t s p i n e
h e r i t a b i l i t y and r e p e a t a b i l i t y of t o l e r a n c e t o
Oq i s t o be found under a low o r medium g e n e t i c
c o n t r o l . However, some provenances have a h i g h e r
h e r i t a b i l i t y of t o l e r a n c e t o 03. Needle i n j u r y
in t h e s e p i n e s was p o s i t i v e l y c o r r e l a t e d w i t h
t h e i n j u r i e s caused by SO2 (Gerhold e t a l . , 1977,
Demeritt, 1977). Bialobok e t a l . , (1978) have
a l s o found a s i g n i f i c a n t c o r r e l a t i o n between t h e
degree of n e e d l e i n j u r y on mother t r e e s and on
t h e i r progenies following treatment w i t h SO?.
However, t h e s e c o r r e l a t i o n s were n o t found follow-
i n g treatment w i t h 03 o r a mixture of 03 and SO2.
The s e l e c t i o n of t r e e i n d i v i d u a l s t o l e r a n t t o
i n d u s t r i a l emissions i s being done by t h e r e l a -
t i v e l y simple methods of mass s e l e c t i o n , without
understanding t h e g e n e t i c b a s i s f o r t h e popula-
t i o n s t r u c t u r e . T h i s s e l e c t i o n i s u s u a l l y being
conducted on s e e d l i n g s in l a b o r a t o r y c o n d i t i o n s .
Sometimes p o s i t i v e r e s u l t s were o b t a i n e d f o r t h e
p r a c t i c e which w i t h some p r o b a b i l i t y a r e being
u t i l i z e d in e s t a b l i s h i n g tree p l a n t a t i o n s o r
producing t r e e s f o r ornamental purposes
(Demeritt, 1977).
 So f a r i t was n o t p o s s i b l e t o s o l v e t h e prob-
l e m of breeding v a r i o u s p i n e s p e c i e s f o r c u l t i v a -
t i o n in f o r e s t s under t h e i n f l u e n c e of i n d u s t r i a l
emissions, and t h e r e i s no cheap way of propaga-
t i o n of s e l e c t e d i n d i v i d u a l s .
Gerhold, (1977) h a s d i s c u s s e d s e v e r a l methods
of breeding trees more t o l e r a n t t o i n d u s t r i a l
e m i s s i o n s , i n which t h e u s e of s e e d s o r c h a r d s
e s t a b l i s h e d from t o l e r a n t c l o n e s i s envisaged.
It appears, however, t h a t i t would be v a l u a b l e
t o expand t h e i n v e s t i g a t i o n s of g e n e t i c v a r i a -
b i l i t y of enzymes in p o p u l a t i o n s of c o n i f e r o u s
trees endangered by i n d u s t r i a l p o l l u t i o n . W e
have c o n s i d e r a b l e information a l r e a d y on t h e
c o u r s e of m e t a b o l i c c y c l e s in t h e tree organism
exposed t o t h e a c t i o n of some gases. I b e l i e v e
t h e s e two s o u r c e s of information could be
combined, (biochemical g e n e t i c s and metabolic
changes) in o r d e r t o i d e n t i f y t h e enzymatic
g e n e t i c marker t h a t would be u s e f u l i n t h e
s e l e c t i o n of t o l e r a n t populations.
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s t r a i n s and a r t i f i c i a l s e l e c t i o n o r produc-
t i o n of such s t r a i n s in o r d e r t o p r o j e c t
v e g e t a t i o n from a i r p o l l u t i o n . Symp. on t h e
e f f e c t s of a i r b o r n e p o l l u t i o n on v e g e t a t i o n ,
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Demeritt, Maurice E. JK.
1977. Genetic Evaluation of Two-Year Height
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s y l v e s t r i s L.) .
A Thesis i n Genetics. The
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Gerhold, Henry G., and Raymond G. Wilhour.
1977. E f f e c t of a i r p o l l u t i o n on Pinus
s t r o b u s L. and g e n e t i c r e s i s t a n c e . A L i t e r a -
t u r e Review. EPA-600/3-77-002 p. 45 Environ-
mental P r o t e c t i o n Agency, C o r v a l l i s , Oregon,
97330.
Mejnartowicz , Leon.
1977-1978. Genetic c h a r a c t e r i s t i c of some
S c o t s p i n e t r e e s s u s c e p t i b l e or somewhat
r e s i s t a n t t o t h e a c t i o n of SO^. Annual
Report from P r o j e c t PL-Fs-74 from U. S.
Department of A g r i c u l t u r e under PL-480.
Mejnartowicz, Leon.
1978. S t r u k t u r a genetyczna p o p u l a c j i sosny
zwyczajnej z terenow przemyslowych zanieczy-
szczonych przez S02. I n Proc. from:
Reakcje b i o l o g i c z n e drzew na e m i s j e
przemyslowe. Kornik 4-5 maj 1978.
Mejnartowicz, L., S. Bialobok and P.
Karolewski.
1978. Genetic c h a r a c t e r i s t i c of S c o t s p i n e
specimens r e s i s t a n t and s u s c e p t i b l e t o SO2
a c t i o n . Arboretum K o m i c k i e . 23: 233-238.
Mejnartowicz , Leon.
1979. Genetic v a r i a t i o n i n some isoenzyme
l o c i i n S c o t s p i n e (Pinus s i l v e s t r i s L.)
population. Arboretum K o m i c k i e . 24: 91-
104.
Niemtur, Stanislaw.
1979. Badania nad wewnatrzgatunkowym
zroznicowaniem o d p o m o s c i sosny zwy-
c z a j n e j (Pinus s i l v e s t r i s L.) n a e m i s j e
przemyslowe. Doctoral T h e s i s , Uniwersytet
&ski, Katowice.
S t e m , K. and P e t e r M. A. T i g e r s t e t .
1974. Okologische Genetic. 98-p. 75 p. G.
Fischer. Stuttgart.
Szmidt, Alfred.
1978. Zmiennosc k a t a l a z y w populacjach
sosny zwyczajnej znajdujacych s i e pod
wplywem zanieczyszczen przemyslowych. In
Proc. from: Reakcje b i o l o g i c z n e drzew n a
e m i s j e przemyslowe. Komik 4-5 maj 1978.
 Interactions of Air Pollutants and Plant

Michael reshow^

Abstract: Each i n t e r a c t i o n between an a i r pollu-

t a n t and d i s e a s e is unique. P o l l u t a n t s may m i t i -
gate d i s e a s e response or i n t e n s i f y i t ; and t h e
presence of c e r t a i n d i s e a s e s can modify the p l a n t s
response t o a p o l l u t a n t . Air p o l l u t a n t s can a c t
d i r e c t l y on a fungus or b a c t e r i a i n h i b i t i n g parasi-
tism. They may a l s o a c t most s t r o n g l y on the h o s t ,
modifying i t s physiology and rendering i t e i t h e r
more or l e s s s e n s i t i v e t o a p l a n t pathogen. Where
t h e p l a n t i s weakened by the p o l l u t a n t s , it tends
t o be more s e n s i t i v e t o weak pathogens, but l e s s
sensitive t o obligate parasites. Where the pollu-
t a n t p h y s i c a l l y i n j u r e s t h e host, i n f e c t i o n may be
facilitated. P o l l u t a n t i n t e r a c t i o n s have been
demonstrated both i n the l a b o r a t o r y and t h e f i e l d
a t ambient p o l l u t a n t concentrations. Interactions
between p o l l u t a n t s and a b i o t i c s t r e s s a r e p a r t i c u -
l a r l y c r i t i c a l . Water regime, temperature r e l a -
t i o n s , mineral n u t r i t i o n and other parameters of
t h e physical environment play a major r o l e i n the
expression of a i r p o l l u t i o n i n j u r y .

Air p o l l u t i o n can k i l l p l a n t s . Even when t h e i n s e c t or o t h e r pathogen. I f some s t a g e of an

concentrations of an a i r p o l l u t a n t a r e not
d i r e c t l y l e t h a l , they may be harmful, adversely
a f f e c t i n g growth, reproduction and myriads of
o t h e r b i o l o g i c a l processes ( M i l l e r and McBride
1975). One such process i s t h e i n t e r a c t i o n t h a t
r e s u l t s between t h e p o l l u t a n t and p l a n t
pathogens.
There a r e many ways i n which such i n t e r -
a c t i o n s might take place. But do they? The most
obvious would be i f the p o l l u t a n t d i r e c t l y
a f f e c t e d the fungus, v i r u s , b a c t e r i a ,
p r e s e n t e d a t t h e Symposium on E f f e c t s of A i r
P o l l u t a n t s on Mediterranean and Temperate Forest
Ecosystems, June 22-27, 1980, Riverside,
C a l i f o r n i a , U.S.A.
2 ~ r o f e s s o rof Biology, University of Utah,
S a l t Lake C i t y , Utah 84112.
organism's l i f e c y c l e , o r some s t a g e of p a r a s i t -
i s m , were adversely impaired by a p o l l u t a n t , t h e
p o l l u t a n t e s s e n t i a l l y would be a c t i n g a s a
fungicide, b a c t e r i o c i d e or o t h e r biocide and
could be expected t o bring about some degree of
control.
The second way an a i r p o l l u t a n t might a c t
would be i n adversely a f f e c t i n g the growth o r
reproduction of the host p l a n t and thereby
influencing i t s s u i t a b i l i t y a s a host. There
a r e two ways i n which such a c t i o n could be
brought about. Most apparently, the p o l l u t a n t
might cause some physical i n j u r y t o the p l a n t
leaving l e s i o n s t h a t could serve a s i n f e c t i o n
c o u r t s f o r a fungus or bacteria. O r , the p l a n t
might simply be weakened, thus lowering i t s
r e s i s t a n c e t o a prospective pathogen. Presum-
ably the weaker pathogens, the f a c u l t a t i v e
p a r a s i t e s , would b e n e f i t most by such
conditions. Obligate parasites, on the other
But such incidents are becoming more infre-

hand, preferring a vigorous host, might respond

quent as ground level SO2 concentrations become

quite differently and their development might

reduced by modern control technology or use of

even be discouraged in the weakened host.

tall stacks. Higher SO2 concentrations, often

exceeding 1 ppm for extended periods, now tend

Conversely, the impact of various diseases or

to be of only historic record. We are now

plant stresses on the plant's response to air

interested in the possible impact of SO2 concen-

pollutants has also been considered. It should

trations in the 0.1 to 0.5 ppm range or even

be quite apparent that adverse climatic or

lower in cases of prolonged duration. Where

edaphic factors might affect a plant's sensi-

such concentrations still exist near smelters,

tivity to a pollutant, but the presence of

power plants, or large urban centers, some

certain biotic diseases may also increase or

pollutant-pathogen interaction might be sought.

decrease a plant's sensitivity.

There is some evidence from laboratory and

<
A major question is not so much do such
field research that certain pathogenic fungi

interactions take place, but what concentrations

respond to SO2 in this concentration range.

of an air pollutant are required to precipitate

Weinstein and others (1975) for instance, have

such effects, and how widely do such concentra-

demonstrated that SO2 concentrations of 0.14 to

tions occur?
0.15 ppm, continuous for 7 days, while not

visibly injuring plant foliage, affected the

Many of the interactions that will be dis-

bean rust fungus, Uromyces. The incidence and

cussed have been demonstrated under laboratory

severity of the rust as measured by the abun-

conditions, often at pollutant concentrations

dance of uredia was decreased as were both the

known to be fairly widespread. Some of the

size and percent germination of the uredo-

interactions have been observed in the field.

spores. Hence the reproductive potential of the

pathogen was reduced. The results suggested

The literature treating such air pollutant-

that the effect on the pathogen was indirect;

plant pathogen interactions has been reviewed

perhaps involving an 802-induced chemical change

most recently in 1978 (Laurence), 1975

in the resistance of the host and its suitabil-

(Treshow), and 1973 (Heagle); it is not the

ity as a habitat for the fungus. The authors

intent of this paper to once again provide an

speculated that production of phytoalexins or

exhaustive review of this literature. Rather, I

supply of metabolites might have been altered,

should like to first bring the literature up to

or compounds derived from SO2 might have accumu-

date, briefly including that delving into the

lated.

interactions of abiotic stresses on pollutant

effects, and discuss the implication and pos-

One of the most comprehensive field studies

sible practical relevance of such interactions,

of the interactions of industrial air pollu-

especially concerning forest ecosystems. Pri-

tants, especially SO2, with plant disease was

marily I should like to develop some of the

conducted by A. Grzywacz and J. Wazny (1973).

principles in pollutant-pathogen interactions.

The study was notable since it dealt with eco-

nomically significant diseases of a coniferous

forest, specifically Scots pine (Pinus

INFLUENCE OF POLLUTANTS ON DISEASE EXPRESSION

sylvestris L.). Interactions with six fungi

Sulfur Dioxide

--
were considered: Armillaria mellea, Fomes

annosus, Lophodermium pinastri, Microsphaera

alphitoides, Melampsora pinitorqua, and

There is no question that sulfur dioxide

Cronartium flaccidium.

interacts with fungi, suppressing their growth

and acting fungicidally. The only question

In an overall study, comparisons were made of

concerns the SO2 concentrations at which such

disease presence in industrial areas and forests

interactions occur.
throughout Poland. In a more detailed approach,

disease presence was compared at increasing

A few examples exist where SO2 has acted in

distances from an industrial town, Torun, with

such a way in the field, limiting the develop-

one major SO2 source.

ment of certain plant diseases. The examples

provided by Scheffer and Hedgcock (1955) in

The general comparisons showed that

which parasitic fungi generally appeared to be

Armillaria was present in 3.7 percent of the

retarded or inhibited near smelters where SO2

trees in the area damaged by SO2 compared with

was present is most often cited (Treshow

only 1.38 percent in all forests. Lophodermium

1975). Suppressed disease development was

also tended to be more serious in the industrial

especially apparent where rust fungi, Melampsora

areas, but the results were indefinite. This

and Puccinia, or needle cast fungi were

was consistent with earlier findings of

involved. Linzon (1973) also discusses the

Donabauer (1966) that high SO2 concentrations

reduced heart rot and blister rust (Cronartium

increased the intensity of Armillaria root rot.

ribicola) near a smelter.

Fomes showed the reverse trend and occurred

in 0.87 percent of the areas affected by indus-

pathogen interactions (Shriver 1974). Although

trial pollution compared with 1.5 percent in the

not involving forest species, the work did

country as a whole. Fomes, as in the overall

demonstrate the way in which plant species might

study, was least prevalent in areas of highest

be affected.

SO2 concentrations. Grzywacz and Wazny (1973)

found that stumps close to the SO2 source were

Kidney beans (Phaseolus vulgaris var. red

sound. Related, decomposition fungi were also

kidney) and willow oak (Quercus phellus) were

less abundant closest to the source. Only 50

exposed to simulated rain acidified to a pH of

percent of the stumps showed signs of decom-

6.0 or 3.2 for 10 minutes each day for 30 to 60

posing, and 10 percent strongly decomposing,

days. The acid "rain" caused an 86 percent

compared with 30 percent strongly decomposing

inhibition in the number of telia produced by

farther out. However, Gryzwacz (1978) later

Cronartium fusiforme on willow oak, a 66 percent

reported that stands influenced by air pollution

inhibition in the reproduction of the root knot

were more subject to decomposition by brown,

nematode (Meloidogyne hapla) on beans in the

white or soft rot fungi. Sufficiently high SO2

field, and a 29 percent decrease in the percent

concentrations might suppress the fungal growth

of leaf area of field "-
erown beans
- -- - - - affected by

- - -

directly. Such was the case where Mejstrik

Uromyces phaseoli. Exposures to pH 3.2 "rain"

(1978) showed SO2 concentrations as low as 84

-
completely destroyed the integrity of the cuti-

ug/m3 (.03 ppm) inhibited growth of Flamulina

cular waxes in both oak and bean. Exposures to

velopes, Nematoloma fasciculare, and Pleurotus

the pH 6.0 solution did not visibly alter the

ostreatus from 34 to 39 percent.

cutin. Shriver postulated that damage to the

cutin facilitated leaching of carbohydrates from

Lophodermium injury was least significant

the leaf including those that might inhibit

near the SO2 source. At distances beyond 1 km

hyphal growth of Botrytis cinerea thus predis-

injury appeared on up to 70 percent of the

posing plants to infection.

needles. There was a slight decrease beyond

this. When annual SO2 concentrations averaged

This is consistent with the findings of F. A.

below 0.2 ppm, disease incidence was essentially

Last and 0. D. Fowler at the Institute of

the same as in the control group. Fungus devel-

Terrestrial Ecology at Penicuik, Scotland (per-

opment was noticeably affected. Within 1 km,

sonal communication) who showed that the cuticle

reproduction was largely by pycnidia; further

of Scots pine was gradually broken down by low

distant, apothecia were common. The numbers of

concentrations,of S02.

needles with both the sexual and asexual stages

increased with distances. Closest to the SO2

Sulfur dioxide, perhaps largely as acid rain,

source, apothecia were only 15 percent the

may have an effect on soil microorganisms

length of controls deformed, collapsed, under-

(Wainwright 1978). Sulfur-oxidizing micro-

developed, closed or even dried up and with ill-

organisms, including Alternaria tenuis,

developed asci. Chiba and Tanaka (1968) however

Auriobasidium pullulans and Cephalosporium sp.,

reported high SO2 concentrations of 2 ppm for 14

were isolated from leaves, litter and soils

hours caused increased infection of injured pine

polluted with high levels of S02. The predomi-

needles (P. densiflora) by Rhizosphoera nant autotroph on leaf surfaces was Thiobacillin

kalkhoffii.
thioparis.

Weidensaul and Darling (1979) inoculated

Hibben and Taylor (1975) found that S02-

Scots pine seedlings with the fungus Scirrhia

pathogen interaction depended on the infection

acicola. The plants were inoculated either 5

stage. Conidial gemination and the appres-

days before or 30 minutes after fumigation for 6

sorial stages of Microsphaera =were most

hours with 533 ug/m3 (0.20 ppm) SO2). After 8

sensitive to S02. Since infection was not

weeks, the seedlings inoculated 5 days before

reduced when leaves were previously fumigated,

fumigations had more lesions incited by the

the effect was thought to be mostly

fungus than those inoculated 30 minutes after

fungicidal.

fumigations in the control group. It was postu-

lated that since the needle blight fungus pene-

trates through the stoma, the ability of the SO2

Ozone

to keep the stoma open facilitated entry of the

fungus.
There is considerable evidence that photo-

chemical pollutants, most notably ozone, affect

The pollutant-disease interaction may vary

plant pathogens and fungus-host interactions.

with the host species. Ham (1971) found that

The interaction may be positive or negative

when loblolly pine served as the host, the brown

depending on the 03 concentrations and the 03

spot fungus (5. acicola) was not measurably sensitivity of the host.

affected by SO2 in the 0.5-0.9 ppm range for 2-3

hours even when SO2 caused visible injury.

In one such study, Miller and Elderman (1977)

showed that fumigation of ponderosa and Jeffrey

Simulated rain, acidified with sulfuric acid

pine seedlings with ozone enhanced infection by

has also been shown to influence pollutant-

Fomes annosus. Average infection of the two

species increased from 57 percent to 78 percent

flavonoids that are toxic to microorganisms.

when the seedlings were exposed to 431.2 p g / ~ 3

According to Laurence and Wood (1978), ozone

(.25 ppm) ozone. They also found that ozone

inhibited infection of soybean by Pseudomonas

injury increased the susceptibility of pine

glycinea presumably because of a bacteriostatic

stumps to colonization by Fomes annosus. In the

compound, possibly an isoflavinoid compound,

laboratory, wood from trees slightly damaged by

produced in response to ozone.

oxidants was more decay-susceptible than wood

from severely damaged trees.

Fluoride

Far more research has involved powdery mildew

(Heagle and Strickland 1972, Heagle 1975,

Fluoride is accumulated in the foliage of

Scheutte 1971), and the negative impact of ozone

plants and affects many metabolic processes.

on disease. Even ozone concentrations in the 5

Consequently, it is reasonable to presume that

to 50 pphn range reduced infection when conidia

pollutant-pathogen interactions can be

were exposed. According to Scheutte, appresoria

influenced, perhaps to a greater extent than

formation was largely affected. Also, some

with any other pollutant. This was confirmed

epidermal cells, hypersensitive to ozone, were

with viruses by some of the early work reviewed

killed thereby reducing infection. Eight hour

by Heagle (1973) and Treshow (1975).

exposure to 50 pphm ozone reduced formation of

secondary hyphae 66 percent and 4 hour exposure

Host-fluoride interactions also have been

reduced 41 percent. Ozone impaired formation of

demonstrated for fungi (McCune and others 1973)

the penetration peg thus inhibiting subsequent

although not on forest species. Studying the

development of a functional host-parasite rela-

interactions of 7 to 10 with 3 diseases

tionship.
of bean and 2 diseases of tomato, HF consist-

ently reduced uredial formation as the foliar

While ozone impacted certain critical stages

fluoride content increased. Tender green beans

of infection under controlled conditions, in the

exposed to HF and inoculated with powdery mildew

field such effects would be less apparent since

conidia developed fewer foliar lesions than non-

all stages of parasitism would be present simul-

fumigated control plants; 4.4 compared with 48.7

taneously.
on fumigated leaves having an average of 399 ppm

F. Powdery mildew was reduced proportionate to

Interactions of facultative parasitic and

the length of the exposure period. Early blight

saprophytic fungi with their hosts may also

lesions on tomato were similarly reduced by the

occur and are most pronounced when the plants

presence of fluoride.

are weakened or injured by ozone (Manning

1975). Obligate parasitism appears to be

retarded by ozone and ozone-injured host

INFLUENCE OF DISEASE ON POLLUTANT SEVERITY

tissue. Colonization of ozone-injured white

pine needles by Lophodermium pinastri was

Not only may the presence of a pollutant

reduced while colonization by the saprophyte

influence pathogenicity, the reverse may be

Aureobasidium pullulans was increased. Ozone-

true~presenceof diseases influence the degree

injured lilac leaves were rarely infected by

of pollutant injury. This has best been demon-

powdery mildew Micosphaera a.

strated with viruses. None of these involved

forest species though, and there has been no new

When the fungus is especially sensitive to

results published since last reviewed by

ozone the impact may be mostly fungicidal there-

Laurence (1978).

by reducing infection. Sporulation and germina-

tion of the weak parasite Botrytis cinerea was

Also, in the forest, variation in pollutant

significantly inhibited by 2, 6 hour ozone

resistance among individual trees would far

exposures at a concentration of 30 pphm (Krause

exceed any disease and pollutant interaction,

and Weidensaul 1978). There was also less

and any impact of a disease on pollutant sever-

infection, based on total lesion area, when

ity probably wouldn't be measurable.

plants were exposed to 15 pphm. Ozonation of

conidia produced in vivo and in vitro decreased

germination of conidia, germ tube length, and

ABIOTIC STRESS

pathogenicity. The authors postulated that the

mode of action was to alter the permeability of

Severity of air pollution injury is deter-

the conidia membrane. Ozone may increase the

mined not only by the dose of the pollutant to

conidiophore respiration causing prematurely-

which plants are exposed but on the predisposi-

formed and non-viable conidia. Ozone also may

tion of the plant. Predisposition is determined

inhibit, directly or indirectly, enzyme activity

by any environmental parameter to which the

of the fungus and cause less maceration of the

plant is subjected. It has been discussed more

host cells resulting in a decreased infection.

in passing than as a major concern (Heck and

others 1965, Heck 1968, Treshow 1970, Taylor

Curtis and others (1976) found that plants

1974), but the interactions of abiotic stress or

exposed to ozone produced peroxidases and iso-

disorders with any pollutant is far more

significant than with biotic pathogens.

conditions allowed only slight to moderate

Moisture stress, the light regime, edaphic

foliar injury on tomato leaves, damage was

factors and temperature relations all influence

enhanced by cadmium treatment especially at

the sensitivity of plants to every pollutant.

higher concentrations.

Unusually high or low temperatures prior to

Along the same line, low concentrations of

exposure tend to reduce plant sensitivity to

SO2 affected metal uptake, increasing zinc and

ozone, for instance. A normal high light inten-

cadmium susceptibility of bean plants. Foliar

sity renders plants more sensitive. High humid-

injury caused by heavy metals was significantly

ity, by encouraging open stomates, also enhanced by SO2 (Krause and Kaiser 1977).

increases sensitivity. In dry air, plants are

more tolerant of ozone than at higher humidi-

Lamoreaux and Chaney (1978) also found sig-

ties, and PAN injury is most severe in the Los

nificant interactions between cadmium and S02.

Angeles area when the relative humidity exceeds

While cadmium or SO2 alone reduced net photosyn-

50 percent.
thesis and transpiration of excised silver maple

leaves, the reduction in cadmium-treated leaves

Any factor favoring moisture stress and

was greatest in the presence of S02.

reduced water uptake would cause stomatal clo-

sure and favor resistance to pollutant uptake

and injury. In this way, plants in more saline

CONCLUSIONS

soils are more tolerant of certain pollutants

(Oertli 1958) and there is little to no ozone or

Virtually every environmental parameter,

PAN injury when soil moisture is deficient.

biotic and abiotic can interact with air pollu-

tants to aggravate or mitigate the extent of

Ozone injury may be negligible when ambient

pollution damage. The significant question then

air temperatures during and after exposure are

becomes, to what extent and at what concentra-

below 90Â F., yet may be severe when tempera-

tions? Is the net impact real, and if so, how

tures exceed 90' F. (Taylor 1974).

meaningful is it under field conditions?

Effects of low atmospheric and soil moisture

All pollutants don't interact the same.

on tolerance or sensitivity of plants to SO2

Sulfur dioxide, for instance, appears primarily

injury are well known; suffice it to note that

to act fungicidally reducing disease and decom-

plants are far more sensitive under higher

position activity. It is also postulated though

moisture conditions.
that SO2 induces chemical changes imparting

resistance to certain pathogens. Ozone tends to

Mineral nutrition also plays arole in pollu-

act more on the host, weakening it and accentu-

tant sensitivity. Vegetable crops are known to

ating disease activity. Research with vegetable

be most susceptible to ozone when nitrogen

crops at low ozone concentrations indicate that

nutrition is adequate although some reports

pollutant-disease interactions do occur with

suggest injury is greatest when nitrogen is

some diseases and very likely occur in forest

low. Sulfur dioxide sensitivity is reported to

stands weakened by photochemical pollutants.

be greatest with increased sulfur nutrition.

Ozone has increased susceptibility to Fomes and

Stomata1 behavior may be involved since a defic-

rendered trees more susceptible to decay. Each

iency of nitrogen, potassium or phosphor,us may

interaction is unique since every fungus has its

decrease stomatal opening. Plants low in sulfur

own sensitivity as does each plant. Thus the

had the lowest stomatal capacity, thus less

potential for interaction is enormous.

capacity for gas absorption (Leone and Brennan

1972).
Pollutants such as fluoride that accumulate

significantly in plants would likely have the

Toxicity from heavy metals also plays an

greatest interaction. While the trend is to

interesting role with air pollutants. Ormrod

inhibit disease development, the concentrations

(1977) has found that ozone toxicity was en-

at which this has been demonstrated are now

hanced on pea plants growing in elevated concen-

infrequent in the field.

trations of cadmium or nickel. However, when

cadmium concentrations were sufficiently high to

The significance of abiotic stress and pollu-

adversely affect growth,100 umol of CdS04, then

tant interactions is often taken for granted,

ozone injury was less than that of control

but it should be emphasized that this is of

plants.
paramount importance. Climatic and edaphic

factors are often determinants in the expression

Harkov and others (1979) found that the

of pollutant damage. While the abiotic environ-

amount of ozone injury depended on the cadmium

ment strongly influences the severity of air

and ozone levels as well as other environmental

pollution damage, air pollutants influence the

conditions. Where environmental conditions were

severity of plant diseases caused by biotic

.
not conducive to ozone injury (e g. bright
sunlight), cadmium had little effect on the

pathogens. It would appear that as in all life

systems, every environmental parameter has some

amount of leaf damage. When environmental

influence on every other. In the case of

pollutants and pathogens, this interaction can

Heck, W. W., J. A. Dunning and I. J. Hindawai.

be significant.
1965. Interactions of environmental factors

on the sensitivity of plants to air

pollution. J. Air Poll. Control Assoc.

LITERATURE CITED
15:511-515.

Chiba, 0. and K. Tanaka.

Hibben, C. R. and M. P. Taylor.

1968. The effect of sulfur dioxide on the

1975. Ozone and sulfur dioxide effects on the

development of pine needle blight caused by

lilac powdery mildew fungus. Environ.

Rhizosphaera kalkhoffii Bubak (I). J. Jap.

Pollut. 9:105-114.

For. Soc. 50: 135-139.

Krause, G. H. M. and H. Kaiser.

Curtis, C. R., R. K. Howell and D. F. Kremer.

1977. Plant response to heavy metals and

1976. Soybean peroxidase from ozone injury.

sulfur dioxide. Environ. Pollut. 12:63-71.

Environ. Pollut. 11:189-194.

Krause, C. R. and T. C. Weidensaul.

Donabauer, E.
1978. Effects of ozone on the sporulation,

1966. Secondary damage to forest caused by

germination, and pathogenicity of Botrytis

industrial exhaust fumes. Mitt. Forst.

cinerea. Phytopathol. 68:195-198.

Bundesvers. 73:lOl.

Lamoreaux, R. J. and W. R. Chaney.

Grzywacz, A.
1978. Photosynthesis and transpiration of

1978. Interactions between air pollutants and

excised silver maple leaves exposed to

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 Studies on Relationship Between Air

Pollutants and Microorganisms in
Japan1

Kiyoshi ~ a n a k a ~

Abstract: A review of the literature on the interaction of air

pollutants with parasitic and epiphytic microorganisms is pre-

sented with a brief outline of air pollution problems in Japan.

Some fungi such as Cercospora sequoiae, Pucciniastrum styracinum,

Puccinia kusanoi, Melampsora coleosporioides, Trichoroma matsutake,

and some epiphytic bacteria disappeared in the areas affected

by air pollutants, and Rhizosphaera kalkhoffii increased

its activity due to SO2 in the air.

Fungi, bacteria, viruses and nematodes, as well

confused with the similarities between the symptoms

as insects are all responsible for bringing a cer-

of many types of pollution damage and those of in-

tain species of higher plants into equilibrium with

fectious diseases sometimes leading to incorrect

its environment (Treshow 1968). The effect of air

diagnoses. Fortunately, each of the common air

pollutants on population trends of various micro-

pollutants produces a characteristic injury pattern,

organisms, therefore, is extremely important in all

and the sensitivity of many species differs suffi-

ecological systems. Further, air pollutants may

ciently for the trained eye to determine the caus-

affect plant disease development in different ways.

ative agent dams and Salzbach 1961) .
Pathogenicity may be influenced through a direct

effect of the pollutants on the parasite, or the

This paper will review the literature on: 1) the

effects may be indirect through pollution-induced

modification of parasitism and the population

changes in physiology of the host plant eagle

changes of epiphytic microorganisms induced by air

1973).
pollutants; 2) attempts to make reliable guides in

distinguishing among symptoms caused by air pollu-

Prior to 19601s, the interaction of microorgan-

tants and parasites, and those occasionally pro-

isms with pollutants in the air had received limit-

duced by saprophyfcic fungi. These subjects are

ed attention in Japan. But during the past decades,

considered in conjunction with a brief outline of

a series of experiments has been conducted in order

air pollution effects on forest ecosystems in Japan

to explain the effect of air pollution on dis-

ease development, and only recently, some attempts

have been made to understand the change of popula-

SULFUR DIOXIDE

tions of microorganisms for use as biological indi-

cators of air pollution.

In Japan, towards the end of 19th century, air

pollution was reported to be harmful to forests

From the beginning of recognition of the signifi-

around ore smelters that emitted large amounts of

cant and sometimes devastating effects of air pol-

SO2. Plume behavior resulted in pollutants disper-

lutants on vegetation, plant pathologists have been

sion in the surrounding area, killing all the veg-

etation (mainly Cryptomeria japonica D.Don and

Pinus densiflora Sieb. et Zucc.) and causing ero-

presented at the Symposium on Effects of Air

sion of topsoils following the death of the plants

Pollutants on Mediterranean and Temperate Forest

(Doi 1919). The concentration of SO2 of the smoke

Ecosystems, June 22-27, 1980, Riverside,

was reported to be 1 ppm at 1 mile from a large ore

California, U.S.A.
smelter located in Ashio, Tochigi Prefecture, and

occasionally reached as high as 8 to 10 ppm at the

2~orestPathologist, Kansai Branch, Forestry ground level (Doi 1919) .

and Forest Products Research Institute, Momoyama,

Fushimi, Kyoto 612 Japan.

 Raising the height of stacks in order to reduce

the concentration of SO at the ground level was

the first and primitive2control measure, new tech-

The analysis of air provides a framework of

nology involves the installation of sulfur recov-

knowledge, and a foundation for developing air

ery devices. In 1915, one ore smelter in Hitachi,

quality management strategies, and for prompting

Ibaraki Prefecture, built a new stack on a hill

the enactment of the Clean Air Act in 1967, and

rising about 350 m above sea level; the stack was

its amendments made in 1970.

nearly 170 m high, so that the smoke was discharged

at an elevation of approximately 520 m. Although

smelter smoke is released from high stacks, it

tends to diffuse rather slowly, and the SO., there-

fore, reaches the ground in harmful concentrations

even after a considerable length of time and dis-

tance. Observations made three years after the

replacement of stacks revealed the fact that the

severity of damage in the area near to the smelter

was markedly reduced,while the total damaged area

increased as large as 24,000 ha ( ~ o i1919).

The reduction of the concentration of S O in the

area in the vicinity of the ore smelter on account

Figure 1. Mean value of annual concentrations

of the change of stack height was good enough to

at 15 strategic locations.

resume the seedling production of Cryptomeria

Japonica in nurseries. Kaburagi (1930) was im-

pressed with the absence of a needle blight of

Figure 1 shows annual changes in average concen-

Cryptomeria ,japonica caused by Cercospora sequoiae

trations of SO at 15 strategic locations between

Ell. et Ev. in the nurseries located in the recov-

2
1965 and 1977, the concentration increased until

ered area close to the higher stack. The needle

1967, and then declined gradually. The decline

blight is one of the most destructive diseases in

has proved the continuing effectiveness of regula-

forest nurseries producing Cryptomeria seedlings,

tion of air pollution sources by the national and

and has spread rapidly throughout Japan except Ho-

the prefectural governments based on the act.

kkaido. The causal fungus was introduced from

North America in the end of 19th century, and Iba-

During the summers in the middle of 19601s,the

raki Prefecture including the ore smelter is con-

worst average concentrations of SO2 were experienc-

sidered to be the epicenter of the introduced dis-

ed. In this period, an unrecorded needle blight

ease (Ito 1976). Kaburagi (1930) suggested that a

of Japanese red pine, Pinus densiflora, occurred

strong correlation existed between SO in the air

throughout the normal range of that species, pre-

and the absence of the disease. This is believed

dominantly in central Japan. Up to that time the

to be the first report on the effect of air pollu-

disease had been little known and of little concern

tion on diseases caused by parasites in Japan. Ka-

to forest pathologists. A specific fungus domi-

buragi's observation is strongly supported by the

nantly found on blighted needles was investigated

following hypotheses that: 1) SO might act direct-

?
by Kobayashi (1967), and he identified it as Rhizo-
ly upon the fungus on the surfaces of host plants;
sphaera kalkhoffii Bubak which had been known as a

and 2) the reduction of disease severity resulted

causal fungus of a needle blight of spruce and fir

in a decrease of inoculum for new infection.

in northern Europe and North America.

Some of the acute type of damage by the emis-

The results of inoculation experiments to Japa-
sions from smelters still continues in areas near
nese red pine seedlings proved that the pathogen-

to sources. In the 19601s,widespread SO, air pol-

icity of the causal fungus was considerably weak

lution damage to ornamental trees had become appa-

for pines, and the disease would scarcely occur if

rent to the public; this increase was due to urban-

pines grew under normal conditions anak aka and Chi-

ization and industrialization. The one most appa-

ba 1971). The unusual incidence of the disease

rent episode in and around Tokyo was the stunting

occurring in the early s m e r of 1.965 might be re-

and death of a large number of Zelkova serrata Mak.,

lated to abnormal weather conditions--extremely

widely used as shade trees throughout Japan (yambe

little rain fell in early spring followed by heavy

1973).
rain in May (Chiba and Tanaka 1968). Although the

disease has not occurred so widely as in 1965,

With a considerable increase in public awareness

severe damage has been observed in the vicinity of

of the chronic type of pollution in many parts,

ore smelters or industrial areas. In addition,

automatic SO recorders were set up by the national

Japanese red pine is highly sensitive to S O (Ino-

2
and the prefectural agencies to determine the con-
ue 1973), and the distinction between the needle

centrations, frequency, and the duration of atmo-

blight symptoms caused by the disease and those

spheric SO exposures. The number of measuring

caused by SO is exceedingly difficult (Tanaka 1975,

locations $ad increased from the beginning at the

Tanita 1976)? Therefore exhaustive experiments
15 strategic locations in 1965 to as many as 1473
were conducted to examine the relationship of SO,,c-

locations in 1979, forming an extensive sampling

with disease development.

system for SO2 (The Japanese Environmental Agency

 A t o t a l of 163 two-year-old seedlings of Japan-
e s e r e d pine were exposed t o 2.0 ppm SO f o r 1,2,3,
and 4 hours i n a fumigation chamber before 2 or after
inoculation i n t h e middle of J u l y , with a water
suspension of conidia of t h e fungus produced on
PDA p l a t e s .
Damage was more severe i n t h e treatment combin-
i n g t h e fumigation with inoculation, e s p e c i a l l y
i n o c u l a t i o n before fumigation, compared with t h o s e
i n t h e treatment of fumigation without inoculation.
Numerous pycnidia of t h e fungus were produced i n
l a r g e l e s i o n s on needles t r e a t e d by fumigation
a f t e r i n o c u l a t i o n a s well a s fumigation f o r longer
duration before i n o c u l a t i o n , no obvious change i n
t h e appearance of needles was detected on t h e seed-
l i n g s inoculated with t h e fungus without fumigation.
Symptoms produced on t h e SO t r e a t e d needles, name-
2 t h e c o n i d i a l spraying of t h e c a u s a l fungus ( f i g u r e
l y , l a r g e l e s i o n s with numerous pycnidia, were
i d e n t i c a l t o symptoms of t h e needle 'blight occur-
r i n g i n t h e f i e l d ( ~ h i b a nd Tanaka 1968).
Most pine seedlings displayed acute symptoms of
i n j u r y following exposure t o 2.0 ppm SO f o r 2
hours. The SO concentration used i n this study
2 d i e b l i g h t e d Japanese red pines were found, and
was s e v e r a l times higher than t h a t found i n t h e
ambient a i r . It was necessary t o e l u c i d a t e t h e
r e l a t i o n s h i p between SO and d i s e a s e development a t
2
lower concentrations because such conditions a r e
f r e q u e n t l y encountered i n r u r a l and f o r e s t regions.
Therefore, three-year- o l d seedlings of Japanese
r e d pine were exposed f o r 2 weeks i n chambers t o
0.2 ppm SO t h e l e v e l of SO i s l i k e l y t o be found
2 t e r s and blowers, and t h e c o n t r o l o r ambient a i r
i n t h e ambient a i r (Tanaka 1576a).
I n t h i s experiment, s p e c i a l emphasis was placed
on t h e e f f e c t of SO fumigation a f t e r inoculation
i n comparison with $he treatment before inoculation.
The t r e a t e d needles with SO2 fumigation a f t e r inocu-
l a t i o n were more s e v e r e l y a f f e c t e d by t h e fungus,
compared with t h o s e fumigated by SO gas before
i n o c u l a t i o n . This r e s u l t i s c o n s i s t e n t with find-
ings of t h e 2.0 ppm SO fumigation (Chiba and Tana-
2 i s , a t o t a l of 300 seedlings were divided i n t o 12
ka 1968). S O i n j u r y appeared t o f a c i l i t a t e t h e
spread and reproduction of t h e fungus within tis-
sues of needles r a t h e r than t o i n c r e a s e t h e i n i t i a l
r a t e of i n f e c t i o n by t h e fungus.
To f u r t h e r examine t h e s e findings from a r t i f i -
c i a l SO exposure of seedlings i n chambers, f i e l d
2 disease i n t h e f i l t e r e d a i r compared with t h a t of
experiments were planned and conducted during t h e
summers of 1972 and 1973 by using a t r a n s p l a n t i n g
method anak aka e t a l . 1974a) and a f i l t e r e d a i r
method (Tanaka e t a l . 1974b) i n Hiroshima Prefec-
t u r e . I n t h e t r a n s p l a n t i n g study, two experimental
p l o t s were e s t a b l i s h e d . One was i n Fukuyama c i t y
c l o s e t o a l a r g e s t e e l r e f i n e r y . Especially i n
t h i s a r e a , d i s t i n c t SO damage markings on t h e
needles of Japanese r e g pine were v i s i b l e . The
o t h e r was i n a r u r a l a r e a i n Miyoshi c i t y . It was
l o c a t e d about 60 km n o r t h of a highly i n d u s t r i a l i z e d
a r e a along t h e Seto Inland Sea including Fukuyama
c i t y . I n t h e r u r a l a r e a , t h e d i s e a s e caused by
Rhizosphaera k a l k h o f f i i had not been noted i n
recent y e a r s . A t o t a l of 900 p o t t e d seedlings of
Japanese r e d pine used i n t h e t r a n s p l a n t i n g study
were cultivated, previously i n t h e n u r s e r i e s i n t h e
r u r a l a r e a i n Miyoshi c i t y . Seedlings were divided
i n t o 12 groups a s shown i n f i g u r e 2 according t o
t h e combination of t r a n s p l a n t i n g and inoculation.
SO concentrations of both a r e a s were determined
2
by means of t h e l e a d peroxide method. Needles were
analyzed f o r sulfate-S and f o r t o t a l - S dry b a s i s
i n order t o determine t h e accumulation of s u l f u r
i n t h e needles. F o l i a r symptoms were r a t e d from
0 t o 5 , according t o t h e degree of injury,and t h e
average of t h e s e i n d i v i d u a l indexes provides t h e
disease s e v e r i t y f o r each group.
After being t r a n s p o r t e d from t h e r u r a l a r e a t o
t h e i n d u s t r i a l i z e d a r e a , s e v e r a l seedlings exhib-
i t e d t y p i c a l symptoms and signs of t h e d i s e a s e ,
while o t h e r s remaining i n t h e r u r a l a r e a displayed
no evidence of i n j u r y , even though they received
2).
In t h e experiment using a f i l t e r e d a i r method,
an experimental p l o t was e s t a b l i s h e d i n an a r e a
i n Mihara c i t y c l o s e t o a powdered-coal-burning
power p l a n t . I n t h i s a r e a , a l a r g e number of nee-
they displayed gradual reduction i n shoot and wood
growth. The previous y e a r ' s annual mean contents
of s u l f u r compounds i n t h e a i r was 1.04 mg SO 1100
3
cm2 PbO2 per day.
P l a n t s were grown i n p l a s t i c covered chambers.
One chamber ( 2 x 4 ~ 2m ) was equipped with carbon f i l -
chamber was provided with blowers. The a i r passed
through each chamber a t a r a t e of about one a i r
change / min. SO2 concentrations of t h e ambient
a i r were determined by means of an automatic re-
corder, and SO concentrations i n chambers were
2
measured, by t h e l e a d peroxide method.
The experimental design was s i m i l a r t o t h e t r a n s -
p l a n t i n g experiment i n Fukuyama and Miyoshi, t h a t
groups according t o t h e combination of exposure t o
SO2 and inoculation of t h e c a u s a l fungus, then seed-
l i n g s were placed i n chambers t o determine whether
some p r o t e c t i o n from i n f e c t i o n could be obtained
when t h e a i r drawn i n t o t h e chamber was f i l t e r e d .
There was a l a r g e reduction i n t h e s e v e r i t y of t h e
t h e disease i n t h e ambient a i r . These findings by
f i e l d s t u d i e s furnished t h e evidence t h a t SO2 i n
t h e a i r was responsible f o r t h e development of t h e
disease. The s u l f u r content e s p e c i a l l y sulfate-S
i n t h e needles of t h e t r e e s exposed t o SO2 was s i g -
n i f i c a n t l y g r e a t e r than t h a t of untreated needles
both i n t h e t r a n s p l a n t i n g and t h e f i l t e r e d - a i r ex-
periments.
An increasing i n disease:severity may.be expected
when a i r p o l l u t a n t s weaken t r e e s , making them more
s u s c e p t i b l e t o i n f e c t i o n by weak p a r a s i t e s , o r re-
duced vigor t o a degree t h a t p l a n t s become predis-
posed t o a t t a c k by f a c u l t a t i v e p a r a s i t e s , such a s
Rhizosphaera k a l k h o f f i i , and B o t r y t i s cinerea Pers.
.
( ~ a n n i n ge t a 1 1970 ) . Conversely, Linzon (1958 )
 On July 18, 1970, Zelkova serrata trees in a

yard of a senior high school located in Suginami

2sp j

Ward in Tokyo showed unusual shedding of their

leaves, and strawberry leaves showed white spots

clearly indicating the effects of oxidants. In

!i 2j,6;/
---
(U addition to the damage to plants, 45 students of

4
CO
the school were hospitalized for treatment of irri-

0 0 tated eyes, sore throats and difficulty in breath-

J A S D J A S D J A S D
ing. From this evidence, the Metropolitan Govern-

5.11 ment issued the statement that the damage to humans

and vegetation was caused by photochemical air pol-

lution (The Tokyo Metropolitan Government 1971),

and this was thought to be the first episode of

transportation
'Los Angeles type smog" in Japan. But as a result

inoculation
of later investigations it was found that there

P
J A S J T ! were differences between Tokyo and Los Angeles air

J : Jun 20
pollution, since SO2 concentration and humidity in

A:Aug 1 Tokyo were much higher than in Los Angeles; fur-

S: Sep 13 thermore, it was confirmed that there was forma-

tion of sulfuric acid in the air (The Tokyo Metro-

politan Government 1971). These findings suggest

that the complex of photochemical oxidants and

sulfuric acid might exert a far greater influence

on humans and vegetation than any other alone at

that time.

In the latter half of 19701s,with continuous

-0-0-0-0 increasing industrialization and transportation,

J A S D JAS
- J D 0-J every major city in Japan has experienced an in-

Figure 2. Effect of SO2 on the development of

creasing in photochemical oxidants even though

needle blight of pine caused by Rhizosphaera

there was a slow but steady decline in the concen-.

kalkhoffii
tration of SO2,in the ambient air (figure 1).

Photochemical oxidants , mainly ozone (0 ) ,con-

found that smelter emissions decreased the inci-
tinue to cause the defoliation of broad leaf trees.

dence of Cronartium ribicola C.J.Fis.,and there are

Especially, in the Kanto district, at the center of

additional reports that aggressive or obligate para-

Tokyo, unusual defoliation of Zelkova serrata

sites, such as Microsphaera alphitoides Griff. et

still continues, since it is sensitive to both SO2

Maubl. ( ~ o c h1935) , and Melampsorella cerastii Wint , (Inoue 1973) and 0 (Kadota and Ohta 1972). While,

Peridermium coloradense Arth. et Kern, and some

in the Kansai dist?ict,at the center of Osaka,

species of Melampsora and Gynmosporangium (Scheffer

there is a high incidence of defoliation of poplar

and Hedgcock 1955) were less abundant in areas ad-

trees in the early summer in the parks, the gardens

jacent to smelter zones or to industrialized areas.

or the yards of schools throughout the district.

ltol found a similar case in Tokyo that a"close re-

lationship existed between the increase of distance

In addition to the measuring system of SO2

from the metropolitan area and the increase of the

throughout the country, a number of automatic oxi-

incidence of willow rust disease caused by

dant recorders were also set up by the national and

Melampsora coleosporioides Dietel and a leaf rust

the prefectural agencies, but there is still a

disease of Styrax japonicum Sieb. et Zucc. caused

shortage of satisfactory apparatus for monitoring

by Pucciniastrum styracinum Hirat. The complex of

the low ambient concentrations. To make up for the

air pollutants was brought about by urbanization

shortage of an instrumented system, plants have

and industrialization, typical of the mixture of

been used extensively as bioindicators of air pollu-

pollutants occurring in and around Tokyo, but SO2

tion. The use of bioinaicators has long been ac-

was considered to be principally responsible for

cepted, since damage to green plants is usually one

reducing the abundance of these fungi.

of the first signs that air pollution is becoming

a serious problem e err^ 1964); their usefulness in

OXIDANTS MAINLY OZONE ( 0 ) this capacity is based primarily on the sensitivity

of selected plant species and varieties for speci-

Beginning in the 19701s, changes from solid to

fic pollutants (Heck 1966).

liquid and gas fuels, and especially increasing use

of motor vehicles, have caused a reduction of the

Many clones, species and varieties of poplars

SO type of urban pollution and an increase of

were selected as bioindicators because of their

"ozone" pollution.
high sensitivity to oxidants and their wide distri-

bution, especially in the Kansai district. They

also have enounnous advantages in that they are

Personal communication from S. Ito, University

of Tokyo, January 15, 1980.

fast growing, easy to propagate vegetatively by

c u t t i n g s , and a r e uniformely s e n s i t i v e t o oxidant
injury.
Despite t h e s e advantages, poplars a r e frequently
plagued by various kinds of i n s e c t s and diseases
t h a t degrade t h e usefulness of poplar clones as
p o l l u t i o n i n d i c a t o r s . For example, i n t e r p r e t a t i o n
of t h e response t o oxidants i s made d i f f i c u l t by
l e a f diseases caused by Marssonina brunnea (El1.et
Ev.) Magnus, S e p t o t i n i a podophyllina Mhetz. and
Melampsora l a r i c i - p o p l u l i n a Kleb. A r e l i a b l e guide
t o d i s t i n g u i s h among symptoms caused by a i r pollu-
t a n t s , i n s e c t s , and diseases has been developed and
published ( Tanaka 1975 ) . The d e f o l i a t i o n caused by
Marssonina brunnea i s one of t h e most serious dis-
eases. It occurs wherever poplars grow, and i s not
r e s t r i c t e d t o urban regions, but frequently occurs
with s e v e r i t y i n urban areas. The disease develops
t i n y spots with l i g h t margins about 1 mm i n dia-
meter over t h e e n t i r e surface of t h e l e a f . These
s u p e r f i c i a l spots a r e not so d i f f i c u l t t o d i s t i n -
guish from t h e symptoms caused by ozone by p l a n t
p a t h o l o g i s t s with a t r a i n e d eye, but many people
who want t o use poplars a s i n d i c a t o r s a r e confused
with t h e s i m i l a r symptoms produced by ozone and t h e
disease (Tanaka 1975). I n addition, t h e disease
causes premature lower l e a f abscission,and t h e de-
f o l i a t i o n progresses upward toward t h e t i p s of t h e
shoots from t h e l a t e spring through t h e summer
months. The unsightly appearance of a f f e c t e d t r e e s
discourages poplars' use as i n d i c a t o r s of oxidants.
A number of i n v e s t i g a t o r s suspected t h a t abscis-
s i o n was not due d i r e c t l y t o t h e pathogen but re-
s u l t e d i n s t e a d from increased r a t e s of ethylene
production from t h e i n f e c t e d p l a n t s (Abeles 19731,
f u r t h e r , ozone-induced abscission of leaves of vary-
ing maturity was c l o s e l y c o r r e l a t e d with r a t e s of
ethylene production ( ~ b e l e s1973). From t h e s e
Table 1. Relationship between c l o n a l s u s c e p t i b i l i t y of poplars t o 0
I S u s c e p t i b i l i t y t o Marssonina brunnea
Highly
Resistant
resistant
resistant
Resistant (R) 1 HE 2, OP 26,
IModerately
resistant
I 72/51, Kamabuchi,
Highly O J 115, OJP 1, OJP 2,
susceptible(ss^ OJP 3, OJP 4, OJP 5,.
OP 4 1
reasons, Tanaka (1976~))hypothesized t h a t increased
r a t e s of ethylene production caused by t h e syner-
g i s t i c e f f e c t s of t h e i n f e c t i o n of t h e fungus and
ozone was responsible f o r t h e incidence of t h e dis-
ease i n t h e urban area.
Fortunately, t h e r e i s s t r i k i n g v a r i a b i l i t y i n
s u s c e p t i b i l i t y of poplar clones t o t h e fungus and
ozone. Clones were s e l e c t e d t h a t a r e s e n s i t i v e t o
ozone and r e s i s t a n t t o t h e disease. The s e l e c t i o n
o f f e r s one of t h e best means of increasing t h e usa-
b i l i t y of poplar clones a s ozone i n d i c a t o r s (Tanaka
1977a). Table 1 presents t h e r e l a t i o n s h i p between
ozone s e n s i t i v i t y and disease r e s i s t a n c e of poplar
clones (Tanaka 1 9 ~ ~ a )The . data of r e s i s t a n c e of
poplar clones t o t h e disease a r e based on t h e t h r e e
years' observation made i n Osaka using 38 clones of
young rooted c u t t i n g s t an aka 1 9 7 ~ a ) ,and t h e d a t a
of ozone s e n s i t i v i t y of them a r e based on t h e re-
s u l t s from t h e exposure experiments with 0.15 ppm
ozone f o r 40 hours i n chambers conducted by Enoki
(1977).
Populus maximowiczii cv. O J 115 and cv. OJP 1,
.
P m a x i m o ~ i ~ c h o c a cv .
r ~ OaP 41, P .maxino-
& z i i x 2.nigra pl&tierens cv.0P 52, and P.nigra
i t a l i c a x ~.maximowiczii,cv.Kamabuchia r e promising
clones f o r ozone i n d i c a t o r s because they a r e sensi-
t i v e t o ozone and r e s i s t a n t t o t h e d i s e a s e ( t a b l e 1 ) .
Yambe (1978) a l s o attempted t o f i n d a possible
usefulness of t h e change of microbial f l o r a on bam-
boo leaves a s i n d i c a t o r s f o r a i r p o l l u t i o n . He
found t h a t Puccinia kusanoi D i e t e l and epiphytic
b a c t e r i a on t h e l e a f surfaces of bamboo decreased
sharply i n number i n t h e urbanized and i n d u s t r i a l -
ized areas.
and t o Marssonina brunnea.
3
Moderately
Susceptible Highly
resistant susce'otible
Serotina
NR 19,
MR 32 I 45/51
Gerlica
I 262
I 154, I 214
I 77/51,
I 488
 OTHE8 POLLUTANTS
I n a d d i t i o n t o SO and oxidants, s i g n i f i c a n t and
2
sometimes devastating e f f e c t s of o t h e r a i r pollu-
t a n t s have long been recognized i n Japan. Fluoride
compounds have a long h i s t o r y a s p o l l u t a n t s i n t h e
v i c i n i t y of ceramic i n d u s t r y , and more r e c e n t l y ,
p l a n t i n j u r y r e s u l t e d from f l u o r i d e s has gradually
increased because of t h e expansion of i n d u s t r i e s
such a s aluminum r e f i n e r i e s m d f e r t i l i z e r manu-
f a c t u r e s ( ~ a t u u r aand Kokubu 1972). Chlorine (Ta-
naka 1977b), ethylene, a c i d i c dust from s t a c k s , and
a l k a l i n e dust from cement k i l n s (Inoue 1972a and
1972b) have a l s o been recognized a s agents of dm-
age t o v e g e t a t i o n y but t h e damaged a r e a s a r e r e l a -
t i v e l y r e s t r i c t e d because most of them a r e emitted
from point sources o r by a c c i d e n t a l s p i l l s .
The i n t e r a c t i o n of t h e s e minor p o l l u t a n t s with
fungi and b a c t e r i a has received only minor a t t e n -
t i o n , but Inoue (1972a and 1972b) found t h a t t h e
number of f r u i t i n g bodies of Tricholoma matsutake
Singer was s i g n i f i c a n t l y reduced i n t h e heavily
dusted f o r e s t s i n t h e v i c i n i t y of cement works com-
pared t o l e s s severely dusted o r non-dusted por-
t i o n s of f o r e s t s . According t o t h e r e s u l t s of
hand-dusted experiments,he concluded t h a t t h e alka-
l i n e s o l u t i o n from t h e dust severely decreased t h e
formation of t h e f r u i t i n g bodiesy and reduced t h e
development of t h e fungus colonies.
CONCLUSIONS
The i n t e r a c t i o n of p o l l u t a n t s with p a r a s i t i c and
epiphytic microorganisms has received l i m i t e d a t -
t e n t i o n i n Japan. The e x i s t i n g r e p o r t s a r e only
a s s o c i a t e d with t r e e s and bamboos and no r e p o r t
with annual p l a n t s was found i n a search of a v a i l -
able l i t e r a t u r e .
Some p o l l u t a n t s a r e d i r e c t l y t o x i c t o microor-
ganisms on t h e l e a f surface. They might possibly
impair t h e i r growth o r reproductiony and change t h e
population and community composition of mCcroorgan-
i s m s on t h e h o s t p l a n t s , s i n c e t h e s e n s i t i v e spe-
c i e s t o a i r p o l l u t a n t s would be gradually replaced
by more t o l e r a n t s p e c i e s (Treshow 1968). Especial-
l y y aggressive o r o b l i g a t e p a r a s i t e s such a s Cerco-
spora sequoiae (Kaburagi 1930), Pucciniastrum m-
racinum and Melampsora coleosporioides ( 1 t o 1 ) ~ and
Puccinia kusanoi (Yambe 1978) a r e l e s s abundant i n
polluted a r e a s y and t h e d i s e a s e s e v e r i t y they induce?
i s s h a r p l y reduced.
On t h e o t h e r hand, i n f o r e s t stands under i n f l u -
ence of i n d u s t r i a l and urban a i r p o l l u t i o n , some
f o l i a r pathogens counted by p a t h o l o g i s t s i n t h e
group of so-called ' f a c u l t a t i v e p a r a s i t e ' a r e of
g r e a t e r importance because a i r p o l l u t a n t s i n j u r e
l e a f t i s s u e and predispose t r e e s t o a t t a c k by
f a c u l t a t i v e p a r a s i t e s . A c l o s e c o r r e l a t i o n has
been found between SO i n t h e a i r and a high i n c i -
dence of needle b l i g h z of pine caused by a weak
' Personal communication from S. It0
of Tokyo, January 1980.
University
p a r a s i t e y Rhizosphaera k a l k h o f f i i (Chiba and Tana-
ka 1968).
Although p l a n t i n j u r y caused by s p e c i f i c pollu-
t a n t s such a s SOpy 0 and f l u o r i d e s has gradually
been abated by enginJering improvements o r by-pro-
ducts recovery, chronic i n j u r y caused by a complex
of urban a i r p o l l u t a n t s has assumed a d d i t i o n a l i m -
portance a s urbanization and t h e use of motor ve-
h i c l e s have increased.
I n j u r y t o green p l a n t s i s usually one of t h e
f i r s t s i g n s t h a t a i r p o l l u t i o n is becoming a s e r i o u s
problem, and s e v e r i t y of s p e c i f i c d i s e a s e s i n pol-
l u t e d a r e a s must be regarded a s an a d d i t i o n a l ex-
pression of unbalanced n a t u r a l conditions. These
s p e c i f i c diseases and t h e change of composition of
l e a f surface microorganisms provide a i d s f o r moni-
t o r i n g a i r p o l l u t a n t s y e s p e c i a l l y i f they a r e sen-
s i t i v e t o very low concentration of a i r p o l l u t a n t s .
These diseases and microorganisms should be consid-
ered a u s e f u l supplement t o t h e more expensive
techniques and instruments used f o r t h e i d e n t i f i -
c a t i o n of chronic a i r p o l l u t a n t e f f e c t s .
Acknowledgments: I thank D r . Paul R. M i l l e r ,
P a c i f i c Southwest Forest and Range Experiment Sta-
t i o n , U.S. Dep. A g r i c a Yf o r h i s c r i t i c a l readings
of t h e manuscript.
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Adms $D,F and C W. Sulzbach .
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a e r a k a l k h o f f i i Bubk. I. J.Jap.For.Soc. 50:
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lg72a. Studies on the influence of smoke-dust

1976b. Tree diseases in air polluted areas and

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Tanaka,K.

1977a. Pest problems in the use of poplar clones

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1972b. Ditto 11. M:215-288. . . .
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TanakayK.,T.Okada and K.Hako.

1976. Blight and canker of Cryptomeria Japonica

1974a. Effect of SO2 on the development of needle

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blight of pine caused by Rhizosphaera kalkhoffii,

.-%-
Everhart. Ann.Phytopath. Soc Jap
2 :234-236. .
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2163pp. Tokyo.
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Kaburagi,T.

1930. (Smoke damage to plants). Iwanami Lectures

Tanaka,K. and 0.Chiba.

for Bioloa. 1-39.

1971. On a needle blight ~f pine caused by Rhizo-

sphaera kalkhoffii--Life historyy physiological

Kobayashi>T.
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fungus. J.Jap .For,Soc 53:279-286.
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a little-known fungus group associated with

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needle disease of conifers. Bul1.Gov.For.Exp.
1976. Histological effects of air pollutants and

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1972. Ozone injury to some Japanese woody plant

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lanzenkr 45 :44-45. lations. Phytopathology 58:1108-1113.

Linzon,S.N.
Ymbe, Y.

1958. The influence of smelter fumes on the growth

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of white pine in the Sudbury region. Can.Dep.

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Agr Publ. ,Ontario Dep.Lands Forests. 45pp.
Ymbe, Y.

Matuura,S. and N.Kokubu.

1978. Decline of trees and microbial florae as the

1972. Fluorine, its distribution and role in

index of pollution in some urban areas. Bull.

nature. 150pp. Tokyo.

For.& For.Prod.Res.1nst. 301:llg-129.

Scheffer,T.C. and G.G.Hedgcock,

19.55. Injury to northern forest trees by sulfur

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1117:49pp.

Tanaka,K.
1975. The similarity between symptoms caused by

air pollutants, insects, and diseases. Agr. and

Hort. 5O:Preface.

Tanaka ,K.

1976~~.
m e effect of low-level SO fumigation on

the development of needle bligh$ of pine caused

by Rhizosphaera kalkhoffii. Anr1.Rep.1975.~m e

range of tolerance of biological indicators to

pollutants :5-9.
 Sensitivity of Lichens to Air Pollution
with an Emphasis on Oxidant Air
Pollutants1

Thomas H. Mash I11 and Lorene L. Sigal^

Abstract: The hypothesis that lichens are sensitive

indicators of air pollution is now well established for

oxidants, sulfur dioxide, hydrogen fluoride and trace ele-

ments. From field studies differential sensitivity of dif-

ferent lichen species around pollution sources is evident.

Laboratory studies with the particular air pollutant general-

ly confirm the same degree of differential sensitivity.

LICHENS AND THEIR ECOLOGY

constraints on photosynthetic activity, primary

productivity assignable to lichens is generally

Lichens, which strictly speaking are called

small compared to higher plants. Nevertheless

lichenized fungi, are composite plants represent-

lichens may be important in ecosystem functioning.

ing a symbiotic union of algae and fungi to form

For example, lichens are generally recognized as

morphological entities unlike either component

important pioneer plants of xeric successional

(Ahmadjian 1967, Hale 1967). Although a few

sequences because of their role in biologically

species survive in aquatic environments, most are

controlled weathering (Syers and Iskandar 1973).

found in terrestrial habitats, occurring on soil,

In addition, lichens are important in mineral

rocks and as epiphytes on other plants. All

cycling, not only because of their ability to alter

lichens are perennial plants with life spans

precipitation chemistry (Ianq and others 19761, but

varying from a few decades to reportedly thousands

also from the ability of blue-green algae contain-

of years (Weber and Andrews 1973). Essentially

ing lichens to fix atmospheric nitrogen. Den son

all lichens are autotrophs because of the photo-

(.1973) has estimated that lichens are respons ble

synthetic activity of the algae. Metabolic

for 50 percent of the nitrogen fixed in the

activity in general is limited to period's when the

Douglas-fir forests of the Pacific Northwest.

lichens are moist, a condition which varies in

concert with atmospheric moisture conditions as

Lichens are morphologically divided into 3 major

lichens have no roots by which moisture might be

growth forms: crustose, foliose and fruticose.

absorbed from the substrate. Thus lichens are

Crustose species are generally small, tightly

prominent examples of poikilohydric plants

attached to (or imbedded in) their substrate and

(Larcher 1973) which also include bryophytes and

poorly differentiated. Foliose species are gen-

some lower vascular plants, such as desert species

erally larger, loosely attached to their substrate

of Selaginella and ferns. Because of moisture

and stratified into several distinct layers. In

contrast to the basically 2-dimensional foliose

species, fruticose s~eciesare strongly 3-dimen-

sional and grow out from their substrate. Most

presented at the Symposium on Effects of Air

fruti'cose species are also differentiated into

Pollutants on Mediterranean and Temperate Forest

distinct layers. Of these morphological forms

Ecosystems, June 22-27, 1980, Riverside,

fruticose species are generally considered to be

California, U.S.A.
the most sensitive to air pollution (Fenton 1964),

'~r. Nash is associate professor of botany,

a fact which may be related to the high surface-

Dept. of Botany, Arizona State University, Tempe,

to-volume ratio of this form.

Arizona 85281 USA; Dr. Sigal is a research scien-

tist, Environmental Sciences Division, Oak Ridge

National Laboratory, P.O. Box X, Oak Ridge,.

LICHENS IN RELATION TO AIR POLLUTION

Tennessee 37830 U.S.A.

Different species of lichens are well-known to

be differentially sensitive to air pollution (Nash

Californian mountains* The study was centered in

1976). A few species actually grow better in

the San Bernardino Mountains, utilizing the sites

urban environments where air pollution levels are

mployed by the University of California (Riverside

high in contrast to their performance in rural

and Davis] ecosystem study of air pollutant effects

areas. Most species, however, exhibit varying

in contrast to sites in a control region, Cuyamaca

degrees of sensitivity to air pollution and some

Rancho State Park in southern San Diego County.

are more sensitive than higher plants. Thus by

In addition, there was an excellent historical

documenting variations in abundance and species

record of the lichens of the San ~ernardinosbased

richness of lichen communities, it is frequently

on collections made by H. E. Hasse at the turn of

possible to identify poliution sources and to

the century. Of the 91 foliose and fruticose

document the magnitude of the air pollution probla,

lichen species reported by Hasse (1913) in his

Patterns within lichen communities may be attribut-

lichen flora of southern California, only 34

able to many environmental factors other than air

species were found during three sunnners of study

pollution, of course, and consequently it is

throughout southern California. Thus there has

extremely important that studies be made within a

been a significant reduction in the number of

multivariate context, including both pollution

species present. To obtain quantitative data on

factors and non-pollution factors.

lichen variation, both conifers (Ab2es concolor,

Pinus ponderosa, Pinus jeffreyi and Pseudotsuqa

The fact that lichens are differentially sensi-

macrocarpa) and California black oak (Quercus

tive to air pollution is based on a series of

kelloggii) were sampled for lichen cover at

field observations and laboratory experiments

breast height.

extending back over the past 130 years. Grindon

(1859) and Nylander (1867) noted that lichens were

In the case of conifers Hasse (1913) reported

disappearing from city centers. Later Arnold

the presence of 16 foliose and fruticose lichen

(1892) documented that lichens transplanted from

species in the San Bernardino Mountains. In our

rural areas to the center of Munich were not able

recent study [Sigal and Nash 19801 only 8 of

to survive. Subsequently this "city effect" has

these species were found in the San Bernardino

been reported for many cities in Europe and North

Mountains and 4 of these were only present in

America (Hawksworth 1971). By air pollution most

vestigial quantities. In contrast, at sites to

workers of the 1800's generally meant coal soot

the north and to the south K'uyamacasl 15 of the

and related particulates. However, over the past

16 species reported by Hasse were found. Thus in

decades a number of invisible gases, such as

the San Bernardinos which lie adjacent to the Los

sulfur dioxide and ozone, have been identified as

Angeles Bash, there has been a SO percent decline

the major causes of air pollutant injury to

in species richness of lkhens on conifers. Of

plants. Experiments with these air pollutants

the species which do occur in the San Bernardinos

over the past two decades have also demonstrated

only two (Letharia Vulpina and Hypogymnia entero-

differential sensitivity among different lichen

momha) occur commonly. The latter species,

species. In general, the sensitivity patterns

although it is common, is definitely showing signs

demonstrated in the laboratory have remarkably

of deterioration (Table 1) when randomly selected

Corresponded to the patterns observed in the field

thalli from the San Bernardinos are compared with

around pollution sources. Perhaps the strongest

a similar set from the Cuyamacas. The San Ber-

evidence supporting the contention that lichens

nardino population has a much higher percentage

are sensitive to air pollution comes from studies

documenting lichen recovery following pollution

abatement. For example, Seaward (19761 has docu-

Table 1-- Morphological characteristics of recent

mented that Lecanora muralis fairly rapidly

collections of Kypogymnia enteromorpha from the

reinvaded an urban complex in southern England

San Bernardinos and the Cuyamacas.

following amelioration of air pollution in the

area.

Percent Bleaching

In the following sections specific evidence

related to various types of air pollutants w211

Forest

be reviewed.

Oxidant Air Pollutants

38 59 41

Oxidant air pollutants, including ozone and

Percent Convoluted

peroxyacetplnitrate (PANI, are a group of gases

characteristic of the oxidizing atmospheres found
Forest

in the Los Angeles type smog (Pitts and Finlayson

1975). Although the effects of these compounds

38 38 47 15

have been studied extensively with higher plants

(National Academy of Sciences 19771, relatively

few studies have been conducted with lichens.

of bleached and convoluted thalli. In addition,

Probably the only field study in which lichens

there was a decrease of approximately 50 percent

have been shown to be sensitive to oxidants is

in overall thallus dimensions in the San Bernardino

that of Sigal and Nash (1980) for the southern

population and a decrease in fertility of 42

percent. Thus there are marked trends between the
San Bernardinos and the Cuyamacas of reduced
species richness and species vitality in the form-
er area. Within the San Bernardinos themselves,
significant variation exists. Below approximately
150 ppm-h oxidant dosage, cover of both Letharia
and Hypogymnia is normal in comparison with other
areas. At higher dosages of up to 285 ppm-h
lichens are almost completely eliminated from
conifers.
Comparisons of trends of lichens occurring on
black oak between the San Bernardinos and Cuyamacas
also indicated that a deterioration was occurring
in the San Bernardinos, but the trends were not as
dramatic as in the case of lichens occurring on
conifers. No direct comparison with the Hasse
material was possible because Hasse did not note
the species of oak in his collections. From our
sampling 39 species of lichens were found on black
oak in the Cuyamacas and 30 specie3 in the San
Bernardinos. Fourteen of the species found in the
Cuyamacas were not found in the San Bernardinos
the San Bernardinos. Among the 20 species which
occurred in common there was a general shift from
high frequency in the Cuyamacas to relatively low
frequency in the San Bernardinos. In the case of
cover of the 6 most common species, no significant
difference was observed between the two mountain
ranges except for Collema nigrescens, the only
nitrogen fixing lichen in the groupi which was
completely absent from the San Bernardinos. Fur-
ther analysis revealed that Parmelia subolivacea,
the most common lichen, was a successional species.
Because older trees occurred in the Cuyamacas, a
significantly higher value for this species in the
Cuyamacas was predictable. Further evidence sup-
porting the hypothesis that the dominant species
was in fact respondhg to oxidants was obtained
by running ordinations (princiwal component analy-
sis) of the sites within the San Bernardinos and
relating the observed variation in species com-
position to oxidant dose estimates and other
environmental variables (fig. 1). An initial
ordination (not shown) included all sites sampled
within the San Bernardinos. It exhibited a small

.
whereas three species were found exclusively in cluster of low cover sites which occurred at the

13.5
12.1
ucc
COO. 12.9
7.6
10.2 12.1

TOTAL % COVER
ALL L I C H E N S
15.7
BF. I % COVER
PARMELIA SUBOLIVACEA
BF
8.2

Bp CA GVC 4
DWA T2 145.8
230.8 186.7
184.9. ,,cc
COO. 134.7
i33.2
175.0

D
SAN BERNARDINO NAT'L FOREST
183.2 STUDY - S I T E LOCATIONS
OXIDANT DOSE ESTIMATES
BF.

Fig. 1--Principal component analysis ordination of 10 sites in the San Bernardino Moun-
tains using cover data for the 5 most important lichen species occurring on Quercus
kelloggii. Eighty percent of the variation is explained by axis one (the abscissa).
Figure 1A shows the positioning of the sites on the ordination plot with respect to to-
tal cover of all lichens; figure lBf the same ordination but with cover values for
melia subolivacea; figure lC, the same ordination, but with oxidant dose estimates
=-
(ppm-h); figure IDf the geographic location of the sites within San Bernardino Nation-
al Forest.
highest elevations where summer fog is infrequently
source of sulfur is sulfur dioxide. The gas

observed. Consequently these sites and one other

readily dissolves in water forming a sulfite or

site on the desert side of the San Bernardinos were

bisulfite solution depending on the pH. As a con-

excluded from the subsequent analysis. The 10

sequence acidification of the substrate frequently

sites plotted in figure 1 are thus homogeneous

occurs, as has been shown in Stockholm where Skye

with respect to altitude and position along the

(19681 found that urban tree bark had a pH of more

Los Angeles side of the San Bernardinos (fig. 1Dl.

than two units less than corresponding trees from

In the subsequent ordination (fig. lA, 1B and lC),

the country-side. Acidification strongly affects

80 percent of the variation is explained by axis 1

the ability of lichens to survive, in part because

(the horizontal axis). From figures 1A and 1B it

sulfur dioxide is much more toxic at lower pH's

is clear that the primary source of variation is

(Tuerk and Wirth 1975). Gilbert (1965 and 1970)

the percent cover of all lichens, which in turn

has clearly demonstrated the ability of lichens

corresponds closely to the percent cover of E. to penetrate into central Newcastle on basic sub-

subolivacea. Furthermore, there is an inverse

strates when they are absent from acidic ones.

relationship between percent cover of E.s- In the laboratory studies, photosynthetic decline

olivacea (fig. 1 ~ )and oxidant dose estimate (1C).

in response to short term sulfur dioxide exposures

This inverse relationship was substantiated by

as low as 0.2 ppm have been demonstrated (Tuerk

running a Speaxman rank correlation between the

and others 19741. In aqueous experiments with

two variables. The test was significant with an

reputedly lower concentrations, effects have also

alpha value of 0.007. For the other 4 species no

been documented Baddeley and others 1973, Puckett

relationship between oxidant dose estimates and

and others 19741.. In addition to reduction of

cover values were found. Thus some species have

photosynthesis, bleaching of lichen thalli may

been completely eliminated from the San Bernardinos,

occur due to phaeophytinisat2on of the chloro-

other species have probably declined in abundance

phyll~ (Nash 1973, Puckett and others 19731. More

and a few species are apparently unaffected.

recently potassium efflux from lichens exposed to

sulfur dioxide has been shown to be the most sen-

Preliminary ozone fumigations have shown dif-

sitive criterion of response (Tomassini and others

ferential responses in photosynthesis reduction

1977). Alteration of membrane permeability may

between Pamelia sulcata and Hypogymnia entero-

thus be an immediate response to sulfur dioxide

morpha with the former species exhibiting greater

exposure. Subsequently it is suggested (Nieboer

sensitivity (Nash and Siyal 19791. The Parmelia,

and other 1976, Puckett and others 1974) that

which grows on black oak, is absent from the San

interference in electron flow in photosystem I and

Bernardinos whereas the Hypogymnia is present, but

I1 will occur'and that proteins will be affected

is exhibiting signs of deterioration. For these

through sulfur dioxide$ ability to cleave disul-

species, at least, the pattern of sensitivity

phide linkages. Thus there is very strong evi-

observed in the field and the laboratory are com-

dence from both field and laboratory studies that

plementary. A similar pattern of response for the

lichens are sensitive to sulfur dioxide.

two species was observed after fumigations with

PAN (Sigal and Taylor L979). Further stud2es with

Hydrogen Fluoride

both ozone and PAN should be run, but the current

evidence strongly supports the assertion that

Hydrogen fluoride is extremely volatile and is

oxidants are a major cause of the decline of the

released into the atmosphere during aluminum and

lichen flora in the San Bernardino Mountains.

rare earth metal refining, fertilizer production

and glass and ceramic manufacturiny. Patterns of

Sulfur Dioxide
lichen decline are well demonstrated around alumi-

num factories (Martin and Dacquard 1968, Gilbert

Sulfur dioxide is released into the atmosphere

1971, LeBlanc and others 1972, Horntredt 1975) and

by combustion reactions involving products con-

around a titanbn plant (Nash 19711 and around a

taining the ubiquitous element sulfur, such as

phosphate plant (Takala and others 19781. In

coal and oil. It has long been recognized as a

these areas fluoride accumulat~onby the lithens

phytotoxic agent in general (Daines 1968).

has been demonstrated with decline in the lichen

Lichens are well known to be sensitive to sulfur

cornunities being associated with elevated fluo-

dioxide (Nieboer and others 19761. Most of the

ride levels. Transplants of healthy lichens into

field studies in northern Europe and North America

the impoverished zones resulted in fluoride accu-

have involved sites where sulfur dioxide was a

mulation and subsequent death of the transplants.

major factor (Hawksworth 1971). One of the

Gilbert suggested that the critical fluoride level

classic studies by Rao and LeBlanc (1967) demon-

lay between 20 and 50 ppm and Nash independently

strated a severe decline in lichen abundance along

suggested that the value lay between 30 and 80 ppm.

a 70+ km transect NNE of Wawa, Ontario where an Schoenbeck (1969) found that his transplants died

iron sintering plant is present. Because there is

near a fluoride source, but that new transplants

essentially no human development in the affected

survived well for several months following cessa-

region, air pollution, and particularly sulfur

tion of the fluoride emissions. In a laboratory

dioxide, is undoubtedly a major cause of the de-

study with exposure to approximately 5 ppb fluoride

cline in the reqionts lichens. Other studies

over a 9-day period Nash (1971) found that lichens

have shown that lichens accumulate sulfur

accumulated 84-115 ppm fluoride in contrast to

(Olkkonen and Takala 1975, Laaksovirta and

control levels of 14-25 ppm and that the fumigated

Olkkonen 1977) in situations where the presumed

samples developed chlorotic and necrotic symptoms

similar to the injury observed in the field trans-

plants. Thus fluoride as the probable cause of

the injury was confirmed.

Trace Elements

Lichens have a well demonstrated ability to

accumulate elements from dilute aqpeous solutions

(Nieboer and others 1977)# a fact probably related

to their apparent dependence on atmospheric

sources for mineral nutrition. As a consequence

of this ability to retain elements# the study of

trace element concentrations in lichens is fre-

quently useful in demonstrating "fallout" patterns

of particulates and aerosols. For exampleI mercury

2oE 10 0 low
DISTANCE FROM EAST SMELTER (kml
is found in elevated concentrations near a chlor-

alkali works in Finland (Lodenius and LaaJcsovirta

Fig. 2 ~ T h e
distribution

of the lichen-impover-

1979); lead is elevated in lichens adjacent to

highways (Laaksovirta and other 1976# Lawrey and

ished zone (Nash 1972) at Palmerton# PennsylvaniaI

Hale 1979); and a number of elements are present

in relation to the zones of modified forest

(Jordan 1975)# to detectable levels of ambient

in high concentrations in citiesI such as Tel Aviv

(Garty and others 1977) and Sendalp Japan (Saeki

sulfur dioxide (Nash l975lI and to elevated levels

of cadmium and zinc in surface soils (Buchauer

and others l977lI near the nickel complex at

SudburyI Ontario (Tomassini and others 1976)I near 1973).

a coal-fired power plant (Gough and Erdman 1977)

and in coal mine ecosystems of Ohio (Lawrey and

company had installed and subsequently contin-

Rudolph 19751. In none of these cases is toxicity

uously used an acid plant at the time that zinc

of the accumulated elements demonstrated. Toler-

sulfide ores were initially processed. In con-

ance to high concentrations of elements may occur

trast# metal concentrations in the soil duff were

if the elements are relatively insoluble and are

as high as 1351000 ppm zinc and 1750 ppm cadmium

localized extracellularly. For exampleI Noeske

at a site adjacent to the smelting complex

and others (1970) showed that iron and copper

Duchauer 1973). The concentration of zinc and

were encrusted on the surface of lichens which

cadmium decreased exponentially with distance from

grew on metal rich substrates in the Harz Moun-

the smelters until background concentrations were

tains of Germany. FurthermoreI Garty and others

found at 20 to 25 km E and 16 km W of the smelters

(1979) have shown that particulates in Tel-Aviv

are incorporated extracellularly within the

(fig. 21. Physiological studies demonstrated

that zinc and cadmium were toxic to the lichens

lichen thallus.

when concentrations reached 300-500 ppm in the

thallus. Since zinc was present in concentrations

In contrast to the above studies "trace" ele-

ments do occasionally occur in toxic concentra-

100 times that of cadmium and since zinc is

approximately as toxic as cadmiumI zinc was in-

tions. Lawrey and Hale a9791 have shown accumu-

ferred to be the most important pollutant. ¶i

lation of approximately 1000 ppm lead in a lichen

points beyond the li'chen impoverishment zone zinc

growing near an expressway in the vicinity of

concentrations ranged from 20-200 ppm and cadmium

WashingtonI D.C. They demonstrated that.lead

from 1-30 ppmI where both sets of values were

accumulation is correlated with reduction in

demonstra5ly non-toxic.

growth of the speciesI but they have neither

demonstrated experimentally that lead was the

cause of the reduced growth nor demonstrated that

LITERATURE CITED

other factors associated with automobile emissions

were unimportant. Probably the most convincing

Ahmadjian, V.

study documenting trace element toxicity is that

1967. The lichen symbiosis. 152 p, Ginn Co.

of Nash (1975) around a zinc smelting complex in

(Blaisdell ) Boston.
eastern Pennsylvania. In the vicinity of the

smelting complexI lichen species richness was

Arnoldy F.

reduced to 7 in contrast to the 77 species found

1892. Zur 1 ichenflora von Muenchen. Ber.

in a control area (Nash 1972). The zone of

Bayer Bot. Ges. 2: 1-76.

lichen impoverishment extended for a distance of

15 km W and 10 km E of the smelting complex

Baddeley, M. S., B. W. Ferry and E. ,.I.Finegan.
(fig- 2). Although detectable and potentially
1973. Sulphur dioxide and respiration in

toxic levels of sulfur dioxide were found adja-

lichens. In Air pollution and lichens.

cent to the smeltersI the distribution of

B. bl. ~ e r r xII. S. Baddeley and D. L. Hawks-
elevated levels of sulfur dioxide did not extend
worthy eds. p. 299-313. Athlone Presc, Lon-

as far as the lichen impoverishment zone extended-

don.

The relative unirnprtance of sulfur dioxide was

further documented by the lack of acidification

of the bark of oak trees and by the fact that the

 Buchauer* M. Hawksworthy D. L.
1973. Contamination of soi 1 and vegetation 1971. Lichens a s litmus f o r a i r pollution:
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135.
Horntvedty R.
Dainesy R. H. 1975. Epiphytic macro1 ichens on Scots pine
1968. Sulfur dioxide and plant response. related t o a i r pollution from industry i n
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Denisony W. C.
1973. Life i n t a l l t r e e s , S c i , h e r , 228: Jordan, M. J .
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f i r e on a chestnut-oak woodland. Ecology
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1964. Atmospheric pollution of Belfast and
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Nat. J . 14: 237-245. 1977. Epiphytic 1ichen vegetation and e l e -
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Garty* J e YM. Galun* C. Fuchs and N. Zisapel. pine needles examined as indicators of a i r
1977. Heavy metals i n the lichen Caloplaca pollution a t Kokk01a~W. Finland. Ann.
aurantia from urbany suburban and rural Bot. Fennici 14: 112-130.
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Water* Air and Soil Pollut. 8: 171-188. >- -and P. Alakuijala.
1976. Observations on the lead content of
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and M. Kessel .
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lichen and bark adjacent t o a highway in
southern Finland. ~ n v i r o n .POI 1u i . li :
elements accumulated i n the lichen thal lus. 247-255.
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New Phytol 82: 159-168.
Lang* G. E a Y W. A. Reiners and R. K. Heier.
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1965. Lichens as indicators of a i r pollution chemi s t r y by epiphytic 1ichens. Oecologia
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1973. Physiological plant ecology. 252 p.
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1971. The e f f e c t of airborne f l u o r i d e s on
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1977. Influence of a coal - f i red ~ o w e ra~n lt a l s from a c i d i c surface substrates of" coal
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Nash, T. H., I 1 1
1971. Lichen s e n s i t i v i t y t o hydrogen f l u o r -
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l i c h e n communities near a z i n c factory.
The B r y o l o g i s t 75: 315-324.
1973. S e n s i t i v i t y o f lichens t o s u l f u r
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t o r y on 1ichens. Ecol Monogr. 45: 183-198.
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1979. Gross photosynthetic response o f 1i c h -
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1977. Ozone and other photochemical oxidants.
719 p. Washington, D. C.
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1975. Total sulphur content o f an e p i p h y t i c
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1968. Lichens and a i r p o l l u t i o n . Acta Phyto-
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1974. CO,,~Gaswechsel-Untersuchunqen z u r SOo-
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Nieboer and D. H. S. Richardson.
1977. The e f f e c t o f time o f exposure t o
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.
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155.

-9 K. J. Puckett, E. Nieboer, D. H. S. Rich- Acknowledgments : This research was supported

ardson and B. Grace. by t h e U.S. National Science Foundation
1976. Determination of copper, iron, n i c k e l , grants DEB-7610244 and DEB-7921953.
and sulphur by X-ray fluorescence i n l i c h e n s
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Can. J. Bot. 54: 1591-1603.
 Influence of Air Pollution on Population
Dynamics of Forest Insects and on
Tree Mortality1
Donald L. Dahlsten and David L. ~owney2

Abstract: Weakened trees are often predisposed to injury

or death by insects, and in forest ecosystems particularly

by bark beetles. In the San Bernardino National Forest the

interaction between photochemical oxidant weakened ponderosa

pine and the western pine beetle (WPB) was examined in de-

tail. The major results from this study suggest that oxi-

dant damaged trees attacked by WPB produce about the same

total brood with lower initial attacks compared to

healthier trees. This higher productivity trend is most

evident in generation 1 trees Generation 2 trees, both

damaged and healthy, are under much greater moisture

stress and produce much less WPB brood than generation 1

trees regardless of oxidant damage.

The implication of these results is that in stands with

a higher proportion of damaged trees, a given population of

WPB could kill more trees and increase at a greater rate

than in a stand with a lower proportion of damaged trees.

Simulation modelling with these results and other factors

that affect ponderosa pine mortality should provide a

basis for predicting long term effects of air pollution

on the WPB population and pine mortality.

Most of the work on air pollution damage has There have been some studies on the direct

focused on direct injury to the plant and effects of air pollution on insects. Feir (1978)

Kozlowski (1980) gives a good review of the im- studied the effects of air pollutants on insect

pact of air pollution on forest ecosystems. By growth and reproduction and Hillmann and Benton

comparison, little has been done on the indirect (1972) looked at the reactions of honey bees to

effects such as the predisposing of plants to sulfur dioxide.

insects or pathogens. Watt (1969) speculated on

the effects of air pollution on population fluc- Bromenshenk (1976, 1978) has studied the

tuations of insects and Heagle (1973) reviewed effects of coal-fired power plant emissions on a

the interaction between air pollutants and plant variety of insects. In another study, Gilbert

parasites. More recently the occurrence of (1971) looked at the indirect effects of air

diseases and insect pests of trees in air pollution on several bark inhabiting insects.

polluted regions of North America has been Air pollution may be the cause for the scarcity

recorded (Dominik 1978). of all orders of insects in New Jersey (Muller

1971).

The effects of insects in pine stands influ-

'presented at the Symposium on Effects of Air enced by air pollution, particularly xylophagous

Pollutants on Mediterranean and Temperate Forest insects, have been studied (Sierpinski 1972,

Ecosystems, June 22-27, 1980, Riverside,

1977). In California on the San Bernardino

California, U.S.A. Mountains, ponderosa pine shown to have advanced

symptoms of oxidant injury were most frequently

professor of Entomology and Statistician in

infested and killed by western pine beetle

Entomological Sciences, respectively, University

of California, Berkeley, Calif. , and mountain pine
(Dendrocbonias'-brevicabis)
beetle (2. ponderosae) (Stark et al. , 1968). The The samples were taken from the tree using a

purpose
- - of this study was to look a little closer
modified ortable 4.5 kg gas powered drill

at the interaction between the western pine beetle

D l P r e c o n Multiple Control, Ridgewood,

and oxidant affected ponderosa pine, to see what

New Jersey). A standard circular hole saw with

effect diseased trees would have on the dynamics

an 11 can diameter blade was used in the chuck of

of bark beetle populations and to look for dif-

the drill. This saw cuts an area of approximately

ferences between beetle population parameters in

88 cm2.

diseased and healthy ponderosa pine.

At each sample height the circumference was

measured, then the two discs were cut. With the

MATERIALS AND METHODS

egg discs it was desirable to take a portion of

the xylem with each disc to protect the galleries.

Field Procedures
After a disc was removed it was examined by the

climber for insects, which were identified and

Beginning about the first week in July, the

called down to the notetaker.

San Bernardino mountain areas with substantial

ponderosa pine stands were searched for first

During the last larval sample emergence car-

generation western pine beetle attacked trees.

tons were placed on the trees at each sample

These areas are generally on the southern side

height. These cartons were not removed until

of the mountains at about 1500-1800 m elevation,

well after brood emergence.

and are areas where high oxidant air pollution

levels occur. Second generation trees were

Laboratory Procedures

located in mid-August to raid-September, depending

on the timing of the first generation. Trees

Once the samples had been returned to Berkeley,

under attack were detected by pitch tubes or frags

they were placed in cool storage until analyzed.

in bark crevices. Attacked trees were checked

X-rayed samples were placed into rearing imme-

with an axe at the base to see which species were

diately after the radiographs were taken.

attacking and to check the staqe. Trees with

Basically there were four laboratory analysis

mixed brood (2. ponderosae and 2. brevicomis) and procedures for sample discs.

trees with strip attacks (one side only) were

relatively uncommon and were not selected for this

Egg Disc Analysis

study. Trees with spp. in the tops were

selected, but sampling for western pine beetle

To determine egg mortality discs were taken at

was stopped at the base of infestation of the

3.0 meter intervals approximately two to three

z. Each suitable tree located was used in the

study until the required number were found (12

weeks after the mass attack period. Sampling at

every 3.0 m interval gave 4 or more heights for

per generation in 1973 and 1974, 6 per generation

each tree. Egg discs were taken only once, at

in 1975 and 1976). In the time period allowed

the time of the first larval sample. This avoids

for sampling the initial WPB stages there was no

an additional climbing of the sample tree,

practical way to locate all the attacked trees in

assures that maximum egg hatch has occurred and

the area and thenpick a random subsample for the

mortality can still be determined.

study.

If xylem tissues still remained on the disc it

Sampling
was carefully removed so that the galleries in the

phloem tissue were not disturbed. Gallery length

Prior to sampling, the major tree character-

was measured with a metric map reader and all

istics were recorded: location, height, DBH,

attacks were recorded. Each disc was then exam-

height of first green branch, and estimated tree

ined with a dissection scope (10-20x) along the

damage due to air pollution. The air pollution

length of all the adult galleries. Each egg

damage was rated by rating upper and lower crown

niche was evaluated asempty or as containing a

needle retention, needle condition, needle length,

viable egg or nonviable egg in addition to re-

and branch mortality. Needle condition and re-

cording larval eclosion. The difference between

tention were difficult to rate on the sample

the total number of egg niches recorded and the

trees because in many cases, fading and needle

total number of larvae hatched was calculated as

loss due to the bark beetle attacks had begun.

egg mortality.

For 1973 only one needle length rating was made

for the entire tree, so the needle length rating

X-ray Analysis

for the other years was combined to be consistent

in the analysis ( in nearly all cases the upper The first samples to be x-rayed were taken con-

and lower crown needle length was the same).

currently with the egg discs (egg discs are des-

troyed during analysis so they can not be placed

Samples for all procedures except where noted

into rearing). For generation one, an additional

later were taken at 1.5 meter intervals over the

x-ray sample was taken late in the brood develop-

length of the western pine beetle infestation.

ment period, usually in mid to late July. For the

Two 88 cm2"discs taken on opposite sides of the

second generation, x-ray samples were removed on

tree at 1.5 meter intervals gave the desired

two later occasions: once in mid-September to

level of precision.
October to check for early emergence of brood and

adult re-emergence, and finally in the following

oblique to the surface of the bark and often have

spring (March to April) after brood development

pitch as a remnant of a pitch tube.

was nearly complete, but before spring emergence.

Field Emergence Cartons

The discs to be x-rayed were tkken from cold

storage as soon as possible and removed from the

As an additional check on emergence, a pro-

plastic bags. The average bark thickness of each

cedure for rearing in situ on the sample trees

disc was measured with a metric caliper and the

was also used.

percentage woodpeckering on each disc was re-

corded. The two 88 cm2 discs from each height

One quart squat ice cream cartons that cover

were placed on one sheet of 8" x 10" Industrial

88 cm2 of bark were painted silver on the outside

~odak@AA-2 film and exposed with a picker@

and prepared with a screened ventilation hole.

radiograph machine. The samples were then re-

Stikem specialawas placed on the inside of the

bagged and transported to a large storage shed f a

cartons to discourage insects from boring out or

rearing (see below).

from going back into the bark. A groove was cut

in the bark with the sampling saw into which the

X-rays were interpreted for western pine

lip of the carton was placed; a nail was driven

beetle, live and dead parent adults, brood

through the bottom of the carton and into the

adults, pupae, and larvae, parasitoids, predators

tree to increase stability.

and miscellaneous. The miscellaneous category

included buprestid and cerambycid larvae,

The cartons were placed on the tree at the

weevils, *,and unknowns, of which there were time the last x-ray or brood sample was taken (to

many judging from the rearing of these discs. A

coincide with the pupal and/or callow adult stages

light table was used for interpretation and all

of the brood) and were left i n place for up to

inclusions were marked with a grease pencil by

three months, until well after beetle emergence

the interpreter, counted and then erased. A sec-

and the collection of all associated insects had

ond interpreter then repeated the process,

taken place. The second generation cartons were

thereby reducing error and bias. The x-rays from

nofc placed on the tree until early spring since

each tree were then stored in envelopes as a per-

cartons do not weather well and therefore cannot

manent record.
be left ort the trees for four to six months over

winter. The cartons were split open and examined

Laboratory Rearing
under the microscope and then discarded. The same

insects that-wererecorded for laboratory rearing

Each x-rayed disc was placed in a 1/2 gallon

were recorded for the sticky emergence carton.

ice cream carton with a small 2 or 3 dram glass

vial pushed through the lid. Since the insects

in the discs are positively phototropic at least

RESULTS AND DISCUSSION

in the adult stage, and since very few beetles or

other insects bore through the sides of the car-

Attacked tree characteristics and variation by

tons, this was a suitable rearing procedure. The

year and generation:

cartons were stored in racks in a large unheated

warehouse where Berkeley's mild, cool climate

The tree heights of the 71 attacked trees

provided an ideal rearing environment.

sampled over the four year period varied from 13

to 44 meters, but most trees were in the 20-30

The emerging insects were collected from the

meter range (Fig. 1). Tree DBH, stem volume, and

vials three to five times per week. Most of the

infested bark area varied more widely, but

specimens could be identified with a lox hand

exhibited no significant trends with regard to

lens and then recorded on a data sheet by collec-

year and generation. Western pine beetle attack

tion date. Only the parasitoids and the western

and emergence density, however, both showed sig-

pine beetle were sexed. All specimens have been

nificant differences (P < -05) by generation.
identified by taxonomic experts and specimens
Year differences were not significant for these

were sent for verification regularly; the col-

variables except for a lower attack density for

lection is updated as necessary. Approximately

generation one trees of 1976 compared to the gen-

130 different insects were recorded. In this

eration one trees of the other three years.

paper, only WPB, 4 predators (Enoclerus lecontei,

Temnochila chlorodia, Aulonium longum, Medetera

Effect of tree oxidant damage on beetle popu-

aldrichii) and 4 parasites (Roptrocerus

-
&
J
X lation :
phagorum, Dinotiscus burkei, Eurytoma conica,

Coeloides sp. nr. brunneri) are considered.

In order to determine if damage to the sample

trees had an effect on the western pine beetle

Sample discs were kept in rearing for nine

populations, the individual components of the

months and at the end of this period the cartons

damage score were analyzed with respect to beetle

were examined for any remaining insects; the

attack density. Each individual component of the

discs were then discarded. The gallery length

score can be regarded as an interval level vari-

and number of attacks were recorded for each disc

able, so multiple regression techniques were used

prior to discarding. Attack holes can be dis-

to determine their relationship to attack density.

tinguished from emergence holea as they are

The technique used was a full screen analysis

 Fig. 1.

Western pine beetle attacked tree characteristics,

attack density and emergence density by generation

and year for 71 ponderosa pines in the San Bernardino

National Forest, California. (dashed lines = mean +

one standard deviation)

Most other generation one beetle variables

were not significantly different between short aid

long NL trees, except for eggs per attack and

emerged brood per attack (Table 1). This prob-

ably indicates that the attacking beetles for

generation one were more productive when the

trees had lower attack density and less compe-

tition within the trees. The short NL generation

1 trees, with lower attack densities, had essen-

tially equal numbers of eggs and emerging brood

compared to the long NL trees.

/,*
The trend toward high productivity in the

lower attack density, short NL trees for

, , L :
-
, generation 1 was not evident for generation 2.

The beetles in the short NL trees had essentially

1 2 1 2 1 2 1 2
74 75 74 75 the same eggs/attack and emerged brood/attack as

GENERATION GENERATION
those in long NL trees (Table 1). Total eggs,

hatched eggs, and late larval densities were all

significantly lower in the lower attack density,

short NL trees. Mortality proportions for each

life stage sampled seemed relatively independent

which calculated p2 values for all possible

of NL although they were higher in generation 2

linear combinations of the score components,

compared to generation 1.

year, and beetle generation for the attack den-

sity dependent variable. The results of this

Table I--Western pine beetle variable means by gen-

analysis indicated that only the needle length

eration and tree a: pollution injury(needle lerytH

(NL) component (upper and lower crown combined)

Generation 1 Generation 2
of the score was significant (P < -05) along with Short Long Short Long
beetle generation. The effect of any other com-
Injury: NL ML NL NL
ponents was insignificant when added to the equa-

tion containing NL and generation. Some of the

No. of trees
other components which have been significant in
~ttacks/~~~
other studies - needle retention and needle con- Gallery length/^^^
dition - probably were unreliable for these trees Total eggs/~M2
as they were examined in a partly faded condition
Eggs/attack
some weeks after being mass attacked by bark
Hatched eggs/Dll
beetles. In subsequent analyses the needle
(1st instar
length (NL) was used to distinguish between trees
larvae/~~2 )
highly affected by air pollution (NL=O) and less
Egg mortality
2
affected trees (NL=l). Table 1 gives mean
Late larvae/DM
values of the attack density and other western
Larval mortality--
pine beetle variables for each generation and
early to late
needle length category. In our sample of trees
larvae (LM)
significantly fewer short NL trees were found in
Emerged WPB
generation one (7 of 36) compared to generation
/Dd
(REAR)
two (16 of 35), but due to the practical limita-
Mortality-late
tions of our sample selection procedure, it is
larvae to emer-
uncertain whether this difference holds true for
gence (REAR) (BPI)
the entire population of attacked trees;
Mortality-eggs to
emergence (REAR)
The attack density was significantly higher
Emerged brood
for long NL trees compared to short NL trees for
per attack

both generations, while generation two trees had

lower attack densities than generation one re-

*significant differences between NL

gardless of ML. classes, P < .05.

 Needle length was also used to compare differ-
The consequences of bark beetle activity in

ences in predator and parasite densities for each

an area are summarized in Table 3 and show that

generation (Table 2). Predators included the

attack rates are lower on oxidant affected trees,

4 common species: Enoclerus lecontei, Temnochila

but the output of brood is essentially the sane

chlorodia, Au1oniw.1 longum, and Medetera

in healthy and diseased trees. Bark beetles,

aldrichii. Parasites included Roptrocerus e-

assuming no direct deleterious effects of air

-
phagorum, Dinotiscus burkei, Eurytoma conica and

Coeloides sp. nr. brunneri. For generation 1,

pollution,should increase in areas with high

ozone damage and tree mortality will also in-

only emerged (REAR) predators/dm2 were signifi-

crease in these areas. The possible interactions

cantly higher for the long NL trees. For genera-

of air pollution, ponderosa pine, and western

tion 2, initial parasitized larvae/dm2 were higher

pine beetle and the effects on forest succession

for short NL trees, but late parasitized larvae/

are shown in Table 4.

dm2 were higher for long NL trees.

A much larger increase

in numbers of

parasitized larvae occurred for the gen. 2, long

Table 4~Interactiontable: air pollution,

NL trees. Increased woodpeckering, along with a

ponderosa pine, and western pine beetle.

higher final larval density for the long NL trees

Causes a change in

may explain the increased numbers of parasitized

larvae. A similar proportion (about 1%) of

VPB 1 Ponderosa Forest
initial larvae were parasitized in both cases.
pop. pine mortal- succes-

level ity caused sion

by WPB

An increase In:

Table 2--WPB predator and parasite means by gen-

air pollution + + +
eration and tree air pollution injury class
soil water ? ? ?
(needle length). foliar injury ++ + +
Generation 1 Generation 2 soil nutrients ? ? ?
Short Long Short Long mature tree growth ? ? ?
Injury: NL NL NL NL 2 root disease + + +
forest succession - - 0
No. trees 7 29 16 19 WPB pop. level 0 ++ +
Initial pred./~~2 0.962 1.910 0.53 0.30 pine mort. caused

Initial para- 0.20 0.36 0.18* 0.08 by WPB ? 0 +

sitized lar./DM
Late predators/~~21.83 2.75 0.80 0.84 Key:

Late parasitized 0.56 0.61 0.26* 0.48 0 = unrelated or not possible

larvae/~ll^ + = moderate increase
Emerged (REAR) 0.73* 1.50 0.25 0.27 ++ = large increase
predators/~~^ - = a large decrease
.
Eraer (REAR)para/D~2 0.48 0.42 0.22 0.28 ? = unknown, needs more investigation
% area samples wood-0.1% 0.8% 7.0% 11.0%
peckered
*Significant difference between means of smog

injury classes, P < .05 . LITERATURE CITED

Table 3--Ozone effects table: western pine

Kozlowski, T. T.

beetle in ponderosa pine.

1980. Impacts of air pollution on forest eco-

systems. BioScience 30:88-93.

Watt, K.E.F.

tree

mort. 1969. Prospective effects of air pollution on

insects. Can. Entomol. 101:1235-1238.

low long 4.0 3.9 consbat constant

Heagle, A. S.

or de- or de-
1973. Interactions between air pollutants and

creasing creasing
plant parasites. Annu. Rev. Phytopathol. 11:

365-388.

high short 2.4 3.7 increaskq increasing

Motes:
Dominik, J.

(1) Effect depends on individual tree character-

1978. Investigations of the occurrence of

istics.
diseases and insect pests of North American

(2) Based on both generations combined and multi-

trees in the regions of different degrees of

plying female adult attacks by two to

air pollution caused by the industry. Final

account for males.

Report of Investigations covering the period

(3) Based on both generations combined.

of August 1, 1973-June 30, 1978. Warsaw:

(4) Assuming constant no. of damaged pines/stand.

Sin., 1978. 91, 21 leaves: 111.

Feir, D.
Bromenshenk, J. J.

1978. Effects of air pollutants on insect

1978. Investigation of the impact of coal-fired

growth and reproduction. Physiologist 21:36.

power plant emissions upon insects: entomo-

logical studies at the zonal air pollution

Hillman, R. C., and A. W. Benton. systems. Ecol. Res. Ser. 600/3-78-021:473-507.

1972. Biological effects of air pollution on

insects emphasizing the reactions of the honey

Sierpinski, 2.
bee, e m e l l i f e r a to sulfur dioxide. J. 1972. The economic importance of secondary

Elisha Mitchell Sci. SOC. 88:195.

noxious insects of pine on territories with

chronic influence of industrial air pollution.

Muller, J.
Mitt. Forstl. Bundes-Versuchsanst Wein. 97:

1971. Is air pollution responsible for

609-615.

melanism in Lepidoptera and for scarcity of

Sierpinski, Z.

all orders of insects in New Jersey, U.S.A.

1977. Economic significance of noxious insects

J. Res. Lepid. 10:189-190.

in pine stands under the permanent impact of

the industrial air pollution. Slyvan. .64:59-71.

Gilbert, 0. L.

stark, R.W., P.R. Miller, F.W. Cobb, Jr., D.L.

1971. Some indirect effects of air pollution on

Wood, and J.R. Parmeter, Jr. 1968. Photochem-

bark living invertebrates. J. Appl. Ecol. 8:

ical oxidant injury and bark beetle (Coleop-

77-84.

tera: Scolytidae) infestation of ponderosa

pine. I. Incidence of bark beetle infestation

Bro~~ienshenk,

J. J.
in injured trees. Hilgardia 39:121-126.

1976. Biological impact of air pollution on

insects. Ecol. Res. Ser. 600/3-76-013:

295-312.

 Air Pollutants and Their Effects on

Wildlife with Particular Reference to
the House Wren (Delichon urbica)'

James R. Newman, P ~ . D . ~

A b s t r a c t : I n j u r y and death t o w i l d l i f e from a i r

emissions have been recorded s i n c e before the
t u r n of the c e n t u r y . A i r p o l l u t i o n can d i r e c t l y
a f f e c t w i l d l i f e ( i . e . , cause d e a t h ) , or can
i n d i r e c t l y a f f e c t w i l d l i f e ( i . e . , cause h a b i t a t
l o s s ) . A r e c e n t i n v e s t i g a t i o n on t h e chronic and
s u b l e t h a l e f f e c t s of a i r p o l l u t i o n on t h e house
m a r t i n (Delichon u r b i c a ) h a s shown t h a t a i r
v
emissions can s i g n i f i c a n t l y a f f e c t t h e n e s t i n g
ecology of t h i s s p e c i e s .

INTRODUCTION emissions ( P r e l l 1936; Hais and Masek 1969).

Eighty-five percent o f t h e w i l d l i f e i n c i d e n t s
S i n c e t h e end o f t h e 19th c e n t u r y , over h a v e been recorded i n the l a s t 25 y e a r s . The
100 a i r p o l l u t i o n episodes have been r e p o r t e d i n c r e a s e i n the number o f w i l d l i f e i n c i d e n t s
i n v o l v i n g i n j u r y and death t o animals (Newnan a p p e a r s t o be due n o t o n l y t o an i n c r e a s e i n a i r
1975, 1979). Although t h e m a j o r i t y o f t h e emissions but t o a b e t t e r understanding o f a i r
i n c i d e n t s involved domesticated animals, a number p o l l u t i o n e f f e c t s , broader communications, and a
o f i n c i d e n t s have been r e p o r t e d f o r w i l d l i f e g r e a t e r i n t e r e s t i n r e p o r t i n g such problems.
s i n c e t h e 1880's ( t a b l e 1 ) . O v e r a l l , t h e few
r e p o r t s i n v o l v i n g w i l d l i f e appear t o be more a Table I-The number of reported a i r p o l l u t i o n
f u n c t i o n o f economic b i a s t o r e p o r t i n j u r y and i n c i d e n t s involving w i l d l i f e and af f e c t e d groups
d e a t h o f domestic animals than some i n n a t e (adapted Newman 1979)
r e s i s t a n c e o f w i l d l i f e t o a i r p o l l u t i o n . For
example, i n t h e e a r l y 1900's a r s e n i c emissions
from a smelter caused t h e i n j u r y and death o f Number of I
s e v e r a l thousand c a t t l e and h o r s e s i n r u r a l W i l d l i f e Groups consumer Groups
Montana arkin ins and Swain 1908). No mention was Period Inci 1Game \ one game IPrim. [secondary
made o f i n j u r y o r d e a t h t o deer o r o t h e r w i l d l i f e dents 1 1
p l e n t i f u l i n t h e a r e a , a 1 though deer and r a b b i t s
a r e known t o b e v e r y s e n s i t i v e t o a r s e n i c Before 1 1 1
1900

1900-1950
p r e s e n t e d a t t h e Symposium on E f f e c t s of A i r
1951-1970
P o l l u t a n t s on Mediterranean and Temperate F o r e s t
Ecosystems, J u n e 22-27, 1980, R i v e r s i d e ,
1971 t o
C a l i f o r n i a , U.S.A.
Present
^sen2ior E c o l o g i s t , P r o j e c t Operations,
Environmental Science and Engineering, Inc
P.O. Box ESE, G a i n e s v i l l e , F l o r i d a 32602.
., TOTAL
Recent Federal l e g i s l a t i o n , namely t h e 1977 Clean
A i r Act Amendments, now r e q u i r e formal
c o n s i d e r a t i o n o f t h e e f f e c t s o f a i r emissions on
w i l d l i f e and o t h e r components o f e c o l o g i c a l
systems.
The purpose o f t h i s paper i s t o summarize some
o f t h e known e f f e c t s o f a i r emissions on w i l d l i f e
and p r e s e n t p r e l i m i n a r y f i n d i n g s on t h e
e c o l o g i c a l e f f e c t s o f a i r emissions on t h e house
m a r t i n ( ~ e l i c h o nu r b i c a ) , an i n s e c t i v o r o u s b i r d .
For t h i s p a p e r , d i s c u s s i o n i s l i m i t e d t o
t e r r e s t r i a l wild1 i f e and v e r t e b r a t e consumers . i r r i t a n t s . Investigators i n ~ z e c h o s l o v a k i a ~
Previous s p e a k e r s h e r e d i s c u s s e d t h e e f f e c t s on
i n v e r t e b r a t e consumers.
GENERAL EFFECTS OF A I R EMISSIONS ON WILDLIFE
One of t h e e a r l i e s t w i l d l i f e i n c i d e n t s (1887)
i n v o l v e s t h e d e a t h of f a l l o w d e e r (Dama dama)
from a r s e n i c emissions from a s i l v e r foundry i n
Germany (Tendron, 1964). As e a r l y a s 1900 t h e
Royal S o c i e t y of London was charged t o i n v e s t i -
g a t e t h e phenomenon of i n d u s t r i a l melanism. T h i s
i n c i d e n t i s recognized more f o r i t s demonstration
o f " n a t u r a l " s e l e c t i o n than a s an environmental
problem. Recently, g e n e t i c changes i n small
mammals because of a i r p o l l u t i o n have been
r e p o r t e d (Newman 1980).
One of t h e ' e a r l i e s t d e t a i l e d d e s c r i p t i o n s of
t h e harmful e f f e c t s of a i r emissions on w i l d l i f e
i s from Germany. Arsenic emissions were the cause
o f widespread d e a t h of game animals i n t h e
Tharandt f o r e s t of Germany i n 1936. S i x t y t o
s e v e n t y p e r c e n t of t h e red deer (Cervus e i e p h u s ) ,
r o e d e e r (Capreolus c a p r e o l u s ) , and wild r a b b i t s
( ~ r ~ c t o l a g uc su n i c u l u s ) d i e d . The d e e r e x h i b i t e d
d e f e c t i v e h a i r growth and a n t l e r formation,
c i r r h o s i s of t h e l i v e r and s p l e e n , and emaciation
( P r e l l 1936).
Adverse e f f e c t s of a i r emissions on w i l d l i f e
have been r e p o r t e d from North America, Euro-pe,
and A s i a , and have included game and nongame
animals; b i r d s and mammals; primary and secondary
consumers; and h e r b i v o r e s , omnivores, and
c a r n i v o r e s ( t a b l e 1 ) . The e f f e c t s on w i l d l i f e
have included d i e - o f f s and o t h e r population
reductions, physiological abnormalities, disease,
p h y s i c a l i n j u r y , and bioaccumulation. The
m a j o r i t y of w i l d l i f e i n c i d e n t s r e p o r t e d involve
d e e r and small b i r d s (~ewman 1979). These groups
a r e not n e c e s s a r i l y more s e n s i t i v e but a r e more
l i k e l y t o be monitored, e s p e c i a l l y d e e r , which
i s a widely managed game s p e c i e s . The s t a t u s of
s m a l l b i r d s i s n o t o n l y of s c i e n t i f i c concern,
b u t a l s o of more widespread p u b l i c i n t e r e s t .
Pathways of contamination by a i r emissions f o r
w i l d l i f e a r e : i n h a l a t i o n , a d s o r p t i o n , and
i n g e s t i o n . Wellings (1970) r e p o r t e d t h e
o c c u r r e n c e of pulmonary a n t h r a c o s i s i n urban
sparrows ( P a s s e r domesticus) compared t o r u r a l
p o p u l a t i o n s i n C a l i f o r n i a . One of t h e most
s t a r t l i n g examples of contamination by i n h a l a t i o n
involved t h e d i e - o f f of 200 t o 500 s o n g b i r d s near
a B r i t i s h Columbia pulp m i l l which emitted high
c o n c e n t r a t i o n s of H2S and o t h e r p o l l u t a n t s .
The dead b i r d s showed i n t e r n a l hemorrhaging i n
t h e l u n g s and l i v e r . 3
Adsorption of a i r emissions i n v o l v e s t h e
adhesion of g a s e s o r p a r t i c u l a t e s t o t h e e x t e r n a l
s u r f a c e s o r e x t e r n a l membranes, e . g . , cornea of
eyes. Light (1973) r e p o r t s a high i n c i d e n c e of
b i i n d n e s s i n bighorn sheep (*canadensis)
found i n mountain a r e a s of C a l i f o r n i a with heavy
o x i d a n t l e v e l s . Oxidants a r e known eye
have observed e a r l y aging of t h e cornea i n h a r e s
( ~ e p u seuropaeus) found i n a r e a s with heavy
SO2 and p a r t i c u l a t e d e p o s i t i o n from power
p l a n t s and o t h e r i n d u s t r i e s .
I n g e s t i o n i s t h e most commonly r e p o r t e d mode
o f contamination f o r w i l d l i f e . I n j u r y and d e a t h
t o w i l d l i f e from i n g e s t i o n o f contaminated food
and water have been r e p o r t e d f o r numerous animal
s p e c i e s i n c l u d i n g r a b b i t s and deer from a r s e n i c
emissions ( P r e l l 19361, d e e r from f l u o r i d e
emissions (Karstad 1967; Robinette and o t h e r s
1956; Newman and Yu 19761, and sparrows from
cadmium emissions (Nishino and o t h e r s 1973).
There a r e many r e p o r t s of t h e bioaccumulation of
a i r p o l l u t a n t s i n w i l d l i f e . With a few
e x c e p t i o n s , t h e e f f e c t s of t h i s bioaccumulation
a r e n o t known (Newman 1980).
A i r emissions may not be l e t h a l t o w i l d l i f e
b u t may lower t h e h e a l t h and r e s i s t a n c e of
w i l d l i f e s o t h a t n a t u r a l s t r e s s such a s cold and
s h o r t a g e of food may r e s u l t i n f u r t h e r i n j u r y o r
d e a t h . S t u d i e s on t h e h e a l t h of wild h a r e s
(Lepus europaeus) l i v i n g i n a r e a s of h i g h a i r
p o l l u t i o n i n Czechoslovakia showed p h y s i o l o g i c a l
r e s p o n s e s s i m i l a r t o animals with i n f e c t i o n s o r
a l l e r g i c r e a c t i o n s . There were a l s o changes i n
t h e normal age s t r u c t u r e of t h e h a r e p o p u l a t i o n s
( ~ o v a k o v a 1969). The g e n e r a l h e a l t h of d e e r
p o p u l a t i o n s a f f e c t e d by f l u o r i d e emissions i s
poor ( ~ o b i n e t t eand o t h e r s 1957; Karstad 1967;
Newman and Yu 1976; Newman and Murphy 1979). The
s u r v i v a l of t h e s e d e e r d u r i n g times of n a t u r a l
s t r e s s i s q u e s t i o n a b l e . Hais and Masek (19691,
r e p o r t t h a t a r s e n i c contamination of red and r o e
d e e r caused emaciation and l o s s of h a i r so t h a t
many of t h e animals f r o z e t o d e a t h d u r i n g t h e
winter.
I n j u r y o r d e a t h t o v e g e t a t i o n caused by a i r
emissions can have s i g n i f i c a n t i n d i r e c t e f f e c t s
on w i l d l i f e . The l o s s of food r e s o u r c e s and
h a b i t a t can occur a s t h e r e s u l t of i n j u r y o r
3 ~ n p u b l i s h e dr e p o r t , R.D. Harris 1971.
B i r d s Collected (Die Off) a t P r i n c e Rupert, B.C.
Canadian W i l d l i f e S e r v i c e , P r i n c e Rupert, B r i t i s h
Columbia.
4 ~ e r s o n a lcommunication from E l i s k a
Novakova, I n s t i t u t e o f Landscape Ecology, Prague,
Czechoslovakia.
 d e a t h t o v e g e t a t i o n which provides c o v e r ,
r e p r o d u c t i v e h a b i t a t , o r food f o r w i l d l i f e . Many
examples e x i s t o f e x t e n s i v e damage t o ecosystems
from a i r e m i s s i o n s . Large a r e a s o f t h e mixed
c o n i f e r o u s f o r e s t o f s o u t h e r n C a l i f o r n i a (over
40,000 h e c t a r e s ) have been k i l l e d o r i n j u r e d from
photochemical o x i d a n t s ( ~ a y l o r1973). No o v e r a l l
assessment h a s been made a s t o t h e l o s s o f
w i l d l i f e whose f o r e s t h a b i t a t h a s been d e s t r o y e d .
S i m i l a r l a r g e w i l d l i f e h a b i t a t l o s s e s have
o c c u r r e d i n Montana, Tennessee, B r i t i s h Columbia,
and O n t a r i o , Canada (~ewman 1980).
ECOLOGICAL EFFECTS OF AIR EM IS S IONS
ON THE HOUSE MARTIN
With few e x c e p t i o n s , t h e r e i s l i t t l e
i n f o r m a t i o n a v a i l a b l e on t h e e f f e c t s o f c h r o n i c
exposure o f w i l d l i f e t o a i r emissions (Newman
1980). Since 1976, RNDr. E l i s k a Novakova, CSC,
from t h e I n s t i t u t e o f Landscape Ecology, Prague,
and myself have been s t u d y i n g t h e e c o l o g i c a l
r e s p o n s e s o f t h e house m a r t i n (Delichon u r b i c a )
t o chronic a i r pollution. In t h i s p o r t i o n of t h e
t a l k I would l i k e t o p r e s e n t some o f our
preliminary findings.
The house m a r t i n b e l o n g s t o t h e family
H i r u n d i n i d a e . It i s an i n s e c t i v o r o u s b i r d lAiich
f e e d s p r i m a r i l y on a p h i d s and d i p t e r e a n s . A
m i g r a t o r y s p e c i e s , t h e m a r t i n summers throughout
Europe and w i n t e r s i n A f r i c a . As a c o l o n i a l
n e s t i n g s p e c i e s , i t b u i l d s mud n e s t s on t h e w a l l s
o f b u i l d i n g s . The n e s t i n g l o c a t i o n s a r e used
e a c h y e a r , o f t e n by t h e same b i r d s .
Previous s t u d i e s have i n d i c a t e d t h a t 2. u r b i c a
i s s e n s i t i v e t o a i r p o l l u t i o n . Fer iancova-
Masarova and Kal ivodova ( 1965) observed changes
i n t h e s p e c i e s d i v e r s i t y o f b i r d s , i n c l u d i n g the
house m a r t i n , i n t h e a r e a o f an aluminum p l a n t i n
Czechoslovakia. Cramp and Gooders (1967)
observed a c o r r e l a t i o n with i n c r e a s e d n e s t i n g of
--
D . u r b i c a and a d e c r e a s e i n smoke p o l l u t i o n i n
London. A follow-up s t u d y (~ewman 1977) i n t h e
v i c i n i t y o f t h e same aluminum p l a n t s t u d i e d by
Feriancova-Masarova and Kalivodova i n 1965 showed
t h a t 2. u r b i c a decreased i t s n e s t i n g d e n s i t y with
increased a i r emissions.
I n 1978 d i r e c t c o u n t s o f a c t i v e n e s t s were
made i n 141 v i l l a g e s and towns l o c a t e d i n
i n d u s t r i a l i z e d (contaminated) and
n o n - i n d u s t r i a l i z e d ( c o n t r o l or. background) a r e a s .
Of t h e 141 n e s t i n g l o c a l i t i e s censused,
101 v i l l a g e s and towns were i n f l u e n c e d by a i r
e m i s s i o n s from v a r i o u s i n d u s t r i a l s o u r c e s .
Power p l a n t s and a s s o c i a t e d s u r f a c e mines,
chemical p l a n t s , l o c a l h e a t i n g p l a n t s ( c o a l
b u r n i n g ) , cement p l a n t s , and g e n e r a l urban
e m i s s i o n s s o u r c e s were t h e major a i r emission
i n f l u e n c e s . Censusing was conducted a t pre-
determined upwind and downwind l o c a t i o n s from an
e m i s s i o n s o u r c e . This census r e p r e s e n t e d
o b s e r v a t i o n s o f over 20,000 b u i l d i n g s . Records
were kept o f t h e number o f a c t i v e n e s t s . A
s t a n d a r d n e s t i n g d e n s i t y measure was d e r i v e d a s
w e l l a s measures o f t h e occupancy o f a n e s t i n g
l o c a t i o n and colony s i z e . The environmental
a t t r i b u t e s o f e a c h n e s t i n g l o c a l i t y was
c h a r a c t e r i z e d . Comparisons o f background n e s t i n g
l o c a l i t i e s were then made t o e c o l o g i c a l l y
s i m i l a r , b u t contaminated, n e s t i n g l o c a l i t i e s .
The mean n e s t i n g d e n s i t y o f 2. u r b i c a from
c o n t r o l a r e a s was 0.56521.94 n e s t s per s i d e . The
r a n g e i n colony s i z e was 1 t o 33 a c t i v e n e s t s per
occupied s i d e , w i t h an average c o l o n y s i z e o f
2.721.1 a c t i v e n e s t s per occupied s i d e . The
a v e r a g e occupancy ( t h e number o f occupied s i d e s
per s u i t a b l e s i d e ) f o r t h e 32 background n e s t i n g
a r e a s was 2 1 . 0 3 . 1 p e r c e n t .
The o v e r a l l n e s t i n g d e n s i t y f o r 1.u r b i c a from
contaminated a r e a s was 0 . 3 9 3 3 . 7 3 4 a c t i v e n e s t s
per s u i t a b l e n e s t i n g s i d e . This n e s t i n g d e n s i t y
( 3 0 p e r c e n t ) was s i g n i f i c a n t l y lower
= 3.674, df = 8968) when compared t o
^0.002
background n e s t i n g s i t e s . Occupancy was o n l y
1 3 . 6 p e r c e n t . Attempts a t n e s t i n g were a l s o
l o w e r , w i t h o n l y 0.74 a t t e m p t s per contaminated
v i l l a g e compared t o 1.12 a t t e m p t s per background
v i l l a g e . Colony s i z e was s i g n i f i c a n t l y lower
-
(2.1+1.5 a c t i v e n e s t s per occupied s i d e ) .
The house m a r t i n responds a l s o t o t h e l e v e l of
a i r emissions. The o v e r a l l n e s t i n g d e n s i t y f o r
downwind l o c a t i o n s was 0.23720.378 (n = 4891)
n e s t s per s i d e compared t o 0.41521.603 (n = 1969)
n e s t s per s i d e f o r upwind l o c a l i t i e s . This
d i f f e r e n c e was s i g n i f i c a n t (P < 0.001; t = 7.263;
d f = 6808). The o v e r a l l upwind n e s t i n g d e n s i t y
was not s i g n i f i c a n t l y d i f f e r e n t from background
areas.
Colony s i z e and occupancy a l s o changed with
p r o x i m i t y t o emission s o u r c e s . Upwind colony
s i z e s averaged 2.521.0 n e s t s per occupied s i d e
compared t o 1.820.8 n e s t s per occupied s i d e f o r
downwind l o c a t i o n s . These d i f f e r e n c e s were
s i g n i f i c a n t (P < 0.001; t = 3.492 d f = 99).
Upwind occupancy (18.828.9 p e r c e n t ) was a l s o
s i g n i f i c a n t l y h i g h e r (P < 0.001, t = 5.70,
d f = 99) when compared t o downwind occupancy
- percent).
(9.1+7.7
The n e s t i n g d e n s i t y f o r downwind l o c a t i o n s
decreases s i g n i f i c a n t l y a s the distance to the
emission s o u r c e d e c r e a s e s ( t a b l e 2 ) . For upwind
locations, the c l o s e r t o the source, the higher
the nesting density; f o r increased distances,
n e s t i n g d e n s i t i e s d e c l i n e d . The upwind n e s t i n g
d e n s i t i e s c l o s e s t t o emission sources were n o t
s i g n i f i c a n t l y d i f f e r e n t from background n e s t i n g
d e n s i t i e s . Colony s i z e was lower a t a l l downwind
d i s t a n c e s , b u t o n l y s i g n i f i c a n t l y lower a t t h e
two c l o s e s t d i s t a n c e s . Occupancy was s i g n i f i -
c a n t l y lower f o r a l l d i s t a n c e s censused downwind.
For t h e two c l o s e s t upwind d i s t a n c e groups ( o u t
t o 6 tan) , b o t h colony s i z e and occupancy were n o t
s i g n i f i c a n t l y d i f f e r e n t from background
conditions .
Table 2--Comparison of n e s t i n g d e n s i t y of
- -u r b i c a a t v a r i o u s d i s t a n c e s upwind and
D.
downwind from emission sources
Sample
Size Me an Percent
( , N ~ . of Nesting D i f f e r e n c e From
Locat i o n Density Background
Sides )
Downwind
I o c c u r . For h i g h s t a c k s o u r c e s , g r e a t e s t e f f e c t s
Upwind
I a f f e c t e d by a i r emissions t h a n w i l d l i f e popula-
S i g n i f i c a n t , P < 0.025.
^ o t S i g n i f i c a n t , P<O.O5
The a i r d i s p e r s i o n p a t t e r n s a s s o c i a t e d with
t h e s t a c k h e i g h t s i n f l u e n c e d the observed
e f f e c t s . For s o u r c e s w i t h h i g h s t a c k s ( g r e a t e r t h a n
100 m e t e r s ) t h e adverse e f f e c t s on n e s t i n g were
observed i n b o t h downwind and upwind l o c a l i t i e s .
For downwind l o c a t i o n s , t h e lowest n e s t i n g
d e n s i t y d i d n o t occur c l o s e s t t o t h e emission
s o u r c e , b u t a t an i n t e r m e d i a t e d i s t a n c e ( 3 t o
6 km). Nesting d e n s i t y was s i g n i f i c a n t l y .
d e p r e s s e d out t o 12 km. For low s t a c k s o u r c e s
( l e s s t h a n 100 m e t e r s ) , t h e lowest n e s t i n g
d e n s i t y , c o l o n y s i z e , and occupancy occurred a t
t h e c l o s e s t d i s t a n c e s ( 0 t o 3 km). Average
n e s t i n g d e n s i t i e s were 20 p e r c e n t o f background.
In c o n t r a s t t o h i g h s t a c k s o u r c e s , t h e n e s t i n g
d e n s i t y i n f l u e n c e d by low s t a c k s o u r c e s was n e a r
normal a t t h e 9 . 1 t o 12 km d i s t a n c e c a t e g o r i e s .
For upwind l o c a l i t i e s , e m i s s i o n s from h i g h s t a c k
s o u r c e s have an adverse i n f l u e n c e , e s p e c i a l l y
c l o s e t o t h e s o u r c e . For low s t a c k l o c a l i t i e s ,
upwind n e s t i n g d e n s i t i e s were e i t h e r a t o r above
background l e v e l s a t a l l d i s t a n c e s censused.
When t h e house m a r t i n n e s t s i n optimum n e s t i n g
c o n d i t i o n s , such a s apartment b u i l d i n g s and
f e e d l o t s i n proximity t o water, the e f f e c t s o f
a i r e m i s s i o n s were m i t i g a t e d . The g r e a t e s t
a d v e r s e e f f e c t s on t h e n e s t i n g ecology o f
--
D . u r b i c a were observed surrounding power p l a n t s
and open s u r f a c e mines. Chemical p l a n t emissions
had t h e n e x t most s i g n i f i c a n t e f f e c t on the house
martin.
In summary, s u b l e t h a l and c h r o n i c l e v e l s o f
a i r emissions from i n d u s t r i a l s o u r c e s , such a s
power p l a n t s and chemical p l a n t s , a d v e r s e l y
a f f e c t t h e n e s t i n g ecology o f 2. u r b i c a ,
including a reduction of the nesting density,
colony s i z e , and occupancy o f t h i s s p e c i e s . The
g r e a t e s t e f f e c t s on t h e n e s t i n g p a t t e r n o f
--
D . u r b i c a were observed i n a r e a s downwind from
t h e emission s o u r c e s and a t d e c r e a s i n g d i s t a n c e s
from the a i r emission s o u r c e s ( f i g . 1 ) . For low
s t a c k emission s o u r c e s , g r e a t e s t e f f e c t s were
observed i n t h e immediate v i c i n i t y o f t h e s t a c k s
where the h i g h e s t emission l e v e l s a r e expected t o
were observed a t i n t e r m e d i a t e d i s t a n c e s from t h e
s t a c k s and a g r e a t e r d i s t a n c e than observed i n
low s t a c k l o c a l i t i e s .
Evidence s u g g e s t s t h a t w i l d l i f e p o p u l a t i o n s
l i v i n g i n optimal h a b i t a t c o n d i t i o n s a r e l e s s
t i o n s l i v i n g i n l e s s t h a n optimal c o n d i t i o n s .
T h i s o b s e r v a t i o n h a s p a r t i c u l a r importance f o r
t h e management and p r e s e r v a t i o n o f endangered
species.
O v e r a l l t h e e f f e c t s o f a i r p o l l u t a n t s on
w i l d l i f e can b e d r a m a t i c , such a s t h e d i e - o f f s ,
o r q u i t e s u b t l e , such a s s h i f t s i n t h e age
s t r u c t u r e of populations. In many c a s e s t h e
e f f e c t s have been d e b i l i t a t i n g i n j u r i e s , o f t e n
contributing to the death of animals during
p e r i o d s o f n a t u r a l s t r e s s . A i r emissions can
a l s o reduce w i l d l i f e p o p u l a t i o n s n o t o n l y
d i r e c t l y , b u t i n d i r e c t l y through l o s s o f
habitat.
Acknowledgments: S p e c i a l r e c o g n i t i o n i s given
t o RNDr. E l i s k a Novakova, CSc f o r c o l l a b o r a t i o n
o n t h e m a r t i n s t u d y . I wish t o thank National
Academy o f Science and t h e Czechoslovak Academy
o f Science f o r support o f t h e b i r d s t u d y and
Environmental Sc i e n c e and Engineering f o r
a s s i s t a n c e i n t h e manuscript p r e p a r a t i o n .
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Chronic Effects of Acidic Precipitation
and Heavy Metals on Forest
Ecosystems

The Acidity Problem-Its Nature,

Causes, and Possible Solutions1

Lowell smith2

Abstract: Interest within the scientific community

in North America and Europe about the nature,

effects, and causes of atmospheric acid deposition

has grown rapidly over the past decade. This

interest has recently intensified because of

the explosion in public awareness of, and concern

over, the acid deposition problem, and a growing

political will to address the problem within

appropriate national and international forums.

his paper sketches the nature of the acid deposi-

tion problem; describes the atmospheric processes

that convert precursor emissions into acidic

compounds as these are transported over distances

ranging from a few to more than a thousand kilometers;

discusses past and possible future trends in geo-

graphical distribution and rate of acid deposition;

and summarizes the governmental activities which

have been initiated to address the problem.

The scientific study of acid deposition is invites reaction from any who take exception

archtypical of many contemporary environmental to the summaries presented in this paper.3

problems, in that it necessarily covers a wide

spectrum of disciplinary interests. Simply The major features of acid deposition are:

listing the many subdisciplines involved

0
would fill more than a page. It is impor- acid deposition results primarily

tant for the active research worker in the from the combustion of fossil

field to recognize the many interconnections fuels which releases sulfur dioxide

between her or his own endeavors and other (SO2) and nitrogen oxides (NOx)

research areas. This involves a careful in the form of nitric oxide (NO)

balance because, at the same time one is and nitrogen dioxide (NO?) to the

encouraging a cross-fertilization of ideas atmosphere;

among various disciplinary efforts, one must

guard against extending scientific judgements depositing acidic material is

beyond one's own limits of competency. Since formed out of these precursor

the author is fully aware of this hazard, he emissions by means of a large

number of chemical reactions as

'presented at the Symposium on Effects of the emissions are transported

Air Pollutants on Mediterranean and Temperate away from their source region;

Forest Ecosystems, June 22-27, 1980, Riverside,

California, U.S.A.
he views expressed in this paper are

those of the author and do not neces-

'~irector, Program Integration and Policy sarily reflect those of the Environmental

Staff, Office of Research and Development, U.S. Protection Agency.

Environmental Protection Agency, Washington, D.C.

 acidic material is deposited

Hicks, 1980). In addition, nitric acid

dry in the form o f fine particu-

depositing onto surfaces out of its

late matter and is incorporated

vapor phase may at times be an important

into all of the possible physical

contributor to dry deposition; but even

forms of precipitation;
less is known about its ambient concen-

tration and geographical distribution

transport distances between

(Altshuller, 1979). Thus, until monitor-

source and receptor regions can

ing techniques are developed for routine

exceed a thousand kilometers,

reliable measurements of dryfall deposition,

although a major source can, on

we are compelled to rely on isolated spot

occasion, significantly affect

measurements and tentative inferences

the rate of acid deposition within

to characterize this important phenomenon.

the first few kilometers downwind

of the source; and

The characterization of wet deposition

is a more tractable problem. While it

the effects of the deposited acid

is doubtful that precipitation at any

can vary markedly, depending upon

place on the globe is entirely free of

the form in which the acid is

anthropogenic contaminants, relatively

deposited, the biologic, geologic

"clean" rainfall is generally found on

and hydrologic pathways between

the windward edge of continental land

deposition site and the receptor

masses and where frontal storm systems

of interest, and the sensitivity

have traversed large expanses of sparsely

of the receptor.
populated land surfaces (Granat, 1978).

Because of the formation of carbonic

This paper explores these major features

acid from the hydrolization of background

of acid deposition.
atmospheric carbon dioxide, such clean

rainfall is thought to have a pH of

Nature of Acid Deposition

about 5.6, approximately 25 times more

acidic than a neutral pH of 7 (Likens,

Acidic compounds may be deposited by

several forms of precipitation (rain,

-
et. g., 1979). Yet even the pH of

rainfall relatively unaffected by anthro-

snow, hail, dew, rime, and mist) as well

pogenic emissions can vary by several pH

as by fine particles that settle out of

units due to the entrainment of alkaline

the atmosphere on to biologic, mineral,

soil particles, reaction with atmospheric

and aquatic surfaces, and on to man made

ammonia, and possibly other little under-

materials, buildings, and artistic objects.

stood factors (Stensland, 1979).

Because monitoring techniques are not

well developed for measuring and character-

The concentration of acidity in precipi-

izing the dryfall component of acid deposi-

tation is observed to be quite episodic

tion, comparatively little is known

(Smith and Hunt, 1979), which combined

about its extent and variability (~urham

with the episodicity of precipitation

and Hicks, 1980). Monitoring data from

rates, leads to great variability over

eastern North America indicates that the

space within a particular rain event and

sulfate ion is the predominant anion

over time at a particular location of the

present in acidic precipitation; the

rate of acid deposition (~ales,1980).

nitrate ion is associated with about

A range of over seven orders of magnitude

half as much acidity as the sulfate ion,

of acidity (pH 2 to pH 9) has been recorded

and chloride and other anions make sub-

for various isolated rainfall events in

stantially lower contributions (Hales,

North America. Averaged over time, spatial

1980). In some areas of the West the

variations tend to vanish at locations

nitrate ion contributes an equal or

distant from large sources. An excep-

larger share than the sulfate ion (McColl,

tion is the orographic effect produced

1980; and Morgan and Liljestrand, 1980).

by high terrain features on precipitation

rates. This effect can be further augment-

Some regional scale atmospheric models

ed by the increasing acidity of cloudwater

estimate that up to half of the sulfate

at higher elevations within a cloud

component may be deposited dry, while

structure (Falconer and Falconer, 1980).

other models estimate a lower percentage

Terrain subject to orographic precipitation

(Whelpdale and Galloway, 1979). The

in North America and Europe, such as the

variance results from a basic lack of

Nothern Alps, that is downwind of high

understanding of the physical, as well

emission areas has been observed to

as biological processes, and their varia-

sustain substantially increased quanti-

tion over space and time, as these processes

ties of deposited acidic material than

transfer gases and particles across the

similarly situated low lying terrain

atmosphere/surface interface (~urhamand

(Schrimpff, 1980).

 Averaging wet acid deposition rates over an

upper soil horizons of forest soils subjected to

annual cycle for monitoring stations in eastern

heavy rates of acid deposition. In extreme cases

North America produces a pattern of depressed pH

ground water quality may be adversely affected

values in the northeast United States (Pack,

(Hultberg and Wenblad, 1980). Abnormally low pH

1980) extending northward into southern Ontario

for drinking water supplies drawn from wells in

and Quebec, westward to the eastern Midwest,

some of these regions has been observed. This

and southward along the Appalachian ridge into

can result in unacceptably high concentrations

Tennessee and the Carolinas. Less depressed

of copper and lead in drinking water in those .
average pH levels extend deep into Florida
homes which are equipped with copper plumbing

(~rezonik,s. &. , 1980), west to Arkansas and systems (~ultbergand Wenbald, 1980).

Missouri, and northward an undetermined distance

into Canada (Likens and Butler, 1980). The

Agricultural crop species cultured in pots and

average acidity of precipitation in the central

subjected to lowered pH simulated acid rain irriga-

portion of this pattern approaches pH 4, approxi-

tion water have displayed mixed responses in

mately a factor of forty more than what many

yield (Lee, z. e., 1980). Some cultivars exhibit

consider to be the "normal" value of pH 5.6

enhanced yields and others depressed yields while

(Likens and Butler, 1980).

many cultivars show no discernible effect on yield.

The effect of acidic deposition on forest growth

Due to the spottiness of available monitoring

rates is more uncertain. Early stages of acidifi-

information less is known about acid deposition

cation can accelerate growth rates for a few

rates west of the Mississippi River. Measurements

years, possibly due to the increased mobilization

made in the Boundary Waters Canoe Wilderness

of nutrients within the soil structure. But

Area (northern Minnesota) (Glass and Loucks,

sustained elevated rates of acid deposition are

19801, the Colorado Rockies (northwest of ~enver)

hypothesized to depress forest growth rates in

(Lewis and Grant, 19801, and California (the

geologically sensitive areas due to nutrient

Sierra Nevada, and the Los Angeles and San Fran-

depletion and elevated concentrations of AI^+
cisco Bay basins) (McColl, 1980; and Morgan and
ion within the root zone. Other ecological effects

Liljestrand, 1980) all suggest that this problem

are discussed elsewhere (Overrein, 1980, Norton,

is not unique to the eastern half of the continent.

et. G . , 1980, and Cowling, 1980).
-
Effects of Acid Deposition
Mechanisms of Atmospheric Formation

Acid deposition creates a public policy problem

As previously stated, acid deposition is primarily

to the extent that potentially sensitive receptors

the result of the anthropogenic release of SO2

are harmed by the deposition rates they sustain.

and NOx to the atmosphere where these react

Research results on acid deposition effects

through several available chemical pathways to

are multiplying rapidly in North America and

sulfuric and nitric acids. To a lesser extent

Europe (International Conference on the Ecological

hydrochloric acid can also be involved. Coal and

Impact of Acid Precipitation, 1980). An impressive

oil fired electric utility generating stations,

body of information now exists as to the impact

predominantly in the central and eastern portions

of acidifying surface waters on the ecosystems

of the U.S. are responsible for two-thirds of the

they support (Gorham, 1976). Some fish species

national SO2 emission inventory (U.S. EPA, 1979a).

show adverse effects below pH 6; while other

Nine-tenths of these emissions are from coal-fired

more tolerant fish species do not evidence serious

stations. Industrial and commerical boilers account

effects until pH 4 or lower has been reached.

for nearly half of the remaining emissions.
SO2

Eggs and especially the fry of sensitive species

emissions in the U.S. are likely to increase slowly

are more susceptible than are adult fish (Johannson,

over the next few decades as emissions from

et. &., 1977).

- Thus, during the initial melting well-controlled new sources slightly overbalance

of a snowpack, acid pulses may be released which

emission reductions expected to be achieved by

abnormally skew subsequent fish population distri-

the control or retirement of existing sources

butions to the extent that one or more generations

(Altshuller and McBean, 1979). Canadian SO2

may be completely absent in some instances.

emissions come predominantly from their non-ferrous

Other portions of aquatic food chains can also

smelter industry at present, but by the end of

be affected to the detriment of wildlife species

the century coal fired utility boiler emissions

at the top of these chains, including fish eating

are expected to about equal those from the smelter

birds (Peakall, 1980) and freshwater wading

industry (Choquette, 1980).

birds (~oucks,1980).

Two-fifths of the nation's NOx emissions come

Recent evidence from Europe suggests that

from motor vehicles. Geographically, these mobile

leaching of calcium and other nutrients from the

sources are clustered around large population

lower horizons of sensitive soils and the concomitant

centers. Nine-tenths of the remaining NOx emissions

mobilization of Al3+ ion within these soils (Ulrich, are released by the same large utility and industrial

1980) is promoted by atmospheric acid deposition.

boilers which are responsible for the predominant

Also observed is a reduction of the decomposition

portion of the SO2 inventory (U.S. EPA, 1979a).
NOx

rate of forest litter and decreased numbers of

emissions are expected to increase significantly in

micro-organisms (Baath, s. &., 1978) within the the U.S. and Canada over the remainder of the century,

unless some promising breakthrough in NOx

inventory has yet to be constructed. Stockyards,

emission controls is achieved and rapidly

municipal waste water treatment plants, certain

implemented by industry (Altshuller and

industrial processes, and decaying vegetable

McBean, 1979).
matter are among its important sources.

Although no emission inventory exists for

Some investigators have hypothesized that fly

chloride ions, these emissions are likely to

ash from coal combustion has historically played

be primarily the result of burning high

a major role in reducing atmospheric acidity and

chloride coals produced from many coal seams

that recent efforts to control fly ash emissions

in the midwestern and eastern U.S. Therefore,

have noticeably worsened the acid deposition

chloride emissions are likely to be colocational

problem (Frohlinger and Kane, 1975). Such an

with large SO2 and NOx emitting sources. Natural

effect is unlikely to have been nearly as important

emissio s of SO2 (Adarns,s. &., 1980) as was once supposed. As will be discussed later

and:ON make only modest additions to the

in this paper it is unclear what the historic

anthropogenic sources in eastern North America,

trends for deposited acid are in the Northeast.

though natural emissions may make an appreciable

Thus, such an ad hoc explanation may not be

contribution to the global background of these relevant to explain what is an inconclusive

atmospheric constituents (Husar, s. al., 1978). trend. More importantly, considerations such

as the size of fly ash particles relative to the

These precursor emissions react in the atmos-

size of sulfate particles, the chemical composition

phere with water vapor, other minor atmospheric

of fly ash from midwestern and eastern coals, and

constituents and sunlight to produce fine

the change from stoker-fired coal boilers to

sulfate particles, nitric acid vapor and

pulverized coal boilers during this period, all

dilute acids in cloud droplets. These submicron

suggest that the neutralizing effect of emitted

particles agglomerate into larger particles

fly ash could only have been important in the

until a natural barrier to further growth is

immediate locality of a relatively few heavy fly

reached at slightly above one micron mean

ash emitting sources.

diameter. The growing particle may be deposited

in dry form onto a surface, incorporated

Mechanisms of Atmospheric Acid Transport

into a cloud droplet (rainout), or scavenged

by a falling raindrop (washout) (Hales, 1980).

Several physical processes are important to

The hydroxyl ion in conjunction with sunlight

the atmospheric transport of acid precursors and

is believed to promote the formation of

their acidic products. Particularly important for

particulate sulfate (Davis,s. &., 19741, while the S~~/sulfuric acidlsulfate

complex is the

the peroxyl ion is believed to promote the conv r-

elevated height of injection into the atmosphere

sion of SO2 to sulfuric acid in cloud droplets.5

level for SO2 emissions from most major power

plant sources. A nocturnal inversion layer

In the summer months, substantially enhanced

frequently isolates these tall stack plumes from

concentration of sulfate ion are found in deposited

the gr~und(Smith, s. &., 1978) until after
rainwater while nitrate ion concentrations tend
sunrise when the incident solar energy begins to

to be more constant throughout the year (Hales,

mix the atmosphere through the activation of

1980). There is some evidence to suggest that

convective cells. In the Midwest during summertime

nitrogen oxides are deposited more rapidly from

conditions, a nocturnal bulge in the wind speed

the atmosphere on the average than are sulfates

vertical profile is frequently observed at normal

(Mueller, s.&. , 1979), so nitric acid -deposi- tall stack plume heights. This condition can

tion may be relatively less important for receptor

transport emissions from a tall stack several

sites far from emission regions than is sulfuric

hundred kilometers overnight (Smith, s. &.,
acid deposition.
1978). The gas to particle conversion of the

emissions in this displaced plume can be greatly

Another complicating feature in the atmospheric

accelerated the next day as these transported

chemistry of acid deposition is the atmosphere's

emissions are mixed with a polluted urban air mass.

ability to partially neutralize its acid load.

Alkaline fine wind blown soil particles, particu-

The highest ambient concentrations of particulate

sulfate are observed under summertime conditions

the acid load (Eisenreich, &..

larly over arid regions, appear to neutralize

s. , 1980),
or create aerosols with basic chemical properties.

when a synoptic scale high pressure system stalls

for more than a day over a region of high SO2 emission

The ammonium ion is also effective in partially

density, such as the Ohio River Basin (Hidy, s. al.,
neutralizing dry and wet atmospheric acidic
1978). In this situation weakly circulating winds can

materials. Many natural and anthropogenic sources

trap a large air mass while it is being continually

produce ammonia, but a complete ammonia emissions

filled with precursor emissions (Vukovich, 1979). The

higher temperature, moisture and sunlight levels

4~ersonalcommunication from Dr. Rudolph

generally encountered under these conditions tend

Husar, August 15, 1980, Washington, D. C.

to increase the chemical reactivity of the atmosphere,

so the higher concentration of precursor emissions

'personal communication from Dr. A. L. Lqzrus,

can more rapidly be converted to particulate sulfate.

May 21, 1980, Washington, D. C.

 A common atmospheric cleansing mechanism

available rainwater chemistry monitoring data is

for this condition is for a cold front to

required in order to establish the level of certainty

approach the high pressure center from the

with which historical trends of acid deposition

north or northwest (Whelpdale, 1978). This

in North America can be determined.

creates a strong pressure gradient which

sweeps much of the polluted air mass parallel

Fortunately, it may be possible to describe

to the line of the front for many hundreds

the gross features of deposition trends by relying

of kilometers, frequently to the northeast.

on other related physical phenomena for corroborative

(LaFleur and Whelpdate, 1977). Frontal

support. Since sulfate aerosols are highly efficient

storm activity can further remove considerable

light scatterers due to their characteristic submicron

amounts of pollutants as rainout and washout.

size, and since sulfate is the dominant component of

Similarly, large convective storms are believed

Eastern aerosols (U.S. EPA, 1979b), visibility trends

to be an efficient mechanism for processing

probably serve as a useful indicator for dry sulfate

and removing pollutants from the large volume

deposition trends. The ratio of dry to wet sulfate

of air they entrain. These storms are capable

deposition should have remained relatively constant

of pumping large quantities of polluted air from

for a particular area unless a climate change has occurred.

the planetry boundary layer to high elevations

where these can be left as isolated patchy layers

Visibility measurements have been routinely made at

to be transported considerable distances and

medium and large size airports in the U.S. for several

eventually deposited.
decades. Recent analysis of these data shows that

summertime visibility has significantly deteriorated

Alternating periods of stagnation and ventila-

throughout large portions of the eastern U.S. (Trijonis

tion over a high emission area produce episodic

and Shapland, 1979). Some regions such as the Tennessee

concentrations of pollutants (~ttar,1978) which

Valley appear to have sustained nearly a factor of two

result in highly variable acid deposition rates in

downwind regions. Likewise, the variability in

wind direction, as it guides and mixes isolated

five year period (Husar,=.

decrease in average summertime visibility over a twenty-

&., 1979). Similar trends

in summertime solar insolation are also suggestive

plumes from major sources and the regional scale

of a increase in atmospheric turbidity during this same

plumes described above, adds to the temporal

period. Further analysis effort is required to

variability of deposition rates at any monitoring

assess the possible causal relationships between

site. Thus, acid deposition must be viewed as a

trends in precursor emission rates and these trends

stochastic process which, in areas not under the

in environmental conditions.

influence of a major source and not affected by

orographic terrain effects, is temporally and

Although the trends of such surrogates for

spatially chaotic over a small scale but is rather

acid deposition may only be used to corroborate an

spatially homogeneous within a given region when

inconclusive record for monitored acid deposited,

averaged over a large number of events.

other studies strongly support the conclusion that

anthropogenic emissions are deposited on ecosystems

Historical Trends in Acid Deposition Rates

far from their point of origin. Dated lake bottom

cores from remote lakes in North America and Greenland

While the routine measurement in a scienti-

icecap cores show a marked increase in deposition

fically reliable manner of wet deposited acid

rates for fossil fuel combustion-related pollutants

has only recently, with the exception of a

shortly after the beginning of the industrial revolution.

very few monitoring stations, been undertaken

in this country, it has been practiced in . A review of emission trends from U.S. sources

several European countries over a longer period

over the past forty years indicates several clear

of time (Granat, 1978). Both the European and

trends. First, SO2 emissions increased about forty

North American experiences have demonstrated

percent. ~ l t h o u ~ hemissions
~ 0 ~
have decreased

the need for strict quality assurance procedures

from most economic sectors during this period, the

for collecting and analyzing the rain water

electric utility sector's emissions increased by

samples. Failure to establish sufficiently

more than a factor of six during this same period

stringent quality assurance procedures early in

(EPA, 1978). Second, the increase in SO2 emissions

monitoring programs has created questions about

from this one sector occurred concurrently with a

the validity of much of the early data (Tyree, 1980).

substantial increase (by approximately a factor of

five) in the stack height for utility sources. Third,

The need to calculate retroactively the acidity

SO2 emissions from coal burning changed from a

of monitored rainfall using one of several ion

balancing procedures, has compounded these

measurement difficulties since acidity or pH

rate (Husar,=.
wintertime peak to a summertime peak in emission

&. , 1979). Fourth, the precursor

emissions for photochemical oxidants increased

was usually not measured directly (Kramer,

markedly during this time (EPA, 1978). At the

1978). Such procedures can propogate the

beginning of the period photochemical smog was hardly

experimental uncertainties, which were introduced

recognizable as a problem, whereas, currently urban

by the collection and laboratory procedures

plumes of photochemical oxidants now frequently blanket

I employed, into rather sizable uncertainties

the Northeast during the summer months (Altshuller,

in the calculated hydrogen ion concentra-

1978). Fifth, total NOx emissions approximately

tion. Further careful analysis of all

quadrupled during this period (EPA, 1978).

 These trends suggest a situation in which

research activities to determine

the atmosphere has become chemically more

more completely the range of

reactive, and for which greater quantities of

environmental effects produced by

acid-forming precursors are added to the atmosphere

acid deposition and the atmospheric

during its most reactive period. Further, a

processes which transport and convert

substantially greater quantity of these emissions

emissions into acid deposition;

is now injected high into the mixed layer where

the emissions and their reaction products have

assessment activities to determine

much longer residence times as they travel to

the potential seriousness of the

areas remote from their point of origin.

acid deposition problem in North

America and the most cost-effective

Possible Solutions
measures that could be employed as

first steps to combat the problem;

At present there are no regulatory requirements

that are primarily directed at reducing emissions

the creation of a ten year inter-agency

to control acid deposition. However, the

Federal Acid Rain Assessment Program

Clean Air Act's requirement to reduce ground

to coordinate the above activities;

level SO2 concentrations has achieved modest

overall reductions in SO2 emissions during

working with the states in order to

the first part of the past decade (u.S. EPA,

promote a mutual understanding of

l979a). But, average stack heights continued

the nature and causes of the acid

to increase, as did the emission rates for

deposition problem, and to encourage

NOv, over this period. Regulatory requirements

the states to engage in collective

established for new coal-fired utility boilers

problem solving on this issue; and

mandate control of seventy to ninty percent of

their SO2 emissions and require NOx emission

becoming a signatory to an inter-

reductions of approximately forty percent

national Convention on Long-Range

(U.S. Federal Register, 1979). EPA is currently

Transboundary Air Pollution, developed

developing new source control performance

under the auspices of the United

standards for industrial boilers which could

Nations Economic Commission for

require similar levels of control for this

Europe, and pursuing bilateral

important source category. From a regulatory

discussions with Canada which are

perspective the principal problem for SO2

expected to evolve into formal

emission control is the control of emissions

negotiations on a U.S.-Canada

from existing sources. In addition, there are

bilateral transboundary air pollu-

large opportunities for improved NOx control

tion agreement.

requirements for new and existing sources.

Such efforts cannot be expected to bring about

Others have maintained that the costs of

immediate and complete relief from the acid deposition

emission control are so high that the only

-
problem nor are they designed to achieve this
cost-effective mitigation measure is to raise
objective. Rather the goals are to establish as

artifically the pH of affected lake water by

quickly as is feasible a scientific basis for under-

the addition of lime (~arnes,1979). Repeated

standing the full range of receptors at risk

treatments would be required as long as acid

and the extent of risk to each receptor category;

deposition rates exceeded the geologically

to determine the extent of the control measures

controlled release rates for a given area

which are required to reduce the acid deposition

(Horn, s. &., 1980). Recent Swedish exper- problem to an acceptable level; to develop new

ience indicates that liming the entire watershed

air pollution control policies and strategies as

for a lake may be necessary, while others

necessary and appropriate; and to promote full

believe that even though it is possible to

cooperation and understanding across interstate

raise the pH of an acidified lake, it is not

and international borders to deal effectively

possible to restore the lake to a natural

with this complex and challenging problem.

condition containing its original food chains.

Others have observed that the potentially

Acknowledgments

sensitive areas in North America cover millions

of hectares, and question the feasibility of

The author is deeply indebted to many colleagues

liming such a vast area for hundreds of years.

and research workers for the opportunity to

learn from and share ideas with them. Particular

Federal government efforts to address

credit is given to Dr. Brand L. Niemann for

the acid deposition problem include:

monitoring activities to establish

^ ~ o t eadded in editing: A Memorandum of Intent

more fully the geographical var-

was signed in Washington, D. C., on Aug. 5, 1980,

iation and temporal trends in

by the two governments in which they agreed to

rainwater chemistry;
initiate formal negotiations by June 1, 1981.

his helpful comments, to Ms. Barbara H. Brandon

-
States A Comprehensive Survey, Report
and Mr. Paul Schwengels for their editorial
prepared by the Pollution Data Analysis

advice, and to Ms. Mable Scales and Ms. Veronica

Division of the Air Pollution Control

Parker for their special efforts in typing the

Directorate, Environment Canada, 24 p.,

manuscript. Finally, I wish to thank Dr. Paul R.

Ottawa, Ontario.

Miller for the extraordinary patience he displayed

while coaxing me to complete this paper.

Cowling, Ellis B.

1980. Effects of Acidic Precipitation

on the Health and Productivity of

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 Acid Precipitation Impact on Terrestrial

and Aquatic Systems in Norway'
2
Lars N. Overrein

Abstract: In r e c e n t decades t h e a c i d i t y of r a i n and snow

has increased sharply over wide areas. The p r i n c i p a l
cause i s t h e r e l e a s e of sulphur and n i t r o g e n oxides by
t h e burning of f o s s i l f u e l s . The a i r q u a l i t y i n any one
European country i s measurably a f f e c t e d by emissions i n
o t h e r European countries. Strong a c i d s have lowered t h e
annual mean pH of p r e c i p i t a t i o n i n much of northern Eu-
rope t o between 4 and 5. In southern c o a s t a l a r e a s of
Norway, t h e annual mean a c i d i t y i n p r e c i p i t a t i o n i s now
4.3 pH-units, o r even more a c i d i c .

Acid p r e c i p i t a t i o n has increased leaching of n u t r i -

e n t s from t h e uppermost s o i l l a y e r s . These l o s s e s of
n u t r i e n t s may be expected t o decrease p l a n t growth, b u t
f i e l d evidence i n Norway and elsewhere, has not y e t
been obtained. I t i s p o s s i b l e t h a t p o l l u t e d a i r and
p r e c i p i t a t i o n over a period of years can influence p l a n t
production.

Atmospheric t r a n s p o r t of sulphur and o t h e r a c i d i f y -

i n g components has l e d t o extensive regional a c i d i f i c a -
t i o n of water courses i n a r e a s with very l i t t l e neutra-
l i z a t i o n capacity. A c i d i f i c a t i o n of watercourses had
had major e f f e c t s on l i f e i n r i v e r s and l a k e s . Lakes
i n an a r e a of 13,000 km2 i n southern Norway have become
empty of f i s h i n r e c e n t decades, and a f u r t h e r a r e a of
approx. 20,000 km2 contains l a k e s with s i g n i f i c a n t l y r e -
duced f i s h stocks.

The e c o l o g i c a l impact of a c i d p r e c i p i t a t i o n has In Norway a c i d p r e c i p i t a t i o n was a t t h a t time seen

been a m a t t e r of growing concern over t h e l a s t de-
cade p a r t i c u l a r l y i n t h e i n d u s t r i a l i z e d c o u n t r i e s
of t h e Northern Hemisphere.
The Norwegian I n t e r d i s c i p l i n a r y Research Pro-
gramme "Acid P r e c i p i t a t i o n -
E f f e c t s on F o r e s t and
Fish", (The SNSF-project) was i n i t i a t e d i n 1972.
a s a p o s s i b l e cause of i n c r e a s i n g a c i d i t y of t h e
watercourses i n t h e southern p a r t of t h e country,
and of t h e gradual disappearence of valuable f i s h
populations from many l a k e s and r i v e r s . I t was a l -
so feared t h a t t h e i n p u t s of a c i d might over time
reduce f o r e s t growth p a r t i c u l a r l y through increased
leaching of n u t r i e n t elements from t h e s o i l .

The SNSF-project has t h i s year marked t h e con-

Presented a t t h e Symposium on E f f e c t s of A i r c l u s i o n of e i g h t years of research by organizing
P o l l u t a n t s on Mediterranean and Temperate F o r e s t t h e I n t e r n a t i o n a l Conference on t h e Ecological Im-
Ecosystems, June 22-27, 1980, Riverside, p a c t of Acid P r e c i p i t a t i o n . This Conference, which
C a l i f o r n i a , U.S.A. was held i n Sandefjord, Norway, March 11 14, 1980, -
2 ~ e s e a r c hD i r e c t o r ,The Norwegian I n t e r d i s c i p l i - a t t r a c t e d more than 300 p a r t i c i p a n t s from some 20
nary Research Program, Acid P r e c i p i t a t i o n - countries.
E f f e c t s on F o r e s t and Fish.
The SNSF-project - 1432 Aas-NLH, Norway. The p r e s e n t r e p o r t i s mainly a summary and b r i e f
discussion of the ecological impact of acid pre-
cipitation in Scandinavia with particular refer-
ence to Norway.
EMISSIONS, TRANSPORT, AND DEPOSITION
The concept of acid precipitation is now used
to denote precipitation with high amounts not on-
ly of the hydronium ion (H+ for short), but with
enhanced concentrations of the acidifying anions
sulphate, 304 and nitrate, NO3, which predominant-
ly stem from anthropogenic sources. "Acid" pre-
cipitation regularly also contains high amounts
of ammonium, NH4, and of various heavy metals
and trace elements including organic micropollu-
tants.
It should also be kept in mind that comparable
amounts of anthropogenic pollution may be deposi-
ted by dry deposition processes from the same air
masses giving acid precipitation. In studies of
effects of acid precipitation the contribution
from dry deposition is often difficult to quanti-
fy. In areas remote from the main industrial
centres, like in Southern Norway, acid precipita-
tion is more or less synonymous with long-range
transported air pollution, but the distinction
should always be observed. In our context, sul-
phur and nitrogen compounds giving rise to the
acidifying properties of precipitation, are of
prime interest, and their sources should be de-
scribed with a view to finding their geographical
distribution, emission rates and seasonal varia-
tion. This is necessary both for enabling mo-
delling of their transport, and for formulating
abatement policies against their negative effects.
Sulphur Emissions
Most of the man-made sulphur emissions occur
as SO2 from combustion of coal and petroleum pro-
ducts. Comparatively less stems from smelting of
sulphur-containing mineral ores and other indus-
trial processes, on a global basis about 10 per-
cent. Knowledge of the location of industrial
and powerproducing units in addition to popula-
tion distribution has allowed quantification of
annual emissions in a 150 km x 150 km grid over
.
Europe (Semb, 1979) The main area of SO2 emis-
sions corresponds to the industrial belt from the
Midlands in U.K., the Netherlands and Belgium,
central and southern parts of Germany and Poland.
Parts of northern France, Czechoslovakia and USSR
also have industrial concentrations with very high
emission rates. On a country basis, total SO2
emissions in 1973 were estimated at e.g. 2.8 lo6
tonnes S from UK, 2.0 from the Federal Republic
.
of Germany and 1.6 from France (OECD, 1977) Nor-
wegian annual emissions were estimated at 91,000
tonnes of S.
Seasonal variations in SO2 emissions occur as
the demand for energy fluctuates through the year.
At high latitudes the variable component of emis-
sions is related to space heating and illumination
during winter. In warmer climates, the maximum
demand may occur during summer due to the use of
air conditioning. In most of Europe, the fuel
consumption will probably be at its peak in Janu-
ary - February. The seasonal variation is about
30 percent of the annual mean emission.
Nitrogen Emissions
Both nitrogen oxides, NOx, and ammonium, NH4,
play important roles for the composition of acid
precipitation. Although global budgets of nitro-
gen compounds are even more uncertain than those of
sulphur, natural sources seem to be larger than the
man-made. However, major man-made emissions also
of nitrogen compounds occur in Europe and thus in-
fluence strongly precipitation chemistry in
Scandinavia. The main anthropogenic source of ni-
trogen oxides is combustion of fossil fuels, by
oxidation of nitrogen compounds in the fuel and
oxidation of nitrogen in the combustion air.
It is well known that the NOx emissions depend
on fuel types, combustion chamber design and oper-
ating conditions. High combustion temperatures
favour the emissions. There is, naturally, a high
spatial correlation between SO2 and NOx emissions.
OECD studies indicated almost a doubling of an-
thropogenic nitrogen oxides emissions in Europe
from 1959 to 19%73,thus increasing more than sul-
phur dioxide emissions in the same period (OECD,
1977).
Trace-Element Emissions
Acid precipitation contains a wide range of
minor and trace elements associated with the ma-
jor chemical constitutents. Heavy metals, other
trace elements and organic micropollutants of man-
made origin are receiving increasing interest as
some of these are enriched in living organisms.
Emission rates for trace elements and micro-
pollutants are largely unknown, but the main sour-
ces are known . Many of the organic micropollu-
tants in the atmosphere are products of human ac-
tivity, including industrial and waste products,
and also chemicals used in industry as solvents
or intermediate products. The emissions are very
complex mixtures of chemical compounds.
Transport and Deposition
Of particular interest for long-range pollu-
tant transport is the build-up of high concentra-
tions in stagnant air near the ground during in-
version situations. Observations show that such
parcels of contaminated air may subsequently move
over long distances without much dilution. A
much used technique in analyzing source areas and
transport directions of air pollutants, is the
sector analysis, grouping together trajectories
belonging to the same sector.
 Of considerable i n t e r e s t f o r t h e p r o p o r t i o n of
s u l p h a t e i n a c i d p r e c i p i t a t i o n t o dry deposited
sulphur components, i s t h e o x i d a t i o n r a t e from
sulphur dioxide t o s u l p h a t e s . The o x i d a t i o n
t a k e s p l a c e both by a b s o r p t i o n of SO2 i n cloud
d r o p l e t s with subsequent o x i d a t i o n , and by oxi-
d a t i o n i n t h e gas phase with oxygen compounds i n
t h e atmosphere. High c o n c e n t r a t i o n s of ozone and
photochemical o x i d a n t s , which a r e observed over
l a r g e a r e a s of Europe, w i l l i n c r e a s e t h e t r a n s -
formation r a t e . The t r a n s f e r of gases and par-
t i c l e s from t h e a i r t o n a t u r a l s u r f a c e s , and t h e
a d s o r p t i o n , a r e u s u a l l y d e s c r i b e d i n analogy with
t h e t h e o r y of e l e c t r i c a l r e s i s t a n c e . The t r a n s -
p o r t from t h e atmosphere t o t h e boundary l a y e r
c l o s e t o t h e s u r f a c e , t a k e s p l a c e by t u r b u l e n t
d i f f u s i o n . The a e r o s o l p a r t i c l e s i n q u e s t i o n ,
i - e . , s u l p h u r and n i t r o g e n a e r o s o l s , a r e mostly
i n t h e 0.1 - 1.0 pm s i z e range with low g r a v i t a -
t i o n a l s e t t i n g , and t h e t u r b u l e n t t r a n s p o r t w i l l
depend on meteorological c o n d i t i o n s .
Model c a l c u l a t i o n s of wet and dry d e p o s i t i o n
p a t t e r n s over Europe show t h a t i n Scandinavia,
p a r t i c u l a r l y i n Norway, t h e wet d e p o s i t i o n out-
weighs t h e e s t i m a t e d dry d e p o s i t i o n . For southern
Norway dry d e p o s i t i o n i s e s t i m a t e d t o account f o r
about 30 p e r c e n t of t h e t o t a l d e p o s i t i o n of ex-
cess sulphate.
The p r e c i p i t a t i o n i n Norway i s l a r g e l y d e t e r -
mined by p o l a r f r o n t lows b r i n g i n g moist maritime
a i r i n w e s t e r l y t o s o u t h - e a s t e r l y d i r e c t i o n s . Of
p a r t i c u l a r importance i s t h e orographic enhance-
ment o f p r e c i p i t a t i o n , caused by t h e l i f t i n g and
subsequent c o o l i n g of t h e a i r masses when flowing
a c r o s s t h e Scandinavian mountain chain. This
g i v e s r i s e t o a maximum zone of e l e v a t e d p r e c i p i -
t a t i o n some 40 - 50 km from t h e c o a s t l i n e . In
this zone, annual mean p r e c i p i t a t i o n exceeds
1000 mm along t h e SE c o a s t i n c r e a s i n g t o an abso-
l u t e maximum of perhaps more than 5000 mrn i n n o r t h
western Norway. S t i l l f u r t h e r n o r t h , i n northern
Norway, annual p r e c i p i t a t i o n i n t h e maximum zone
exceeds 2000 mm.
The group of macrocomponents t y p i c a l of a c i d
p r e c i p i t a t i o n , i . e . H+, NH4, SO4 and NO3, has a
marked north-south g r a d i e n t . The c o r r e l a t i o n s
between s u l p h a t e , n i t r a t e and ammonium a r e high,
and t h e r e a r e roughly e q u i v a l e n t amounts o f t h e
p r i n c i p a l c a t i o n s H + NH4+ and anions SO4
-. and
NO3 The c o n t e n t of s t r o n g mineral a c i d i n pre-
c i p i t a t i o n i s s t r o n g l y c o r r e l a t e d with excess
s u l p h a t e . The c o r r e l a t i o n c o e f f i c i e n t i s 0.7 -
0.9 a t Norwegian s t a t i o n s . A t Norwegian s t a t i o n s
No3 makes about 30 p e r c e n t o f t h e sum SO4 + NO3.
Ammonium and n i t r a t e occur i n Norwegian p r e c i p i -
t a t i o n i n about e q u i v a l e n t c o n c e n t r a t i o n s , lowest
a t mountain s t a t i o n s . The c o n c e n t r a t i o n s of
~ 0 4 ~ and - H+ i n p r e c i p i t a t i o n a r e h i g h e s t along
t h e south-east c o a s t . The mountain p l a t e a u i n
northernmost Norway i s a f f e c t e d by a i r t r a n s p o r t
from t h e s o u t h , which g i v e s a c i d p r e c i p i t a t i o n
(pH 4.5 - 5.0) a s f a r n o r t h a s 7 0 O ~ . l a t . In
southernmost Norway about 10 p e r c e n t of t h e pre-
c i p i t a t i o n h a s a pH below 4.0 and about 5 p e r c e n t
above pH 5.0. P r e c i p i t a t i o n a c i d i t y below 3.5
has been observed s e v e r a l times a t s e v e r a l p l a c e s ,
Dovland and Semb, (1980) .
P r e s e n t knowledge o f atmospheric d e p o s i t i o n of
inorganic t r a c e elements i n Norway i s compiled by
Semb (1978). They a r e found i n t h e w a t e r s o l u b l e
f r a c t i o n of a e r o s o l s , o r absorbed on t h e o t h e r
p a r t i c l e s , and a r e mostly contained i n t h e a e r o s o l
s i z e f r a c t i o n with aerodynamic mass diameter be-
low 2 urn. Available d a t a f o r l e a d , z i n c and cad-
m i u m i n p r e c i p i t a t i o n shows a d e p o s i t i o n p a t t e r n
s i m i l a r t o t h a t of excess s u l p h a t e . Also antimo-
ny, a r s e n i c , selenium and vanadium seem t o have
s i m i l a r d e p o s i t i o n p a t t e r n s . Influence from me-
t a l l u r g i c a l i n d u s t r i a l centres is evident f o r in-
s t a n c e f o r chromium (western Norway) and a r s e n i c ,
selenium, n i c k e l , chromium and copper (smelting
i n d u s t r y i n t h e Murmansk a r e a , U.S.S.R.). I n nor-
t h e r n Norway t r a j e c t o r y a n a l y s i s shows t h a t h i g h l y
p o l l u t e d episodes a r e o f t e n a s s o c i a t e d with t h e
r e t u r n flow p a t t e r n d i s c u s s e d by Rahn and McCaff-
r e y (1980). Several s t u d i e s of o r g a n i c micropol-
l u t a n t s i n p r e c i p i t a t i o n and i n a e r o s o l s have
been performed w i t h i n t h e SNSF p r o j e c t . They
have i d e n t i f i e d a wide range of compounds i n t h e
same a i r masses b r i n g i n g a c i d p r e c i p i t a t i o n t o
Norway.
ECOLOGICAL IMPACT
The e f f e c t s of a i r p o l l u t i o n have h i s t o r i c a l l y
been considered l o c a l problems, o c c u r r i n g n e a r pol-
l u t a n t s s o u r c e s , u s u a l l y urban a r e a s . This con-
c e p t of p o l l u t e d c i t i e s versus c l e a n r u r a l a r e a s
i s no longer a p p l i c a b l e . The i n c r e a s e i n anthro-
pogenic emission sources coupled with t h e i n c r e a s -
ed h e i g h t of emissions have enhanced t h e pheno-
mena of a i r p o l l u t i o n e f f e c t s on r u r a l a r e a s . The
most s t a r t l i n g e f f e c t s discovered s o f a r of t h e
long-range transmission of p o l l u t a n t s , have ap-
peared i n r e l a t i v e l y remote, p r i s i t i n e a r e a s of
Norway, Sweden and t h e Eastern United S t a t e s and
Canada.
Despite t h e f a c t t h a t sulphur dioxide emissions
t o a large extent contribute t o acid precipitation
t h e two p o l l u t a n t s show g r e a t d i f f e r e n c e s i n e f -
f e c t s . Sulphur dioxide i s a primary a i r p o l l u t a n t
a s well a s a primary t o x i c a n t . Acid p r e c i p i t a t i o n
on t h e o t h e r hand, i s a secondary p o l l u t a n t caus-
i n g mainly i n d i r e c t e f f e c t s on ecosystems.
F o r e s t Ecosystems
The e f f e c t s of a i r p o l l u t a n t s on p l a n t s i s ex-
tremely d i v e r s i f i e d ; it depends upon s p e c i e s -
l i n k e d t o l e r a n c e o r s u s c e p t i b i l i t y , and i s a func-
t i o n of many exposure parameters (frequency, time,
concentration, e t c . ) . Many t y p e s of response have
been d e s c r i b e d on t h e b a s i s of l a b o r a t o r y e x p e r i -
ments, where known chemicals were t e s t e d under
c o n t r o l l e d c o n d i t i o n s with d i f f e r e n t p l a n t s p e c i e s .
However, t h e i n v e r s e o p e r a t i o n - namely i d e n t i f y -
i n g and e s t i m a t i n g t h e n a t u r e and importance of
an e x i s t i n g source on t h e b a s i s of response symp-
toms - i s o f t e n d i f f i c u l t , except i n t h e case of
a c u t e i n j u r y and when a p o l l u t i o n source i s known
o r suspected i n t h e v i c i n i t y .
Vegetation damage due t o t h e emission of a c i d
and poisonous substances has long been observed
i n t h e v i c i n i t y of emission sources. V i s i b l e
symptoms have been decribed and a r e o f t e n asso-
c i a t e d with t h e decrease i n growth. Recently,
however, concern has been expressed t h a t f o r e s t
growth may a k o be a f f e c t e d f a r away from emission
sources. Even i f t h e d i r e c t evidence i s meagre,
T a m , (1976), t h e r e i s f a i r l y s u b s t a n t i a l i n d i -
r e c t evidence t h a t continued exposure t o a c i d
r a i n has a growth-decreasing e f f e c t . The most
s i g n i f i c a n t i n d i r e c t evidence i s t h e p o s i t i v e
c o r r e l a t i o n between f o r e s t y i e l d and t h e s o i l
base s t a t u s . Jonsson and Sundberg (1972) c l a s s i -
f i e d a r e a s i n southern Sweden a s r e l a t i v e l y r e -
s i s t a n t t o a c i d r a i n and r e l a t i v e l y s u s c e p t i b l e
t o a c i d r a i n , r e s p e c t i v e l y , and compared t h e
growth t r e n d s i n both a r e a s by measuring annual
r i n g s on increment c o r e s from groups of t r e e s
which were otherwise a s i d e n t i c a l a s p o s s i b l e .
They found a s t a t i s t i c a l l y s i g n i f i c a n t d i f f e r e n c e
and "found no reason f o r a t t r i b u t i n g t h e reduc-
t i o n i n growth t o any cause o t h e r than a c i d i f i -
c a t i o n . " These r e s u l t s however, have n o t been
confirmed by Norwegian r e s e a r c h e r s , Abrahamsen
and o t h e r s , (1976) ; Abrahamsen, (1980), S t r a n d ,
(1980).
When e v a l u a t i n g t h e e f f e c t of a c i d p r e c i p i t a -
t i o n on t h e supply of p l a n t n u t r i e n t s i n a f o r -
e s t , a b a s i s could be t o c o n s i d e r t h e n u t r i e n t
c y c l e i n a t e r r e s t r i a l ecosystem. Plant avail-
a b l e n u t r i e n t s a r e g e n e r a l l y s u p p l i e d t o t h e sy-
stem from two s o u r c e s ; from t h e atmosphere, a s
f o r N and S, and from t h e m i n e r a l s , a s f o r Ca,
Mg, P I K, S, and t h e m i c r o n u t r i e n t s . In natural
systems n o t h a r v e s t e d by man, n u t r i e n t s a r e a l s o
l o s t i n two ways: To t h e atmosphere by v o l a t i l i -
z a t i o n and t o t h e s e a by leaching. Evaluation of
t h e e f f e c t of a c i d p r e c i p i t a t i o n on t h e amount of
p l a n t n u t r i e n t s i n a f o r e s t ecosystem can t h e r e -
f o r e be r e s t r i c t e d t o t h e c o n s i d e r a t i o n of f o u r
p r o c e s s e s ; d e p o s i t i o n from t h e atmosphere, wea-
t h e r i n g , v o l a t i l i z a t i o n and l e a c h i n g from t h e
s o i l . Obviously many p r o c e s s e s i n t h e s o i l and
t h e p l a n t s can a f f e c t t h e a c c e s s i b i l i t y of p l a n t
nutrients. Acid r a i n may a f f e c t some of t h e s e
processes.
Experimental s t u d i e s on t r e e growth i n r e l a -
t i o n t o a c i d r a i n have been conducted i n r e c e n t
y e a r s i n s e v e r a l c o u n t r i e s . A s y e t no conclu-
s i v e evidence of decreased growth has evolved.
On t h e c o n t r a r y , a s l i g h t l y p o s i t i v e growth e f -
f e c t on t h e s e e d l i n g , which was e x p l a i n e d a s a
n i t r o g e n f e r t i l i z e r e f f e c t , was r e p o r t e d by
Wood and Bormann, (1975). Such i n c r e a s e s , though,
a r e l i k e l y t o be temporary, a s d e p l e t i o n of nu-
t r i e n t c a t i o n s through a c c e l e r a t e d l e a c h i n g
should e v e n t u a l l y r e t a r d growth. Experiments on
t h e e f f e c t of a r t i f i c i a l a c i d i f i c a t i o n on f o r e s t
growth under f i e l d c o n d i t i o n s have been c a r r i e d
Out i n Sweden and Norway. The Swedish experiments
have shown t h a t i n c r e a s i n g a p p l i c a t i o n of
d i l u t e H2S04 has s i g n i f i c a n t l y i n c r e a s e d t h e ba-
s a l a r e a growth, Tanun e t a l . , (1980) . The Norwe-
g i a n s t u d i e s c o n s i s t of f i v e f i e l d p l o t e x p e r i -
ments where a r t i f i c i a l r a i n has been produced by
mixing groundwater and H2S04 t o pH values from 6
t o 2. I n one experiment with Scots p i n e , i n c r e a s -
ed h e i g h t and diameter growth was observed i n
1976 and 1977 a t t h e p l o t s s u p p l i e d with 250 mm
of water p e r y e a r of pH 3 , 2.5 and 2. I n 1979
however, t h e most a c i d i f i e d p l o t s showed s i g n i f i -
c a n t l y l e s s growth than t h e o t h e r experiments.
(See Abrahamsen 1980).
The experiments t h u s show i n c r e a s e d growth t h e
f i r s t couple of y e a r s i n t h e a c i d i f i e d p l o t s ,
followed by decreased growth t h e l a s t y e a r . Sim-
i l a r p a t t e r n s , though n o t s i g n i f i c a n t , have been
found i n some of t h e o t h e r experiments. Chemical
analyses of t h e f o l i a g e have revealed t h a t t h e
most l i k e l y e x p l a n a t i o n of t h e i n c r e a s e d growth
i s i n c r e a s e d N uptake. The decrease i n growth
observed i n 1979 might be r e l a t e d t o reduced a-
v a i l a b l i t y of Mg a s t h e f o l i a r c o n c e n t r a t i o n i s
c l o s e t o values g i v i n g v i s u a l d e f i c i e n c y symp-
toms.
Short-term growth r e s u l t s from a c i d i f i c a t i o n
experiments must be t r e a t e d with c a u t i o n . They
i n d i c a t e , however, t h a t t r e e growth may be rea-
sonably s t a b l e when t h e p l a n t - s o i l system i s
s t r e s s e d by a c i d r a i n . Another d i f f i c u l t y t o be
k e p t i n mind i s t h a t p a r t of t h e a c i d i t y of r a i n
i s due t o n i t r i c a c i d o r n i t r o g e n o x i d e s , which
means t h a t t h e p o s i t i v e f e r t i l i z e r e f f e c t of n i -
trogen may p a r t l y o r f u l l y compensate f o r any
harmful e f f e c t s .
T h e o r e t i c a l l y t h e r e might be c a s e s where a c i d -
i t y caused by sulphur oxides i s c o u n t e r a c t e d by
f e r t i l i z e r e f f e c t s , s i n c e sulphur i s an indispen-
sable plant nutrient. However, d e f i c i e n c y i n
sulphur has never been observed i n f o r e s t t r e e s
under n a t u r a l c o n d i t i o n s i n Scandinavia, and con-
sidering the rather t i g h t nutrient circulation
i n t h e f o r e s t ecosystem, it i s n o t l i k e l y t o occur
except p o s s i b l y on very extreme s i t e s .
A number of p o s s i b l e e f f e c t s of a c i d r a i n on
b i o l o g i c a l p r o c e s s e s i n t h e f o r e s t s o i l has been
considered by T a m , (1976) , Abrahamsen, (1980) .
Most f o r e s t s o i l s have a considerable b u f f e r ca-
p a c i t y . Therefore, we may assume t h a t t h e supply
of a c i d i t y , measured e i t h e r a s hydrogen i o n s
o r a s " s t r o n g a c i d " of i n d u s t r i a l o r i g i n , cannot
y e t have a f f e c t e d t h e e n t i r e s o i l p r o f i l e , ex-
c e p t p o s s i b l y i n t h e immediate v i c i n i t y of emis-
s i o n sources. S t i l l , e f f e c t s may be found on
processes o c c u r r i n g i n t h e t o p - s o i l o r on t h e
s u r f a c e of s o i l p a r t i c l e s . S o i l organisms, i n -
cluding r o o t s , i n t h e upper s o i l horizons may
a l s o be a f f e c t e d . Soil respiration, nitrogen
turnover which i s intimately connected t o organic
matter decomposition i n s o i l , n i t r i f i c a t i o n , ni-
trogen f i x a t i o n and nitrogen immobilization a r e
some of the processes apparently affected by in-
creasing s o i l a c i d i t y . There a r e a number of
other biological processes which may be affected
by a change i n s o i l a c i d i t y o r sulphur supply
which have not y e t been studied.
Several comparative and experimental investi-
gations have yielded evidence i n support of the
t h e o r e t i c a l assumption t h a t a c i d i f i e d precipi-
t a t i o n , l i k e any other change i n climate, w i l l
r e s u l t i n changes i n the properties of s o i l . In-
fluences from t h i s changed chemical climate on
s o i l conditions have been indicated through de-
creases i n pH and base s a t u r a t i o n a s well as in-
creased leaching. From the studies performed up
t o now, however, it i s d i f f i c u l t t o draw any de-
f i n i t e conclusions on the time required f o r the
reactions and t h e i r i n t e n s i t i e s . Many s o i l s a r e
f a r from the s t a b l e and mature stage, and it i s
a well-known f a c t t h a t considerable changes due
t o f a c t o r s other than acid p r e c i p i t a t i o n may si-
multaneously be a f f e c t i n g the properties of the
s o i l . The g r e a t variation i n s o i l types and i n
t h e i r s u s c e p t i b i l i t y t o acid p r e c i p i t a i t o n make
detection even more d i f f i c u l t before the expec-
t e d e f f e c t s have become extensive.
The r e l a t i v e significance of strong acids and
associated heavy metals found i n heavily pollu-
t e d areas has not been c l e a r l y established i n
terms of t o x i c e f f e c t s on p l a n t s and s o i l organ-
i s m s . The most serious consequence of regional
a c i d i f i c a t i o n a t currently observed l e v e l s may
be the increased r a t e of leaching of major ele-
ments and t r a c e metals from f o r e s t s o i l s and veg-
e t a t i o n . This i s t r u e f o r t h e f o r e s t ecosystem
and a l s o has a bearing on the aquatic systems
receiving these e f f l u e n t s .
Aquatic Ecosystems
Freshwater bodies i n many areas of northern
Europe and eastern North America, t h a t today l i e
i n and adjacent t o the areas where p r e c i p i t a t i o n
i s most a c i d , a r e threatened by the continued
deposition and f u r t h e r expansion of acid preci-
p i t a t i o n . Many of these bodies of fresh water
a r e poorly buffered and vulnerable t o acid in-
puts. These ecosystems appear fated t o s u f f e r
a c i d i f i c a t i o n and l o s s of f i s h populations. E-
qually a s serious a s damage t o f i s h a r e the l e s s
conspicuous e f f e c t s of the a c i d i f i c a t i o n of
fresh water including changes occurring i n commu-
n i t i e s of aquatic organisms such a s microdecom-
posers, algae, aquatic macrophytes, zooplankton
and zoobenthos.
Water chemistry
The composition of the lakes which a r e dis-
cussed i n connection with a c i d i f i c a t i o n depends
on three principal sources of chemical components
- atmospheric inputs of sea-water s a l t s , atmospher-
i c inputs of acid p r e c i p i t a t i o n and t e r r e s t r i a l
inputs of chemical-weathering products.
Unpolluted, s o f t water lakes a r e generally
d i l u t e solutions of Ca and Mg bicarbonate. The
bicarbonate system c o n s t i t u t e s the main buffering
system i n the water. Lakes i n regions underlain
by highly r e s i s t a n t , carbonate-poor rocks have
lower buffer capacities, and a r e vulnerable t o
the input of acid p r e c i p i t a t i o n . A major number
of the lakes i n Scandinavia f a l l within t h i s ca-
tegory, especially above the postglacial marine
l i m i t , where the bedrock over large areas i s
covered by only t h i n g l a c i a l deposits. A contin-
uous supply of acid substances t o lakes and
streams eventually leads t o the depletion and l o s s
of the normal buffer system. The pH f a l l s t o be-
low 5.0, and sulphate becomes the major anion.
Such lakes have only minimal capacity t o neural-
i z e additional inputs of acid; and new inputs of
acid cause sharp drops i n pH, Wright and Gjessing,
(1976) , Henriksen, (1980) .
Acid p r e c i p i t a t i o n a l s o causes other changes i n
lake water chemistry a s well. The a c i d i c , high
sulphate lakes a l s o have high aluminum concen-
t r a t i o n s . Since p r e c i p i t a t i o n contains very
l i t t l e A l l the A 1 i n the l a k e water must come from
the drainage basins. That has been shown t o be
the case i n investigations conducted on 9 small
drainage basins i n southern Norway. I t has been
shown t h a t l o s s of calcium, magnesium and aluminum
from the basins is p a r t l y due t o natural wea-
thering processes, but a major f r a c t i o n probably
r e s u l t s from the massive inputs of acid precipi-
t a t i o n . In g r a n i t i c basins there i s approximate-
l y equivalence between net H+ input and Ca + Mg +
A 1 output.
The deposition of acid p r e c i p i t a t i o n occurs epi-
sodically. Acid p r e c i p i t a t i o n generally causes
two seasons of increased a c i d i t y i n streams and
rivers - the f a l l , a season of frequent r a i n , and
the spring, when p o l l u t a n t s stored i n the snow-
pack a r e released i n the f i r s t p a r t of snowmelt.
Laboratory and f i e l d studies of polluted snow have
shown t h a t the f i r s t f r a c t i o n s of meltwater due
t o concentration e f f e c t s within the snowpack con-
t a i n higher concentrations of p o l l u t a n t s than the
bulk snow. The f i r s t 30 per cent of the meltwater
contains up t o 70 - 80 per cent of the t o t a l a-
mount of H', ~ 0 and ~ "s ~ ~ ~ .
The episodic deposition of a i r p o l l u t a n t s and
a c e r t a i n temporary accumulation of sulphate i n
the summer r e s u l t s i n major short-term increases
i n the a c i d i t y of lakes and r i v e r s and these
changes are most frequent i n t h e f a l l and spring.
From a biological point of view these periods a r e
often c r i t i c a l because they a r e spawning and hatch-
ing seasons f o r many aquatic organisms.
Regional surveys
A large number of surveys have been conducted
t o give a p i c t u r e of the a c i d i f i c a t i o n of Scan-
dinavian lakes and r i v e r s . A systematic survey
of 155 lakes i n southern Norway was conducted i n
October 1974, and repeated every year since, on
varying number of lakes.
Excess sulphate i s sulphate t h a t does not come
from sea water s a l t s . Although i n some lakes,
t h i s excess sulphate comes from a t e r r e s t r i a l
source i n the drainage basin, the r e g u l a r i t y of
a SE-NW gradient most probably i s due t o chronic
inputs of anthropogenic sulphur through precipi-
t a t i o n and dry deposition. Indeed, the d i s t r i -
bution of excess sulphate i n lakes i s remarkably
similar t o the weighted average concentration of
excess sulphate measured i n p r e c i p i t a t i o n over
southern Norway.
The pH l e v e l s i n the lakes can be largely ex-
plained by inputs of acid p r e c i p i t a t i o n i n the
SE-NW gradient superimposed upon the variations
i n buffer c a p a c i t i e s due t o the geology of the
drainage basin.
Aquatic organisms
Acid s t r e s s on l i f e i n r i v e r s and lakes has
e f f e c t s on a l l stages i n the food chain. Prima-
r y producers communities, l i k e phytoplankton,
a r e simplified by a reduction i n the number of
species. The composition of a community may
a l t e r with a s h i f t t o more acid-tolerant species.
Macrophytic vegetation has been observed t o
change i n a c i d i f i e d lakes. In some lakes Sphag-
num occurs i n dense mats, and epiphytes a r e well
developed. Generally, t h e vegetation i n acidi-
f i e d lakes i s poor i n species.
The same tendency of an acid-tolerant " s h i f t i s
observed i n diatom communities i n 7 locations i n
southern Norway described i n 1949 and r e v i s i t e d
i n 1975. There was an increase i n the proportion
of species which p r e f e r o r require acid water.
In some a c i d i f i e d lakes, an increased accumu-
l a t i o n of organic bottom sediment has been ob-
served, indicating a reduced r a t e of decomposi-
t i o n . There a r e strong suggestions t h a t micro-
b i a l decomposition i s reduced i n acid water and
t h a t slow-acting fungi take over. This w i l l in-
fluence t h e n u t r i e n t exchange with t h e lake sed-
iments.
In the invertebrate freshwater fauna the
same trends a r e observed: a reduced number of
species and a t o t a l reduction i n biomass i n the
acid locations. Experiments on the tolerance
of c e r t a i n crustaceans which a r e important a s
f i s h food (Gammarus l a c u s t r i s and Lepidurus
t i c u s ) have shown a d i r e c t mortality of eggs a t
pH 5.5 o r lower. There a r e a l s o delays i n the
development from stage t o stage of surviving ani-
mals. Exposure below pH 5.5 w i l l k i l l a majority
of adult individuals even a f t e r a short time, 1
- 2 days. In Norwegian lakes, s n a i l s a r e r a r e a t
pH below 5.8 and disappear below 5.2.
In s p i t e of the strong e f f e c t s on f i s h food
organisms, the indications a r e t h a t changes i n
f i s h food supply play a small r o l e i n the elimi-
nation of f i s h from acid r i v e r s and lakes. In-
stead,lack of recruitment seems t o be the dominant
factor. The tolerance against acid water i s low-
e s t i n newly hatched larvae. This f a c t makes the
spring flood a p a r t i c u l a r l y c r i t i c a l time f o r the
f i s h population.
The physiological mechanism o r mechanisms re-
sponsible f o r f i s h death i n acid water a r e not
f u l l y understood,, but it has been well estab-
lished t h a t acid s t r e s s i s accompanied by a f a i l -
ure i n s a l t regulation within t h e f i s h body.
Metabolism and osmotic c e l l regulation seem t o
be affected. I t i s a l s o q u i t e c l e a r t h a t the
s a l t uptake and l o s s i s influenced by the ion
content of the water. Elevated concentrations
of aluminum, manganese, zinc, cadmium, lead, cop-
per, and nickel have frequently been observed i n
a c i d i f i e d lakes. The abnormally high concentra-
t i o n s are apparently due i n p a r t t o d i r e c t depo-
s i t i o n with p r e c i p i t a t i o n a s well a s t o increase
release ( s o l u b i l i t y ) from the sediments. These
metals may represent a major physiological s t r e s s
f o r various aquatic organisms.
Fish population s t a t i s t i c s from nearly 1000
lakes i n southern Norway show t h a t when both pH
i s low (e.9. pH 4.7) and s a l i n i t y i s low (e.g.
K 10 IJS cm) almost a l l lakes a r e empty of f i s h .
A t higher s a l i n i t y (K 20pS/cm) several lakes
have sparce populations and a few even good.
The recent a c i d i f i c a t i o n of freshwater i n
p a r t s of Europe and eastern North-America has
profound impacts on aquatic l i f e . It can be
s t a t e d with r e l i a b i l i t y t h a t a l l trophic l e v e l s
are affected. Of immediate concern t o the people
l i v i n g i n the a c i d i f i e d regions i s the major de-
c l i n e i n f i s h populations. In the four southern-
most counties i n Norway more than half of the
f i s h populations have been l o s t during the 1940 -
1980 period. Today, lakes i n more than 13,000
km2 of south Norway a r e p r a c t i c a l l y devoid of
f i s h , and i n an additional 20,000 km2 the f i s h
stocks a r e reduced. Continued water a c i d i f i c a -
t i o n i s a t h r e a t t o hundreds of lakes s t i l l har-
bouring valuable f i s h populations(Muniz and
Leivestad, 1980).
 Semb, A.
1979. Sulphur emissions i n Europe.
LITERATURE CITED A t m . Env. 12 p. 455-460.

Abrahamsen, G.
1980. Acid p r e c i p i t a t i o n , p l a n t n u t r i e n t s and
f o r e s t growth. I n Proc. I n t e r n a t i o n a l con-
ference on t h e e c o l o g i c a l impact of a c i d
p r e c i p i t a t i o n , Norway, March, 1980.
SNSF,1432 Aas-NLH, Norway ( I n p r e s s )
Abrahamsen, G. , Bjor, K. , Horntvedt, R. and
B. Tveite.
1976. E f f e c t s of a c i d p r e c i p i t a t i o n on coni-
f e r o u s f o r e s t . I n SNSF-project FR 6/76,
SNSF, 1432 as-NL~ Norway
Dovland, H. and A. Semb.
1980. Atmospheric t r a n s p o r t of p o l l u t a n t s .
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I n Proc. I n t e r n a t i o n a l conference on t h e
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 The Impact of Acidic Precipitation and
Heavy Metals on Soils in Relation to
Forest Ecosystems
2

Stephen A. Norton, Denis W. Hanson, and Richard J. Campana

Abstract: Normal terrestrial cycling of metals in eastern

North America and the Pacific Coast states has been altered

by the increasing acidity of precipitation, and associated

heavy metal deposition and mobilization. Pb and chemically

similar metals are accumulating in soils. Al, Ca, K, Mg, and

Mn are being leached from soils. The mobilities of Fe, Zn,

and P vary with site characteristics. Biological recycling

of nutrients by decomposition and uptake is impeded by lowered

pH and elevated levels of toxic metals in soils. Increased

leaching of nutrients in the 0 and A horizonscaused by in-

creased H"1" inputs, decreases percent base saturation and thus

decreases nutrient pools for shallow rooted plants, especially

seedlings. Deeper rooted plants are subjected to elevated,

potentially toxic, concentrations of dissolved metals (e.g.,

A1 and Mn).

In many contemporary forest ecosystems, nutrient availability

is barely adequate for sustained yield with bole harvesting

techniques. Our work indicates that nutrient pools are dimin-

ishing in the northeastern United States, suggesting that de-

creases in forest productivity will occur.

Large areas of the northern hemisphere are re- ranged up to about 8.2 (corresponding to semi-

ceiving precipitation which is more acidic than arid to arid regions where CaC03 dust, or its

would be predicted by equilibration of rain and equivalent, dominatesthe rain chemistry).

atmospheric CO2:

Atmospheric concentrations of CO2 should result

in a pH of about 8.2. In eastern North America,

where vegetation cover minimizes particulate in-

jection into the atmosphere, unpolluted precipi-

tation would probably have a pH of about 5.6.

Associated with the precipitation are numerous

The resulting pH should be about 5.6. This may be metals and plant nutrients (e.g., Na, K, Ca, Mg,

modified by the hydrolysis of particulates or the NO;, NQ, H2P04, Pb, Zn, etc.).

addition of naturally occurring.organic and inor-

ganic acids. Precipitation pH's in the U.S., Precipitation (wet and dry) is one of three

prior to pollution of the atmosphere, probably inputs into the nutrient budgets for forest eco-

systems. The others are chemical weathering of

inorganic soil and nutrient cycling within the

presented at the Symposium on Effects of Air canopy/root space. Both are closely linked to

Pollutants on Mediterranean and Temperate Forest precipitation chemistry.

Ecosystems, June 22-27, 1980, Riverside,

California, U.S.A. Simply put, from a nutritional point of view:

Professor and Chairman of Geological Sciences; Input - Output = Net accumulation of organic
M.S. degree candidate in Geological Sciences; material.

Professor of Botany and Plant Pathology; Univer-

sity of Maine at Orono, Orono, Maine 04469. Output consists of leaching to groundwater (below

the root zone), injection of particulates into the

atmosphere, volatilization of certain elements,

and loss of particulates "downstream", including

harvesting. If the mass balance (above) is posi-

tive for all limiting nutrients, growth and accu-

mulation of organic matter will occur as living

biomass or as soil organic matter. If the mass

balance is negative, growth can occur only so long

as the decreasing reservoirs of organic matter and

nutrients from precipitation and chemical weather-

ing can supply necessary nutrients; organic matter

must decrease. The amount of organic material on

the forest floor and contained in the rooting zone

of the mineral soil is generally of the same order

of magnitude orlarger than the organic material

contained in the above ground biomass. Removal of

this biomass disrupts the recycling of nutrients.

Continued growth of new biomass must occur from

the reservoir within the soil, forest floor, and

from precipitation.

A forest ecosystem can subsist on nutrients de-

livered solely by precipitation, both wet and dry

(Art et al. 1974). However, it is doubtful that

this precipitation-based forest ecosystem could be

harvested periodically and still have sustained

growth. If there are changes in the chemistry of

Figure I~GeneralizedpH (dotted) (NADP 1979,

precipitation, specific inputs/outputs of the

1980; CANSAP 1979),Pb (solid) and Zn (dashed)

forest nutrient budget may be altered so as to

isopleths (Davis and Galloway 1980) for eastern

affect both sustained growth yields and net orga-

U.S. precipitation. Fluxes are g/h/mo.

nic matter accumulation.

area receiving acidified precipitation. Recent

Many forest ecosystems are not forced to sub-

data (NADP 1979; NADP 1980 a,b; CANSAP 1980) shows

sist on atmospheric inputs alone. They receive

a continued decrease in the pH to average values

additional primary inputs of nutrients from chemi-

of less than 4.0 for the "bull's eye"; the eastern

cal weathering of mineral matter. However, in

half of the U.S. and the eastern half of southern

unglaciated areas of the eastern U.S. where chemi-

Canada is receiving precipitation with a pH less

cal weathering dominates over mechanical weather-

than 5.0 (fig. 1).

ing or in glaciated granitic (nutrient poor) ter-

rains characteristic of large areas of eastern

Most of the lowering of the pH of precipitation

North America, nutrient pools are largely con-

from "normal" to present levels occurred over the

tained within the organic litter of the forest

last 40 years. In North America, there are no

floor. Thus, these areas are the most vulnerable

long term regional studies of the acidification

to depletion of nutrient pools due to acidic pre-

of soils showing trends related to regional pH

cipitation.
gradients. Linzon and Temple (1980) (in Ontario)

found acidification of soils over a 16 year ob-

Nutrient availability is commonly classified as

servation period. Many studies have been made of

deficient (where addition of the limiting nutrient

soil acidification and nutrient status in areas

elicits a positive response), adequate (where ad-

heavily affected by point sources.

dition of a nutrient does not elicit a response),

and excessive (where addition of the nutrient eli-

Expected changes in the inorganic aspects of

cits a negative response). This paper investigates

soil chemistry associated with acidification in-

some of the consequences of increased acidity and

clude desorption of metals from organic and inor-

metal availability, associated with acidic precipi-

ganic cation exchange surfaces, increased solu-

tation to nutrient availability (and thus to for-

tion of "mineral" colloids and crystalline min-

est productivity). Excellent reviews of potential

erals, and changes in metal speciation and thus

problems are given by Voigt (1979) and in Hutchin-

biological availability.

son and Havas (1980).

Desorption of metals

EFFECTS OF DECREASED pH

Regardless of the nature of the substrate, ion

Although early precipitation chemistry is sparse

exchange (for a monovalent ion) may be represented

for North America, Cogbill and Likens (1974) con-

as

structed 3 pH isopleth maps spanning 1952 to 1972

which suggested increasing acidity for precipita-

tion in the eastern U.S. and a broadening of the

where X i s a n exchange
m e t a l , H+ i s a p r o t o n ,
specie. This reaction
r e c t i o n by changing H"1'
+
s i t e , R i s any monovalent
and aq r e f e r s t o a n aqueous
can be f o r c e d i n e i t h e r d i -
o r R+ a c t i v i t i e s .

P r e - p o l l u t i o n p r e c i p i t a t i o n i n e a s t e r n North
America e n t e r e d t h e s o i l w i t h a pH probably i n t h e
r a n g e 5 . 5 t o 6.0. M i c r o b i a l a c t i v i t y i n t h e or-
g a n i c l i t t e r produces molecular CO2 which can r e -
duce t h e pH c o n s i d e r a b l y (as low a s 4.5 t o 5 . 0 ) .
The p r o d u c t i o n of v a r i o u s o r g a n i c weak a c i d s (e.g.
f u l v i c and humic) may d e p r e s s t h e pH f u r t h e r t o
4 . 0 t o 4.5. To m a i n t a i n e l e c t r i c a l n e u t r a l i t y i n
t h e s e s o l u t i o n s t o o f f s e t t h e ff4' p r o d u c t i o n , e i -
t h e r a n i o n s must be gained ( H C Oa~c t i v i t y i s r e -
duced by t h e l o w e r i n g of pH; o r g a n i c a n i o n s may
be produced) o r c a t i o n s must be l o s t from t h e so-
l u t i o n . T h i s i s most e f f e c t i v e l y accomplished i n
t h e l i t t e r and a t r o o t s u r f a c e s where H+ i s ex-
changed f o r p e r c o l a t i n g c a t i o n s . Thus n u t r i e n t s
a r e gained by t h e s o i l .

I f t h e p r e c i p i t a t i o n h a s a pH o f 4.0 due t o t h e
s t r o n g a c i d s H2S04 and HN03, t h e exchange r e a c t i o n
i s forced strongly t o the right, stripping cations
( p a r t i c u l a r l y Ca and Mg) from t h e l i t t e r , r e d u c i n g
p e r c e n t of b a s e s a t u r a t i o n . Continued p r o d u c t i o n F i g u r e 2--Location of s o i l l o c a l i t i e s r e p o r t e d i n
of CO2, o r g a n i c a c i d s , by m i c r o b i a l a c t i v i t y as- Table 1 (Hanson 1980) and l o c a t i o n of l a k e s f o r
s u r e s t h a t t h e c a t i o n s a r e l o s t from t h e system. which F i g u r e 4 i s developed. pH i s o p l e t h s a r e
f o r 197516 (Likens e t a l . 1979).
No l o n g term s t u d i e s of s o i l s e x i s t t o demon-
s t r a t e t h e s w i t c h from accumulation t o l o s s o f a l s o suggesting t h a t f o r e s t l i t t e r is being leached
c a t i o n i c n u t r i e n t s a s a r e s u l t of low pH p r e c i p i - before i t i s transported t o t h e l a k e and/or t h a t
tation. I n d i r e c t evidence f o r t h e switch c o n s i s t s l e a c h i n g c o n t i n u e s w h i l e t h e sediment i s i n con-
of l o n g term changes i n s u r f a c e w a t e r q u a l i t y s u c h t a c t with a c i d i c lake water.
a s c o n d u c t i v i t y (Malmer 1976 [ i n wede en] ) , a l k a l i
and a l k a l i e a r t h c o n c e n t r a t i o n s (Malmer 1976), and
t h e commonly observed r e l a t i o n s h i p between non- S o l u t i o n o f Metals from M i n e r a l s o r C o l l o i d s
d y s t r o p h i c low pH w a t e r s and e l e v a t e d Ca, Mg, Al,
Mn, and o t h e r m e t a l s . U l r i c h (1980) and Linzon The r e l e a s e of c a t i o n s from m i n e r a l s due t o
and Temple (1980) have shown l o s s o f b a s e s a t u r a - a c i d i c s o i l w a t e r w e a t h e r i n g may b e r e p r e s e n t e d
t i o n i n s o i l s o v e r 8 and 1 6 y e a r s , r e s p e c t i v e l y .
Abundant e x p e r i m e n t a l e v i d e n c e ( e . g . , Hutchinson
1980; Abrahamsen and S t u a n e s 1980) and s t u d i e s o f Table I ~ C h e m i s t r yo f f o r e s t l i t t e r from h i g h
s o i l s adjacent t o l a r g e point source e m i t t e r s of a l t i t u d e f i r f o r e s t s . Sample s i t e s a r e shown
SOx and NOx s u g g e s t what l o n g t e r m r e s u l t s might on f i g u r e 2. Note: S i t e 6 i s anomalous f o r a l l
look l i k e . Ca, Mg, K, Zn, Cd, and Mn a r e r e a d i l y p a r a m e t e r s . S i t e 1 2 had abundant admixed m i n e r a l
l e a c h e d from l i t t e r . However, t h e pH l e v e l s em- s o i l and t h e bedrock i s v e r y low i n MnO. D e t a i l s
ployed f o r e x p e r i m e n t a l work a r e commonly w e l l be- of c o l l e c t i o n and a n a l y s i s a r e i n Hanson (1980).
low what we might e x p e c t on a r e g i o n a l b a s i s .
P r o c e s s e s o p e r a t i n g a t a pH <4.0 may n o t b e e f f e c -
t i v e even o v e r l o n g p e r i o d s of time a t pH>5.0. CaO MnO
Site A ~ ~ O ?~ 1 . 7 0 3 Pb ( P P ~ ) Zn ( P P ~ )
A t r a n s e c t of " e q u i v a l e n t " s o i l s i t e s a c r o s s a 1 7.62 0.21 189 72
r e g i o n a l pH g r a d i e n t ( f i g . 2 ) , i n e f f e c t , i s a
t i m e s t u d y of t h e e f f e c t s of low pH p r e c i p i t a t i o n .
Table 1 i n d i c a t e s a p r o g r e s s i v e d e c r e a s e i n
and -i n a s o u t h w e s t e r l y d i r e c t i o n , to-
A1203 A1203
ward lower pH p r e c i p i t a t i o n . We i n t e r p r e t t h i s
a s a p r e f e r e n t i a l l e a c h i n g ( d e s o r p t i o n ) of Ca and
Mn from t h e l i t t e r .

Organic m a t t e r i n l a k e sediment i s d e r i v e d from

b o t h t h e w a t e r s h e d and t h e l a k e w a t e r column. O r -
ganic-rich l a k e sediments i n a c i d i f i e d l a k e s i n
New England (Williams 1980) a r e d e p l e t e d o f CaO,
 Overall, acidic precipitation accelerates pod-

solization, expanding depths of upper soil hori-

zons, and depleting thenutrient pool in the upper

soil.

Changes in Speciation of Dissolved Metals

Biological uptake of nutrients and toxicants

is via molecular diffusion through semi-permeable

membranes which are somewhat specific to the spe-

cies involved. Biological response is highly

specific to the dissolved form ("biological avail-

ability") of the element.

Decreasing pH affects speciation in two ways.

Elements which complex with OH groups will be

preferentially partitioned into a less hydroxylated

form. For example:

Lowered pH also results in protonation of weak

acid radicals (e-g. HC03, fulvic , humic ) causing

a decrease in ligands for the cation of interest.

For example:

PH

Figure 3--Eh-pH diagram for the system Al-Te-Mn-

0-H20 at 2 5 " ~
and
1 atmosphere. Solubility in
Because toxicity is generally greater for the

moles/I.
uncomplexed metal (Hg is an exception), acidifi-

cation of soil waters should result in greater

direct toxicity to roots and micro-organisms or

foliage after uptake, and on "downstream" ecosys-

tems.

where R is a metal, n is the valence of the metal,

Xxl is the stoichiometric formula for the mineral

INCREASING TRACE METAL MOBILITY/AVAILABILITY

(minus R), H+ is a proton, and aq refers to aqueous

species. Reactions of this type are forced by

Trace metal levels, particularly heavy metals,

elevated H+ levels. MII+~-, ~e+2-, F~-^-Y and ~ l + ~ - are

intimately affected by acidic precipitation

bearing minerals are somewhat unique in that the

in that their flux to the forest ecosystem is

respective metal's mobility is a function of [@I2

greatly increased over pre-pollution values and

or even [&I3 activity and the transition from geo-

their mobility is in some cases greatly altered.

chemical immobility to mobility under oxidizing

conditions takes place at a pH between 4 and 6 Historic data to evaluate the changing atmos-

(fig. 3).
phericflux of heavy metals in North America is

absent. Changes in fluxes with time in remote

Increased mobilization of A1 from inorganic

regions have been evaluated from snowlice cores

soils has been documented by Cronan and Schofield

(Cragin et al. 1975). Anthropogenic emission

(1979). Both experimental and field studies

rates may be compared with natural emission rates

(Table 1) suggest that Mn is leached from upper

to obtain an estimated percentage increase or

levels of soils. Data are sparse but indicate

mobilization factor. For Cd, Mn, Pb, and Zn the

that acidic non-dystrophic surface waters have

factors are 13, 0.48, 180, and 13 respectively.

elevated Mn levels. A1 and Mn have toxic effects However, these factors may not relate closely to

on plants via root effects, and on aquatic animals.

deposition values (Galloway et al. 1980), because

Fe, although leached somewhat by acidic percolating

of spatially non-homogeneous emission, dispersion,

solutions (Cronan and Schofield 1979), is relatively

and deposition.

enriched in the litter in leaching experiments and

in the field (Hanson 1980). Downstream reduction

Limits on pre-pollution metal concentrations

in concentration of metals may be accomplished by

in precipitation may be established using modern

dilution (Ca, Mg, K), precipitation (Fe, Al, Mn),

data for precipitation chemistry from modern re-

and adsorption (P, Zn). mote sites. Metal levels for eastern North Ameri-

can must have been between the remote (Antarctica)

 MRRRNR GRRNITE SPECK UNNRMED e s t l i t t e r (Table 1 ) a l t h o u g h b i o l o g i c a l l y a v a i l -

a b l e B-horizon accumulation o c c u r s (Conrad, p e r s .
corn.). This appears t o be r e l a t e d t o increased
l e a c h i n g of Zn from l i t t e r due t o d e c r e a s e d p H o f
Zn
p r e c i p i t a t i o n ; l a k e sediments from New England
e x h i b i t s i m i l a r behavior ( f i g . 4 ) . A c i d i f i e d
watersheds have l a k e sediments b e i n g d e p o s i t e d
u

which have l e s s Zn t h a n sediments d e p o s i t e d p r i o r

PPÃ ICN. P M IGN. PPU IGN. t o their acidification.

SUMMARY

Acidic p r e c i p i t a t i o n , e x p e r i m e n t a l l y and em-

PPU IGN. PPH IGh. PPK IGh.
CONC. VS. DEPTH (CM)
F i g u r e A--Pb and Zn p r o f i l e s f o r sediment from a
circum-neutral (Maranacook), s l i g h t l y a c i d i c (pH,
5-6) ( G r a n i t e ) , and a c i d i c (pIK5) (Speck) l a k e
w i t h s u r f a c e i n l e t s and o u t l e t s and an a c i d i c (pH
< 5 ) k e t t l e pond ("unnamed" Pond). Ponds a r e l o -
c a t e d on F i g u r e 2.
and North A t l a n t i c v a l u e s . (We know t h a t Green-
l a n d [ c r a g i n e t a l . 19751 h a s been r e c e i v i n g
p o l l u t e d p r e c i p i t a t i o n f o r approximately 200
y e a r s . ) Modern d e p o s i t i o n r a t e s f o r Pb and Zn a r e
roughly known ( f i g . 1 ) . However, o u r poor know-
l e d g e of p r e - p o l l u t i o n d e p o s i t i o n r a t e s d o e s n ' t
permit a good assessment of t h e i n c r e a s e s t h a t
have o c c u r r e d f o r e a s t e r n North America.
p i r i c a l l y , a c c e l e r a t e s p o d s o l i z a t i o n and d e p l e t e s
n u t r i e n t p o o l s i n f o r e s t l i t t e r and s h a l l o w i n o r -
g a n i c s o i l s . Some heavy m e t a l s (e.g. Pb) accumu-
l a t e t o c o n c e n t r a t i o n s which may impede b i o l o g i c a l
s o i l p r o c e s s e s and immobilize phosphorus. Other
m e t a l s a r e mobilized (Al, Mn, Zn) and may a f f e c t
ecosystems "downstream". Paleolirnnological and
s o i l s d a t a from New England i n d i c a t e t h a t a c i d i -
f i c a t i o n of d r a i n a g e b a s i n s ( i n c l u d i n g s o i l s ) h a s
o c c u r r e d and n u t r i e n t d e p l e t i o n i s underway.
P a r t i a l s u p p o r t f o r t h i s work came from t h e
U.S. N a t i o n a l Science Foundation g r a n t //DEB-78-
10641 t o S.A. Norton; t h e U.S. Dept. of t h e In-
t e r i o r g r a n t #14-31-001-4240, and t h e U.S. Dept.
of t h e I n t e r i o r g r a n t #AO26-W ( O f f i c e of Water
Resources and Research).

Lake sediments r e c o r d changes i n atmospheric

d e p o s i t i o n b u t normally n o t i n a s t r a i g h t forward
manner because of watershed e f f e c t s , sediment fo-
c u s i n g , and d i a g e n e s i s (Norton e t a l . 1980). . F i g -
u r e 4 (unnamed Pond p r o f i l e ) s u g g e s t s a minimum
of a n 800% and 200% i n c r e a s e f o r t h e d e p o s i t i o n
r a t e f o r Pb and Zn, r e s p e c t i v e l y , o v e r t h e l a s t
100 y e a r s . These f i g u r e s assume t h a t t h e back-
ground l e v e l s (below 15-30 cm, depending on t h e
l a k e ) a r e due t o atmospheric d e p o s i t i o n . However,
most of t h e background c o n c e n t r a t i o n s a r e probably
bedrock c o n t r i b u t i o n s . Based on i n c r e a s e s f o r Pb
and Zn c o n c e n t r a t i o n s i n sediments from remote
l a k e s i n New England, we e s t i m a t e t h a t atmospheric
d e p o s i t i o n r a t e s i n New England have i n c r e a s e d a t
l e a s t by a f a c t o r of 30X. P r e - p o l l u t i o n concentra-
t i o n s of heavy m e t a l s i n s o i l s a r e unknown. S h o r t
term s t u d i e s (Siccama e t a l . 1980) of heavy m e t a l s
i n s o i l s i n d i c a t e t h a t concentrations a r e increas-
i n g w i t h time. Our d a t a ( f i g . 2, Table 1 ) shows
a s t r o n g r e l a t i o n s h i p between t h e pH g r a d i e n t and
Pb accumulation, c o n s i s t e n t w i t h t h e Pb d e p o s i t i o n
g r a d i e n t ( f i g . 1 ) . These c o n c e n t r a t i o n s a r e of
concern w i t h r e s p e c t t o t o x i c e f f e c t s f o r s o i l
m i c r o b i a l a c t i v i t y and n u t r i e n t c y c l i n g . High Pb
concentrations i n l i t t e r a r e a l s o considered a s
p o s s i b l e c o n t r o l s on t h e b i o l o g i c a l a v a i l a b i l i t y
of phosphorus i n s o i l s (Cox and Raisons 1972).

However, even w i t h e l e v a t e d d e p o s i t i o n r a t e s
i t a p p e a r s t h a t Zn (and o t h e r elements w i t h s i m i -
l a r chemical b e h a v i o r ) i s n o t accumulating i n f o r -
 LITERATURE CITED
Hutchinson, Thomas C., and M. Havas, eds.

1980. Effects of Acid Precipitation on Terres-

Abrahamsen, Gunnar, and A.O. Stuanes.

trial Ecosystems. 654 p. Plenum Publishers,

1980. Effects of simulated rain on the effluent

New York.

from lysimeters with acid, shallow soil, rich

in organic matter (abs.). Int. Conf. on The

Lazrus, A.L., E. Lorange, and J.P. Lodge, Jr.

Ecological Impact of Acid Precipitation, The

1970. Lead and other metal ions in United

SNSF Project. 1:27.

States precipitation. Envir. Sci. Tech. 4:55-

58.

Art, Henry W., F.H. Bormann, G.K. Voigt, and G.M.

Woodwell.
Likens, Gene E., R.F. Wright, J.N. Galloway, and

1974. Barrier island forest ecosystem: Role of

T.J. Butler.

meteorologic nutrient inputs. Science. 184:

1979. Acid Rain. Scient. Amer. 241:43-51.

60-62.

Linzon, S.N., and P.J. Temple.

Bunzl, K.
1980. Soil resampling and pH measurements after

1974. Kinetics of ion exchange in soil organic

an 18-year period in Ontario (abs). Int. Conf.

matter. I1 Ion exchange during continuous

on The Ecological Impact of Acid Precipitation,

addition of ~bZ+-ions to humic acid and peat.

The SNSF Project. 1:42.

Jour. Soil. Sc. 25: (3).

Malmer, Nils.

Cox, W.J., and D.W. Raisons.

1976, Acid precipitation: chemical changes in

1972. Effect of lime and lead uptake by five

the soil. Ambio. 5:231-234.

plant species. Jour. Envir. Qual. 1:167-169.

National Atmospheric Deposition Program.

Cragin, J.H., M.M. Herron, and C.C. Langway, Jr.

1979. NADP first data report, July 1978 through

1975. The chemistry of 700 years of precipita-

February 1979. Fort Collins, Col. Natural

tion at Dye 3, Greenland: CREEL Research

Resource Ecol. Lab., Col. State University.

Report 341, 18 p.

National Atmospheric Deposition Program.

Cronan, Christopher S., and C.S. Schofield.

1980a,b. NADP Data Report. Fort Collins, Col.

1979. Aluminum leaching response to acid pre-

Natural Resource Ecol. Lab., Col. State Uni-

cipitation: Effects on high-elevation water-

versity, I and 11: (1)&(2).

sheds in the northeast. Science. 204:304-305.

Norton, Stephen A., C.T. Hess, and R.B. Davis.

Davis, Anthony O., and J.N. Galloway.

1980. Rates of accumulation of heavy metals in

1980. Atmospheric trace metal deposition into

pre- and post-European sediments in New Eng-

lakes of the eastern United States. In Input

land lakes. 2 Input of Atmospheric Pollutants
of Atmospheric Pollutants to Natural Waters.
to Natural Waters. Steven J. Eisenreich, ed.

Steven J. Eisenreich, ed. Ann Arbor Science

Ann Arbor Science Publishers, Michigan.

Publishers, Michigan.

Siccama, T.G., W.H. Smith, and D.L. Mader.

Environment Canada.
1980. Changes in lead, zinc, copper, dry weight,

1979. CANSAP data summary. 15 p.

and organic matter content of the forest floor

of white pine stands in central Massachusetts

Galloway, James N., H.L. Volchok, D. Thornton, S.

over 16 years. Envir. Sci. Tech. 14:54-56.

A. Norton, and R. McLean.

1980. Trace metals: a review and assessment.

Ulrich, B.

In Toxic substances in atmospheric deposition.

- 1980. Deposition, production and consumption of

James N. Galloway, S.J. Eisenreich, and B.

hydrogen ions in a beech and spruce ecosystem

Scott, eds. National Atmospheric Deposition

in the Soiling District (abs.). Int. Conf. on

Program, Fort Collins, Col.

the Ecological Impact of Acid Precipitation,

The SNSF Project. 1:50.

Hanson, Denis W.

1980. Acidic precipitation-induced changes in

Voigt, Garth K.

sub-alpine fir forest organic soil layers. M.

1979. Acid precipitation forest ecosystems and

S. unpublished thesis, University of Maine.

intensive harvesting. In Impact of Intensive

Harvesting on Forest ~utrientCycling. p. 33-

Hutchinson, Thomas C.
48. College of Environmental Science and

1980. Effects of acid leaching on cation loss

Forestry, School of Forestry, Syracuse, N.Y.

from soils. In Effects of acid precipitation

on terrestrial ecosystems. T.C. Hutchinson

Williams, John S.

and M. Havas, eds. p. 481-497. Plenum Press,

1980. The relative contributions of local and

New York.
regional atmospheric pollutants to lake sedi-

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lished thesis, University of Maine at Orono.

 Impact of Heavy Metals on Terrestrial

and Aquatic Ecosystems1

2
Tom C. Hutchinson

Abstract: The high t o x i c i t y of many m e t a l s and m e t a l l o i d s

t o a wide range of b i o t a , coupled w i t h t h e i r long r e s i d e n c e
times i n t h e s o i l s , i n sediments and i n t h e oceans has l e d
t o r e a l concern about t h e i r r o l e i n environmental d e t e r i o r a -
t i o n . Residence times i n watersheds a r e commonly measured
i n hundreds of y e a r s , w h i l e r e s i d e n c e of m e t a l s i n a i r i s
r a r e l y a s long a s s e v e r a l days. O v e r a l l i n d u s t r i a l a c t i v i t y
and t r a n s p o r t a t i o n l e a d s t o widespread m e t a l d i s p e r s i o n .
Major e l e v a t i o n s i n many m e t a l s occur around mines and
s m e l t e r s and f o r l e a d e s p e c i a l l y , alongside highways. Coal-
burning and a p p l i c a t i o n s of f e r t i l i s e r s and p e s t i c i d e s add
m e t a l s t o a g r i c u l t u r a l s o i l s and t o n a t u r a l ecosystems. The
s u r f a c e o r g a n i c l a y e r s of b o t h s o i l s and sediments a c t a s
a d s o r p t i o n and exchange s i t e s s o t h a t major accumulations
may occur. Yet, t h i s shallow o r g a n i c l a y e r is t h e c r i t i c a l
s i t e f o r many m i c r o b i a l a c t i v i t i e s , i n c l u d i n g those e s s e n t i a l
f o r n u t r i e n t c y c l i n g , n i t r o g e n f i x a t i o n and f o r pathogens.
Genetic and p h y s i o l o g i c a l t o l e r a n c e s a r e shown i n a wide
a r r a y of d i f f e r e n t organisms which have s u r v i v e d i n metal-
s t r e s s e d h a b i t a t s . 'Most r e c e n t l y , a c i d p r e c i p i t a t i o n has
mobilised A l , Mn, Fe and Zn from t h e s o i l and sediment.
These a r e now producing p a r t i c u l a r s t r e s s e s f o r a q u a t i c b i o t a .

The c o n s i d e r a b l e t o x i c i t y of many m e t a l s t o systems a r e t h e r e c i p i e n t s of i n c r e a s i n g quanti-

b i o t a i s w e l l known e.g. l e a d , mercury, cadmium,
a r s e n i c . The q u a n t i t i e s mined and smelted f o r
innumerable uses c o n t i n u e s t o r i s e each y e a r . It
is a p p a r e n t t h a t b o t h n a t u r a l and man-made eco-
p r e s e n t e d a t t h e Symposium on E f f e c t s of Air
P o l l u t a n t s on Mediterranean and Temperate F o r e s t
Ecosystems, June 22-27? 1980, Riverside,
C a l i f o r n i a , U.S.A.
~ r o f e s s o rof Botany and F o r e s t r y , Chairman,
Dept. of Botany, U n i v e r s i t y of Toronto, Toronto,
Ontario, Canada. M5S 1 A 1 .
t i e s of t h e s e elements and t h a t we have t o be
continuously v i g i l a n t t o ensure t h a t we do n o t
e i t h e r poison o u r s e l v e s , our f e l l o w b i o t a , o r our
a g r i c u l t u r a l and n a t u r a l ecosystems. Major human
poisonings have occurred such a s t h a t by mercury
i n Japan and i n I r a q , and t h a t of cadmium i n
Japan. The beer-deaths i n Birmingham, England a t
t h e t u r n of t h e century were a l s o b e l i e v e d t o b e
metal-related, being variously ascribed t o arsenic
and/or selenium. Concern h a s a l s o been expressed
about n o t only t h e p o t e n t i a l f o r food c h a i n con-
tamination l e a d i n g t o man b u t a l s o t o d i r e c t a e r i a l
i n p u t s t o man v i a t h e r e s p i r a t o r y t r a c t . Lead
from automobile emissions where i t i s used a s an
anti-knock i n g a s o l i n e , and from primary and
secondary s m e l t e r s , a s w e l l a s many o t h e r s m e l t e r -
emitted metals such as a r s e n i c , copper, n i c k e l ,
zinc, cadmium, antimony and selenium, a l l have
p o t e n t i a l l y harmful consequences due t o t h e i r
p e r s i s t e n c e i n t h e body, and t h e i r a b i l i t y t o
i n t e r f e r e with s p e c i f i c enzyme systems. It
should a l s o be noted t h a t many instances a r e
known of s y n e r g i s t i c and a n t a g o n i s t i c i n t e r a c t i o n s
between metals, both i n a q u a t i c and t e r r e s t r i a l
systems and i n t h e human body. Notable amongst
these a e t h e a m e l i o r a t i v e e f f e c t s of selenium
f
and a r s e n i c on both mercury and cadmium t o x i c i t y
i n mammals and t h e r e c e n t l y described s y n e r g i s t i c
e f f e c t s of ozone on cadmium and n i c k e l t o x i c i t y
i n c e r t a i n crop p l a n t s , e.g. Parizek 1978,
Levander 1977, Groth and o t h e r s 1973, Czuba and
Onnrod 1974. Nickel and copper synergisms Rave
been described f o r a v a r i e t y of b i o t a , including
freshwater u n i c e l l u l a r a l g a e , f l o a t i n g a q u a t i c
p l a n t s , and t r e e s e e d l i n g s , Hutchinson 1973,
Hutchinson and Stokes 1975, Hutchinson and Czyrska
1972, and Hutchinson and Whitby 1974.
Despite what sometimes seems t o be a gloomy
p i c t u r e with r e s p e c t t o metal accumulations i n
the environment, i t ought t o be borne i n mind
t h a t concentration of a i r b o r n e p a r t i c u l a t e s i n
urban and i n d u s t r i a l a r e a s of Europe and North
America have o f t e n been much worse i n t h e p a s t .
Cohen and Ruston (1925) r e p o r t e d very high
a r s e n i c l e v e l s i n t h e a i r of Leeds, England i n
1902-1910 due t o c o a l burning, w h i l e t h e o v e r a l l
l e v e l s of SO2 and a c i d i c a e r o s o l s were much
h i g h e r than p r e s e n t l y occur.
Residence Times and Watershed Loss
Rather l i t t l e a t t e n t i o n has been paid t o the
residence times of metals i n components of t h e
biosphere. The s t r o n g r e t e n t i o n of metals on
the organic s u r f a c e l a y e r s of the s o i l and of t h e
sediments is of g r e a t importance, i n t h a t i t
causes long residence times a s w e l l a s i n allow-
i n g accumulation of metals t o p o t e n t i a l l y t o x i c
concentrations. Since i t i s p r e c i s e l y i n these
s u r f a c e zones t h a t t h e major populations of
microbes a r e l o c a t e d , and where the e s s e n t i a l
processes take p l a c e of decomposition of organic
m a t t e r , of n i t r o g e n f i x a t i o n and of elemental
cycling e t c . Thus, t h e p e r s i s t e n c e and
accumulations of t o x i c elements i s of r e a l
concern. L i t t e r i n urban a r e a s contains e l e v a t e d
l e a d concentrations.
The r e t e n t i o n of t h e metals themselves i s an
exchange process, with the elements behaving i n
a reasonably p r e d i c t i v e way, based on such
p r o p e r t i e s a s i o n i c r a d i u s and e l e c t r o n e g a t i v i t y .
Acid leaching can cause a downward movement of
heavy metals through t h e p r o f i l e , s o t h a t they
may e n t e r ground water o r watershed streams.
(e.g. Abrahamsen, Stuanes and Bjor 1979, Cronan
and Schofield 1979, Hutchinson 1980, Bacon and
Maas 1979 .) Both rock s u r f a c e s and s o i l s can
c o n t r i b u t e and such c l a y -mineral c o n s t i t u e n t s a s
aluminium, manganese, z i n c and f e r r i c i r o n e n t e r
drainage water i n t h i s way. Sediments can
s i m i l a r l y l o s e these same elements t o t h e water
bodies of l a k e s under a c i d i f y i n g conditions
(Schindler and o t h e r s 1980). The damaging e f f e c t
of t h e r e s u l t a n t aluminium concentrations t o f i s h
have been described by various a u t h o r s , including
Schofleld 1976, Baker and Schofield 1980, a t l e v e l s
as low a s 0.1-0.2 mg/1. Increased t r a n s p o r t of
aluminium I n t o a q u a t i c systems can a l s o a f f e c t
phosphorus a v a i l a b i l i t y (Cronan and Schofield 1979).
The residence time of metals i n the a i r i s
always very much s h o r t e r than t h a t i n s o i l , w a t e r ,
sediments o r oceans. This i s I l l u s t r a t e d by Table
1, which emphasises t h e r a t h e r l a r g e r residence
time of l e a d i n a i r than t h a t of a wide range of
o t h e r m e t a l s . This p a r t l y explains t h e e l e v a t i o n s
of l e a d noted a t remote l o c a t i o n s , such a s i n t h e
a r c t i c , i n g l a c i a l i c e i n Greenland and a t moun-
t a i n tops i n C a l i f o r n i a (see National Academy of
Sciences Lead Review 1980). It i s a l s o a f u n c t i o n
of p a r t i c u l a t e s i z e and p a r t i a l vapour pressure.
Table 1. Residence times1 of metals i n t h e atmos-
phere a t La J o l l a and Ensenada, from Hodge,
Johnson and Goldberg, 1978.
days
L a Jolla ' '1' Ensenada
7 8
0.7 0.5
0.2 0.1
0.4 0.3
0.5 1
3 0.8
1.2 0.2
1.0 0.4
0.8 0.2
0.8 0.4
0.6
1.0 0.2
5 ---
1 ---
Standing rop of metals on p a r t i c u l a t e s i n
3
1,000 m X 1 an column of a i r f i l t e r data--Table
$
1 ) divided by the f l u x t o 1 cm of ground s u r f a c e
("bucket d a t a --Table 2 ) . F i l t e r d a t a averaged
over period during which buckets exposed. F i l t e r
concentrations of Co, Fe, Mn, C r and A l h a v e been
m u l t i p l i e d by 2 i n order t o account f o r t h e
d i s c r i m i n a t i o n a g a i n s t l a r g e p a r t i c l e s by t h e H i -
Vol sampler.
The c o n t r a s t i n g d a t a f o r watershed s o i l s a r e
i l l u s t r a t e d by Table 2, taken from Bowen 1975, and
from which i t is apparent t h a t soil-watershed
residence t i m e s a r e measured i n hundreds of y e a r s .

Table 2 . Inputs and outputs i n mg X/m y r , and
residence times i n years, f o r nine elements i n
s o i l s of the Upper Thames b a s i n .
Rain Fertilizer Rock
X input input input
Clearly t h e p o t e n t i a l f o r accumulation t o t o x i c
l e v e l s i s much g r e a t e r . The excess of l e a d and
chromium i n i n p u t over output i s a f e a t u r e of
systems s u b j e c t e d t o i n d u s t r i a l deposition. The
v o l a t i l i z a t i o n of some of these heavy metals
from t h e f o l i a g e of v e g e t a t i o n e.g. zinc, mercury,
and selenium (Beauford and o t h e r s 1975) may
re-mobilise s m a l l q u a n t i t i e s of t h e s e metals and
i n c r e a s e atmospheric r e s i d e n c e times b u t i t w i l l
n o t i n f l u e n c e l o s s i n t o drainage waters. Allen
and Steinnes (1979) determined t h e r e g i o n a l
d i s t r i b u t i o n of l e a d , z i n c , cadmium, copper,
a r s e n i c , antimony and selenium i n Norwegian
s u r f a c e s o i l s , u t i l i s i n g 500 humus samples. Lead
l e v e l s were 10-fold h i g h e r i n t h e south than i n
t h e a r c t i c a r e a s and a l s o h i g h e r along t h e c o a s t
than i n l a n d . Cadmium, a r s e n i c , antimony and
selenium showed a s i m i l a r north-south trend. A l l
of these elements a r e h i g h l y v o l a t i l e , low b o i l i n g
p o i n t components of t h e atmospheric load from
i n d u s t r i a l and urban c e n t r e s .
Residence times i n s o i l water a r e a f f e c t e d t o
a g r e a t e x t e n t by pore s i z e . The water i n t h e
l a r g e pores i n f i l t r a t e s i n t o lower l a y e r s and a i r
e n t e r s a g a i n behind i t . The residence t i m e i n
these pores i s not more than hours. Meanwhile,
t h e w a t e r i n t h e narrow pores i s displaced only
centimetres o r m i l l i m e t r e s . An example i s shown
i n Table 3 from t h e work of F r i s s e l (19781, w i t h
residence times a s high a s 5000 y e a r s . The
i m p l i c a t i o n s f o r ground water contamination a r e
apparent.
Targets f o r H
+ and Heavy Metals
I should l i k e t o emphasize t h a t one very
u s e f u l way of considering t h e p o t e n t i a l t h r e a t s
t o ecosystems i s through a c o n s i d e r a t i o n of t a r -
g e t s i n the ecosystem. Clearly, a l l s u r f a c e
i n t e r f a c e s f a l l i n t o t h i s category. Surfaces
p r e s e n t a r e a s of p o t e n t i a l accumulation o r
,
residence. I n t e r r e s t r i a l ecosystems including
z
Drainage Cropping Residence time/
output output years
a g r i c u l t u r a l ones, the s u r f a c e of the l e a f i s one
such i n t e r f a c e . Higher p l a n t s a r e covered by a
r a t h e r impermeable waxy c u t i c l e , which reduces
g a s and water flow t o a minimum b u t i s p e r f o r a t e d
by numerous stomata, o f t e n on t h e underside of t h e
l e a f e s p e c i a l l y . While p a r t i c u l a t e s can accumulate
on such a s u r f a c e , they a r e a l s o r a t h e r e a s i l y
washed off by r a i n o r blown o f f by wind.
Frequently, however, small p a r t i c l e s can b e
incorporated i n t o the c u t i c l e o r e n t e r the stomata.
The l e a f s u r f a c e of many p l a n t s a l s o a r e covered
By numerous f i n e branched h a i r s o r glands. These
can a c t as t r a p s f o r p a r t i c u l a t e s s o t h a t very
d i r t y l e a f s u r f a c e s can occur i n a r e a s of h i g h
d u s t f a l l . Nevertheless, the d i r e c t l y t o x i c e f f e c t s
of p a r t i c u l a t e metal c o n s t i t u e n t s a r e l i m i t e d a s
they a r e k e p t away from m e t a b o l i c a l l y a c t i v e s i t e s .
I n the mosses, l i v e r w o r t s and l i c h e n s , the
c u t i c l e i s e f f e c t i v e l y absent. The exposed c e l l
w a l l s u r f a c e i s a t t h e a i r i n t e r f a c e and i t
c o n s i s t s of charged s i t e s , which can exchange both
c a t i o n s and anions. The metals a r e s e l e c t i v e l y
exchanged onto t h i s s u r f a c e and a r e h e l d t h e r e .
Large accumulations can take p l a c e . Lichens i n
p o l l u t e d regions a t t e s t t o t h i s , a s do mosses
Ce.g. RUhling and Tyler 1970) . The use of
Sphagnum moss bags a s a i r monitors is based on
t h i s c a t i o n exchange capacity. The s p e c i a l s e n s i -
t i v i t y of -many l i c h e n s t o a i r p o l l u t a n t s is due
t o t h e ready e n t r y of t h e p o l l u t a n t t o metabolically
active s i t e s .
S o i l Surface Layers a s Target Areas a t Risk
The s u r f a c e of t h e s o i l a s a c r i t i c a l s i t e f o r
accumulation of a i r b o r n e m e t a l l i c contaminants has
already Been r e f e r r e d t o h e r e . It is i n t h i s
upper few centimetres of t h e s o i l t h a t n u t r i e n t
uptake i n t o p l a n t r o o t s takes p l a c e and i n which
new r o o t h a i r s a r e developed. Seeds germinate i n
t h i s l a y e r amongst t h e f o r e s t l i t t e r and s e e d l i n g s
e s t a b l i s h t h e r e . The m i c r o b i a l sequences e s s e n t i a l
 Table 3. Residence times of water i n t h e s a t u r a t e d
zone of t h e s o i l .

Discharge System Estimated r e s i d e n c e time Watershed

per year and pathways depending on t h e p l a c e of
bud i n f i l t r a t i o n and on p o r o s i t y
of t h e s o i l o r rock

About c o n s t a n t 100-500 y e a r s Merkenf r i t z b a c h

slow discharge 17 ( F e d e r a l Republic of
p a r t l y 12 Germany, Land
slow discharge 1 2 1-500 years Hessen, Main a r e a )
f a s t discharge 10 1 dayÑ1 y e a r s 1600 h a
p a r t l y 12

10Ñ100 y e a r s Okkenbroek
1 d a y ~ 1 0y e a r s (The Netherlands,
1 h o u r ~ dl ay I J s s e l area)
443 h a

f o r e f f e c t i v e l i t t e r decomposition t a k e p l a c e , i n copper a r e now h i g h , abnormal accumulations of

t h i s zone pathogens s t r i v e t o i n f e c t s e e d l i n g s o r
r o o t systems, and t h e mycorrhizal f u n g i e s s e n t i a l
f o r e f f e c t i v e n u t r i t i o n of many f o r e s t t r e e s ,
e s p e c i a l l y i n t h e b o r e a l f o r e s t , develop h e r e .
The r h i z o b i a l b a c t e r i a which a c t a s n i t r o g e n
f i x e r s i n legumes and t h e actinomycetes and blue-
greens which f u l f i l l t h i s r o l e i n o t h e r shrubs
and g r a s s e s a l s o have t o i n f e c t r o o t s i n t h e s e
upper few c e n t i m e t r e s of t h e s o i l . Yet, onto t h i s
s u r f a c e is b e i n g d e p o s i t e d i n c r e a s i n g loads of
t o x i c heavy m e t a l s , of a c i d i f y i n g substances and
a l s o of gaseous p o l l u t a n t s . The t h r e a t t o t h e
s a f e f u n c t i o n i n g of such ecosystems and t o t h e
well-being of man a r e focused on t h i s zone.
Indeed, we can c o n s i d e r t h a t t h e r e d u c t i o n o r
e l i m i n a t i o n of j u s t a few key processes could p u t
t h e whole system a t r i s k . The enzyme a r y l
s u l p h a t a s e which produces t h e p l a n t - a v a i l a b l e
s u l p h a t e from t h e non-available o r g a n i c s u l p h u r
i n s o i l s i s one such s t e p , and i t i s known t h a t
soil-extracted a r y l sulphatases a r e susceptible
t o a wide range of heavy metals i n c l u d i n g
aluminium. The a b i l i t y of r h i z o b i a l b a c t e r i a l
t o i n f e c t legume r o o t s i s a l s o known t o be a c i d
s e n s i t i v e and heavy metal s u s c e p t i b l e . The condi-
t i o n s f o r s e e d l i n g e s t a b l i s h m e n t might be a f f e c t e d
i f t h e atmospheric i n p u t s of wet and dry d e p o s i t i o n
were t o a c i d i f y s u r f a c e s o i l s s o a s t o favour f u n g i
a t t h e expense of b a c t e r i a .
The t h r e a t t o l i t t e r decomposition may b e a
long time i n developing i n most a r e a s b u t i n those
where i n t e n s e heavy metal accumulations have
occurred from s m e l t e r emissions, examples of t h i s
have a l r e a d y been demonstrated. I n both t h e
remnant f o r e s t i n t h e major Sudbury s m e l t i n g a r e a ,
where n i c k e l and copper c o n c e n t r a t i o n s have reached
up t o 2000 ppm i n t h e p a s t and i n t h e New Lead
B e l t of Missouri, where l e a d , z i n c , cadmium and
l i t t e r on t h e f o r e s t a r e r e p o r t e d (Freedman and
Hutchinson 1980, Watson and o t h e r s 1976). A t
t h e z i n c s m e l t e r of Palmerton i n Pennsylvania,
S t r o j a n (1978) r e p o r t e d reduced decomposition of
t h e f o l i a g e of a number of s p e c i e s and a s c r i b e d
i t t o e l e v a t e d z i n c and cadmium c o n c e n t r a t i o n s .
The r e p o r t s have a l s o included d e t r i m e n t a l
e f f e c t s on a number of s o i l enzyme a c t i v i t i e s , on
o v e r a l l m i c r o b i a l r e s p i r a t o r y a c t i v i t y and on
micro-arthropod and earthworm a c t i o n . Indeed,
many r e p o r t s a r e now a v a i l a b l e which show s e n s i -
t i v i t y of earthworms t o heavy metal accumulations.
Sediments a s S i t e s of Risk
I t should b e noted t h a t a r a t h e r s i m i l a r b u t
p a r a l l e l c a s e can b e made f o r e f f e c t s on s u r f a c e
sediments. Again, i n t h i s zone, much of t h e
microbial a c t i v i t y takes place, the aquatic p l a n t s
have t o r o o t , t h e b e n t h i c organisms l i v e and
reproduce and t h e important gas exchanges t a k e
p l a c e w i t h t h e w a t e r column. This i s t h e zone of
d e p o s i t i o n of t h e dead p l a n k t o n i c organisms, of
incoming p a r t i c u l a t e m a t t e r and of p o l l u t a n t
m a t e r i a l e q u a l l y . The sediments a r e o f t e n h i g h l y
organic and have a l a r g e c a t i o n exchange c a p a c i t y .
Toxic l e v e l s can develop (mercury i n p o l l u t e d
sediments of t h e D e t r o i t River and Lake S t . C l a i r ) ,
and benthic-feeding f i s h , clams, c r a y f i s h e t c .
come i n c o n t a c t w i t h p e s t i c i d e s , PCB's and heavy
m e t a l s which have been i n i t i a l l y t r a n s p o r t e d by
a i r . The methylation t r a n s f o r m a t i o n which c r e a t e s
o r g a n i c mercury compounds n i n e t o t e n times more
t o x i c than i n o r g a n i c e q u i v a l e n t s take p l a c e h e r e
i n t h e sediment s u r f a c e l a y e r s .
 Reproduction: c r u c i a l s t e p s a t r i s k
A number of e s s e n t i a l s e q u e n t i a l s t e p s can b e
i d e n t i f i e d , t h e i n t e r f e r e n c e with which places t h e
whole reproductive process a t r i s k . These include
a ) the h e a l t h of p o l l i n a t o r s , e s p e c i a l l y the
i n s e c t s e s s e n t i a l f o r those p l a n t s with s p e c i a l i s e d
mechanisms and a p p r o p r i a t e f l o r a l guides, t r i p
mechanisms, n e c t a r production, p o l l e n p o s i t i o n i n g
e t c . , b) t h e a b i l i t y of the p o l l e n t o germinate
s u c c e s s f u l l y on t h e stigma and t o then success-
f u l l y produce a p o l l e n tube which can reach t h e
u n f e r t i l i s e d ovules c ) t h e a b i l i t y of sperm t o
s u c c e s s f u l l y reach and f e r t i l i s e t h e egg d) the
a b i l i t y of p l a n t s and animals t o s u c c e s s f u l l y
d i s p e r s e t h e i r progeny t o s u i t a b l e new h a b i t a t s .
I n t e r e s t i n g l y , i t i s known t h a t s t e p s a-c
can a l l b e a f f e c t e d by e l e v a t e d l e v e l s of heavy
metals. For example, bees a r e known t o b e very
s u s c e p t i b l e t o a i r b o r n e a r s e n i c . I n the region
of t h e Novatny power s t a t i o n i n Czechoslovakia
bee h i v e s have been wiped out. The power-station
burns c o a l with high l e v e l s of a r s e n i c i . e .
s e v e r a l hundred ppm, and consequent elevated a i r
l e v e l s of As203 occur. Bees a l s o w i l l pick up
and accumulate selenium when p o l l i n a t i n g high
selenium p l a n t s , such a s some of t h e loco-weeds
.
(As t r a g a l u s s p e c i e s )
The s u c c e s s f u l growth of t h e p o l l e n tube i s
r e p o r t e d t o b e pH-dependent and a l s o t o b e
inÂluenced by t h e presence of t o x i c heavy metals
such a s z i n c and copper. The f e r t i l i z a t i o n of
f e r n archegonia has been r e p o r t e d t o b e i n h i b i t e d
by acid s o l u t i o n s and a c i d r a i n , L. Evans,
(personal communication) . L. S c h l i c t e r , a
graduate s t u d e n t i n botany a t University of Toronto,
has r e c e n t l y shown t h a t t h e s u c c e s s f u l post-
f e r t i l i z a t i o n s t e p s i n embryo development of f r o g
eggs i s i n h i b i t e d by even q u i t e minor decreases
i n pH below 6.0 and t h a t m u l t i p l e - f e r t i l i z a t i o n of
an egg, which normally a r e precluded, can occur
under t h e s e a c i d i c conditions. The consequence
i s an e a r l y aborted embryo. E f f e c t s on t r o u t
eggs, on New J e r s e y f r o g s and on reproductive
success i n planktonic crustaceans have been
reported by a number of workers, e.g. Krishna
(1953), Gosner and Black (1957), Havas (1980).
Other Factors which Inf h e n c e t h e
Outcome of Metal Impacts
I t does seem t o b e t h e c a s e t h a t the damage
t o ecosystems from a i r b o r n e metals, a s w e l l a s
from gaseous p o l l u t a n t s , i s g r e a t e s t when t h e
i n d i v i d u a l s i n t h e ecosys tem a r e metabolically
most a c t i v e . Thus, t h e damage t o f o r e s t eco-
systems i n temperate zones i s c l e a r l y g r e a t e s t
i n t h e summer growing season. Damage during t h e
day when stomata a r e open Is g r e a t e r than a t n i g h t
when they a r e closed. The l i c h e n s and mosses a r e
most s u s c e p t i b l e when they a r e moist and photo-
s y n t h e s i z i n g a c t i v e l y . I n dry o r a r i d h a b i t a t s ,
such a s t h e d e s e r t s of Arizona, t h e damage from
l a r g e smelter-emitted SO2 and copper p a r t i c u l a t e s
i s minimal i n c o n t r a s t t o t h a t of w e t t e r a r e a s
such a s Palmer ton Pennsylvania, Ducktown
Tennessee o r Sudbury, Ontario. This probably
p a r t l y r e l a t e s t o metabolic a c t i v i t y including
t h e percentage of time stomata remain open, b u t
a l s o t o the a c t i v i t y of the r o o t systems. I f one
follows through with t h i s g e n e r a l i z a t i o n , then we
can p r e d i c t t h a t a r c t i c regions w i t h very s h o r t
growing seasons and very long dormant periods
w i l l b e l e s s a f f e c t e d by e q u i v a l e n t metal pollu-
t a n t inputs ( o r SO?, O3 o r F l i n p u t s ) than would
Temperate o r e s p e c i a l l y Tropic Rain Forest
systems. Equally, one can p r e d i c t t h a t the more
a r i d an a r e a , t h e l e s s s u s c e p t i b l e t h i s a r e a ' s
vegetation w i l l b e t o t o x i c damage.
The importance of t h e longevity of t h e
i n d i v i d u a l a l s o needs t o be s t r e s s e d . Damage t o
long-lived trees- may t a k e a long time t o become
apparent and f i n a l l y perhaps only through t h e i r
i n a b i l i t y t o reproduce. Equivalent reproductive
f a i l u r e i n a n annual w i l l obviously be very
r a p i d l y apparent. Metal s t r e s s e s o f t e n seem t o
favour p e r e n n i a l p l a n t s w i t h l a r g e l y v e g e t a t i v e
reproduction, such a s grasses and sedges. The
equivalent a q u a t i c examples would be t h a t of
f i s h compared with planktonic a l g a e o r crustaceans.
F i n a l l y , i t must b e emphasised t h a t t h e
i n i t i a l s e n s i t i v i t i e s of t h e s p e c i e s , populations
and i n d i v i d u a l s of an a r e a when f i r s t s u b j e c t t o
m e t a l s t r e s s , a r e n o t t h e f i n a l response. While
pre-adaptation o r p r e - s e n s i t i v i t y may allow a n
i n i t i a l s e l e c t i o n and s o r t i n g , t h e s t r e s s e d
environment r e p r e s e n t s a changing h a b i t a t i n
which evolutionary change occurs. The occurrence
of m e t a l - t o l e r a n t g r a s s e s on mine waste s i t e s i s
w e l l known. The a b i l i t y of some of these g r a s s e s
t o evolve multiple-metal tolerances and co-
tolerances is now a l s o r e c e i v i n g a t t e n t i o n
(Tatsuyama- and o t h e r s 1975, Cox and Hutchinson
1980J. Even i n h a b i t a t s such a s t h e Smoking H i l l s
of a r c t i c Canada, where pond water pH's may reach
a s low a s 1.8 and s o i l pH's t o < 3.0 from t h e
i n i t i a l v a l u e s of pH > 7.0, some organisms do
survive. These even have the a b i l i t y t o t o l e r a t e
t h e high a c i d i t y , low n i t r o g e n and phosphorus
a v a i l a b i l i t y and t h e extremely e l e v a t e d l e v e l s of
normally t o x i c metals such a s aluminium, manganese,
f e r r i c i o n and zinc, Hutchinson and o t h e r s (1978).
A t t h e extreme, some organisms seem t o have
evolved s p e c i f i c a l l y on a r e a s of very elevated
metal l e v e l s . The occurrence of a legume Becium
homblei on copper m i n e r a l i z a t i o n s i s an example
of t h i s , i n which the p l a n t accumulates enormous
l e v e l s of copper and a l s o r e q u i r e s concentrations
which would b e l e t h a l t o o t h e r p l a n t s ( R e i l l y
1967).
While t h i s allows a c e r t a i n re-assurance about
t h e a b i l i t y of l i f e to t h r i v e under even most
adverse conditions, i t does n o t a t a l l i n f l u e n c e
t h e f a c t t h a t we must b e extremely concerned w i t h
t h e a c c e l e r a t i n g l i b e r a t i o n of heavy metals t o o u r
a g r i c u l t u r a l and n a t u r a l environments. Economic
and s o c i a l p r e s s u r e s do compel us t o t a k e s t o c k .
Proposals f o r d i s p o s a l of metal-contaminated
sewage s l u d g e on farm o r f o r e s t l a n d s have a l r e a d y
caused agencies and s c i e n t i s t s t o t h i n k i n a much
longer time frame than we a r e used t o do.
Acknowledgments: I wish t o thank D r . A 1 Page
of t h e S o i l Science Dep., U n i v e r s i t y of C a l i f o r n i a ,
R i v e r s i d e , f o r most u s e f u l d i s c u s s i o n s i n connec-
t i o n w i t h t h i s paper.
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metals and r a d i o n u c l i d e s from t h e sediments
of a f r e s h w a t e r l a k e . Can. J. F i s h . Aquatic
s c i . 37:373-377.
C.L. S c h o f i e l d . 1976. Acid p r e c i p i t a t i o n :
e f f e c t s on f i s h . Ambio. 5-6: 228-230.
S t r o j a n , C.L. 1978. F o r e s t l e a f l i t t e r
decomposition i n t h e v i c i n i t y of a z i n c
s m e l t e r . Oecologia 32: 203-212.
Tatsuyama, K., H. Egawa, H. Yamamoto and
H. Senmaru. 1975. Tolerance of cadmium
r e s i s t a n t microorganisms t o o t h e r m e t a l s .
Trans. Mycol. Soc. Jap. 16: 79-85.
Watson, A.P., R.I. Van Hook, D.R. Jackson and
D.E. Reichle .
1976. Impact of a lead-
mining complex on t h e f o r e s t - f l o o r
arthropod fauna i n t h e New Lead B e l t
r e g i o n of S.E. Missouri. Oak Ridge N a t l . Lab.
Environ. S c i . Div. Publ. 881. 1 6 1 pp.
 Effects of Acidic Precipitation on Health
and the Productivity of Forests1

Ellis B. Cowling and Leon S. Dochinger2

Abstract: Acid precipitation has become a dominant feature

of man-induced change in the chemical climate of the earth.

But acid precipitation is only one special feature of the

changing chemistry of atmospheric deposition in developed

and developing regions throughout the world. In recent

decades, human activities (mainly increased combustion of

fossil fuels and decomposition or combustion of waste

products) have greatly increased the total emissions and

deposition of beneficial nutrients and injurious substances

(such as strong mineral acids) from the atmosphere. Pro-

jected increases in the use of fossil fuels, and especially

in the use of coal, will add still further to the total

burden of beneficial and injurious substances deposited on

forest and rangeland ecosystems from the atmosphere. The

purpose of this brief paper is to summarize certain impor-

tant principles concerning the phenomena of acid precipi-

tation and atmospheric deposition and their beneficial and

detrimental effects on the health and productivity of

forests.

The supply of both beneficial nutrientelements transpiration and the maintenance of cell

and injurious substances in the atmosphere influ- turgor.

ence the health and welfare of forests. Plant

life as we know it would be impossible without Some epiphytic plants, such as orchids,

atmospheric sources of carbon dioxide for photo- spanish moss, and certain lichens, obtain essen-

synthesis, nitrogen for biological fixation and tially all their nutrients and water from the

proteins synthesis, oxygen for respiration and atmosphere. Although these plants represent an

synthesis of carbohydrates, and water for extreme case of dependence on atmospheric resources,

many forest trees and some herbaceous plants also

derive a significant portion of their nutrients

from the atmosphere.

presented at the Symposium on Effects of Air

Pollutants on Mediterranean and Temperate Forest Plants suffer when the concentrations of inju-

Ecosystems, June 22-27, 1980, Riverside, rious substances in the atmosphere exceed the

California, U.S.A. amounts they can tolerate. Injurious gases can

enter through the stomata of leaf tissues and

poison the photosynthetic system of living cells.

~ssociateDean for Research, School of Toxic particles can accumulate on plant surfaces

Forest Resources, North Carolina State Univer- and injure plant cells. Strong acids can dissolve

sity, Raleigh. N. C.; and Project Leader, in rain drops or adsorb to snowflakes and then be

Northeastern Forest Experiment Station, Forest deposited in precipitation. Dissolved substances

Service, U. S. Department of Agriculture, can accumulate in snow where they may be concen-

Delaware, Ohio. trated and released with the first meltwater. In

all these ways substances transferred from the

cause very sudden increases in acidity of surface

atmosphere to the biosphere can influence plant

soils, vegetation, and surface waters. Thus a

growth, beneficially (as in the case of beneficial

given plant may be subject to beneficial atmos-

nutrient elements) or harmfully (as in the case of

pheric influences at one time and to negative

toxic gases, aerosols, dry particulate matter, and

influences at another time within a given day,

injurious substances dissolved in precipitation).

month, growing season, or the years of its

development in the case of perennial plants

The growth and productivity of forests are

and animals. Even a given substance, such as

determined by the availability of sixteen elements

sulfur or nitrogen dioxides, may be absorbed and

that are essential for growth and a few that are

utilized as a beneficial nutrient at one concen-

toxic to plants. The essential elements include

tration in the atmosphere. At another, higher

nine major elements: carbon, hydrogen, oxygen,

concentration, even on the same day, however,

nitrogen, phosphorous, potassium, sulfur, calcium,

the same substance may be absorbed and found to

and magnesium and seven minor elements: iron,

be toxic or even injurious to the very same plant.

copper, zinc, manganese, molybdenum, boron, and

chlorine.
Forests and rangelands cover a larger fraction

of the total land area of the United States than

Some elements are both essential and injurious

all other uses of land combined. For this reason,

to plants. For example, sulfur and nitrogen are

terrestrial vegetation, soils, and surface waters

needed for synthesis of protein, nucleic acids,

are the primary deposition sites for precipitation

and other substances; hut gaseous sulfur and

and airborne particulate matter of all types.

nitrogen oxides and sulfuric and nitric acid

Trees develop very large canopies of leaves and

aerosols are also injurious to plants at very

branches that extend high into the air. Thus,

low concentrations. Similarly, excess amounts

forests and range plants provide an extremely

of the minor nutrient elements also can injure

large surface for deposition and assimilation of

plants. Atmospheric fluoride is toxic to plants

both beneficial nutrient elements and injurious

at 25-50 ppm. Aluminum is the most abundant

substances dispersed in the atmosphere.

potentially toxic element in soils. Its avail-

ability (and thus its toxicity) is influenced

Direct injury to vegetation is most likely

greatly by the acidity of soils, which in turn

when a particularly vulnerable life form is

is influenced by the abundance of acid precipi-

exposed at a particularly vulnerable life stage,

tation.
and is growing in a poorly buffered environment

during a season of the year when acid precipita-

Uptake of nutrients from atmospheric sources

tion is most likely. For example, a tender young

is especially important in natural ecosystems

plant, at the earliest stage of reproduction,

such as lakes, estuaries, wetlands, forests, and

growing on a poorly buffered sandy soil, during

rangelands where nutrients from other sources are

a heavy spring rain is especially vulnerable to

scarce and where fertilization is not a normal

acid rain.

management procedure. But this capacity also

increases the vulnerability of terrestrial and

Both herbaceous and perennial plants are

aquatic organisms to injury by acid precipitation

subject to changes in atmospheric deposition

and toxic aerosols and gases (Galloway and others

within a given growing season. In addition,

1978).
perennial shrubs and trees live in the same

environment for many years or even decades. As

a result, they are subject to very long-term

ACID PRECIPITATION AS PART OF A GENERAL

changes in the chemistry of the air and precipi-

PHENOMENON 0F.ATMOSPHERICDEPOSITION
tation.

Air-borne substances that influence terrestrial

The effect of acid precipitation on plants is

plants include sea spray from oceans and large

only one facet of the much larger subject of

lakes; dust resulting from wind erosion of soil

atmospheric/plant/soil interactions. Acidity

as well as from volcanic and cosmic sources;

in precipitation should be understood as a

gases such as CO2, NH3, SOy, H2S, CH4, released

reflection not only of the amounts of substances

from decomposing organic matter and volcanoes;

yielding hydrogen ions (such as sulfuric, nitric,

biogenic particles such as spores, hyphal frag-

hydrochloric, and organic acids) but also of

ments, bacteria, and pollen; particulate matter,

the total balance between all the cations and

aerosols, and gases produced by wild fires and

anions dissolved in precipitation. These major

controlled burning of agricultural, forest and

anions and cations include W-, NH4+, NO3-,
urban wastes as well as from industrial, agricul-
SO4= and many others including K+, Na+, Ca*,

tural residential and commercial heating, and

Mg*, C03-, Cl-, and PO^.

transportation operations (Tam 1958).

For all of the above reasons, it is difficult

Rain and snow change in chemical composition

to assess the effects of acids in rain or snow

within, as well as between, precipitation events.

in isolation from the general chemistry of

In cold climates, acid substances accumulate in

precipitation and atmospheric deposition. Also,

the snowpack where they are released in concen-

the effects of a given "acid rain" or a prevailing

trated form with the first melt water and thus

condition of "acid rains" are very complex,

variable in time, and involve significant inter-

listed above can be taken up readily through

actions that are only partially understood.

foliar organs as well as by absorption from the

soil solution (Tam 1958; Wittwer and Bukovac

1969).

POSSIBLE DETRIMENTAL EFFECTS OF ACID

PRECIPITATION ON VEGETATION
Much larger amounts of essential nutrients

are required for sustained-yield agriculture

A partial list of theoretical effects of acid

than for sustained-yield range management or

precipitation on vegetation was developed earlier

hardwood or softwood forestry. This is true

by T a m and Cowling (1977) and is reproduced in

in rangelands because biomass yields are very

Table 1. The effects are classified as either

low and in forests because the parts of trees

direct or indirect, although most direct effects

that usually are harvested(thewood and bark

will have many indirect consequences as well.. A

of treestems) contain much less of most essential

decreased rate of growth is the expected conse-

elements than the seeds and fruits that are

quence of most of the effects postulated in Table

commonly harvested in agriculture. This is a

1 but unequivocal evidence of significant growth

major reason why fertilization is so common in

effects have yet to be demonstrated in forest or

agriculture and so rare in forestry and range

range ecosystems. Specific biological effects

management. In some forested regions, atmos-

that have been proven to occur in at least one

pheric deposition alone is more than adequate

experimental plant are marked with an asterisk

to permit harvesting of crop after crop of trees

(*) in Table 1. without fertilizing the forest. This is much

less likely to remain so, as more and more of

Many factors (i.e., genetic composition, biotic

the nutrient-rich branches, foliage, and roots

and abiotic stress factors, dose of pollutant, and

of trees are harvested in so-called "whole-tree

pollutant combinations) affect the impact of acid

chipping" 2nd other modern harvesting practices.

precipitation and other pollutants on terrestrial

plants and animals. Variation fn any one factor

Some scientists believe that acid rain and

can result in variation in the nature and magni-

snow are deposited directly onto soils where

tude of pollutant effects. This is shown simplis-

acid substances can be neutralized in well-

tically in Figure 1.
buffered soils or by applications of lime. This

is true on some agricultural lands, especially

Previously, it was believed that the essential

after harvest, but is not true in forests,

and potentially toxic elements listed above were

rangelands, or even on most agricultural lands

taken up by plants almost entirely from the soil

during the growing season. Most raindrops are

solution. Now, it is recognized that airborne

intercepted by the foliage of plants where sub-

gases, particulate matter, and aerosols signifi-

stances dissolved in rain can induce various

cantly augment the supply of both essential and

physiological changes before reaching the soil

injurious elements. All of the substances

(see Table 1). In a mature forest, for example,

rain will wash over at least three tiers of

-I
POLLUTANT
CONCENTRATION foliage before reaching the soil.

SOURCES, AMOUNTS, AND DISTANCES OF

NUMBER OF DURATION OF TRANSPORT OF BENEFICIAL AND INJURIOUS

EXPOSURES DOSE -*ÑÑ'à EACH EXPOSURE

SUBSTANCES IN THE ATMOSPHERE

Forest, range, and aquatic biologists are

becoming increasingly concerned about atmospheric

transport and deposition of both nutritionally

EDAPH IC FACTORS Ñ^ beneficial and potentially injurious substances

for three major reasons:

(1) vegetation, soils, and surface waters are

the primary deposition sites for precipi-

tation and airborne particulate matter;

MECHANISM OF ACTION
7 (2) atmospheric deposition constitutes an

important source of nutrients and poten-

tially injurious substances that affect

the productivity and stability of agricul-

tural, forest, and aquatic ecosystems; and

ACUTE CHRONIC SUBTLE (3) human activities are steadily increasing

Figure 1. Conceptual model of factors involved

the amounts and variety of substances

in air pollution effects on vegetation (Heck

dispersed in the atmosphere (Oden 1968;

and others 1977).

Bolin and others 1972).

 Table I - - P o t e n t i a l E f f e c t s of Acid
P r e c i p i t a t i o n on T e r r e s t r i a l Vegetation

DIRECT EFFECTS

*1. Damage t o p r o t e c t i v e s u r f a c e s t r u c t u r e s such a s c u t i c l e . Damage t o s u r f a c e s t r u c t u r e s

may occur due t o a c c e l e r a t e d e r o s i o n of t h e c u t i c u l a r l a y e r t h a t p r o t e c t s most f o l i a r organs.
It a l s o could r e s u l t from d i r e c t i n j u r y t o s u r f a c e c e l l s by high c o n c e n t r a t i o n s of s u l f u r i c
a c i d and o t h e r harmful substances t h a t a r e concentrated by evaporation o r adherence of s o o t
p a r t i c l e s on p l a n t s u r f a c e s .

*2. I n t e r f e r e n c e w i t h normal f u n c t i o n s of guard c e l l s . Malfunction of guard c e l l s w i l l l e a d

t o l o s s of c o n t r o l of stomata and t h u s a l t e r e d r a t e s of t r a n s p i r a t i o n and gas-exchange p r o c e s s e s
and p o s s i b l y i n c r e a s e d s u s c e p t i b i l i t y t o p e n e t r a t i o n by l e a f - a t t a c k i n g p l a n t pathogens.

*3. Poisoning of p l a n t c e l l s a f t e r d i f f u s i o n of a c i d i c substances through stomata o r c u t i c l e .

This could l e a d t o development of n e c r o t i c o r senescent s p o t s on f o l i a r organs i n c l u d i n g
l e a v e s , f l o w e r s , t w i g s , and branches.

*4. Disturbance of normal metabolism o r growth processes without n e c r o s i s of p l a n t c e l l s .

Such d i s t u r b a n c e may l e a d t o decreased p h o t o s y n t h e t i c e f f i c i e n c y , a l t e r e d r e s p i r a t o r y p a t t e r n s
and i n t e r m e d i a r y m e t a b o l i s m , a s w e l l a s abnormal development o r premature senescence of l e a v e s
o r o t h e r organs.

*5. A l t e r a t i o n of l e a f - and root-exudation processes. Such a l t e r a t i o n s may l e a d t o changes

i n p o p u l a t i o n s of l e a f - s u r f a c e and root-surface microorganisms, i n c l u d i n g n i t r o g e n - f i x i n g
organisms.

*6. I n t e r f e r e n c e w i t h reproduction processes. Such i n t e r f e r e n c e may be achieved by d e c r e a s i n g

t h e v i a b i l i t y of p o l l e n , i n t e r f e r e n c e w i t h f e r t i l i z a t i o n , decreased f r u i t o r seed production,
decreased g e r m i n a b i l i t y of s e e d s , e t c .

7. S y n e r g i s t i c i n t e r a c t i o n w i t h o t h e r environmental s t r e s s f a c t o r s . Such r e i n f o r c i n g i n t e r -
a c t i o n s may occur w i t h gaseous s u l f u r d i o x i d e , ozone, f l u o r i d e , s o o t p a r t i c l e s , and o t h e r a i r
p o l l u t a n t s a s w e l l a s drought, f l o o d i n g , e t c .

INDIRECT EFFECTS

*l. Accelerated l e a c h i n g of substances from f o l i a r organs. Damage t o c u t i c l e and s u r f a c e

c e l l s may l e a d t o a c c e l e r a t e d l e a c h i n g of m i n e r a l elements and o r g a n i c substances from l e a v e s ,
twigs, branches, and s t e m s .

2. I n c r e a s e d s u s c e p t i b i l i t y t o drought and o t h e r environmental s t r e s s f a c t o r s . Erosion of

c u t i c l e , i n t e r f e r e n c e w i t h normal f u n c t i o n i n g of guard c e l l s , and d i r e c t i n j u r y t o s u r f a c e
c e l l s may l e a d t o i n c r e a s e d e v a p o t r a n s p i r a t i o n from f o l i a r organs and v u l n e r a b i l i t y t o drought,
a i r p o l l u t a n t s , and o t h e r environmental s t r e s s f a c t o r s .

* 3 . A l t e r a t i o n of symbiotic a s s o c i a t i o n s . Changes i n l e a f - and root-exudation processes and

a c c e l e r a t e d l e a c h i n g of o r g a n i c and i n o r g a n i c substances from p l a n t s may a f f e c t t h e formation,
development, balance, and f u n c t i o n of symbiotic a s s o c i a t i o n s , such a s mycorrhizae, nitrogen-
f i x i n g organisms, l i c h e n s , e t c .

*4. A l t e r a t i o n of h o s t - p a r a s i t e i n t e r a c t i o n s . Resistance and/or s u s c e p t i b i l i t y t o b i o t i c

pathogens, p a r a s i t e s , and i n s e c t s may be a l t e r e d by s u b j e c t i n g p l a n t s t o any environmental
stress. Acid p r e c i p i t a t i o n may i n c r e a s e t h e s u s c e p t i b i l i t y of p l a n t s t o t h e s e i n j u r i o u s
a g e n t s , a l t e r t h e i r c a p a c i t y t o t o l e r a t e d i s e a s e o r i n j u r y , o r a l t e r t h e v i r u l e n c e of pathogens.
The e f f e c t s of a c i d i c p r e c i p i t a t i o n may vary w i t h t h e following: t h e n a t u r e of t h e pathogen
involved (whether a fungus, bacterium, mycoplasma, v i r u s , nematode, p a r a s i t i c seed p l a n t ,
i n s e c t , o r multiple-pathogen complex); t h e s p e c i e s , age and p h y s i o l o g i c a l s t a t u s of t h e h o s t ;
and t h e s t a g e i n t h e d i s e a s e c y c l e i n which t h e a c i d i c s t r e s s i s appled, f o r example, a c i d i c
r a i n might d e c r e a s e t h e i n f e c t i v e c a p a c i t y of b a c t e r i a b e f o r e i n f e c t i o n and i n c r e a s e t h e
s u s c e p t i b i l i t y of t h e h o s t t o d i s e a s e development a f t e r i n f e c t i o n .

Source: Tamm and Cowling, 1977

 Recent increases in the deposition of sub-
very large, as shown in table 3. Most gases,

stances on terrestrial vegetation are due mainly

carbon oxides, and aerosols result from the com-

to increases in combustion of fossil fuels in

bustion of fossil fuels. A very large part of

industrial enterprises, residential heating,

these global emissions are produced in the United

transportation, and agricultural operations.

States.

Previously, it was believed that most of these

materials fell out of the atmosphere near the

If the United States continues to add to the

site of emission. Now it is recognized, particu-

amount of substances dispersed i n the atmosphere

larly with increased use of tall stacks at power

and deposited into the biosphere of the earth, it

plants, that atmospheric processes can lead to

is essential that we measure the amount and chemical

extensive mixing and both chemical and physical

form of the deposited matter and understand the

interactions and transformations of atmospheric

biological consequences of that deposition. Regret-

particles, aerosols, and gases. Furthermore,

tably our understanding of these processes in the

these substances and their reaction products are

United States is very fragmentary. Fortunately,

dispersed by meteorological processes and finally

however, more extensive measurements of atmospheric

are deposited on vegetation or soils as much as

deposition and its biological consequences have

several hundreds of kilometers from the original

been made in Europe, where an atmospheric-deposition

sources of emission. The recent fallout of

network has been maintained since the late 1940s

radioactive materials in the eastern United States

(Oden 1968).

as the result of atomic explosions in the Peoples

Republic of China provides a dramatic reminder of

The European Air Chemistry Network began in

the long-distance transport and deposition of

Sweden and has gradually spread to include most

pollutants.
of western Europe and parts of eastern Europe,

including Poland and the Soviet Union. Since the

The amounts of substances introduced delib-

mid 19501s,a network of about 100 stations has

erately or inadvertently by man are becoming so

made monthly measurements of changes in the

large that man is becoming a major force in the

chemistry of precipitation. The substances

biogeochemistry of the earth (Kovda 1975). This

analyzed at most of these stations include the

is shown in table 2 which contains a tabulation

following major cations and anions: NH4, Na,

of data on annual outpuG of fertilizers, indus-

Ca, K, Mg, 804, NO3, PO4, Cl as well as pH,

trial dusts, garbage and other urban wastes and

conductivity, and titratable acidity and akla-

by-products, mine refuse, and discharges of

Unity. These data have shown various long-term

aerosols and gases mainly from the combustion

trends. for example, the amount of nitrate

of fossil fuels. All these categories of matter

nitrogen in precipitation (an important fertilizer

are becoming comparable in magnitude to the

element) increased markedly in many parts of

discharges of dissolved and suspended substances

Europe during the fifteen years between 1955 and

in all the rivers of the world, the annual yield

1970. Nitrate nitrogen helps plants grow. Thus,

of photosynthetic products, or the cycling of

the nitrogen added in precipitation probably

inorganic elements in the earth as a whole.

increased yields of agricultural and forest crops.

Man-made emissions into the atmosphere are also

But not all the substances detected in precipi-

tation were beneficial. Long-term trends of in-

Table 2--Biogeochemical and Technological Forces

jurious sulfate and hydrogen ions also were detected

in the Biosphere of the Earth

from 1955-1970. The latter changes were attributed

to strong acids formed in the atmosphere, mainly

Biosphere Components Tons/Year:. from oxides of sulfur and nitrogen produced during

combustion of fossil fuels. More recent data show

Biogeochemical processes:
that thesetrends of Increasing acidity are

Global yield of photosynthesis

Cycle of inorganic elements

River discharges:
Table 3--Anthropogenic Emissions into the

Dissolved substances
Atmosphere
Suspended substances

Types of Emissions I~onsf~ear

Anthropogenic sources:
I
Output of fertilizers
Dust 2.5 x lo8
Industrial dust
Gases (mainly .SOg,HC and N O ) 6.5 x lo8
Garbage, urban wastes and
Carbon oxides (CO + C02) 2.0 lo9
byproducts
Aerosols 1.0 x 109
Mine refuse

Aerosols and gas discharges

Note: Doubling about every 7-10 years

Source: Kovda, 1975

Source: Kovda, 1975

continuing although t h e r e l a t i v e c o n t r i b u t i o n of
s u l f u r i c and n i t r i c a c i d s i s changing (Likens
1976).
CHANGES I N THE CHEMISTRY OF
PRECIPITATION I N THE UNITED STATES
Some monitoring of t h e chemistry of p r e c i p i -
t a t i o n has been c a r r i e d on i n t h e United S t a t e s
(Feth and o t h e r s 1964; Lodge and o t h e r s 1968).
Many of t h e s e s t u d i e s provide e x c e l l e n t and
r e l i a b l e information about t h e a c i d i t y of p r e c i p i -
t a t i o n . But most s t u d i e s i n t h i s country have
s u f f e r e d from t h r e e major shortcomings:
1. The d a t a were c o l l e c t e d f o r a l i m i t e d land
a r e a ~ t y p i c a l l yonly a s i n g l e p o i n t o r a
few p o i n t s i n one o r two s t a t e s (Gambell
and F i s h e r 1966);
2. The d a t a were c o l l e c t e d f o r very l i m i t e d
p e r i o d s of t i m e ~ t y p i c a l l yonly one o r
two y e a r s ;
3. Very few d i r e c t measurements of a c i d i t y
have been made.
There i s only one l o c a t i o n i n t h e United
States~aH t ubbard Brook Experiment F o r e s t i n
New H a m p s h i r e ~ w h e r et h e a c i d i t y of r a i n has been
measured d i r e c t l y and c o n s i s t e n t l y f o r a s long a s
10 y e a r s . The longest-term n a t i o n a l monitoring
program was o p e r a t e d by t h e U.S. P u b l i c Health
S e r v i c e f o r 6 y e a r s , from 1960 t o 1966 (Lodge
and o t h e r s 1968). These d a t a showed t h a t
p r e c i p i t a t i o n g e n e r a l l y i s a c i d i c e a s t and
g e n e r a l l y a l k a l i n e west of t h e M i s s i s s i p p i River,
t h e l a t t e r because of a l k a l i n e d u s t i n t h e a i r .
Using fragmentary b i t s of information,
o b t a i n e d i n d i r e c t l y and i n l i m i t e d a r e a s and
p e r i o d s of time, C o g b i l l and Likens (1974)
managed t o c a l c u l a t e t h e probable changes i n
average a c i d i t y of r a i n f a l l i n v a r i o u s p a r t s of
t h e e a s t e r n United S t a t e s from 1955-1973. ' P r e c i p i -
t a t i o n i n a l a r g e p o r t i o n of t h e e a s t e r n United
S t a t e s was l e s s than pH 5.6 i n 1955-56; t h e zone
of g r e a t e s t a c i d i t y (lowest pH) was g e n e r a l l y
c o n s i s t e n t w i t h t h e zone where s u l f u r emissions
were high--parts of Ohio, Pennsylvania, West
V i r g i n i a , New York, and New England. By 1973,
however, t h e a r e a w i t h an average pH of r a i n
below 4.5 had extended t o i n c l u d e p a r t s of
Missouri, Arkansas, M i s s i s s i p p i , Alabama,
Georgia, South C a r o l i n a , V i r g i n i a , Kentucky,
I l l i n o i s , Michigan, and f u r t h e r n o r t h i n t o New
England and Canada. E s s e n t i a l l y , i t embraces
most of t h e a r e a e a s t of t h e M i s s i s s i p p i River.
I n d i v i d u a l r a i n s t o r m s w i t h pH v a l u e s between 2.1
and 3.6 have been r e p o r t e d i n New York, I l l i n o i s ,
Indiana, New Hampshire, Massachusetts and North
C a r o l i n a ~ i nsome c a s e s many hundreds of k i l o -
meters from major s o u r c e s of a i r p o l l u t i o n
(Likens 1976).
The r e l a t i v e c o n t r i b u t i o n of s u l f a t e and
n i t r a t e t o t h e t o t a l a c i d i t y of p r e c i p i t a t i o n
a p p a r e n t l y changed markedly during t h e y e a r s s i n c e
1964-65. A t Hubbard Brook, New Hampshire, t h e
r a t i o of s u l f a t e t o n i t r a t e changed from 83:15
i n 1964 t o 66:30 i n 1974. During t h i s same
decade, t h e t o t a l i n p u t of hydrogen i o n s i n c r e a s e d
by 36 p e t . Thus, most of t h i s i n c r e a s e appears
t o be due t o i n c r e a s e d d e p o s i t i o n of n i t r i c a c i d .
EFFECTS OF ACID PRECIPITATION ON
TERRESTRIAL ECOSYSTEMS
Cowling (1980a, 1980b) has r e c e n t l y completed
2 h i s t o r i c a l a n a l y s e s of p r o g r e s s i n s c i e n t i f i c
and p u b l i c understanding of a c i d p r e c i p i t a t i o n
and i t s b i o l o g i c a l consequences. S e v e r a l pub1i.-
c a t i o n s a r e worthy of s p e c i a l n o t i c e i n t h i s
connection. The p i o n e e r i n g r e s e a r c h e s by Robert
Smith. E v i l l e Gorham and Svante Oden d e a l t w i t h
e f f e c t s on l a k e w a t e r s , a q u a t i c v e g e t a t i o n ,
t e r r e s t r i a l v e g e t a t i o n , andhuman h e a l t h (Smith
1872: Gorham 1958, 1976: Oden 1968). I n 1971.
Bolin and h i s co-workers completed t h e Swedish
Case Study Contribution t o t h e United Nations'
Conference on t h e Human Environment (Bolin and
o t h e r s 1972). I n 1972, t h r e e Norwegian r e s e a r c h
organizations established a s p e c i a l research
p r o j e c t c a l l e d Acid P r e c i p i t a t i o n : E f f e c t s on
F o r e s t s and Fish, w i t h an annual budget of
10,000,000 Norwegian kroner (U. S. $2,000,000).
The f i r s t I n t e r n a t i o n a l Conference on Acid
P r e c i p i t a t i o n and t h e F o r e s t Ecosystem was h e l d
a t Ohio S t a t e U n i v e r s i t y a t Columbus i n May of
1975 (Dochinger-and S e l i g a 1976a). I n June of
1976, an I n t e r n a t i o n a l Conference on E f f e c t s of
Acid P r e c i p i t a t i o n was h e l d a t Telemark, Norway,
and t h e major papers assembled f o r t h i s meeting
published by Braekke (1976) and i n a s p e c i a l
i s s u e of Ambio (1976). I n November, 1976, Gene
Likens published h i s summary r e p o r t i n Chemical
and Engineering News (Likens 1976). I n May. 1978,
a NATO Advanced Research I n s t i t u t e on E c o l o g i c a l
E f f e c t s of Acid P r e c i p i t a t i o n was held a t
Toronto, Canada (Hutchinson and Havas 1980). In
September, 1978, t h e C e n t r a l E l e c t r i c i t y Generating
Board i n England and t h e E l e c t r i c Power Research
I n s t i t u t e i n t h e United S t a t e s sponsored an
i n t e r n a t i o n a l symposium on t h e b i o l o g i c a l e f f e c t s
of a c i d p r e c i p i t a t i o n (Howells 1979). I n March,
1980, t h e Norwegian s p e c i a l p r o j e c t on a c i d
p r e c i p i t a t i o n sponsored an I n t e r n a t i o n a l Conference
on E f f e c t s of Acid P r e c i p i t a t i o n i n Sandefjord,
Norway (SNSF 1980).
The e f f e c t s of a c i d p r e c i p i t a t i o n on t e r r e s t r i a l
ecosystems g e n e r a l l y have been l e s s w e l l documented
than t h o s e on populations of f r e s h w a t e r f i s h and
o t h e r a q u a t i c organisms (Ambio 1976; Braekke 1976).
Nevertheless, c e r t a i n d e f i n i t e e f f e c t s have been
r e p o r t e d . The most s t r i k i n g of t h e s e e f f e c t s was
t h e development of p e a t moss (Sphagnum s p . ) a s a
submarine, r a t h e r than a t e r r e s t r i a l p l a n t i n
a c i d i f i e d l a k e s and streams i n Sweden. Dense
mats of Sphagnum and heavy f e l t s of a l g a e develop
on t h e bottom of t h e s e l a k e s i n water a s deep a s
1 8 m. This growth i s r e p o r t e d by Grahn and
o t h e r s (1974) t o induce o l i g o t r o p h i c a t i o n
(opposite of eutrophication)--a s e l f - a c c e l e r a t i n g
process that leads to a substantial nutrient
published by Dochinger and Seliga (1976a, 1976b).

impoverishment of lake waters.

At Congressional hearings in July, 1975, Cowling

(1976) testified on the inadequacy of research in

Analyses of forest growth in southern Sweden

the United States on Acid Precipitation and its

from 1896 to 1965 showed a 2 to 7 percent decrease

biological consequences. Specifically, the lack

in growth between 1950 and 1965. Johnsson and

of a coordinated program of research on ecological

Sundberg (1972) "found no good reason for attrib-

effects and lack of a stable monitoring network

uting [this] reduction in growth to any cause other

were recognized as primary causes of our profound

than acidification." Similar attempts to quantify

ignorance of acid precipitation. In the spring

possible effects on growth of forests in the

of 1976, however, a cadre of scientists in various

United States have been inconclusive.

institutions and agencies throughout the United

States began the process of creating the National

Both direct and indirect damage to crops and

Atmospheric Deposition Program (NADP) to meet

forests have been reported by various investiga-

these two critical needs (Kennedy 1977; Galloway

tors in laboratory, greenhouse, and field experi-

and Cowling 1978).

ments in which synthetic rain equivalent in chemical

composition and rate of deposition to natural rains

In the fall of 1977; the President's Council on

has been applied. The biological effects recorded

Environmental Quality contracted with the NADP for

in these experiments include the fol3.owing

the drafting of "A National Program for Assessing

(Cowling 1980c) : the Problem of Atmospheric Deposition (Acid Rain)."

--Induction of necrotic lesions on foliage;

This publication (Galloway and others 1978) provided

--Loss of nutrients due to leaching from

the basis for a Presidential Initiative on acid

leaves and other foliar organs;

precipitation which President Carter announced on

--Predisposition of plants to infection by

August 2, 1979 in his Second Environmental Message

bacterial and fungal pathogens;

(Carter 1979). This initiative calls for a 10-year

--Accelerated erosion of waxes on leaf surfaces;

long, $10,000,000 per year program of research on

--Inhibition of nodulation of legumes leading to

the causes and consequences of acid precipitation.

decreased fixation of nitrogen by symbiotic

A standing Acid Rain Coordinating Committee was

bacteria; and
established by the President to plan and manage

--Reduced rates of decomposition of leaf litter

the program. Leadership for the Committee is

leading to decreased mineralization of

provided by co-chairmen from the Department of

organically-bound nutrients.
Agriculture and the Environmental Protection

Agency. At the present time, the Acid Rain

Abrahamsen (1980) has recently summarized many

Coordinating Committee is drafting a coherent

years of research showing both positive and

program of research on atmospheric chemistry and

negative effects of acid precipitation on forest

transport, chemical and biological monitoring,

growth. He concludes with the following general

ecological and materials-damage effects, economic

statements: "Apart from possible direct effects

assessments, and public-policy options for control

of acid precipitation on forest trees, the effects

of acid precipitation and/or amelioration of its

on forest growth can be considered a nutrition

ecological effects.

problem ... increased deposition of N and S can

be regarded as a ... fertilization effect, and Wetstone (1980) has recently summarized the

the increased leaching of nutrient cations ... biological and materials-damage effects of acid

as an oligotrophication or acidification effect

precipitation in relation to the pollution-

... the general hypothesis that acid precipita- control laws in North America.

tion significantly will decrease forest production

over large areas must be revalued. The deposition

In conclusion, the Presidential Initiative on

of N and to some extent S ... is likely to Acid Precipitation, coupled with growing Congres-

increase forest production. Reduced growth may

sional, public, and private-industrial interest in

be expected where or when nutrients like Mg and

acid precipitation research, provide a basis for

possibly K are the growth limiting elements."

increasing hope that the United States will do its

part, together with Sweden, Norway, England,

Canada and other nations, to meet the challenge

RECENT INITIATIVES DEALING WITH ACID

of continuing economic development with adequate

PRECIPITATION AND ITS BIOLOGICAL EFFECTS

safeguards for the quality of life and the long-

term productivity of ecosystems on which our

In 1975, the National Academy of Sciences'

good life critically depends.

Committee on Atmospheric Sciences published its

report on Atmospheric Chemistry: Problems and

Scope (NAS 1975). Growing awareness of important

LITERATURE CITED

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Abrahamsen, G.

plants led the U. S. Forest Service to sponsor

1980. Acid precipitation, plant nutrients and

the First International Symposium on Acid Precipi-

forest growth. In Proc. Int. Conf. on

tation and the Forest Ecosystem at Columbus,

Ecological impacts of Acid Precipitation

Ohio in May, 1975. The proceedings of this

[Sandefjord, Norway].

Symposium and the Associated Workshop Report were

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1976. Report of t h e I n t e r n a t i o n a l Conference on
t h e e f f e c t s of a c i d p r e c i p i t a t i o n i n Telemark,
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Braekke, F. H., ed.
1976. Impact of a c i d p r e c i p i t a t i o n on f o r e s t
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1979. The P r e s i d e n t ' s Environmental Program 1979.
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Kennedy, V. C.

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Swanson, K. C. Hill, E. Lorange, and A. L. Lazrus.

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p. 235-261. Springer-Verlag, New York.

Simulation Modeling of the Effects of

Chronic Pollutant Stress on Plant
Processes and Plant Community
Dynamics
Modeling Pollutant Uptake and Effects
on the Soil-Plant-Litter System '
R. J. Luxmoore2

Abstract: Five coupled models of water, carbon, and chemi-

cal dynamics in a soil-pl a n t - l i t t e r system are outlined.
Algorithms defining gaseous and particulate pollutant
uptake are described along with functions for chemical
effects on plant growth and l i t t e r decomposition. Some
simulation results of a deciduous forest i l l u s t r a t e the
importance of diurnal and annual cycles of environmental
conditions on pollutant movement in vegetation. This
modeling approach has provided (1) insights into plant
physiological processes and t h e i r interactions, ( 2 ) identi-
f i cati on of plant properties important i n pollutant uptake,
( 3 ) a1ternati ve hypotheses about pollutant effects, and
( 4 ) a unified basis f o r assessment of diurnal and long-term
pollutant impacts on plant communities.

'and a l l the king's horses and a l l the king's
men coul dn I t p u t Humpty together again.''
from Humpty Dumpty, Anon.
The discouraging words of the nursery rhyme
suggest that the synthesis of bits of an egg t o
a whole will not happen at l e a s t while horses
and men are in charge! Our task of trying to
couple together b i t s and pieces of mechanistic
information about the physiology of trees and
responses to soil and atmospheric environments
i s no less awesome a challenge. Simulation
modeling is a remarkable tool f o r meeting t h i s
challenge, since through mathematics coupled
relationships may be quantified. In t h i s paper,
presented a t the Symposium on Effects of Air
Pollutants on Mediterranean and Temperate Forest
Ecosystems, June 22-27, 1980, Riverside,
California, U.S.A.
research staff member, Environmental
Sciences Divisi on, Oak Ri dge National
Laboratory, Oak Ridge, Tennessee 37830.
Operated by Uni on Carbi de Corporati on under
contract W-7405-eng-26 with the U .S. Department
of Energy. Publication No. 1553, Environmental
Sciences Division, ORNL.
paper, an outline i s presented of five models
that link together and provide a framework f o r
study of pollutant uptake and effects in the
whole plant environment complex. Some appl ica-
tions are shown and the use of models in
analysis of experiments i s explored. Lastly,
some specul ati ons are presented about pol 1utant
impacts on whole plants and t h e i r diurnal
metabolism.
MODELING THE SOIL-PLANT-LITTER SYSTEM
The development of a unified approach t o the
modeling of t e r r e s t r i a1 processes has been
undertaken a t Oak Ridge. Five component models
of water, carbon, and chemical dynamics in a
soil-pi ant-1 i t t e r system were constructed and
linked together (Baes e t a1. 1976). The models
(table 1) are deterministic. The flow processes
are dependent on gradient terms calculated by
the models t o provide the flow driving forces
and empirical inputs are used to represent path-
way resistances or conductivities. Flow
directions are not predetermined and the models
can be applied t o a range of different soil-
plant systems (e.g., coniferous, deciduous
f o r e s t ) by changing the empirical properties in
the input data. The reader i s referred t o the
documentation reports (table 1) f o r further
details.
Tab1 e 1--Some attributes of coup1ed models g i s gas density (Ug/ml)
describing carbon, water, and chemi cal dynamics
in the soil -pi ant-li t t e r system. ra i s boundary layer diffusion
resistance (seclcm)
CoMFONErn WATER SOIL EHCHANBE M O T mLUTl UPTAKE SOLUTES
rs i s stomata1 resistance (seclcm)
NAME PROSPER ITEHMI SCEHM CERES DIFMAS DRYADS
w m;".
TIME STEP
ATTRIBUTES
1 6 0 8 60 mi".
EVAPOTRANsflRATIOt
I S OR 00 mi".
USES EMPIRICAL
wm h
W2 DIFFUSION
IS OR
IMPLEMENTS MODEL
I 6 OR 80 mi".
SOLUTEUPTAKEBY
rm i s mesophyll resistance (sec/cm)
BY COMBINATION DISTRIBUTION EOUATION FOR 3F DIFFUSION AND ROOTS AND LEAVES.
EOUATION. COEFFICIENT NET PHOTOSYN. MASS FLOW OF

SOIL WATER FLOW

BY DARCV FLOW
IKdl FOR SOIL
OF INTEREST.
THESIS.

SUBSTRATE GHADt:
SOLUTES TO ROOTS
BY BALDWIN. NYE
AND TINKER 119731.
DIFFUSIVE GAS UPTAKE
BY LEAVES. Ug i s uptake (iig/cm* leaf/sec)
EOUATION. ENT EOUATION FOR GRADIENT EOUATION
lRANSLOCAT10N FOR PHLOEM TRANS-
USES EMPIRICAL
RELATIONSHIP USES INPUT VALUES
LOCATION
The value of gi i s made to vary between zero
BETWEENSURFACE
RESISTANCE AND
SURFACE WATER
FOR POTENTIAL
GROWTH OF LEAF.
STEM. BOOT, FRUIT
TRANSPIRATION FLUX
USED FOR XYLEM and ge depending on the level of pollutant in
POTENTIAL.
EMPIRICAL LITTER
TRANSPORT.

P U N T DEMAND FUNC-
leaf storage (Ei) as follows,
EMPIRICAL DATA DECOMPOSITION TION DETERMINED BY
FOR SOIL HYDRAU- RELATIONSHIPS. POTENTIAL SOLUTE
LIC PROPERTIES. CONCENTRATION INPUT
VALUES.

REFERENCE MUFF fI,11977; BEGOVICH AND OIXON rn i l 119781

JACKSON 118751

The coupling between models (fig. 1) shows Em i s the maximum allowable level of pollu-
that every model has informati on transfer with tant in leaf storage, an input parameter.
at least two other models, and these take place Operationally this i s the pollutant level at
on either an hourly time step or every 15 min- which the leaf tissue becomes necrotic.
utes during storm events. Hourly values of sto- ORNL-DWG 75-15812R2

matal resistance and plant water potential from JroweRI

PROSPER are used in CERES to determine photo-
synthesis and growth respectively. Leaf and
root growth in t u r n influence transpiration and
thus soil water flow. During rainfall, i n f i l -
tration and the movement of water between soil
layers (calculated in PROSPER) i s used in the
soil chemistry model (SCEHM) to calculate chemi-
cal f l uxes. Chemi cal concentrati on and root
water uptake information are used in DIFMAS t o
calculate chemical uptake into root by diffusion
and mass flow. Chemicals within the plant are
moved up in the transpiration stream and down in
the phloem pathway.
This set of models can be run for simulation
periods of several years and annual budgets for
water, carbon and chemicals can be evaluated as
well as detailed results for hourly periods of
interest. The algorithms defining gaseous and
parti cul ate pollutant uptake and effects on
plant growth and l i t t e r decomposition are out-
lined in the next two sections along with
example simulation results.
AIR POLLUTANT UPTAKE
The uptake of a i r pollutants by vegetation
may occur directly through leaves (gaseous and
parti cul ate) or indirectly through roots after
the pollutants have been incorporated into soil.
Gaseous uptake i s represented by a diffusion PROSPER soi 1-pi ant-atmosphere water f l ow model
equation (same form as the photosynthesis
equati on ) Thus . CERES
SCEHM
carbon dynamics of vegetation and l i t t e r
soil chemistry model
DIFMAS diffusion andmass flow of chemicals t o
roots
DRYADS chemi cal dynami cs of vegetation and
1i t t e r
where g i s the external pollutant concentration
(ml/ml) Figure 1--Coupling of five process models that
describe hourly carbon, water, and solute
gi i s the internal pollutant concentration dynamics of the soil-plant-litter system.
(ml/ml)
 Sulfur dioxide uptake by an oak-hickory distributed and has one of two f a t e s . I t may be
f o r e s t in the v i c i n i t y of a lead mining and transported t o other plant parts or be incorpo-
smelter complex in southeastern Missouri was rated in the leaf in an immobile form. The cu-
simulated and r e s u l t s i l l u s t r a t e the behavior of t i c u l ar uptake process i s considered r e v e r s i b l e
the model. Cumulative s u l f u r levels in leaves in the model. Thus during r a i n f a l l , wash-off
( f i g . 2) show a rapid increase on t h e 25th of occurs and i f Se becomes l e s s than S i , then
August, a day in which the atmospheric SO2 leaching of pollutant out of leaves will occur.
level was increased 10 f o l d above ambient. The
translocation of s u l f u r from leaf t o stem ORNL- DWG 7 6 - 13390R
( f i g . 2 ) c l e a r l y shows a diurnal pattern and a t
elevated r a t e s on the 25th of August. Some of
the s u l f u r material t h a t was transported t o t h e
roots, leaked i n t o the t r a n s p i r a t i o n stream and
returned from t h e roots t o the stem, a l b e i t in
t r a c e amounts. The phloem and xylem transport
pathways can a l l ow considerable mobi 1i t y of sol-
'06 6
utes between plant t i s s u e s according t o the
simulation. The cumulative s u l f u r l e v e l s in t h e
l e a f , stem and root components ( f i g . 3 ) show
t h a t t h e majority of s u l f u r remained in t h e
leaves. The value of 8 x 105 p g ~ / m 2 i s
equivalent t o a 1eaf concentration of 180 ppm.
ORNL- DWG 80-11126 €

10'
22 23 24 25 26 27 28 29
DAYS O F AUGUST
-
21 23 25 27 29 31
Figure 3--Simul ated s u l f u r elemental accumul a-
DAYS IN AUG t i o n in l e a f , stem, and root t i s s u e (pg/m2)
Figure 2--Simul ated cumulative s u l f u r dioxide r e s u l t i n g from gaseous uptake.
uptake by vegetation ( g/m2) and leaf t o stem
phloem translocation r a t e ( pg/m^/h) f o r 11
days in August. S e n s i t i v i t y analysis of the leaf c u t i c l e con-
d u c t i v i t y ( f i g s . 4a, b) shows t h a t g r e a t e r
conductivity i s associated with greater chemical
The uptake of p o l l u t a n t s from p a r t i c u l a t e s (zinc in t h e example) uptake by leaves and a
deposited on leaves (Ui ) i s represented by a s l i g h t l y reduced uptake of zinc from t h e s o i l
gradient equation using empirical input values solution (Begovich and Luxmoore 1979). This
f o r t h e c u t i c u l a r conductivity ( k l ) and thick- l a t t e r and more s u b t l e e f f e c t i s induced by the
ness ( W ) . Thus, higher zinc level in t h e plant with higher con-
ducti vi t y which feeds back a reduced chemical
demand in t h e root uptake algorithm. I t i s pos-
s i b l e t h a t s u b t l e e f f e c t s may become s i g n i f i c a n t
when integrated over long time periods. Cuticu-
l a r conductivity and t h e equivalent property a t
t h e root-soil i n t e r f a c e ( r o o t conductivity, k c )
where S i s t h e external p o l l u t a n t on leaf were shown t o be very s e n s i t i v e parameters in
surface (g/m2 land) t h e model, and yet these are perhaps t h e l e a s t
well characterized experimentally. Results from
S i i s t h e i n t e r n a l p o l l u t a n t within a s e n s i t i v i t y analysis of root conductivity on
f o l i a g e (g/m2 land) lead uptake ( t a b l e 2) show large increases in
uptake by roots and lead c o n c e n t r a t i o ~ i n t r e e
The amount of dissolved pollutant on leaf t i s s u e s with increase in kr from 10" cm/sec
surfaces i s calculated as the l e s s e r of e i t h e r t o 10-6 cmlsec. The simulations a l s o show
t h e product of sol ubi 1i t y and the water volume t h a t pollutants accumulate p r e f e r e n t i a l l y in t h e
on leaves ( i n t e r c e p t i o n ) or the current amount
,
leaf and root, the s i t e s of pollutant entry. A
of p o l l u t a n t on leaves. The soluble p o l l u t a n t modification has subsequently been added t o t h e
within leaves ( S i ) i s assumed t o be uniformly model t o allow chelation of chemical within t h e

176
plant (Luxmoore and Begovich 1979) which has the
effect of i ncreasing the mobi 1i t y of pollutant
within the plant. Thus, the s i t e of pollutant
entry may not be the s i t e of accumulation.
ORNL-OWG 80- 11127 ESD
The monthly pattern of lead uptake by roots
and foliage simulated for an oak forest near a
mine-smelter complex during the f i r s t year of
operation shows that uptake corresponds with the
growing season (table 3). The major proportion
(88%) of root uptake occurred during the day
chiefly due to two compl ementary transportation
processes; the mass flow of pollutant t o roots
and mass flow of pollutant from roots to shoots.
The l a t t e r was the controlling process in the
simulations. Overall, leaf uptake was more than
double that simulated for roots for the f i r s t
year of smelter operati on.

Tabl e 3--Simul ated root and leaf

uptake, (mg pb/m2 landlmonth) of
lead by oak vegetation in the
vicinity of a mine-smel t e r complex.

2.0 - Month Night

I
1.8 - -
- - --- lo-'
YT -
- -..-....-. /
I- 10-9
-
a!?
1.6
-- !o-~~
10-13
Jan.
2
'
2 1.4 -
1 -
0
- Feb.
.-.--*' March
*.---

- April
- May ,

June
-
July
Aug.
0 5 10 15 20 25 30 Sept.
DAYS IN JULY
Oct.
Figure 4--a. Influence of leaf cuticle perme-
a b i l i t y on zinc uptake by leaves. Nov.
b. Influence of leaf cuticle penne- Dec.
a b i l i t y on zinc uptake by roots.
Total

Tabl e 2--Sensi t i vity of annual root lead uptake and tissue concentration (prior t o leaf
f a l l ) in an oak forest to change in the root solute conductivity parameter ( k r ) .

I Annual
I September tissue concentration (ppm)
Stem
I
Root
I
root uptake
( pg/cm2/year) Leaf Sapwood Heartwood Sapwood Heartwood Fruit
Pollutant Impacts
Simple ramp functions are used to determine
pollutant effects on the growth and decomposi-
tion of leaf, stem, root and f r u i t components.
Separate ramp functions for either growth ef-
fects ( f i g . 5) or control of decomposition in
the l i t t e r (same form as for growth effects)
represent ranges of chemi cal def i ciency,
sufficiency, and toxicity as the chemical con-
centrati on increases. Hypotheses concerning
benefici a1 ( f e r t i l i z e r ) and toxic pollutant
effects can thus be examined. The product of
the growth coefficient and tissue growth r a t e
(from CERES) provides a modified growth rate due
to pol 1u tant effects.
Figure 5--The re1 ati onship between the growth
coefficient (Gc) and the amount of element in
tissue (Ei) used t o represent deficiency
(Ei < E1)m sufficiency ( E l < Ei < E?),
and toxicity (Ei > E 2 ) effects of the
elements on tissue growth rate.
A six-year simulation of heavy metal deposi-
tion, transport, and uptake in an oak-hickory
forest in southeastern Missouri showed that the
lead accumulation was greatest in the l i t t e r
(Luxmoore et a1. 1978). Root uptake of lead
i ncreased through the six-year period, whereas
leaf uptake was a constant for the repetitive
annual deposition of 25 g Pb/m . Due to the
buildup of lead in the plant tissues, the mor-
t a l i t y of pl ant parts returned increasingly
greater amounts of lead to the l i t t e r system
The l i t t e r dry weight increased through the six-
year period by 949 g/m2. This compares
reasonably with a difference of 1130 g/m2
between the l i t t e r mass at a control s i t e and a
s i t e exposed to equivalent heavy metal deposi-
tion (Watson e t a1. 1976). The simulation
results pose an alternative hypothesis to the
experimental inference of reduced rates of l i t -
t e r decomposition at the elevated levels of
heavy metal accumul a t i on (Jackson and Watson
1977), by showing that the same effect could be
obtained with increased mortality of plant parts.
Next Step
The previous sections outline one particular
s e t of models and show some simulation results
including s e n s i t i v i t y analysis of selected
parameters. The work presented i s best viewed
as "equipment" ; the subrouti nes being component
parts which collectively form a package of
hypotheses, theories, or knowledge in mathemati-
cal form. We need t o thoroughly t e s t models
through applications t o experimental studies as
much as possible to, hopefully, invalidate parts
of the model structure. The deviations of model
predictions from experimental findings provide
the key to new insights - in t h i s way models
f a c i l i t a t e the analysis and synthesis of complex
interactions. Putting models to work in t h i s
way requires data from well-documented experi-
ments. For example, the uptake and physiolog-
ical effects of gaseous pollutants have been
documented for several t r e e species (Jensen and
Kozlowski 1975, Thompson e t a1. 1967, Roberts
1974, Lawhon 1973, Houston and S t a i r s 1973), and
these experimental data can be used in leaf
physiological models (Kercher 1977) or in the
models outlined in the e a r l i e r sections. A con-
siderable body of experimental data has been
developed for a i r pollutant effects on plants,
and i t i s timely to apply modeling techniques in
the research analysis of impacts. An alterna-
t i ve approach i s one of conceptual extrapol a t i on
of the model behavior. Some speculations are
presented in the next section.
Pollutants and the Diurnal Cycle
The modeling of water, carbon, and chemicals
as coupled components in soi 1-pl ant-1 i t t e r
systems has stimulated the development of a con-
ceptual framework for the diurnal cycle in
plants ( f i g . 6) that can be used t o invent
hypotheses of pollutant effects on whole
plants. In the diurnal cycle, plants change
between two relative states: ( a ) lowest
sucrose, metabolite, and solute reserves at max-
imum hydration (dawn s t a t e ) ; and ( b ) highest
sucrose, metabol i t e , and solute reserves a t min-
imum hydration (dusk s t a t e ) . These states are
relative and apply to a given day. Photosyn-
thesis recharges the plant with sucrose and
increases starch storage (or equivalent) during
the day. A t the same time, the plant i s also
. recharging with nutrients and undergoing dehy-
dration. The loss of water can reduce the r a t e
of cell expansion processes during the day with
greater growth being favored with rehydrati on.
Thus plants may need t o solve a timing imbalance
between carbon gain and uti 1ization by changes
in internal storage. The higher internal carbon
status of leaves during the afternoon may reduce
the significance of pollutant impacts on leaves
during t h i s part of the day. Photosynthesis may
be already slowed by product accumulation, or
alternatively detoxification mechanisms using
readily avai 1able carbon metabolites and/or
energy may more easily cope with pollutant in-
s u l t than during early morning when internal
carbon status i s lower.
The diurnal pattern of behavior (fig. 6) also
suggests that root exudation of carbon compounds
could be f a c i l i t a t e d during the day. In the
same way, the carb6n supply t o mycorrhizae and SUMMARY

root nodules may be f a c i l i t a t e d . Disruption of

these processes through the impact of a i r pol- Modeling of pollutant i n t e r a c t i o n s with whole
l u t a n t s may be of great importance t o under- pl ant processes has provi ded :
1
standing who1 e pl ant responses. Pollutant Insights about t h e processes and t h e i r
s t r e s s t h a t causes reduced ,photosynthesis and/or i n t e r r e l a t i o n s h i p s , e.g.,
greater r e s p i r a t i on in f o r e s t ecosystems may ( a ) Transpiration may f aci 11i t a t e pol -
decrease t h e carbon 1eakage t o mycorrhizal asso- l u t a n t uptake by transporting chemical
c i a t i o n s with roots, p o t e n t i a l l y decreasing the from roots t o stem thus maintaining a
extent and e f f i c i e n c y of t h e fungi in supplying favorable chemical gradient f o r continu-
n u t r i e n t s back t o the t r e e . T h u s , i t may be ing uptake.
f u r t h e r hypothesized t h a t phytotoxic a i r pollut- ( b ) Phloem and xylem may provide ready
ants may cause f o r e s t ecosystems t o be l e s s transport pathways f o r pollutant movement
e f f i c i e n t in nutrient r e t e n t i on ( i .e., become between plant p a r t s ( f i g . 3 ) .
more leaky, see also OINeill et a1. 1977) and I d e n t i f i c a t i o n of plant properties im-
conversely beneficial a i r p o l l u t a n t s may in- portant in pollutant uptake. In particu-
crease n u t r i e n t r e t e n t i on of f o r e s t ecosystems. 1a r , 1eaf and root chemical conductivity
Elevated atmospheric CO2 l e v e l s may be an ex- have great influence on pollutant uptake
ample of t h e l a t t e r . ( f i g . 5, t a b l e 2 ) .
Alternative hypotheses, e.g., increased
f o r e s t 1i t t e r in areas polluted with
heavy metals could be due t o increased
I PHOTOSYNTHESISRECHARGESPL.hT WITH SUCROSE
2 'LOWER WAlEm POTEhTIAL REDUCES CELL EXPANStON IMINIMUM G R M H RATE1
mortality of plant p a r t s in addition t o
3. +ACTIVE SOLUTE UPTAKE FROM MASSFLOW POOL reduced decomposition r a t e .
5. ROOT EXUDATION PROMOTED. LESS ROOT SLOUGHING
& FASTERPHLOEMTRANSLOCATIONOFSUCROSEFROMLEAVESTOSTEMSANDROOTS
A basis f o r short-term ( d i u r n a l ) and
7 FASTER XYLEM TRANSPORT OF SOLUTES AND METABOLLTES FROM ROOTS TO STEMS AND LEAVES long-term speculation or pollutant
impacts, e.g.,

SUCROSE

CYTOPLASM
- STARCH
( a ) Hourly changes in water, carbon, and
nutrient s t a t u s of pl ants may inf 1uence
physiological s e n s i t i v i t y t o pollutant
insult.
( b ) Pollutant disruption of carbon a l l o -
cation t o be1 owground processes may have
long-term n u t r i e n t cycling impacts.

I'
/
,
I SUCROSEP
SUELL
-
-

SOLUTES IN -^----P
CYTOPLASM
STARCH
SHRINK
SOLUTES IN
VACUOLE
\,
\
\,
LITERATURE CITED
Baes, C. F., C. L. Begovich, W. M. Culkowski,
K. R. Dixon, D. E. Fields, J. T. Holdeman,
D. D. Huff, D. R. Jackson, N. M. Larson,
NIGHT CONDITION
R. J. Luxmoore, J. K. Munro, M. R. Patterson,
R. J. Raridon, M. Reeves, D. C. S t e i n ,
J. L. Stolzy, and T. C. Tucker.
1976. The unified transport model. In
-
4. METABOLITE CYTOPLASM
S. ROOT SLOUGHING PROMOTED. LESS EXUDATION Ecology and analysis of t r a c e contaminants
-
6. SLOWER PHLOEM TRANSLOCATION REDUCED SUCROSEGRADIENT
progress report October 1974-December 1975.
7. ,SLOWER XYLEMSOLUTE AND METABOLITE FLUX - REDUCEDTRANSPIRATION
R. I. Van Hook and W. D. Shults, eds.,
.OOMINANT EhERGV UTIL#.?ATsONFOR T M E PERIOD
pp. 13-62. ORNL/NSF/EATC-22. Oak Ridge
N D L C E D B Y ROOT RESISTANCE TO TRANSPIRATION National Laboratory, Oak Ridge, Tennessee
37830. 200 pp.
Baldwin, J. P., P. B. Nye, and P. B. Tinker.
Figure 6--Diurnal pattern of carbon, water, and 1973. Uptake of solutes by multiple root
solute dynamics showing re1 a t i ve tendencies and systems from s o i l , 111. A model f o r calcu-
r e l a t i v e s t a t e s i n vegetation. l a t i n g the solute uptake by a randomly
dispersed root system developing in a f i n i t e
volume of s o i l . Plant Soil 38:621-635.
Begovich, C. L., and D. R. Jackson.
Perhaps, l i k e Humpty Dumpty, these attempts 1975. Documentati on and appl i c a t i on of SCEHM.
a t deriving whole system understanding from the A model of s o i l chemical exchange of heavy
pieces involved shows many cracks and flaws. metals. ORNL/NSF/EATC-16. Oak Ridge
Nevertheless, we give i t a go! The key t e s t i s National Laboratory, Oak Ridge, Tennessee.
our answer t o the question "Did we learn some- 67 PP.
thing t h a t we d i d n ' t know before?"
Begovich, C. L., and R. J. Luxmoore.
1979. Some s e n s i t i v i t y studies of chemical
transport simulated in models of the s o i l -
plant- 1i t t e r system. ORNL/TM-6791. Oak
Ridge National Laboratory, Oak Ridge,
Tennessee. 97 pp.
Dixon, K. R., R. J . Luxmoore, and C. L. Begovich.
1978. CERES - A model of f o r e s t stand biomass
dynamics f o r predicting t r a c e contaminant,
nutrient and water e f f e c t s . I. Model
description, 11. Model documentation. Ecol.
Model. 5: 17-38. 93-114.
Houston, D. B., and G. R. S t a i r s .
1973. Genetic control of s u l f u r dioxide and
ozone tolerance i n eastern white pine. For.
.
Sci 19: 267-271.
Huff, D. D., R. J. Luxmoore, J . B. Mankin, and
C. L. Begovich.
1977. TEHM. A t e r r e s t r i a1 ecosystem
hydrology model. ORNL/NSF/EATC-27. Oak
Ridge National Laboratory, Oak Ridge,
Tennessee. 153 pp.
Jackson, D. R., and A. P. Watson.
1977. Description of n u t r i e n t pools and
transport of heavy metals i n a forested
watershed near a lead smelter. J. Environ.
.
Qual 6: 331-338.
Jensen, K. F., and T. T. Kozl owski.
1975. Absorption and trans1 oeati on of s u l f u r
dioxide by seedlings of f o u r f o r e s t t r e e
species. J. Environ. Qual. 4:379-382.
Kercher, J. R.
1977. GROW1: A crop growth model f o r
assessing impacts of gaseous p o l l u t a n t s f rom
geothermal techno1 ogi es. UCRL-52247.
Lawrence Li vermore Laboratory, Cal i f orni a.
Lawhon, W. T.
1973. Radial growth and wood density of white
pi ne in re1 a t i on t o coal -deri ved envi ron-
mental pollutants. Ph.D. d i s s e r t a t i o n .
Graduate program in ecology, University of
Tennessee, Knoxvi 11e , Tennessee. 110 pp.
Luxmoore, R. J., and C. L. Begovich.
1979. Simulated heavy metal fluxes in t r e e
microcosms and a deciduous f o r e s t . Internal.
Soc. Ecology. Model1 ing. J. 1:48-60.
Luxmoore, R. J., C. L. Begovich, and K. R. Dixon.
1978. Modeling solute uptake and
incorporation i n t o vegetation and l i t t e r .
Ecol. Model. 5:137-171.
O'Neill, R. V., B. S. Ausmus, D. R. Jackson,
R. I. Van Hook, P. Van Voris, C. Washburne, and
A. P. Watson.
1977. Monitoring t e r r e s t r i a1 ecosystems by
analysis of nutrient export. Water, Air,
and Soil Pollut. 8:271-277.
Roberts, B. R.
1974. Fol i ar sorption of atmospheric s u l f u r
dioxide by woody plants. Environ. Pollut.
7:133-140.
Thompson, C. R., D. C. Taylor, M. D. Thomas, and
J. 0. Ivie.
1967. Effects of a i r pollutants on apparent
photosynthesis and water use by c i t r u s
t r e e s . Environ. Sci. Technol. 1:664-650.
Watson, A. P., R. I. Van Hook, D. R. Jackson,
and D. E. Reichle.
1976. Impact of a lead mining-smelting
complex on the f o r e s t f l o o r l i t t e r arthropod
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 Data-Based Ecological Modeling of
Ozone Air Pollution Effects in a Southern
California Mixed Conifer Ecosystem1

Ronald N. K i c k e r t and Barbara emm mill^

A b s t r a c t : The purpose of t h i s r e s e a r c h was t o determine t h e

e f f e c t s of ozone a i r p o l l u t i o n on a mixed c o n i f e r f o r e s t
ecosystem i n t h e San Bernardino National F o r e s t , C a l i f o r n i a .
We used an e c o l o g i c a l systems modeling approach i n con-
c e r t w i t h v a r i o u s b i o l o g i o a l s p e c i a l i s t s . This r. q u i r e d
conceptual model development, computer programming, and
t h e a n a l y s i s of o r i g i n a l p r o j e c t d a t a f o r model c a l i b r a t i o n .
fie found t h a t t h i s p r o c e s s l e d t o t h e i n v e s t i g a t o r s con-
d u c t i n g new r e s e a r c h o f an i n t e g r a t i v e n a t u r e . A s t r u c t u r e
f o r complex i n t e r a c t i o n s of f o r e s t e f f e c t s was produced.
I n s i g h t s on changes i n ecosystem dynamics and a worst-case
s c e n a r i o of f u t u r e f o r e s t changes were d e r i v e d . ,
We conclude t h a t sudden q u a l i t a t i v e changes i n c o n i f e r
f o r e s t composition can occur under t h e i n f l u e n c e of ozone
a i r p o l l u t i o n and t h e e x c l u s i o n of n a t u r a l f i r e e v e n t s .

I f i t were known t h a t a i r p o l l u t a n t s d i d n o t on The b i o l o g i c a l , e c o l o g i c a l , and p h y s i c a l e n v i -

a f f e c t people and t h e i r environments, s o c i e t y
would be l i k e l y t o have l i t t l e i n t e r e s t i n t h o s e
p o l l u t a n t s . The c e n t r a l i s s u e i s "What a r e t h e
effects?".
INSTITUTIONAL SETTING
ronment, t h e secondary s t a n d a r d . Recently, t h e
s t a n d a r d s were r a i s e d from 0.08 t o 0.12 ppm f o r one
hour p e r y e a r (u.s. Environmental P r o t e c t i o n Agency
1979). I n view of t h e f a c t t h a t knowledge o f p o l l -
u t a n t e f f e c t s c o n t i n u e s t o develop, t h e c r i t e r i a
f o r j u s t i f y i n g t h e l e g a l s t a n d a r d i s expected t o
be re-evaluated e v e r y few y e a r s .

I n t h e United S t a t e s , N a t i o n a l Ambient A i r
Q u a l i t y S t a n d a r d s f o r ozone have been l e g a l l y e s -
t a b l i s h e d w i t h a view f o r e f f e c t s on humans, t h e
primary s t a n d a r d , and s e p a r a t e l y f o r t h e e f f e c t s
p r e s e n t e d a t t h e Symposium on E f f e c t s o f A i r
P o l l u t a n t s on Mediterranean and Temperate F o r e s t
Ecosystems, June 22-27, 1980, R i v e r s i d e ,
California, U.S.A.
s e n i o r Systems Analyst and A s s i s t a n t
S p e c i a l i s t , r e s p e c t i v e l y , D i v i s i o n of
B i o l o g i c a l C o n t r o l , U n i v e r s i t y of C a l i f o r n i a ,
Albany, C a l i f .
THE PROBLEM
I n e v a l u a t i n g c r i t e r i a f o r d e c i d i n g upon t h e
secondary s t a n d a r d f o r ozone, i t has been recog-
nized throughout t h e 1970's t h a t b i o l o g i c a l and
e c o l o g i c a l e f f e c t s i n f o r m a t i o n was b i a s e d toward
t h e more r e d u c t i o n i s t i c l e v e l s , i . e . , biochemis-
t r y , p l a n t s c i e n c e , p l a n t physiology, and, because
of l o g i s t i c a l problems w i t h l a r g e r s p a t i a l and
time s c a l e s , biased a g a i n s t , o r a t l e a s t f a i l i n g
t o c o n s i d e r , e f f e c t s on " n a t u r a l " e c o l o g i c a l s y s -
tems i n t h e landscape. B i o l o g i c a l e f f e c t s c r i t e r i a
have been based on d a t a f o r i n d i v i d u a l organisms,
but t h e d i r e c t and i n d i r e c t e f f e c t s on p l a n t and
animal communities have been mostly s p e c u l a t i v e
(u.s. Environmental P r o t e c t i o n Agency 1978).
 H i s t o r y of The Study
O b j e c t i v e s of t h e E n t i r e P r o j e c t
I n 1973, t h e EPA e s t a b l i s h e d a s e v e r a l - y e a r
s t u d y of o x i d a n t e f f e c t s on t h e mixed c o n i f e r eco-
system i n t h e San Bernardino National F o r e s t . The
i n t e r p r e t a t i o n s d e r i v e d t o d a t e from t h i g p r o j e c t
have c o n s t i t u t e d t h e major s o u r c e of i n f o r m a t i o n
f o r t h e ecosystem c h a p t e r on a i r q u a l i t y c r i t e r i a
published by t h e EPA; however, t h e m a j o r i t y of t h e
d a t a i n t e g r a t i o n remains t o be completed. A s a f u r -
t h e r f o c u s o f t h e p r o j e c t , two y e a r s a f t e r i t was
i n i t i a t e d , t h e s e n i o r a u t h o r was brought i n t o
i n t r o d u c e computer s i m u l a t i o n a s a t o o l i n g u i d i n g
t h e c o l l e c t i o n , i n t e g r a t i o n and i n t e r p r e t a t i o n of
d a t a on f o r e s t r e s p o n s e s t o o x i d a n t s t r e s s . The
p o t e n t i a l use of t h i s s t u d y i n f u t u r e policy-making
r e q u i r e d an emphasis i n t h e modeling e f f o r t p a r t i c -
u l a r l y on long-term e f f e c t s , p r o j e c t e d e f f e c t s a t
d i f f e r e n t t h e o r e t i c a l l e v e l s of o x i d a n t f l u x , and
e f f e c t s on t h e b e h a v i o r of t h e n a t u r a l community
a s opposed t o i n d i v i d u a l organisms.
O b j e c t i v e s o f t h e Modeling A c t i v i t y
S t r u c t u r a l Simplification--The g o a l of t h e ecosys-
tem modeling e f f o r t i s twofold, and e q u a l l y d i -
v e r s e i n each d i r e c t i o n . Due t o t h e n a t u r e of t h e
SBNF p r o j e c t , i t was r e q u i r e d t h a t t h e modeler be-
g i n w i t h a l o c a l i z e d , real-world s i t u a t i o n and
make e x t e n s i v e u s e o f t h e l a r g e data-base i n con-
s t r u c t i n g t h e model. The real world s i t u a t i o n ,
from which t h e d a t a a r e d e r i v e d i s extremely v a r -
i a b l e , c o n s i s t i n g of an east-west t r e n d i n g moun-
t a i n range which i n c r e a s e s i n e l e v a t i o n and changes
i n s p e c i e s composition a l o n g t h e same g r a d i e n t of
o x i d a n t f l u x , such t h a t e s s e n t i a l l y no c o n t r o l a r e a s
a r e p o s s i b l e . Given such a complex system, The
f i r s t g o a l of t h e modeling a c t i v i t y was t o break
down t h i s system s t r u c t u r a l l y i n t o i t s s i m p l i f i e d ,
b a s i c components and d r i v i n g f a c t o r s .
Experiments and Model Behavior-- The o t h e r h a l f of
t h i s g o a l was t o p r o v i d e answers t o t h e q u e s t i o n :
how might one u s e a s i m u l a t i o n model f o r computer
experiments t o a s s e s s t h e t o t a l i t y of t h e s e e f f e c t s ,
a c t i n g a l o n e o r s y n e r g i s t i c a l l y , on ecosystem s t r u c -
t u r e and f u n c t i o n ? A l i s t of e f f e c t s does n o t h e l p
p o l i c y makers v e r y much when t h e y a r e i n t h e p o s i -
t i o n of making d e c i s i o n s i n t h e f a c e of u n c e r t a i n -
t y - even l e s s does i t inform b i o l o g i c a l l y know-
l e d g e a b l e people who r i g h t f u l l y s u s p e c t t h a t i n t e r -
a c t i o n s o c c u r between i t e m s on t h e l i s t t h a t w i l l
a f f e c t f u t u r e outcomes a s much o r more t h a n a sum-
mary of s i m p l e e f f e c t s could e v e r e x p r e s s . Thus,
t h e modeling e f f o r t h a s been developed t o a d d r e s s
the following questions:
I
The EPA/SBNF P r o j e c t h a s attempted t o e s t a b -
l i s h e f f e c t s o f ozone a i r p o l l u t i o n on t r e e
stem growth, f o l i a r i n j u r y , t r e e m o r t a l i t y ,
r e g e n e r a t i o n , cone p r o d u c t i o n , n u t r i e n t c y c l -
i n g , and i n s e c t and d i s e a s e occurrence.
What i s t h e consequence of t h e s e e f f e c t s when
combined t o g e t h e r i n a s i m u l a t e d ecosystem?
What time s c a l e i s n e c e s s a r y t o u s e t o s e e t h e
f u l l long-term ecosystem e f f e c t s ? Is t h e r e
p o t e n t i a l i n t h i s system f o r sudden jumps and
i r r e v e r s i b l e trends?
The modeling methods and philosophy used i n t h e
p r o j e c t have been d e s c r i b e d i n p r e v i o u s p u b l i c a t i o n s
( ~ i c k e r t1977a, 1977b, 1980).
RESULTS
Because t h e modeling a c t i v i t y i s s t i l l b e i n g
conducted, t h e r e s u l t s p r e s e n t e d h e r e a r e n o t based
on experiments performed on t h e computer u s i n g t h e
models. R a t h e r , t h e y a r e based on i n s i g h t s gained
d u r i n g t h e model development p r o c e s s , from concep-
t u a l i z a t i o n , t o mathematical f o r m u l a t i o n , t o compu-
t e r coding, and a n a l y s i s of o r i g i n a l d a t a toward t h e
g o a l of c a l i b r a t i n g t h e models f o r t r e e s p e c i e s and
s i t e s w i t h i n t h e SBNF and t h e n a p p l y i n g t h o s e mod-
e l s i n experiments of a i r p o l l u t i o n e f f e c t s .
How t h e Systems Modeling P r o c e s s Aided t h e P r o j e c t
There a r e two ways i n which t h e t o t a l s t u d y was
improved. Model development a i d e d t h e p r o j e c t i n -
v e s t i g a t o r s i n viewing t h e i r own work a s a p a r t o f
a n i n t e g r a t e d conceptual s t r u c t u r e . Also, w i t h t h e
d e s i g n of a g r a p h i c model of v a r i o u s subsystems,
discussions with investigators led t o t h e i d e n t i f i -
c a t i o n of q u e s t i o n s s u b s e q u e n t l y t u r n e d i n t o r e -
s e a r c h which o t h e r w i s e would n o t have been done.
A mixed t r e e s p e c i e s p o p u l a t i o n dynamics approach
l e d t o s e e d l i n g e s t a b l i s h m e n t experiments, s t u d y
p l o t s e e d l i n g r e g e n e r a t i o n s u r v e y s , and a compre-
h e n s i v e p e s t damage i n v e n t o r y , t o determine m o r t a l -
i t y p a t t e r n s . Data needed f o r c a l i b r a t i n g a s t a n d
moisture model l e d t o a seismograph survey f o r p l o t
s o i l d e p t h s which i n d i c a t e d s o i l w a t e r m o n i t o r i n g
p r o f i l e s were t o o s h a l l o w on s e v e r a l p l o t s . I n f o r -
mation r e q u i r e d f o r r o o t d i s e a s e and b a r k b e e t l e dy-
namics l e d t o more c a u t i o u s u s e of t h e smog i n j u r y
s c o r i n g procedure, a s well a s t o t h e dendrochrono-
l o g i c a l a n a l y s i s o f t r e e r a d i a l growth.
A S t r u c t u r e f o r Complex I n t e r a c t i o n s
The e f f e c t s o f a i r p o l l u t i o n i n t h e f o r e s t eco-
system a r e n o t o n l y t h e d i r e c t v i s i b l e e f f e c t s
t h a t a c a s u a l o b s e r v e r might n o t i c e by d i s c o l o r e d
f o l i a g e on t h e t r e e s , b u t a l s o l e s s a p p a r e n t , b u t
nonetheless r e a l , i n d i r e c t e f f e c t s t h a t a r e s u b t l y
t r a n s f e r r e d through t h e system. Such i n d i r e c t c h a i n
r e a c t i o n s can o c c u r a t t h e l e v e l of i n d i v i d u a l
t r e e s , a t t h e p o p u l a t i o n l e v e l of t r e e s of a c e r -
t a i n s p e c i e s , and because of changes i n t h e m i x t u r e s
o f t h e l a t t e r over t h e long-term, changes which
occur a t t h e whole community l e v e l . A s a map of how
such changes can be t r a n s f e r r e d throughout t h e s y s -
tem, f i g u r e 1 d i s p l a y s some s i g n i f i c a n t p o r t i o n s
of a f o r e s t ecosystem which must be c o n s i d e r e d . The
r e f e r e n c e numbers a s s o c i a t e d w i t h each component i n
t h i s diagram p e r t a i n t o v a r i o u s k i n d s of environ-
mental c o n d i t i o n s and b i o l o g i c a l organisms impor-
t a n t t o understanding changes o c c u r r i n g i n a f o r e s t
ecosystem. These numbers a l s o r e f e r e n c e p a r t i c u l a r

Figure 1--Components of t h e f o r e s t ecosystem d i r e c t l y and i n d i r e c t l y
a f f e c t e d by photochemical a i r p o l l u t i o n .
r e s e a r c h t o p i c s which have been s t u d i e d between
1973 through 1980 i n t h e San Bernardino National
Forest.
The purpose of t h i s overview i s t o p r e s e n t a n
i n t e g r a t e d , s i m p l i f i e d frame of r e f e r e n c e w i t h i n
which t h e d i s c o v e r i e s , r e s u l t s , and c o n c l u s i o n s from
t h e p r o j e c t may be viewed a s a whole.
While a f o r e s t i s more t h a n simply a group of
t r e e s , t h e l a t t e r i s by d e f i n i t i o n t h e dominant
l i f e form of such a system. Reference w i l l be made
t o t h e numbers i n v a r i o u s p a r t s of f i g u r e 1 . To
view t h e e f f e c t of a i r p o l l u t i o n ( 1 ) on a community
of t r e e s , i t i s n e c e s s a r y t o c o n s i d e r s e v e r a l r e l a -
t i v e l y s t a t i c s i t e and s o i l p r o p e r t i e s ( 7 ) , a s w e l l
a s v e r y dynamic m e t e o r o l o g i c a l c o n d i t i o n s such a s
a i r temperature and p r e c i p i t a t i o n ( 3 ) . s i n c e a l l of
t h e s e may c o n t r i b u t e t o a s y n e r g i s t i c e f f e c t of a
long-term, c h r o n i c a i r p o l l u t i o n exposure i n terms
of t r e e response. Some of t h e p r e c i p i t a t i o n ( 3 ) ,
depending on s i t e and s o i l c h a r a c t e r i s t i c s ( 7 ) ,
e n t e r s t h e s o i l a s a v a i l a b l e s o i l water ( 2 ) f o r
t r e e growth ( 3 , 9 ) . F o r l a c k of b e t t e r d a t a , we en-
v i s i o n a i r temperature ( 3 ) a s a rough index o f h e a t
a v a i l a b l e f o r e n a b l i n g a v a i l a b l e s o i l water t o be
d e p l e t e d through w a t e r l o s t from t r e e l e a v e s t o t h e
atmosphere through t r a n s p i r a t i o n .
Over time, and depending on s e n s i t i v i t y between
and w i t h i n v a r i o u s t r e e s p e c i e s , some of t h e g r e e n
f o l i a g e ( 4 , 5 ) on t r e e s becomes i n j u r e d , d i s c o l o r e d
f o l i a g e ( 4 , 5 ) , and some of t h a t i s dropped from t h e
t r e e s . T h i s , added t o normal amounts o f n e e d l e shed
a f f e c t e d by t h e a v a i l a b i l i t y of s o i l m o i s t u r e ( 2 ) ,
becomes a p a r t of ground l i t t e r ( 6 ) .
OIHER L E M A L DEAD r R E E S
The r e l a t i v e amount o f f o l i a g e t h a t changes
from green t o i n j u r e d i s thought t o have a b e a r i n g
on t h e r a t e of stem wood growth ( 8 , g ) o f t r e e s .
These t h r e e responses a r e thought t o be a s s o c i a t e d
with t h e r a t e of production of cones and t h e r e f o r e
seeds (13) f o r r e g e n e r a t i o n of new t r e e s . A s p o l l u -
t a n t s l e a d t o a g r e a t e r degree o f f o l i a g e i n j u r y
f o r some t r e e s p e c i e s , and stem growth i s reduced,
otherwise mature i n d i v i d u a l s o f t h e s e s p e c i e s pro-
duce l e s s and l e s s cones, i f any.
It h a s been mentioned how a i r p o l l u t i o n can i n -
c r e a s e t h e ground l i t t e r depths ( 6 ) . This i s s i g n i -
f i c a n t because s e e d s (1 3) of some t r e e s p e c i e s
have a b i o l o g i c a l behavior which i s adapted t o l i t -
t l e o r no ground l i t t e r f o r s p r o u t i n g and s u r v i v -
i n g 3s s e e d l i n g s (14,15) d u r i n g d r y summers ( t h e
e f f e c t o f a v a i l a b l e s o i l water ( 2 ) once a g a i n ) .
Many cones, s e e d s , and small s e e d l i n a s a r e l o s t t o
w i l d l i f e of v a r i o u s forms under n a t u r a l c o n d i t i o n s .
Any f u r t h e r r e d u c t i o n i n t h i s r e p r o d u c t i o n c h a i n
because of a i r p o l l u t i o n e f f e c t s can make continued
replacement o f some t r e e s p e c i e s a v e r y p r e c a r i o u s
circumstance.
Those s e e d l i n g s t h a t do s u r v i v e t h e f i r s t few
y e a r s e v e n t u a l l y grow t o a l a r g e r s i z e o f t e n c a l l e d
s a p l i n g s (1 6) i n t h e p o p u l a t i o n s t r u c t u r e . The ex-
t e n t t o which t h e e f f e c t o f a i r p o l l u t i o n r e t a r d s
stem growth (8.9) simply t e n d s t o keep t r e e s i n t h i s
s i z e range f o r a l o n g e r t i m e , s u b j e c t t o t h e many
causes o f d e a t h which can occur. E v e n t u a l l y , some
s a p l i n g s grow i n t o l a r g e r s i z e s which a r e mature
(16) and p o t e n t i a l l y capable of producing cones, a s
w e l l a s b e i n v a l u e d f o r e s t h e t i c purposes and a s
p o t e n t i a l l y merchantable timber.
Those mature t r e e s t h a t develop a s i g n i f i c a n t
degree of v i s i b l e f o l i a g e i n j u r y ( 4 , 5 ) a r e some-
t i m e s c u t down whether l e g a l l y o r by poaching. T h i s
produces stumps. I t h a s been discovered t h a t t h e
d e g r e e of f o l i a g e i n j u r y ( 4 , 5 ) , namely t h e younger
t h e age of t h e o l d e s t n e e d l e s , i s s i g n i f i c a n t l y
c o r r e l a t e d with t h e i n c i d e n c e of i n f e c t i o n and c o l -
o n i z a t i o n of such stumps by t h e r o o t r o t d i s e a s e
( 1 0 ) Fomes annosus, i f t h e f r e s h stumps a r e n o t
t r e a t e d e n t h e l i v e t r e e i s f i r s t c u t . One would
h a r d l y be concerned about t h i s i f i t were n o t f o r
t h e o b s e r v a t i o n t h a t such d i s e a s e s can spread from
dead stump r o o t systems i n t o a d j a c e n t l i v e t r e e r o o t
systems, from s e e d l i n g s , up t o l a r g e , mature, p o l l -
u t i o n - r e s i s t a n t t r e e s , depending on t h e s p a t i a l
d e n s i t y of t h e s t a n d . When t h i s does happen w i t h
t h e l a t t e r , and t h o s e t r e e s a l r e a d y have a c o n s i d e r -
a b l e d e g r e e of i n j u r e d f o l i a g e ( 4 , 5 ) , t h e n , espec-
i a l l y on ponderosa p i n e , bark b e e t l e s (1 1 ) appear
b e t t e r a b l e t o s u c c e s s f u l l y a t t a c k and k i l l such
t r e e s . M o r t a l i t y s u r v e y s show numerous o t h e r p a t -
t e r n s o f d i s e a s e and i n s e c t combinations (1 2) a l s o
a c t t o k i l l t r e e s . Reduced growth cannot be s u s -
t a i n e d i n d e f i n i t e l y , and t h e l e n g t h of t i m e of r e -
duced growth p r i o r t o d e a t h a p p e a r s t o be age-
dependent.
Over t i m e , t h e complexion of t h e f o r e s t ecosys-
tem w i l l change a c c o r d i n g t o which t r e e s p e c i e s a r e
b e s t a b l e t o r e s i s t t h e agents t h a t cause death
( 1 6 ) , whether n a t u r a l o r human-caused, and a r e a l s o
c a p a b l e of p r o v i d i n g new young s e e d l i n g s (1 4,15)
a b l e t o s u r v i v e t o m a t u r i t y . I n many c a s e s , t h e
evidence seems t o i n d i c a t e t h a t t h e b a l a n c e between
various t r e e species i s s h i f t i n g dramatically i n
t h e San Bernardino N a t i o n a l F o r e s t .
Given t h r e e d i r e c t e f f e c t s of a i r p o l l u t a n t s on
f o r e s t growth, namely f o l i a r i n j u r y and a c c e l e r a t e d
n e e d l e c a s t , woody growth r e d u c t i o n , and i n d i r e c t l y
i n c r e a s e d m o r t a l i t y , t h e s e e f f e c t s might be t i e d
t o g e t h e r i n t h e f o l l o w i n g s c e n a r i o s of ecosystem-
level effects.
Ecosystem Dynamics Under a S i n g l e S t r e s s
F o l i a r I n j u r y Consequences-- F o l i a r i n j u r y and pre-
mature shedding of p a s t y e a r ' s n e e d l e s w i l l slow
down t h e n a t u r a l development of canopy c l o s u r e i n a
s t a n d . It h a s been shown and r e p e a t e d l y confirmed
t h a t a s a s t a n d grows, l e a f a r e a expands u n t i l i t
r e a c h e s a p l a t e a u , a t which i t remains f o r t h e r e -
mainder o f t h e l i f e of t h e s t a n d ( ~ r i e rand o t h e r s
1978) ( f i g . 2 ) . Stand growth, developmental p a t t e r n s
and time t o m a t u r i t y a r e e n t i r e l y dependent
upon t h e r a t e o f canopy c l o s u r e . Net p r o d u c t i o n by
coniferous f o r e s t s is related t o l e a f area
( W h i t t a k e r and N i e r i n g l 9 7 5 ) , and a l l o t h e r t h i n g s
being equal, t h e g r e a t e r t h i s l e a f area, the g r e a t e r
i s t h e p r o d u c t i v i t y . Once maximal l e a f a r e a (cano-
py c l o s u r e ) i s o b t a i n e d , o t h e r ecosystem f u n c t i o n s
b e g i n t o make major q u a l i t a t i v e changes, a s d i s c u s s -
ed below. I f time t o canopy c l o s u r e i s i n c r e a s e d ,
n o t o n l y a r e p r o d u c t i v i t y r a t e s reduced, b u t a l s o
q u a l i t a t i v e e c o l o g i c a l changes may occur.
S i n c e f o r e s t f o l i a g e always t e n d s toward form-
i n g a c o n t i n u o u s , complete s u r f a c e a r e a , t h e degree
of completion r e p r e s e n t s t h e d e g r e e of occupancy of
t h e s i t e . A vigorous, healthy stand of t r e e s w i l l
STAND AGE- YEARS
Figure 2--Natural f o r e s t s t a n d canopy c l o s u r e .
achieve occupancy o f a n open s i t e a t about 1 2 y e a r s
of age, e s t a b l i s h i n g dominance o v e r competing vege-
t a t i o n , and w i l l m a i n t a i n occupancy u n t i l a t l e a s t
middle age ( s m i t h 1962). A given g e n e r a t i o n o f
t r e e s u l t i m a t e l y l o s e s command of t h e s i t e , g i v i n g
way t o younger members a n d / o r o t h e r s p e c i e s . A'for-
e s t canopy e x p e r i e n c i n g p o l l u t i o n - c a u s e d i n j u r y
might n o t be a b l e t o f u l l y e s t a b l i s h occupancy of
t h e s i t e . The amount o f accompanying v e g e t a t i o n ,
e s p e c i a l l y of an u n d e r s t o r y n a t u r e , might i n d i c a t e
t h e d e g r e e t o wh'ich t h e main s t a n d f a l l s s h o r t of
f u l l occupancy.
Growth Reduction Consequences-- Leaf a r e a i s d i r e c t -
l y r e l a t e d t o woody p r o d u c t i o n i n a f o r e s t s t a n d ,
a s mentioned. Impediments t o l e a f a r e a expansion
and canopy c l o s u r e might a f f e c t t h e o v e r a l l p a t t e r n
of woody growth i n a s t a n d . S t u d i e s have shown t h a t
second-year n e e d l e s a r e more i m p o r t a n t i n p r o v i d i n g
photosynthate f o r stem growth, while c u r r e n t y e a r
n e e d l e s c o n t r i b u t e p r i m a r i l y t o s h o o t and n e e d l e
e l o n g a t i o n (walker and o t h e r s 1972); t h e second
y e a r n e e d l e s a r e t h e most impacted by a i r p o l l u -
t i o n . Trees might c o n t i n u e t o p u t on h e i g h t growth
a t t h e expense of d i a m e t e r growth f o r a l o n g e r per-
i o d of time t h a n with a normally c l o s i n g canopy,
both because of a l a c k of p h o t o s y n t h a t e f o r stem
growth and because t h e open canopy f o s t e r s r a p i d
h e i g h t growth r a t e s . However, because growth and
p r o d u c t i v i t y i s s u p p r e s s e d , both h e i g h t and diame-
t e r growth r a t e s i n g e n e r a l would be much s l o w e r
t h a n normal.
I t i s a t e n e t o f s i l v i c u l t u r e t h a t changes i n
s t a n d d e n s i t y do n o t s i g n i f i c a n t l y a l t e r t h e t o t a l
amount of d r y m a t t e r o r stem wood produced by a
s t a n d . Thus, precommercial t h i n n i n g s do n o t marked-
l y change production r a t e s b u t r a t h e r add t h e same
amount of wood t o a l e s s e r number o f t r e e s ( s m i t h
1962). Mathematically, mean p l a n t s i z e m u l t i p l i e d
by d e n s i t y t e n d s toward a c o n s t a n t . T h i s r e l a t i o n -
s h i p h a s been confirmed t o hold t r u e f o r many veae-
t a t i o n t y p e s , i n c l u d i n g f o r e s t s t a n d s (cooper
1961). I n a p o l l u t i o n s t r e s s e d f o r e s t , however, one
might expect t h e r e l a t i o n s h i p e i t h e r t o be weak, o r
t o break down a l t o g e t h e r ( f i g . 3 ) . A t low d e n s i -
 I -unstressed forest
STAND DENSITY - TREES/ ACRE
Figure 3--Relation between stand d e n s i t y and average
t r e e s i z e with and without a i r p o l l u t i o n .
t i e s , mean p l a n t s i z e i s not remarkably l a r g e a s
t h e t r e e s a r e n o t i n good v i g o r ; a t high d e n s i t i e s ,
p l a n t s i z e i s suppressed even more than would be
expected due t o d e n s i t y e f f e c t s alone. I n e s p e c i a l -
l y damaged and open s t a n d s , t h e r e l a t i o n s h i p might
be expected t o break down. The functioning of t h e
r e l a t i o n s h i p i s dependent upon t r e e s e x e r t i n g an
i n f l u e n c e over each o t h e r . I f a l l t r e e s a r e gener-
a l l y weakened i n competitive a b i l i t y , t h e i r growth
w i l l depend more on l i m i t a t i o n s imposed by t h e phy-
s i c a l environment and l e s s on i n t e r - t r e e i n f l u e n c e s .
An open stand with highly v a r i a b l e t r e e s i z e s might
be t h e r e s u l t .
Tree M o r t a l i t y Consequences-- T y p i c a l l y , a stand of
t r e e s achieves a d e n s i t y i n balance with t h e physi-
c a l environment by means of d e a t h of l e s s competi-
t i v e t r e e s i n t h e s t a n d . Thus, t h e r e i s g e n e r a l l y
a continuous d e c l i n e i n stand d e n s i t y with i n c r e a s -
i n g stand development. I n a p o l l u t i o n - s t r e s s e d f o r -
e s t , one f i n d s t h e phenomenon of s e l e c t i v e m o r t a l i -
t y . However, t h e magnitude of t h e m o r t a l i t y can be
g r e a t e r and t h e cause i s o t h e r than simple competi-
t i v e s t r e s s . A s young t r e e s prematurely age, t a p e r
o f f i n growth, and d i e , more openings a r e c r e a t e d
i n t h e canopy. While i n a h e a l t h y f o r e s t t h i s pro-
vides room f o r t h e dominant t r e e s t o expand, i n
p o l l u t i o n - s t r e s s e d f o r e s t s t h e open canopy i s open-
ed f u r t h e r . Even i f t h e p o l l u t i o n should be removed
from t h e system, a lower stand d e n s i t y because of
increased m o r t a l i t y r a t e s r e q u i r e s a g r e a t e r time
t o develop a f u l l canopy than would a stand with a
more normal s t o c k i n g r a t e .
P o t e n t i a l Responses Under Multiple S t r e s s e s
Dolan and Hayden (1978) c l a s s i f i e d types of
changes i n n a t u r e r e s e r v e park ecosystems a s e i t h e r
steady s t a t e , eddy s t a t e , o r t r e n d s t a t e . Steady
s t a t e changes i n c l u d e d i u r n a l and seasonal environ-
mental changes under which t h e system has evolved.
Eddy s t a t e changes a r e d i s c r e t e pulses of environ-
mental d i s t u r b a n c e s . Trend s-cate changes a r e long-
term changes t h a t a r e o f t e n t h e most s u b t l e t o de-
t e c t , a s well a s t h e most d i f f i c u l t from which t o
p r o t e c t n a t u r a l landscape ecosystems i f those chan-
ges a r e a s s o c i a t e d with human i n f l u e n c e s . E f f e c t s
of a i r p o l l u t i o n on f o r e s t s can have t h i s trend
s t a t e nature.
A Worst-case Scenario of Future Forest Change--
I t could be i n s i g h t f u l t o i n t e g r a t e t h e present
s t a t e of e c o l o g i c a l understanding on combinations
of trend s t a t e changes i n an attempt t o s e e what a
p o s s i b l e worst-case scenario might be f o r v e g e t a t i o n
i n western coniferous f o r e s t s . The r e l a t i v e o r d e r
of s e n s i t i v i t y , conceived a s p r o b a b i l i t y of mortal-
i t y a s s o c i a t e d with a p a r t i c u l a r environmental
s t r e s s , i s o f t e n d i f f e r e n t (even opposite) between
various s p e c i e s f o r one s t r e s s , compared t o another.
Ranking t r e e s p e c i e s of mature i n d i v i d u a l s i n terms
of l i k e l y m o r t a l i t y t o f i r e would place ponderosa
and lodgepole pine a s "low", while white f i r and
incense cedar would o f t e n be r a t e d "high". The l a t -
t e r would be k i l l e d by a moderate i n t e n s i t y s u r f a c e
f i r e ( n o t a prescibed burn n e c e s s a r i l y ) .
I n c o n t r a s t , research on ambient oxidant a i r
p o l l u t i o n s e n s i t i v i t y has shown ponderosa pine a s
very s e n s i t i v e , while white f i r and incense cedar
might have a low s e n s i t i v i t y t o t h i s s t r e s s . A i r
p o l l u t i o n weakens c e r t a i n t r e e s p e c i e s which a r e
subsequently h i t by b i o t i c d i s e a s e s and i n s e c t s ,
and produces a decreased competitive advantage,
compared t o l e s s s e n s i t i v e s p e c i e s . I n t h e commun-
i t y , t h i s can lead t o decreased l o n g e v i t y of t h e
sensitive species.
A s a worst case condition, one could envision
t h a t our present western f o r e s t h e r i t a g e from p r i s -
t i n e decades ago under a n a t u r a l f i r e frequency
s h i f t e d t h e balance of t r e e s p e c i e s composition such
t h a t i t was h e a v i l y proportioned with what a r e now
a i r p o l l u t i o n s e n s i t i v e s p e c i e s . I f t h e a i r pollu-
t i o n problem i n t e n s i f i e s over t h e y e a r s , t h e s e
s p e c i e s can be expected t o be decimated. The combin-
ed e f f e c t of both of t h e s e trend s t a t e changes,
each working on d i f f e r e n t t r e e s p e c i e s , micht make
it impossible t o preserve and p r o t e c t coniferous
f o r e s t s i n c e r t a i n l o c a t i o n s ( f i g . 4).
AIR 4
SENSITIVE,
FIRE-
TOLERANT
SPECIES
ABUNDANCE
v
0
A I R POLLUTION-TOLERANT.
FIRE- SENSITIVE SPECIES A B U N D A N C E
Figure 4--Possible f u t u r e course of f o r e s t s p e c i e s
composition under combined s t r e s s of a i r p o l l u t i o n
and n a t u r a l f i r e exclusion.
 CONCLUSIONS
I n t h e c a s e of a i r p o l l u t i o n , t h e r e could be t h e
g r a d u a l e l i m i n a t i o n of many f i r e - t o l e r a n t t r e e spe-
c i e s from f o r e s t s . Whenever a n a t u r a l f i r e does oc-
c u r under such a f u t u r e s c e n a r i o , t h e p r o p o r t i o n
of f o r e s t s t a n d s p e c i e s i n t h e f i r e - s e n s i t i v e c a t e -
gory could be much h i g h e r t h a n normal, and sudden
q u a l i t a t i v e changes, o r ecosystem c a t a s t r o p h e s , i n
c o n i f e r f o r e s t s p e c i e s composition could be expected.
There i s a s t r o n g l i k e l i h o o d t h a t c o n i f e r s t a n d s
might change i n t o mixtures of deciduous t r e e and
shrub communities a t mid-elevations, and perhaps
s c r u b f i e l d ecosystems a t h i g h e r e l e v a t i o n s which
p r e s e n t l y c o n t a i n c o n i f e r f o r e s t s . This would repre-
s e n t a q u a l i t a t i v e change from one s u c c e s s i o n a l p a t -
t e r n t o a n o t h e r , and i s a p o s s i b i l i t y which f o r e s t
management has a r e s p o n s i b i l i t y t o t r y t o e v a l u a t e .
Acknowledgments:
This s t u d y was funded i n p a r t with f e d e r a l funds
from t h e Environmental P r o t e c t i o n Agency under
Contract Numbers 68-03-0273, 68-03-2442, and
Grant Number R805410. The c o n t e n t of t h i s paper
i s not t o be construed a s r e p r e s e n t i n g views o r
p o l i c i e s of t h e EPA, n o r a s a concurrence of t h e
Agency with t h e r e s u l t s presented. Mention of
t r a d e names o r commercial products i n t h i s paper
does n o t c o n s t i t u t e e i t h e r an endorsement o r a
recommendation f o r t h e i r use. This paper does n o t
r e p r e s e n t EPA p o l i c y , p o s i t i o n , o r f i n d i n g s .
LITERATURE CITED
Cooper, C.F.
1961. Equations f o r t h e d e s c r i p t i o n of p a s t
growth i n even-aged s t a n d s of ponderosa pine.
For. Sci.:72-80.
Dolan, R. and B.P. Hayden.
1978. Environmental dynamics and r e s o u r c e
management i n t h e U.S. National Parks:Environ.
Manag. 2 ( 3 ) :249-258.
G r i e r , C.C., R.L. Edmonds, R.H. Waring and D.W.
Cole. 1978. F o r e s t management i m p l i c a t i o n s of
p r o d u c t i v i t y , n u t r i e n t c y c l i n g and water
r e l a t i o n s r e s e a r c h i n western c o n i f e r s . p.96-
106 I n : Proc., J n t Conv. of S.A.F. and Can.
I n s t . For., 1978.
K i c k e r t , R.N.
1977a. Toward f o r e c a s t i n g a l t e r n a t i v e f u t u r e
ecosystem responses: ecosystem modeling.
p.63-85 I n Photochemical A i r P o l l u t a n t E f f e c t s
on Mixed'Tonifer Ecosystems. Progress Report
1974-75. EPA-600/3-77-058. U. S. Environmental
P r o t e c t i o n Agency,.Corvallis Environmental
Research Laboratory, C o r v a l l i s , Oregon.
K i c k e r t , R.N.
1977b. D e f i n i t i o n of t h e c o n i f e r f o r e s t ecosys-
tem a s a group of coupled e c o l o g i c a l models.
p. 71 -1 05 I n Photochemical Oxidant A i r
P o l l u t i o n ' I T f f e c t s on a Mixed Conifer F o r e s t
Ecosystem--A Progress Report, 1976. Paul R.
M i l l e r and Michael J. Elderman, eds.
EPA-600/3-77-1 04. U. S. Environmental Protec-
t i o n Agency, C o r v a l l i s Environmental Research
Laboratory, C o r v a l l i s , Oregon.
K i c k e r t , R.N.
1980. Ecosystem s i m u l a t i o n modeling. Chpt. 2
- I n Photochemical Oxidant A i r P o l l u t i o n E f f e c t s
on a Mixed Conifer F o r e s t Ecosystem. O.C.
.
Taylor (ed. ) U.S. Environmental P r o t e c t i o n
Agency, Environmental Research Laboratory,
C o r v a l l i s , Oregon. EPA-600/3-80-002. 1 95 p
Smith, D.M.
1962. P r a c t i c e of S i l v i c u l t u r e . John Wiley and
Sons, New York. 578p.
U.S. Environmental P r o t e c t i o n Agency.
1978. A i r q u a l i t y c r i t e r i a f o r ozone and o t h e r
photochemical oxidants. EPA-600/8-78-004.
Volume 11. O f f i c e of Research and Development.
Washington, D.C. 341p.
U.S. Environmental P r o t e c t i o n Agency.
1979. EPA changes ozone s t a n d a r d t o 0.12 ppm.
Environmental News, January 26, O f f i c e of
P u b l i c Awareness, Washington, D.C. 3p.
Walker, R.B., D.R.M. S c o t t , D . J . S a l o and K.L. Reed.
1972. T e r r e s t r i a l process s t u d i e s i n c o n i f e r s : a
review. pp.211-225 In: Proc., Res. on Coniferous
F o r e s t Ecosystems Symp., Bellingham, WA.
March 23-24, 1 972.
Whittaker, R.H. and W.A. Niering.
1975. Vegetation of t h e Santa Cruz Mountains,
Arizona. V. Biomass, production and d i v e r s i t y
along t h e e l e v a t i o n g r a d i e n t . Ecol. 56:771-790.
 Response of Plant Communities to Air
Pollution1

R. Guderian and K. ~ u e ~ ~ e r s *

A b s t r a c t : Under t h e i n f l u e n c e of a i r p o l l u t i o n two
r e t r o g r e s s i v e p r o c e s s e s a r e s e t i n motion i n p l a n t com-
m u n i t i e s : By means of d i r e c t and i n d i r e c t e f f e c t s , chan-
ges occur i n s t r u c t u r e and f u n c t i o n of t h e community
l e a d i n g up t o t o t a l d e s t r u c t i o n . P a r a l l e l t o t h i s de-
g r a d a t i o n ( r e t r o g r e s s i o n ) i s a spontaneous o r man i n i -
t i a t e d process d u r i n g which t h e p r i g i n a l a d a p t i v e r e -
s i s t a n t members of t h e e x i s t i n g community a s w e l l a s
new a r r i v a l s undergo secondary succession. The causes
and mechanisms f o r a i r pollution-induced changes i n
p l a n t communities a r e demonstrated by means of l i t e r -
a t u r e a n a l y s i s and t h e i n t e r a c t i o n of dose response de-
termining f a c t o r s a r e summarized. I n o r d e r t o empha-
s i z e t h e e x i s t i n g p o t e n t i a l danger and t o s e t remedial
procedures i n motion, r e s e a r c h themes a r e pointed o u t
t h a t must r e c e i v e immediate a t t e n t i o n .

Thus f a r r e s e a r c h on t h e e f f e c t s of a i r p o l -
l u t a n t s on p l a n t s has been c e n t e r e d on homotyp-
i c a l p o p u l a t i o n s of economically important
s p e c i e s , With t h e development of long-term
l o a d i n g of e x t e n s i v e a r e a s e n t i r e ecosystems
a r e a l s o i n c r e a s i n g l y being i n f l u e n c e d by a i r
p o l l u t a n t s . From t h i s , t h e q u e s t i o n a r i s e s
a s t o t h e p o s s i b l e r e a c t i o n s of phytocoenoses
t o changed a i r q u a l i t y a s a new h a b i t a t f a c t o r .
c e r t a i n i n h e r e n t p a t t e r n s of t h e c o n f r o n t a t i o n
between p l a n t communities and a i r p o l l u t i o n
may be deduced.
For t h e following comparative s t u d y of t h e
i n f l u e n c e of v a r y i n g c o n c e n t r a t i o n s of a i r
p o l l u t a n t s on v e g e t a t i o n , a d i v i s i o n i n t o t h e
f o l l o w i n g l e v e l s , derived from Smith's c l a s -
s i f i c a t i o n (1974) seems p r a c t i c a l : h i g h , i n -
t e r m e d i a t e and low dosage e f f e c t s .

REACTIONS OF PLANT COMMUNITIES RELATED TO High P o l l u t i o n Dosage

AIR POLLUTANT CONCENTRATIONS
Single or repeated observations i n the v i -
c i n i t y of s i n g l e s o u r c e s a s w e l l a s w i t h i n and
o u t s i d e of extended r e g i o n s s u b j e c t e d t o a i r
p o l l u t i o n load can only provide momentary r e -
c o r d s o r sequences o f changes under t h e r e s p e c -
t i v e l o c a l c o n d i t i o n s . I n g e n e r a l , however,
~ r e s e n t e da t t h e Symposium on E f f e c t s of A i r
P o l l u t a n t s on Mediterranean and Temperate F o r e s t
Ecosystems, J u n e 22-27, 1980, R i v e r s i d e ,
C a l i f o r n i a , U.S.A.
2 ~ e s p e c t i v e l y ,Biowissenschaften, Universi-
t a t Essen, Gesamthochschule, 4300 Essen 1,
West Germany
A c h a r a c t e r i s t i c of t h e r e l a t i o n s h i p be-
tween high dosage and t h e r e a c t i o n of a p l a n t
community is a breakdown of community s t r u c -
ture -- more o r l e s s obvious depending on t h e
complexity of t h e ecosystem. The d e g r a d a t i o n
of t h e system is c h a r a c t e r i z e d by a r a p i d change
i n s t r u c t u r e , i n c l u d i n g composition. It is
accompanied and f i n a l l y r e p l a c e d by a second-
a r y s u c c e s s i o n which can lead t o a new e q u i l -
ibrium under s u s t a i n e d load.
D i r e c t a c u t e and c h r o n i c i n j u r y appearing
e s p e c i a l l y on l e a v e s , b u t n o t always c o r r e -
l a t e d w i t h t h e i r p o l l u t a n t c o n t e n t , (Guderian,
1970; Linzon, 1979) w i l l f i r s t a f f e c t t h e most
s e n s i t i v e s p e c i e s of t h e t r e e s t r a t u m i n a
f o r e s t and can l e a d t o t h e t o t a l d e s t r u c t i o n of
t h e canopy. Without p r o t e c t i o n from t h e f r e e l y
e n t e r i n g a i r masses of p o l l u t e d a i r (Bennett and
H i l l , 1975) s h r u b , h e r b and moss o r l i c h e n l a y -
e r s a r e d e s t r o y e d one a f t e r a n o t h e r , u n t i l a
b a r r e n zone r e s u l t s (Gordon and Gorham, 1963;
Woodwell, 1970). As an example of such condi-
t i o n s , t h e z o n a t i o n under t h e i n f l u e n c e of ap-
proximately 10 t o n s of SO2 per day from an i r o n
o r e r o a s t i n g f u r n a c e i n B i e r s d o r f (Germany) w i l l
be d e s c r i b e d b r i e f l y .
The denuded zone i n t h e immediate v i c i n i t y of
t h e emission s o u r c e i s surrounded by t h e t r a n s i -
t i o n zone w i t h i s o l a t e d c l u s t e r s of g r a s s (Des-
champsia f l e x u o s a ) and r e s i s t a n t ground c o v e r
( E r i c a c i n e r e a , Galium mollugo, Veronica o f f i c i n -
alis, -
- .
Rumex a c e t o s a , and C o n v a l l a r i a m a j a l i s )
Now and t h e n one a l s o e n c o u n t e r s t h e r e l a t i v e l y
r e s i s t a n t Sambucus racemosa and Rhamnus f r a n g u l a
i n t h i s zone. I n t h e g r a s s cover of v e g e t a t i o n
c o n s i s t i n g mostly of Deschampsia f l e x u o s a t h e
f i r s t s h o o t s of Quercus p e t r a i a a r e e s t a b l i s h e d
i n t h e s h e l t e r of t h e herbaceous v e g e t a t i o n .
The p o p u l a t i o n s of t h i s oak and Fagus s i l v a t i c a
which a d j o i n t h e s t u n t e d f o r e s t zone show
s i g n s of d i s i n t e g r a t i o n s t a r t i n g a t t h e p e r i -
phery. As SO2 l o a d i n g d e c r e a s e s t h e s p e c i e s
d i v e r s i t y increases, u n t i l f i n a l l y i t reaches
t h e combination t y p i c a l f o r t h e a c i d i c s o i l -
mixed f o r e s t (Fago-Quercetum).
A comparison w i t h o t h e r s t u d i e s (Treshow,
1968; Smith, 1974; M i l l e r and McBride, 1975;
and, Linzon, 1978) shows t h a t t h i s i s t h e typ-
i c a l p i c t u r e of t h e break-down of a p l a n t com-
munity, t h a t i s , a change i n s p e c i e s composition
toward a s i m p l i f i c a t i o n of t h e system. I n
p r i n c i p l e i t does n o t d i f f e r from t h a t caused
by gamma r a d i a t i o n (Woodwell, 1963, 1970).
The secondary s u c c e s s i o n which s e t s i n a s soon
a s t h e o r i g i n a l v e g e t a t i o n b e g i n s t o change
l e a d s i n time under c o n s t a n t l o a d i n g t o t h e
f o r m a t i o n of new, l e s s complex s t a b l e s t r u c t u r e s .
Thus, i n an o l d manufacturing d i s t r i c t of Upper
S i l e s i a (Poland), i n l o c a t i o n s once stocked w i t h
n a t i v e c o n i f e r o u s o r mixed deciduous f o r e s t s ,
Wolak (1977 and 1979) d e s c r i b e d a s t a b l e zon-
a t i o n i n r e l a t i o n t o d i f f e r e n t l o a d s of S02,
z i n c , and l e a d . Under heavy l o a d s an i n d u s t r i a l
w a s t e l a n d i s followed by a g r a s s zone w i t h
a s s o c i a t i o n s dominated by Deschampsia f l e x u o s a
on o l i g o t r o p h i c sand, by C a l a m a g r o s t i s e p i g e i o s
on mesotrophic s i t e s , and by C a l a m a g r o s t i s a-
l o s a on damp o r g a n i c s o i l . I n t h e a d j a c e n t
- i n g on dosage (Borgsdorf, 1960; Gordon and
shrub zone one f i n d s b o t h c u l t i v a t e d and v o l -
u n t e e r s c r u b t r e e s p e c i e s . I t i s remarkable
t h a t P i n u s s i l v e s t r i s can t a k e on t h e shape of
a c r e e p i n g shrub o r of a t r e e w i t h b r a n c h e s
p r o j e c t i n g h o r i z o n t a l l y up t o 5 m from t h e
stem. These dwarf forms a r e no more than 2 m
h i g h a t an age of 30 t o 50 y e a r s . On low g r a d e
sands g r o u p s of t h e d e s c r i b e d P i n u s s i l v e s t r i s
forms and Solanum dulcamara were found which
were n o t found i n s i m i l a r l o c a t i o n s w i t h o u t
t h e s t r o n g i n f l u e n c e of a i r p o l l u t i o n . Those
p l a n t communities a r e c a l l e d i n d u s t r i o - c l i m a x
communities (Wolak, 1971). They r e p r e s e n t spon-
taneous a s s o c i a t i o n s w i t h r e l a t i v e l y c o n s t a n t
s p e c i e s composition which have developed grad-
u a l l y through i n d u s t r i o g e n o u s (secondary)
s u c c e s s i o n , b o t h from s p e c i e s p r e s e n t b e f o r e
p o l l u t a n t l o a d i n g a s w e l l a s from new a r r i v a l s ,
under t h e combined i n f l u e n c e of t h e h a b i t a t
f a c t o r s ; c l i m a t e , s o i l , i n s e c t s and p a r a s i t e s - -
dominated by t h e f a c t o r a i r p o l l u t i o n .
I n t e r m e d i a t e P o l l u t i o n Dosage
I n t e r m e d i a t e a i r p o l l u t i o n dosage c o n d i t i o n s
a r e e c o l o g i c a l l y s i g n i f i c a n t because t h e i r sub-
t l e , d i r e c t and i n d i r e c t e f f e c t s on t h e i n d i v i -
d u a l s p e c i e s can s e t t h e s t a g e f o r changes i n
t h e s t r u c t u r e of t h e community w i t h p o s s i b l y ir-
r e v e r s i b l e consequences. I n p l a n t communities
e x p e r i e n c i n g i n t e r m e d i a t e p o l l u t i o n dosage, i n -
t e r r u p t i o n of growth and r e p r o d u c t i o n p r o c e s s e s
a s w e l l a s impairment of t h e v i t a l i t y o f i n d i -
v i d u a l p l a n t s , among o t h e r f a c t o r s through i n -
c r e a s e d v u l n e r a b i l i t y t o a b i o t i c and b i o t i c
s t r e s s , become p a r t i c u l a r l y important (Wentzel,
1965; Huttunen, 1979; and Laurence, 1980).
I n p i n e and s p r u c e p o p u l a t i o n s i n t h e Lower
Main Region (Germany), which indeed show an i n -
creased sulfur content i n the leaves, but did
n o t y e t d i s p l a y an abnormal l o s s o f i n d i v i d u a l s ,
morphological changes such a s t h i n crowns coup-
l e d w i t h s h o r t e r n e e d l e s were d e t e c t e d i n o l d e r
s t a n d s (Wentzel, 1979). Such changes occur slow-
l y and o n l y t h e accumulation of annual e f f e c t s
g r a d u a l l y l e a d s t o h i g h e r m o r b i d i t y (Wentzel,
1980). I n t h i s c o n t e x t t h e c a r r y - o v e r of ac-
cumulated t o x i c a n t s i n t h e new s h o o t s o f t h e
n e x t growing season should b e mentioned ( K e l l e r ,
1978; P r e s t o n , 1979). A s l i g h t change of t h e
h o r i z o n t a l s t r u c t u r e i n t h e canopy w i l l i n f l u -
ence such h a b i t a t f a c t o r s a s t h e s u p p l y of l i g h t
and p r e c i p i t a t i o n f o r t h e lower-lying v e g e t a t i o n .
F r e q u e n t l y , t h e i n t e r a c t i o n of changed s o i l r e -
action--pH--and t o x i c a n t c o n t e n t b r i n g s about
a r e s t r u c t u r i n g of t h e shrub and h e r b s t r a t a
o v e r extended a r e a s sometimes i n f l u e n c i n g na-
t u r a l r e p r o d u c t i o n of woody s p e c i e s (Lux, 1964;
Wentzel, 1971; Harward and Treshow, 1975)
Such changes i n t h e composition of p l a n t
communities were d e t e c t e d through v e g e t a t i o n
surveys caused by a complex of f a c t o r s . F o r
example, c e r t a i n s p e c i e s were found i n d e n s e
c l u s t e r s , w h i l e o t h e r s were e v e n l y d i s t r i b u t e d
and s t i l l o t h e r s were t o t a l l y a b s e n t , depend-
Gorham, 1963; N i k l f e l d , 1967; Ionescu, e t a l . ,
1971; Trautmann, e t a l . , 1971). Hajduck (1961)
t a l k s about p o s i t i v e o r n e g a t i v e p h y t o i n d i c a t o r s ,
w h i l e Anderson (1966, quoted i n Treshow 1968)
employs t h e terms " i n c r e a s e r " o r " d e c r e a s e r 'I .
The concept of p h y t o i n d i c a t o r s i s e s s e n t i a l l y
t h e same a s b i o i n d i c a t i o n w i t h l i c h e n o r moss
s p e c i e s (Le Blanc and Rao, 1975; Taoda, 1977;
and P i l e g a a r d , 1978). K a l e t a (1972) was a b l e
t o demonstrate i n a d d i t i o n t h e dynamics of
change of whole p l a n t a s s o c i a t i o n s under t h e
i n f l u e n c e of magnesite.
Brandt and Rhoades (1972, 1973) took t r e e
s p e c i e s of s e v e r a l s t r a t a i n t o c o n s i d e r a t i o n
i n t h e i r s t u d y on t h e i n f l u e n c e of l i m e s t o n e
d u s t on a f o r e s t community. T h i s method made
i t p o s s i b l e t o a s s e s s t h e t r e n d of f u t u r e suc-
c e s s i o n , e s p e c i a l l y through s h i f t s d e t e c t e d
i n t h e s p e c i e s d i v e r s i t y of t h e s e e d l i n g and
s p r o u t d a t a . Thus, Quercus c o c c i n e a , Quercus
v e l u t i n a o r T i l i a americana oan drop o u t a s
members of t h e oak-chestnut a s s o c i a t i o n and
L i r i o d e n d r o n t u l i p i f e r a , & saccharinum
and p o s s i b l y Quercus muehlenbergii could be-
come dominant s p e c i e s . Through t h i s example i t
a l s o becomes e v i d e n t how a i r p o l l u t a n t s can
i n f l u e n c e t h e makeup of phytocoenoses by i n -
f l u e n c i n g r e p r o d u c t i o n (Wentzel, 1963; Karnoskv,
and S t a i r s , 1974; K e l l e r , 1976).
McClenahen (1978) u t i l i z e d community comp-
o s i t i o n a s a means t o i n v e s t i g a t e changes i n
p l a n t communities a l o n g a p o l l u t i o n g r a d i e n t
i n t h e Ohio V a l l e y (USA). I n t h i s s t u d y , a n
e a s t e r n deciduous f o r e s t e x p e r i e n c i n g interme-
d i a t e dosages was shown t o d e c l i n e i n s p e c i e s
r i c h n e s s , evenness, and Shannon d i v e r s i t y i n -
dex w i t h i n a l l s t r a t a of t h e community, p a r t -
i c u l a r l y i n those locations experiencing t h e
h i g h e s t r e l a t i v e dose. Simultaneously, t h e
s i m i l a r i t y i n composition d e c r e a s e d w i t h i n -
c r e a s i n g dosage. Thus, t h e r e l a t i v e importance
of saccharinum, a s p e c i e s s l i g h t l y s t i m -
u l a t e d by l i m e s t o n e d u s t (Brandt and Rhoades,
1972), showed d i s t i n c t d e c l i n e i n a l l s t r a t a ,
w h i l e t h e importance of Aesculus o c t a n d r a i n -
c r e a s e d . Opposing t e n d e n c i e s i n d e n s i t y were
observed i n some s t r a t a . A d e c l i n e i n t h e t r e e
and h e r b s t r a t a was accompanied by a n i n c r e a s e
of t h e subcanopy and t h e s h r u b s t r a t a . This
can be a t t r i b u t e d t o b e t t e r l i g h t c o n d i t i o n s i n
t h e lower s t r a t a combined w i t h a r e l a t i v e i n -
c r e a s e of h e r b s i n t o l e r a n t t o shade.
Low P o l l u t i o n Dosage
The e f f e c t s of low dosages on v e g e t a t i o n l i e
i n t h e b o r d e r zone between t h e f l u c t u a t i n g
s t a t e s of normal, i . e . , u n a f f e c t e d v e g e t a t i o n
on t h e one hand, and s i g n i f i c a n t i n j u r i o u s e f -
f e c t s on t h e o t h e r hand. Depending upon t h e
r e s p e c t i v e p o l l u t a n t , i t s c o n c e n t r a t i o n and
d u r a t i o n of a c t i o n , a s w e l l a s t h e a f f e c t e d ob-
j e c t and t h e l o c a l c o n d i t i o n s , t h e s e e f f e c t s
can r a n g e from i n c r e a s e s t o r e d u c t i o n s i n growth,
r e p r o d u c t i v e c a p a b i l i t y o r q u a l i t y of p l a n t s .
Under p r a c t i c a l c o n d i t i o n s such e f f e c t s can be
d e t e c t e d t o o n l y a c e r t a i n d e g r e e of t h e a c t u a l
i n t e n s i t y . The d e t e c t i o n l i m i t h a s been low-
e r e d through t h e development of new exposure
systems w i t h f i l t e r e d and u n f i l t e r e d a i r (Mandl,
e t a l . , 1973; Lee, e t a l . , 1973; M i l l e r , e t a l . ,
1979; Shinn, e t a l . , 1979). However, measure-
ment of p o l l u t a n t e f f e c t s on p l a n t communities
occupying l a r g e r e g i o n s p r e s e n t s p a r t i c u l a r
d i f f i c u l t i e s because t h e n e c e s s a r y p o l l u t i o n -
f r e e c o n t r o l a r e a s w i t h comparable s o i l and
climate a r e not available.
Before d e t e c t a b l e r e d u c t i o n s occur i n pro-
d u c t i v i t y o r a l t e r a t i o n of environmental con-
d i t i o n s c a n b e observed, t h e r e a r e v a r i o u s
changes t h a t a r e induced a t t h e p l a n t biochem-
i c a l , physiological o r substructural l e v e l
( K e l l e r , 1974; J a g e r and K l e i n , 1977; Horsman
and Wellburn, 1977; and, Raabe and Kreeb, 1979).
Of c o u r s e t h e q u e s t i o n of how much such f i n d -
i n g s r e v e a l a b o u t t h e economic and e c o l o g i c p e r -
formance of a p a r t i c u l a r p l a n t s p e c i e s remains
of c e n t r a l importance h e r e . Some of t h e r e -
a c t i o n s undoubtedly have no e f f e c t s on t h e t o t a l
organism; even i f s i g n i f i c a n t e f f e c t s a r e found,
it is very d i f f i c u l t t o e s t a b l i s h a causal l i n k
t o t h e primary r e s p o n s e s mentioned above. The
p o s s i b l e e f f e c t s of low dosage on p l a n t com-
m u n i t i e s , f o r example, through changes i n i n -
t e r s p e c i f i c competition, a r e almost t o t a l l y
unresolved. The f i l t e r i n g e f f e c t s of v e g e t a t i o n
i s a n important p r o c e s s b u t t h i s t o p i c w i l l
only be introduced h e r e .
A s shown i n t h e S o i l i n g p r o j e c t ( U l r i c h , e t
a l . , 1978) o r t h e Hubbard Brook s t u d y (Bormann
and Likens, 1979) v e g e t a t i o n can f i l t e r l a r g e
amounts of p o l l u t a n t s o u t of t h e atmosphere
w i t h o u t showing s i g n s of e x t e r n a l i n j u r y o r
growth d e p r e s s i o n . P a r t i c u l a t e and gaseous
p o l l u t a n t s e n t e r a n ecosystem through a d s o r p -
t i o n and a b s o r p t i o n mainly on l e a f s u r f a c e s a s
w e l l a s s o i l and w a t e r s u r f a c e s ( H i l l , 1971;
Bennett and H i l l , 1975; and, Olsen, 1976). The
s p e c i f i c behavior of t h e s u b s t a n c e i s i m p o r t a n t
f o r t h e p o s s i b l e long-term e f f e c t s of low do-,
s a g e on p l a n t communities. P o l l u t a n t s which a r e
s u b j e c t t o r a p i d decomposition such a s ozone
o r PAN t a k e e f f e c t through t h e summation of
d i r e c t e f f e c t s . NOx, NH3, o r s u l f u r compounds
can be channeled i n t o t h e n u t r i e n t c y c l e and
may d e s t r o y t h e b a l a n c e of e s p e c i a l l y s e n s i t i v e
ecosystems, such a s moors, through e u t r o p h i -
c a t i o n ( P o r t e r , e t a l . , 1972; Cowling and Lock-
y e r , 1976). The importance of a c i d p r e c i p i t a t i o n
i n t h i s c o n t e x t i s n o t y e t c l e a r (Braekke, 1976;
and, T a m , 1976). Accumulating s u b s t a n c e s such
a s heavy m e t a l s , r e p r e s e n t a s p e c i a l danger f o r
ecosystems (Kraemer, 1976; and Guderian, 1980).
S o i l samples and a n a l y s e s of moss specimens have
r e v e a l e d t h a t a c o n s t a n t i n p u t i n t o ecosystems
i s o f f s e t by o n l y a l i m i t e d e x p o r t , and t h a t
t h i s i s now o c c u r r i n g over wide a r e a s (Huckabee,
1973; Ruhling and T y l e r , 1973; and Grdzinska,
1978). Such components can a l s o endanger t h e
n u t r i e n t c y c l e (Mags, 1977 and Uba, 1977);
f u r t h e r , t h e y r e d u c e t h e number and a c t i v i t y
of decomposers, t h e r e b y i m p a i r i n g r e m i n e r a l -
i z a t i o n a s a requirement f o r u n i n t e r r u p t e d b i o -
geochemical c y c l e s ( T a y l o r , 1975; and, G r e s z t a ,
e t a l . , 1979). E s p e c i a l l y , w i t h accumulating
s u b s t a n c e s and under continuous l o a d i n g , it i s
only a q u e s t i o n of time b e f o r e t h e d i r e c t and
i n d i r e c t e f f e c t s d e s c r i b e d above begin t o i n -
t e r r u p t t h e s t r u c t u r e and f u n c t i o n of p l a n t
communities.
THE INFLUENCE OF POLLUTANTS ON THE
FUNCTION OF PLANT COMMUNITIES
Changes i n p l a n t communities caused by p o l -
l u t a n t s can l e a d t o more o r l e s s l a s t i n g im-
pairment of economic and e c o l o g i c f u n c t i o n s de-
pending on t h e dosage. The damage t o a g r i c u l t u r e
through growth r e d u c t i o n , l o s s of q u a l i t y and
h i g h e r l a b o r c o s t s have long drawn c o n s i d e r -
a b l e a t t e n t i o n , b u t only now i s an a t t e m p t be-
ing made t o t a k e t h e e f f e c t s on performance
of ecosystems i n t o account. I n t h i s connec-
t i o n , an e s p e c i a l l y important q u e s t i o n i s how
p o l l u t a n t s a f f e c t such f u n c t i o n s of v e g e t a t i o n
a s f i l t e r e f f e c t , s t a b i l i z a t i o n of c l i m a t e ,
r e g u l a t i o n of water and n u t r i e n t c y c l e s , s o i l
c o n s e r v a t i o n a s w e l l a s t h e p r e s e r v a t i o n of
l i v i n g space f o r polymorphic zoo- and phyto-
ceonoses. Extensive changes i n v e g e t a t i o n
cover a r e probably l i n k e d t o i n t e r r u p t i o n o r
even t o t a l breakdown of a l l t h e above-men-
tioned f u n c t i o n s . For example, t h e f u n c t i o n
of p l a n t communities a s p r o t e c t i o n a g a i n s t
erosion o r a s a f a c t o r i n counter balancing
e x c e s s i v e temperature f l u c t u a t i o n s and t h e
accompanying danger from l i g h t f r o s t d u r i n g
bud b r e a k i s c o n s i d e r a b l y more impaired i n
a r e a s experiencing h i g h p o l l u t a n t dosage where
woods have been r e p l a c e d by s p a r s e v e g e t a t i o n
than i n unpolluted a r e a s . C u r r e n t l y , t o what
e x t e n t i n t e r m e d i a t e and low dosages a f f e c t
t h e f u n c t i o n s of v e g e t a t i o n mentioned, can,
a t b e s t , b e deduced t o an o r d e r of magnitude
(Materna, 1980) from t h e known r e a l e f f e c t s on
vegetation.
Causes f o r t h e Observed Responses
of P l a n t Communities
The e f f e c t s of a given p o l l u t a n t on p l a n t
communities, a s i l l u s t r a t e d by s e v e r a l exam-
p l e s , a r e determined by: t h e g e n e t i c a l l y pre-
determined d e g r e e of r e s i s t a n c e of t h e com-
panion s p e c i e s (Dochinger e t a l . , 1965; Rohmeder
e t a l . , 1965), t h e modifying i n f l u e n c e of en-
vironmental c o n d i t i o n s of r e s i s t a n c e , and t h e
changes i n i n t r a - and i n t e r s p e c i f i c r e l a t i o n s
caused by p o l l u t a n t s .
P o p u l a t i o n s of c e r t a i n p l a n t s p e c i e s , va-
r i e t i e s , and c l o n e s , a s w e l l a s i n d i v i d u a l s
w i t h i n t h e r e s p e c t i v e populations s t u d i e d ,
r e a c t t o a given a i r p o l l u t i o n s t r e s s w i t h
v a r y i n g d e g r e e s of s e n s i t i v i t y . I n c o n t r a s t
t o c e r t a i n phytopathogenic organisms (Baumann,
1951; Grossmann~1970), t h e r e i s no a b s o l u t e
r e s i s t a n c e , a s demonstrated by t h e e x i s t e n c e
of v e g e t a t i o n - f r e e zones. The schematic
diagram (Figure 1 ) i s an a t t e m p t t o demonstrate
which f a c t o r s i n f l u e n c e t h e response t o a i r
pollution stress.
I n d i v i d u a l and S p e c i e s S p e c i f i c Responses
According t o L e v i t t (1972), two mechanisms
determine a p l a n t ' s r e s i s t a n c e t o s t r e s s :
s t r e s s avoidance and s t r e s s t o l e r a n c e . I n
t h e f i r s t c a s e t h e s t r e s s , caused h e r e by a
s p e c i f i c p o l l u t a n t dose, i s prevented from
t a k i n g e f f e c t - - i t i s excluded. A m u l t i t u d e
of f a c t o r s determines t h e r e s i s t a n c e of a
p l a n t organism t o t h e e n t r y of p o l l u t a n t s i n t o
t h e c e l l . Morphological p r o p e r t i e s such a s
shape and s u r f a c e s t r u c t u r e i n c l u d i n g wax
l a y e r s ( R e n t s c h l e r , 1973; S h r i n e r , 1980) a s
w e l l a s t h e number, d i s t r i b u t i o n , and a p e r t u r e
of t h e stoma (Meidner and Mansfield, 1968)
must be mentioned. According t o Taylor (1978),
whose d e f i n i t i o n was taken i n t o c o n s i d e r a t i o n
i n t h e corresponding s e c t i o n of F i g u r e 1,
s t r e s s t o l e r a n c e presupposes t h e e n t r y of t h e
r e s p e c t i v e p o l l u t a n t i n t o t h e c e l l . As long
a s t h e e n t e r i n g substance i s t o l e r a t e d , a s -
s i m i l a t e d o r b u f f e r e d , and consequently no
morphological o r p h y s i o l o g i c a l change t a k e s
p l a c e , one speaks of " s t r a i n avoidance." Above
c e r t a i n i n t r a c e l l u l a r concentrations, for
example, a f t e r c e r t a i n biochemical t h r e s h o l d
v a l u e s have been exceeded, i n j u r y o c c u r s which
i s e i t h e r r e v e r s i b l e ( e l a s t i c s t r a i n ) , such
a s s u b t l e changes i n p h o t o s y n t h e t i c performance
( S i j and Swanson, l 9 7 4 ) , o r i r r e v e r s i b l e ( p l a s -
t i c s t r a i n ) , such a s i n j u r y t o l e a v e s i n t h e
form of n e c r o s i s .
Thus, t h e biochemical t h r e s h o l d v a l u e s ,
which c h a r a c t e r i z e t h e t r a n s i t i o n s from s t r a i n
avoidance t o s t r a i n t o l e r a n c e a s w e l l a s from
e l a s t i c s t r a i n t o p l a s t i c s t r a i n , determine
t h e t o l e r a n c e of a p l a n t . It follows t h a t t h e
i n d i v i d u a l response can m a n i f e s t i t s e l f i n
terms of i n d i f f e r e n c e , m o d i f i c a t i o n o r d e a t h
of t h e a f f e c t e d p l a n t depending on t h e l e v e l
of ambient s t r e s s , and t h e r e s p e c t i v e r e s i s t a n c e .
As i s a p p a r e n t from F i g u r e 1, a m u l t i t u d e of
organismal and environmental f a c t o r s b e f o r e ,
d u r i n g , and a f t e r t h e p o l l u t a n t impact i s r e -
s p o n s i b l e f o r t h e sometimes v e r y g r e a t d i f -
f e r e n c e s i n t h e " r e s i s t a n c e s e r i e s " o r "re-
s i s t a n c e groups" of v a r i o u s a u t h o r s ( S t o k l a s a ,
1923; Bredemann, 1956; Thomas, 1961; Garber,
1967; Mooi, 1974; Davis and Wilhour, 1976;
Guderian, 1977). A l l c l i m a t i c f a c t o r s , f o r
example, t h a t r e g u l a t e t h e number, s i z e and
a p e r t u r e of t h e stomata (Bronte and Conguet,
1975; H a l l and Kaufmann, 1975), such a s l i g h t ,
optimal water supply, h i g h r e l a t i v e humidity
o r adequate temperature, determine t h e r a t e
a t which p o l l u t a n t s a r e absorbed (Guderian,
1970; Jones and Mansfeld, 1970; McLean and
Schneider, 1971).
The i n f l u e n c e of edaphic f a c t o r s i s demon-
s t r a t e d w i t h two examples: Copper-beech (Fagus
-
s i l v-a t i c a ) i s c o n s i d e r a b l y more r e s i s t a n t on
s o i l s w i t h high lime c o n t e n t than on sandy
s o i l low i n n u t r i e n t s ; elm (Ulmus c a m p e s t r i s )
proved t o b e one of t h e most r e s i s t a n t s p e c i e s
i n a l l u v i a l forests, but i n l e s s suitable
h a b i t a t s i t was one of t h e most v u l n e r a b l e
of a l l t h e deciduous s p e c i e s (Wentzel, 1968).
T h i s shows t h e d i f f i c u l t y i n s e t t i n g up gener-
a l l y accepted r e s i s t a n c e s e r i e s , a s r e c e n t
s t u d i e s w i t h v a r i o u s soybean c u l t i v a r s under
changing environmental c o n d i t i o n s have c l e a r l y
shown (Heagle, 1979a, b ) .
The l a r c h s e r v e s a s a t y p i c a l example f o r
changes i n r e s i s t a n c e i n r e l a t i o n t o l e v e l s
of c o n c e n t r a t i o n (Guderian and Stratmann, 1962;
Wentzel, 1963). Under h i g h , a c u t e SO2 con-
c e n t r a t i o n s , both L a r i x europea and L a r i x
l e p t o l e p i s show s i g n s of n e c r o s i s b e f o r e spruce
 -
momentary dosage
- ..
momentary
low intermediate high I
air oollution stress
momentary momentary

succession
interspecific

relationship

areal and areal and

abundance.

alteration alteration

low dosage response intermediate dosage response high dosage response

no significant first alterations in extensive simplification

alteration of structure and compositions up to total destruction

plant communities of plant communities of plant communities

Figure 1: Effect-determining-factors'for various responses of plants on individual-,

species- and community-levels.

 ( P i c e a a b i e s ) and p i n e (Pinus s i l v e s t r i s ) . On
t h e o t h e r hand, t h e l a r c h i s among t h e most
r e s i s t a n t c o n i f e r s under continuous low l e v e l s
of c o n c e n t r a t i o n (Wentzel, 1963), and i t i s
widely used t o r e e s t a b l i s h t r e e p o p u l a t i o n s
a f t e r t h e d e g r a d a t i o n of spruce and p i n e f o r -
e s t s i n r e g i o n s of c h r o n i c s t r e s s .
Regarding organismal f a c t o r s , t h e s i g n i f i -
cance of age f o r s e n s i t i v i t y should be s t r e s s e d .
According t o o b s e r v a t i o n i n t h e f i e l d , c o n i f e r s
such a s P i c e a a b i e s and P i n u s s i l v e s t r i s r e -
main p a r t i c u l a r l y s u s c e p t i b l e from t h e l a t e
p o l e timber s t a g e ( a t t h e time of accumulating
growth) through t o t h e t r e e timber s t a g e . Dur-
i n g t h e s e p e r i o d s of development, e s p e c i a l l y
s t r o n g r e d u c t i o n s i n growth and t h e widespread
d e g r a d a t i o n of e n t i r e s t a n d s occur (Wentzel,
1962; Materna e t a l . , 1969), w h i l e deciduous
p o p u l a t i o n s respond i n a much weaker form. I n
p l a n t i n g s , however, under mostly c h r o n i c SO2
c o n c e n t r a t i o n s , t h e c o n i f e r s mentioned and t h e
copper-beech (Fagus s i l v a t i c a ) and pedunculate
oak (Quercus pedunculata) e x h i b i t e d n e a r l y
e q u a l r e s i s t a n c e (Guderian and Stratmann, 1968).
One a s p e c t n o t o f t e n taken i n t o account when
judging t h e r e s i s t a n c e of p l a n t s , b e s i d e s en-
vironmental, p o l l u t a n t , and o r g a n i c i n f l u e n c e s ,
i s t h e c r i t e r i a used t o i n t e r p r e t t h e e f f e c t .
Various deciduous t r e e s p e c i e s , such a s l i n d e n
( T i l i a c o r d a t a an& T i l i a p l a t y p h l l o s ) and beech
(Fagus s i l v a t i c a ) respond t o a c u t e SO2 con-
c e n t r a t i o n s w i t h l e a f n e c r o s i s e a r l i e r than
s p r u c e (Picea a b i e s ) o r S c o t s p i n e (Pinus s i l -
v e s t r i s ) . N e v e r t h e l e s s t h e deciduous s p e c i e s
can s t i l l grow i n p o l l u t e d r e g i o n s where spruce
and S c o t s p i n e d i e o u t (Wentzel, 1968). Thus
r e s i s t a n c e must f i r s t of a l l b e c h a r a c t e r i z e d
by t h e d i f f e r e n c e s i n t h e r e d u c t i o n of growth
and y i e l d of t h e s p e c i f i c s p e c i e s . The func-
t i o n s of t h e p l a n t s p e c i e s being considered
h e r e determine t h e c r i t e r i a used t o e v a l u a t e
t h e e f f e c t s (Guderian, 1977). Through t h e
s h o r t d e s c r i p t i o n of f a c t o r s determining t h e
r e s i s t a n c e of a n i n d i v i d u a l o r a s p e c i e s i t
can b e seen what d e g r e e s of v a r i a t i o n must be
expected i n t h e responses. The use of t h e s e
r e s u l t s t o f o r e c a s t t h e behavior of s i n g l e
s p e c i e s under a i r p o l l u t i o n s t r e s s i s n e c e s s a r -
i l y v e r y d i f f i c u l t , e s p e c i a l l y when s t u d y i n g
p l a n t communities.
I n t h e following model, supported by exper-
i m e n t a l r e s u l t s , an a t t e m p t i s made t o i l l u s -
t r a t e t h e p o s s i b l e responses of two p l a n t
species t o increasing a i r pollution s t r e s s
(Fig. 2 ) . A s p e c i f i c response of two p l a n t
s p e c i e s (A and B) i s shown i n p e r c e n t of con-
t r o l . Up t o a s p e c i f i c c o n c e n t r a t i o n l a b e l e d
A 1 and B3 no s i g n i f i c a n t d e v i a t i o n i n t h e r e -
sponses of t h e exposed p l a n t s and t h e c o n t r o l
p l a n t s could b e d e t e c t e d . Important q u a l i t a -
t i v e d i f f e r e n c e s i n response between t h e two
s p e c i e s e x i s t above c o n c e n t r a t i o n A1. With
Species A, t h e s p e c i f i c response i s i n i t i a l l y
s t i m u l a t e d by sulphur d i o x i d e ; a r e d u c t i o n of
performance o n l y occurs a t a h i g h e r concentra-
t i o n , w h i l e t h i s Species B l a c k s s t i m u l a t i n g
e f f e c t . The f u r t h e r s l o p e of t h e c u r v e shows
t h e degree of r e d u c t i o n i n response. The
c o n c e n t r a t i o n A 4 / ~ 4should be emphasized, a s
h e r e t h e r e s i s t a n c e r e l a t i o n s h i p of t h e two
s p e c i e s changes (Wentzel, 1963). I n t h e con-
c e n t r a t i o n range A1 t o A 4 / ~ 4s p e c i e s A would
have an advantage over s p e c i e s B, even under
p o l l u t a n t c o n c e n t r a t i o n s which do n o t y e t
have a n adverse e f f e c t on s p e c i e s B. Accord-
i n g l y , changes i n t h e composition of p l a n t
communities must be expected even i f SO2 con-
c e n t r a t i o n s a r e s o low t h a t they do not y e t
have a d i r e c t harmful e f f e c t . This i s a
s i g n i f i c a n t aspect f o r ecological research.
Community S p e c i f i c Responses
The p r e v i o u s l y demonstrated r e l a t i o n s h i p s
between h e r e d i t y , environment and r e s i s t a n c e
i n i n d i v i d u a l s o r homotypical p o p u l a t i o n s a r e
n a t u r a l l y a l s o v a l i d f o r p l a n t communities.
Community s p e c i f i c a s p e c t s must be given ad-
d i t i o n a l c o n s i d e r a t i o n when a s c e r t a i n i n g p o l -
l u t a n t e f f e c t s . The importance of t h e r e -
l a t i o n s h i p between two o r more p o p u l a t i o n s
shown i n F i g . 1 i s underlined by t h e few ex-
i s t i n g r e s u l t s from experiments on t h e i n f l u -
ence of a i r p o l l u t a n t s t o p l a n t communities.
Thus, according t o experimental a n a l y s i s of
pure and mixed seedings, c o n s i s t i n g of r y e
g r a s s (Lolium multiflorum) h a i r y v e t c h (Vicia
v i l l o g a ) and crimson c l o v e r ( T r i f o l i u m i n c a r -
natum) s h i f t s i n t h e composition of p l a n t com-
m u n i t i e s cannot b e explained e x c l u s i v e l y through
t h e d i r e c t e f f e c t of p o l l u t a n t s on v a r i o u s
s p e c i e s of d i f f e r e n t s e n s i t i v i t y (Guderian,
1966, 1977). Under SO2 t h e i n f l u e n c e of i n t e r -
s p e c i f i c competition was a l t e r e d . As a r e s u l t ,
t h e primary e f f e c t on t h e more s u s c e p t i b l e mem-
b e r s was magnified t o such a degree t h a t they
could no longer compete e f f e c t i v e l y f o r v i t a l
growth-determining f a c t o r s . As a r e s u l t of
changed competition i n t h e community, t h e
d e c l i n e of t h e more s e n s i t i v e members allowed
improved growth of t h e more r e s i s t a n t s p e c i e s .
The t o t a l community y i e l d decreased l e s s than
would have been expected from t h e l o s s of
t h e more s u s c e p t i b l e s p e c i e s . S i m i l a r r e s u l t s
were found under t h e i n f l u e n c e of ozone (Bennett
and Runeckles, 1977), u l t r a v i o l e t r a d i a t i o n
(Fox and Caldwell, 1978) and i o n i z i n g r a d i a t i o n
(McCormick, 1963). The l a s t of t h e s e s t u d i e s
shows t h e importance of s t r e s s d u r i n g t h e seed-
l i n g and s p r o u t s t a g e s .
The e x t e n t of s h i f t s i n p l a n t communities a s
a r e a c t i o n t o a given load i s a l s o dependent
t o a l a r g e degree on t h e c o n d i t i o n of t h e com-
munity i t s e l f . The importance of t h e b u i l d i n g
of s t r a t a , of morphological s t r u c t u r e s , a s w e l l
a s r e l i e f and uniformity of t h e v e g e t a t i o n cover
were a l r e a d y pointed o u t a s was t h e i n t e r c o n -
n e c t i o n between s t a g e s of succession and system
responses. The s t a b i l i t y of a community g r e a t -
l y i n f l u e n c e s t h e response of i t s i n d i v i d u a l
members a s w e l l a s t h e whole t o a given p o l l u -
t a n t load. I n t h e presence of small d i s t u r b a n c e s ,
h i g h l y productive, complex systems can u s u a l l y
Special responses of
p l a n t species A and B
control = 100%

Typical reactions of two p l a n t species d e p e n d i n g

on t h e s u l f u r dioxide content of a i r
r e g a i n t h e i r s t a t e of b a l a n c e q u i c k l y because
of t h e i r complex feedback systems. Under
h e a v i e r l o a d s on t h e o t h e r hand, d r a s t i c chan-
ges must be e x p e t t e d p a r t i c u l a r l y i f c e r t a i n
key s p e c i e s a r e v e r y s e n s i t i v e t o t h e r e -
s p e c t i v e a i r p o l l u t a n t . But even v e r y low
p o l l u t a n t c o n c e n t r a t i o n s can produce q u i t e
c o n s i d e r a b l e e f f e c t s i n p l a n t communities,
e s p e c i a l l y i f s y n e c o l o g i c a l amplitudes of t h e
n a t u r a l community s p e c i e s a r e f a r a p a r t . Ad-
d i t i o n a l s t r e s s through p o l l u t a n t s of lower
dosage can lead t o d r a s t i c r e d u c t i o n i n v i -
t a l i t y of p l a n t s a l r e a d y l i v i n g o u t s i d e t h e i r
e c o l o g i c optimum. The r e l a t i v e l y h i g h sus-
c e p t i b i l i t y of spruce ( P i c e a a b i e s ) i n t h e
Erz mountains (Materna, 1972) and i n t h e b o r e a l
c o n i f e r o u s f o r e s t s i n F i n l a n d (Huttunen, 1979)
might w e l l b e caused by t h e i r u n s u i t a b l e hab-
i t a t s . Keeping t h i s i n mind, i t seems prob-
lematic t o t r a n s f e r those dose-effect re-
l a t i o n s h i p s determined from "production eco-
systems1'--in which t h e food p l a n t s g e n e r a l l y
encounter f a v o r a b l e c o n d i t i o n s - - t o n a t u r a l
ecosystems.
The r e l a t i o n s h i p s enumerated up t o t h i s p o i n t
show c l e a r l y why emissions induce d e g r a d a t i o n
i n p l a n t communities. On t h e o t h e r hand spon-
taneous a d a p t a t i o n t o a i r p o l l u t i o n s t r e s s
may be observed--adaptation which does n o t
e n s u r e t h e s u r v i v a l of t h e s p e c i e s through
s t u n t i n g , b u t r a t h e r seems t o have g e n e t i c
o r i g i n s . When Marchantia polymorpha (Briggs ,
1972) was exposed t o l e a d , and v a r i o u s g r a s s
s p e c i e s (Bradshaw, 1971, 1972, 1976) were a f -
f e c t e d by copper and z i n c , more t o l e r a n t pop-
u l a t i o n s developed i n a s h o r t time through
d i r e c t e d s e l e c t i o n . B e l l and Clough (1973),
B e l l and Mudd (1976), and Horsman and Wellburn
(1977) mention s i m i l a r p r o c e s s e s w i t h Lolium
perenne and Rumex o b t u s i f o l i u s s u b j e c t e d t o
decades of SO2 loading. F i n a l l y , t h e r e s u l t s
of long-term fumigation of n a t i v e g r a s s l a n d
( P r e s t o n and B u l l e t t , 1978) a l s o p o i n t t o t h e
f a c t t h a t under a n y t h i n g l e s s than a c u t e con-
c e n t r a t i o n s spontaneous a d a p t a t i o n may occur
i n t h e c o u r s e of t h e formation of a new sec-
ondary e q u i l i b r i u m , which may a l s o i n t e r r u p t
p o s s i b l e long-term i n j u r y ( P r e s t o n , l979a).
Accordingly, t h e dose and i t s r a t e of change
should b e a d j u s t e d over t h e long-term such t h a t
p l a n t communities r e t a i n t h e i r c a p a b i l i t y - -
even by t h e e v o l u t i o n a r y method mentioned a-
bove--to f u l f i l l t h e i r f u n c t i o n i n n a t u r a l and
a g r a r i a n ecosystems t o t h e f u l l e s t (Guderian
and Kueppers, 1979).
The g e n e t i c a l l y f i x e d v a r i a t i o n i n popu-
l a t i o n s which i s expressed i n t h e d e s c r i b e d
spontaneous a d a p t a t i o n , p r o v i d e s t h e b a s i s
f o r b r e e d i n g of p o l l u t a n t r e s i s t a n t p l a n t s
through s e l e c t i o n and r e p r o d u c t i o n of r e l a -
t i v e l y r e s i s t a n t i n d i v i d u a l s (Bialobok, 1979).
The r e s p o n s e s of i n d i v i d u a l s and homotypic
p o p u l a t i o n s t a k i n g i n t o account i n t e r s p e c i f i c
r e l a t i o n s discussed i n t h i s section, lead t o
t h e community s p e c i f i c r e s p o n s e s shown i n F i g .
1 which range from i n s i g n i f i c a n t changes t o
t h e t o t a l d e s t r u c t i o n of p l a n t communities.
CONCLUSIONS
Contamination of e x t e n s i v e a r e a s due t o
i n c r e a s i n g emissions and c o n t r o l s t r a t e g i e s
u s i n g t a l l s t a c k s f o r d i l u t i o n h a s made t h e
s t u d y of p l a n t communities and ecosystems
e s p e c i a l l y n e c e s s a r y . To a i d i n r e c o g n i t i o n
of p o s s i b l e r i s k s and i n making d e c i s i o n s r e -
garding c o n t r o l measures a t t h e s o u r c e and i n
t h e a f f e c t e d a r e a , t h e f o l l o w i n g p o i n t s must
be c l a r i f i e d :
1. Under which doses do changes occur
i n s t r u c t u r e and f u n c t i o n of p l a n t communities
of d i f f e r e n t complexity?
2. To what e x t e n t do p l a n t communities
show more s e n s i t i v e r e s p o n s e s than t h e i n -
d i v i d u a l s p e c i e s composing them?
3. What a r e t h e mechanisms of t h e s e
changes?
- P o i n t s of impact f o r a i r p o l l u t a n t s
i n t h e ecosystem.
- Location of p o l l u t a n t s i n t h e eco-
system ( a s s i m i l a t i o n , accumulation,
b r e a k down).
- Direct stimulatory o r injurious
e f f e c t s on t h e i n d i v i d u a l s p e c i e s of
t h e community.
- Causes and mechanisms of changes i n
competition e q u i l i b r i u m .
- Secondary succession w i t h p a r t i c u l a r
a t t e n t i o n t o a d a p t a t i o n and compen-
sation.
4. How a r e r i s k s determined f o r p l a n t
communities?
- Development of experimental d e s i g n s
and i n t e n s i f i c a t i o n s of epidemio-
l o g i c a l studies f o r the determination
of e f f e c t s t o h i g h l y s t r u c t u r e d sys-
tems.
- Establishment of permanent s t u d y
a r e a s t o i n v e s t i g a t e succession.
- Use of model p l a n t communities a s
indicators i n eco-toxicological t e s t s .
- A n a l y s i s of t h e c o n d i t i o n of ecosys-
tems b e f o r e and a f t e r s t a r t - u p of
a p o l l u t a n t source.
5. What measures a r e n e c e s s a r y f o r t h e
p r o t e c t i o n of v e g e t a t i o n ?
- Determination of dose-response r e -
l a t i o n s f o r p l a n t communities a s a
b a s i s f o r r i s k p r e d i c t i o n s and t h e
e s t a b l i s h m e n t of s t a n d a r d s f o r eco-
systems.
- C o l l e c t i o n of g e n e t i c r e s o u r c e s i n
n a t u r a l r e s e r v e s and i n gene banks.
- P r o t e c t i o n of endangered p l a n t com-
munities, especially i n existing
n a t u r a l r e s e r v e s , from e f f e c t s of'
a i r pollutants.
- Development and maintenance of a i r
p o l l u t a n t c o n t r o l s t r a t e g i e s allow-
i n g p o l l u t a n t dose and i t s r a t e of
change be s o c o n t r o l l e d t h a t t h e
s t r u c t u r a l d i v e r s i t y , and t h e eco-
l o g i c and economic f u n c t i o n s of t h e
vegetation, a s w e l l a s i t s function
a s a gene pool, a r e f u l l y p r o t e c t e d .
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 Forecasting Effects of SO2 Pollution on
Growth and Succession in a Western
Conifer Forest
2

J.R. Kercher, M.C. Axelrod, and G.E. Bingham

Abstract: A simulator has been developed for the mixed conifer forest type

of the Sierra Nevada, California to forecast the effects of SO2 on forest

growth and succession. The model simulates recruitment, growth, and death

of each tree and is based on a northeastern USA simulator with extensive

modifications. These modifications include the introduction of fire ecology,

temporal seed crop patterns unique to the Sierra, and water stress. Pollutant

stress is modeled as an effect on tree growth. The model simulates the shift

from the ponderosa pine dominated forest type to the white fir dominated mixed

conifer type as elevation increases from 5000 to 6000 ft. It also simulates

the fire-suppression of white fir and the fire-climax of ponderosa pine. For

a 10% growth reduction of ponderosa pine from pollutant stress and with growth

reductions in other species as determined by their relative sensitivities,

standing crops of ponderosa pine were reduced and white fir increased.

It is anticipated that extensive fossil fuel

JABOWA, a simulator of forests of the northeastern

energy development will occur in the United States

USA. For a case study, SILVA has been applied to

over the next several decades with increased emis-

the ponderosa pine and mixed conifer forest types

sions of phytoactive effluents. It has long been

of the Sierra Nevada, California, USA. The asso-

recognized that many of these pollutants have de-

ciated species in these forests are ponderosa pine

leterious effects on the growth and behavior of

(Pinus ponderosa), white fir (Abies concolor),

vegetative communities. The effects of SO2 in

Douglas-fir (Pseudotsuga menziesii), sugar pine

particular have been extensively studied and occur

(Pinus lambertiana), incense-cedar (Libocedrus

at all levels of resolution from the metabolic pro-

or Calocedrus decurrens), California black oak

cess level to the ecosystem level. In the work

(Quercus kelloggii), and Jeffrey pine (Pinus

reported on here, we wanted to predict the effects

Jeffrey!).

at the population and community levels given the

results of the effects at the whole plant level.

MODEL DESCRIPTION

We have developed other models to forecast effects

at the process level (Kercher 1977; Kercher 1978).

SILVA calculates environmental parameters of

The model, SILVA, uses an empirical dose-response

the stand and initializes number and sizes of the

relationship for the effects of pollutants at the

trees from environmental and control data respec-

tree-level. By virtue of the ecological interac-

tively. A table of good and bad seed crop years

tions contained in the model, the effects at the

and a list of fire years is generated. The effect

tree level are translated into effects at the

of pollution on trees is calculated. The

community level.
number of new seedlings for that year, the growth

of each tree,,and mortality are then determined for

We have followed the modeling approach devel-

each year. Growth is modeled as a difference

oped by Botkin and others (1972) who developed

equation in the tree dbh and as a function of en-

vironmental variables. The killing is done sto-

chastically depending on the probability of death

presented at the Symposium on Effects of Air

as determined by ecological risk, lack of growth,

Pollutants on Mediterranean and Temperate Forest

and fire damage. The dynamics of fuel accumulation

Ecosystems, June 22-27, 1980, Riverside,

(litter and brush) are also modeled.

California, U.S.A.

Temporal Seed Crop Patterns--For the conifers

of the Sierra Nevada, there can be significant

temporal variations in the annual cone production.

~nvironmentalScientist; Electrical Engineer;

and Environmental Scientist, Lawrence Livermore

We modeled the phenomenon of high and low yield

Natl. Laboratory (LLNL), Livermore, Calif. opera-

seed years as .a Bernoulli random process with

ted by the University of California for the Dept.

blocking. If the species is in an unblocked state,

of Energy under contract number W-7405.

the probability of a good crop is p and of a poor

crop is (1-p). If a good seed crop occurs, the

the process is assumed to be blocked for r-1 years.

dels the probit of the effect being proportional

The parameters p and r were taken from cone crop

to the log of the generalized dose.

data.

SIMULATION RESULTS

Fire Ecology--Fire is a critical factor in the

population dynamics of western forests. The most

Fire Ecology--Figure 1 shows the response of

important aspect of fire is fire-induced mortality.

ponderosa pine and white fir with fire occurring

The occurrence of fire was also modeled as a

at the natural frequency and with complete fire

Bernoulli random process with blocking and p and

suppression. Ponderosa pine is well adapted to

r are based on fire incidence data. The blocking

fire and dominates where undergrowth is thinned by

in this case arises from the time required for

fire. The model reproduces this result and indi-

fuel to build back up to levels capable of sup-

cates white fir would eventually outcompete pon-

porting fire propagation. Fire kills by raising

derosa pine in the absence of fire. The model

the temperature inside the tree and by damaging

suggests that a significant factor in the fire

the crown. Fire intensity is calculated in kilo-

adaptation of ponderosa pine is its growth rate

watts/meter of fireline length using FIREMOD

and growth form which allow it to evade fire by

(Albini 1976) and probability of death is deter-

minimizing the time that the crown is exposed to

mined as a function of dbh, bark thickness, and

fire. The effects of fire on tree mortality is

scorch height. Scorch height is calculated from

shown in figure 2a. Note the shift in age of

fire intensity, ambient temperature, and windspeed.

death to the lower ages in the presence of fire.

Moisture Stress--The effects of moisture stress

Pollution Simulations--As an example of effects

are modeled by multiplying the difference equation

of pollution, consider the minimally significant

for growth by a moisture stress factor. Parame-

case of 10% growth reduction in ponderosa pine.

ters for this function are taken from published

We scaled the response of the remaining species

ranges of tolerance data. The moisture stress

according to their published relative sensitivi-

factor is a function of the ratio of actual evapo-

ties. These calculations used the seasonal aver-

transpiration to potential evapotranspiration.

age model. The results for ponderosa pine and-

white fir (fig. 3) indicate that while white fir

MODELING SO2-POLLUTANT EFFECTS

undergoes a nominal growth reduction of about 1 to

2% per tree with pollution, total basal area actu-

It has long been held that chronic injury re-

ally shows a dramatic increase. This is due to

sults from sulfate accumulation in plant tissues.

the much greater growth retardation that the do-

Guderian (1977) has suggested that in most cases

minant species experiences. Tree mortality of

involving a single point source, chronic injury

ponderosa pine (fig. 2b) indicates the trees are

results from the "short-term action of relatively

at higher risk at higher ages under pollution.

high concentration peaks". Thus the long-term

The older, slower growing, pollution-stressed

average air concentration can be quite low due to

trees have size-dependent risks comparable to

the large number of pollution-free time periods.

those of the younger unstressed trees. We can

Because two different perspectives exist, i.e.,

summarize (fig. 4 ) the results for ponderosa pine,
(1) measuring average annual concentration or

accumulated dose or (2) regarding injury as aris-

4 0 , 1 , 1 , I '
ing from episodes and making detailed measurements
-
35
of episode parameters, we have two different pol-

lutant-effects submodels.
30 -

25 -
Seasonal Average Submodel--This approach as-

sumes that growth reduction is a simple function

Without fire
-
of the SO2 concentration averaged over the growing
I
season, or equivalently, of the integral of SO2
- 10 -1 -
.c
concentration over time. We use a dose-response
E 5 -
function in which growth decreases linearly with
So L ~ l l l l t l ~
increasing accumulated dose based on the prelimi-

nary study of the tree-ring data of Lathe and

14
McCallum (1939) for ponderosa pine grown near the
Without fire
smelter at Trail, B.C.
12
10

Successive Episode Model--An alternative ap-

proach is to calculate the accumulated damage

caused by successive short episodes separated by

4 With fire
time intervals with no or negligible pollution.
2 ^-,,
One method to implement this approach would be to

0 100 200 300 400 500

use a process model (Kercher 1978). The method
Time from clearcut (yr)
used here is an empirical dose-response where the

dose is that accumulated from successive episodes

Figure I--Average of basal area from 25 simula-

(Kercher and Axelrod 1980). We use the empirical

tions showing effects of fire. (a) Ponderosa pine

dose-response of Larson and Heck (1976) which mo-

(b) White fir.

 0.080

0.070

0.060
Ponderom Pine

EÑÑ
Ponderom Pine
NO SO,
With SO,

- 0
Ponderosa Pine White Fir Douglas Fir

Figure 2--Fraction of trees which died in simula-

Figure 4--Boxplots of polluted and unpolluted

tions of figure 1 plotted against age at death.

cases. Median is line at notches. Top of box is

(a) With and without fire. (b) With and without

75th percentile; bottom of box is 25th. Range is

pollution.
vertical line. Non-overlapping notches indicate

significance at 95% level.

Botkin, D.B., J.F. Janak, and J.R. Wallis.

1972. Some ecological consequences of a compu-

ter model of forest growth. J. Ecol. 60:

849-872.

Guderian R.

1977. Air pollution. 127 p. Springer-Verlag,

New York.

Kercher, J.R.

-
l ' l r
White Fir
l T l ' 1977. GROW1: A crop growth model for assessing

- impacts of gaseous pollutants from geothermal

technologies. UCRL-52247, 33 p. Lawrence

Livermore Natl. Laboratory, Livennore, Calif.

Kercher, J.R. ,
1978. A model of leaf photosynthesis and the

effects of simple gaseous sulfur compounds

(H2S and SO2). UCRL-52643, 37 p. Lawrence

Livennore Natl. Laboratory, Livermore, Calif.

1 , l ~ I ~ l ~ Kercher, J.R. and M.C. Axelrod.

0 100 200 300 400 500

Time from clearcut lyr)
1980a. A model for forecasting the effects of

SO2 pollution on succession in a western

Figure 3--Basal area growth with and without pol-

coniferous forest: Interim Report. UCID-

lution for (a) pine and (b) fir.

18537. 58 p. Lawrence Livennore Natl. Labora-

tory, Livermore, Calif.

white fir, and Douglas-fir by using boxplots of

Kercher, J.R. and M.C. Axelrod.

the distributions of the 500 annual data points of

1980b. SILVA: a model for forecasting the

each species fraction of the total basal area.

effects of SO2 pollution on growth and suc-

Note the decrease in pine and the increase in fir

cession in a western coniferous forest:

with pollution. The basal area of Douglas-fir is

Final report. UCID (to be published). Law-

extremely reduced. The environmental conditions

rence Livermore Natl. Laboratory, Livennore,

were poor for Douglas-fir even in the absence of

Calif.

SO?. The competitive disadvantage for Douglas-fir

Larson, R.I. and W.W. Heck.

is made worse by pollution because Douglas-fir is

1976. An air quality data analysis system for

sensitive to SO->and carries its needles longer

interrelating effects, standards, and needed

than ponderosa pine. Thus the growth reduction

source reductions: Part 3. Vegetation injury.

for an individual tree (greater than that for pon-

J. Air Poll. Control Assoc. 26:325-333.

derosa pine) translates into a much larger effect

Lathe, F.E. and A.W. McCallum

on basal area.
1939. The effect of sulphur dioxide on the dia-

meter increment of conifers. 3 Effect of

LITERATURE CITED
sulphur dioxide on vegetation. National Re-

search Council of Canada. p. 174-206. N.R.C.

Albini, F.A.
No. 815. Ottawa, Canada.

1976. Computer-based models of wildland fire

behavior: a user's manual. 68 p. USDA For.

Serv., Intermt. For. and Range Stn., Ogden,

Utah.

 Forest Models: Their Development and

Potential Applications for Air Pollution
Effects Research1

H. H. Shugart, S. B. McLaughlin, and D. C. west2

Abstract: As research tools for evaluating the effects of

chronic a i r pollution stress, forest simulation models
offer one means of i ntegrati ng forest growth and develop-
ment data with generalized indices of pollution stress.
This approach permits consideration of both the competitive
interactions of trees in the forest stand and the influ-
ences of the stage of stand development on sensitivity of
component species. A review of forest growth models,
including tree, stand, and gap models, i s provided as a
means of evaluating re1 ati ve strengths, weaknesses, and
1imi t s of appl i cabi 1i t y of representati ve examples of each
type. Data from recent simulations with a gap model of
eastern deciduous forest responses to a i r pol 1uti on stress
are presented to emphasize the potential importance of
competition in modifying individual species' responses in a
forest stand. Recent developments in dendroecology are
discussed as a potential mechanism for model validation and
extended appl i cati on.

Atmospheric emissions from widespread indus-
t r i a l and urban sources have now significantly
a1 tered the a i r qua1 i t y of extensive forested
regions of the world. Wolak (1971) described
the influence of industrial emissions on
forested areas of Poland as an abiotic para-
natural ecological factor. He viewed the
results of these emissions on forest succession
as the establishment of a new final sera1 stage
termed the industrio-climax. Assessing the
impacts of these changes and those which may
ensue as we rely increasingly on fossil fuels in
'presented a t the Symposium on Effects of Air
Pollutants on Mediterranean and Temperate Forest
Ecosystems, June 22-27, 1980, Riverside, Califor-
nia, U.S.A.
Z~esearch Staff Members of the
Environmental Sciences Division, Oak Ridge
National Laboratory, Oak Ridge. Research
supported by the National Science Foundation's
Ecosystem Stud i es Program under Interagency
Agreement No. DEB77-25781 with the U.S.
Department of Energy under contract W-7405-eng-26
with Union Carbide Corporation. Publication No.
1545, Environmental Sciences Division, ORNL.
the future i s a challenge made comsiderably more
d i f f i c u l t by the complex nature of forest eco-
systems. The perennial growth habit of forest
trees and the nature of their competitive inter-
actions in a forest community make d i f f i c u l t the
evaluation of chronic exposures of forests t o
atmospheric pol 1utants. Treshow (1970) pointed
out that t e r r e s t r i a1 ecosystems are del i cately
balanced with a structure that may depend on a
few c r i t i c a l species. He indicated the response
of vegetation may be slow, b u t once natural
balances are sufficiently disrupted, subsequent.
alterations may occur much more rapidly because
of irreversible a1 t e r a t i ons of essenti a1 system
functions or species interactions.
Traditionally, studies of responses of forest
trees t o a i r pollution stress have focused pri-
marily on species level responses, seedlings, a
few selected physiological processes, and gen-
eral ly simp1 i s t i c exposure regimes. While valu-
able informati on has been gained on specific
pl ant-pol lutant interactions, we s t i 11 know very
1 i t t l e about the potenti a1 effects of pollutants
on forest communities. For instance, how are
individual species effects integrated over space
and time into responses of the forest community?
What are the probable limits of impacts on
forests based on our current knowledge of
s e n s i t i v i t y of individual species responses?
To address these questions necessitates that
we combi ne both autecologi cal and synecological
approaches. The former we can derive in large
part from dose-response data for individual
species. In the l a t t e r task, we can derive from
the experiences of two decades of experimenta-
tion with mathematical simulation of the growth
and development of forest communi t i es (Rei chl e
and others 1973, Munro 1974, Shugart and West
1980). The purpose of t h i s paper i s to review
the basic components of these models with a view
toward understanding their strengths and weak-
nesses and t h e i r potential u t i l i t y as tools for
studying comnunity-1 eve1 responses to a i r pol 1u-
tion stress.
Computer Models of Forest Dynamics
In the mid-19601s, foresters and ecologists
independently began to develop extremely
detailed computer models of forest growth and
development. Foresters realized that certain
changes in f o r e s t practice (e.g., change in
trees due to genetic improvement, use of fer-
t i l i z e r in f o r e s t s ) would render less useful the
stand yield tables that had been laboriously
developed over the prior several decades. Some
foresters began to develop models of forest
growth and yield that could be calibrated on the
extant, stand-table data s e t s and could also be
used to incorporate some of the changes in
forestry practice ( f i g . 1). A t the same time,
ecologists became dissatisfied with the s t a t i c
notion of forest typology and developed inten-
s i ve investigations (e.g., the International
Biological Program) of the dynamic aspects of
ecosystems. This increased interest in eco-
system dynamics led naturally t o the development
of f o r e s t models. By the mid-1970's (fig. I ) ,
three approaches evolved to modeling the long-
term dynamics of f o r e s t s (table 1). We will
discuss the u t i l i t y of each of these approaches
in terms of i t s applicability to assessing the
consequences of a i r pollution effects over long
time scales. The approaches are:
(1) Forest models consider the forest as the
focal point of the simulation model. For-
e s t r y yield tables constitute a highly data-
dependent subset of these f o r e s t model s.
( 2 ) Tree models take the individual t r e e as
the basic unit of a f o r e s t simulator. The
degree of complexity ranges from simp1e
tabu1 ati on of the probabi 1i t i es of an
individual t r e e of one kind being replaced by
an individual of another kind t o extremely
detailed models that include 3-dimensional
geometry of different species at different
sizes.
( 3 ) Gap models dynamically simulate particu-
l a r attributes of each individual tree on a
prescribed spatial unit of relatively small
ORNL DWG 80 H U B ESD
M I D 1960's
TABLES 1 COMMUNITY
CLASSIFICATION 1
FORESTERS RECOGNIZE ECOLOGISTS BECAME
LATE 1960's
THE POSSIBILITY OF CHANGE INCREASINGLY AWARE OF ECOSYSTEM
I N TREE GENETICS AND FORESTRY
PRACTICE
FOREST DYNAMICS AND OF
USE OF COMPUTERS
M I D - 1970's
MODELS MODELS MODELS
4
WORK
PROPOSED
"I
** PUBLISHED
42
LATE 1970 '1 APPLICATION OF DEVELOPED MODELS TO
NEW PROBLEMS INCLUDING POLLUTION EFFECTS
Figure I--Recent historical origins of computer
models used for pollution effects assessment a t
the forest ecosystem level.
size. The spatial unit i s usually either a
gap in the forest canopy or a sample quadrat.
In general, the model type used i s based on
the problem considered, the data available, and
the desire to develop a flexible model. The
t r e e and forest model categories correspond t o
the t r e e and stand model categories used in a
recent review of forestry models (Munro 1974).
In the present review, gap models (which might
be considered a special case of three models)
are recognized as a category developed exclu-
sively for use in studying ecological succession.
Forest Models
Yield tables used in forestry management are,
in f a c t , empirical models of expected responses
of an even-aged forest of (usually) a single
species. In t h i s context, a forest i s taken as
a larger spatial dimension than either single
t r e e or gap models considered explicitly.
Comparable succession models have been
developed using a variety of mathematical
approaches. Most of these models consider the
landscape to be composed of a number of mosaic
elements that chanae in response to success.iona1
processes. These changes may be viewed as proba-
-a---
listit5 (e.g., Wilkins 1977, Hool 1966) or
continuous (Shugart and others 1973), depending
on modeling assumptions relating to the actual
size of the landscape considered. Forest models
tend to be data-dependent concerning changing
rates of the mosaic elements assumed to comprise
the forests, and the actual mechanisms that
cause the changes in the forests do not appear
explicitly in the models. All of the forest
models listed (table 1) require l i t t l e computer
time and can be solved analytically in many
 Table 1. C l a s s i f i c a t i o n and c h a r a c t e r i z a t i o n o f f o r e s t s i m u l a t i o n models as t o o l s f o r e v a l u a t i n g s t r e s s e f f e c t s .

Model

categor
Age-structure 1 Space 1 Examples
Limitations
Assessment p o t e n t i a1

Advantages

Forest Even ( u s u a l l y ) Nonspatial Most y i e l d t a b l e s i n use i n Usual 1y c a l i b r a t e d on High degree o f

f o r e s t r y today long-term data s e t s on r e a l ism because o f
many d i f f e r e n t s i t e s . data input.
Slow t o develop.
Familiar t o the
f o r e s t r y industry.
Mixed Nonspati a1 ~ 0 0 11966 U s u a l l y r e q u i r e s data Provide a r e g i o n a l
Olson and C h r i s t o f o l i n i 1966 o r i n s i g h t s t h a t are inventory o f effects.
Moser and H a l l 1969 c o l l e c t e d over a l o n g
Shugart and o t h e r s 1973 t i m e period. Mathematically
Johnson and Sharpe 1976 simple and c o u l d be
W i 1k i n s 1977 coupld w i t h eco-
nomic models.
Newnham 1964 Require extremely Tremendous d e t a i l .
Lee 1967 d e t a i l e d growth data
M i t c h e l l 1969 and o t h e r d e t a i l e d Economic v a r i aoles
L i n 1970 parameters. (e.g., board f e e t ,
B e l l a 1971 products) simulated
Tree Even Hatch 1971 C m e r c i a l forests only directly.
Hegyi 1974 are considered.
L i n 1974
Establishment may n o t
be considered.
Nonspati a1 C l u t t e r 1963 Require extremely Economic v a r i a b l e s
C u r t i s 1967 d e t a i l e d growth data. (e. g., board f e e t )
Dress 1970 simulated d i r e c t l y .
Goulding 1972 Commercial f o r e s t s o n l y
S u l l i v a n and C l u t t e r 1972 are considered. F a s t computationally;
Burkhart and Strub 1974 c o u l d be i n t e r f a c e d
Solomon 1974 Establishment may n o t w i t h economic models.
C l u t t e r 1974 be considered.
E l f v i n g 1974

S p a t i a1 Adlard 1974 Detailed data Tremendous d e t a i l .

Arney 1974 requirement.
Mixed Ek and Monserud 1974 Have been proposed
M i t c h e l l 1975 Computationally slow. f o r use i n long-term
p o l l u t i o n assessment.
Nonspati a1 Leak 1970 Lack o f d e t a i l i n Models nave oeen
Bosch 1971 output. explored f o r t h e i r
Namkoong and Roberts 1974 t n e o r e t i c a l aspects.
F o r c i e r 1975 Require c l e v e r va1 i- Level o f a b s t r a c t i o n
Suzuki and Umemura 1974 d a t i on procedures. i s b o t h an advantage
Horn 1976 and disadvantage.
Noble and S l a t y e r 1978
Waggoner and Stephens 1970

Gap Mixed Vertical B o t k i n and o t h e r s 1972 Spatial i n the v e r t i c a l Have been used i n
Shugart and West 1977 dimension only. 1ong-term p o l 1u t i on
M i e l k e and o t h e r s 1978 assessment.
Tharp 1978 Require c l e v e r v a l i-
Shugart and Noble 1980 d a t i o n procedures. Complex parameters
Shugart and others 1980 can be i n f e r r e d from
Doyle and o t h e r s 1980 ecological principles.

cases. All of these models could be used for spacing, and harvesting schemes in commercial
assessing the consequences of some inferred forests. These models produce information used
pollution effect on a region's forests assuming primarily by 1arge governmental or industri a1
that the primary problem of estimating the for- land managers which i s as a consequence, nor-
e s t stand response could be overcome. mally communicated by direct means that do not
necessarily involve the s c i e n t i f i c l i t e r a t u r e
Spati a1 l y Explicit Tree Models e . g . , internal reports). The models we l i s t e d
i n these categories (table 1) are probably only
Two categories of models in table 1 (even- a subsample of such models that are actually in
aged or mixed age) are used almost exclusively use.
in sophisticated evaluations of planting,
 These models function by incrementing indi-
vi dual trees (usually tree diameter, crown
volume, and various form and shape parameters)
periodically and are usually solved in 1- t o
5-year time steps. To i l l u s t r a t e the degree of
detail used in such models, Mitchell's (1969)
model of white spruce (Picea glauca) uses
branch-pruning of trees that overlap to deter-
mine competition interaction.
The models explicitly consider the crowding
of trees and can be easily adapted to either
even- or mixed-age stands. In f a c t , Hegyi's
(1974) even-aged model i s derived from Arney's
(1974) mixed-age model, and Mitchell Is (1969,
1975) models are derived in the converse man-
ner. The models are designed for commercial
forestry operations and do not include phenomena
that ecologists would expect in a succession
simulator. They generally ignore establishment
of invading seedlings and often use functions
for geometry of trees that could only be
expected to hold in young, vigorously growing
trees. The models sometimes use thinning or
harvest as a surrogate for mortality. Because
of the level of detail needed, these models
synthesize great amounts of autecological data
that are usually only available for commercial
species and are difficult to extend to mixed-
species forests. Nonetheless, the FOREST model
( E k and Monserud 1974) does simulate mixed-
species, mixed-age northern hardwood forest in
Wisconsin. This model i s also being considered
for use in a pollution effects assessment pro-
blem (fig. 1). There i s also a potential t o
apply the other models of the commercial species
that should be expl ored.
Even-aged, Nonspatial Tree Models
Even-aged, nonspati a1 models have been used
in commercial forestry also and are logical non-
spatial a1 ternatives to models in the previous
category. Nonspatial models have been used
almost exclusively in pine (Pinus spp.) planta-
tions and are usually in the form of differen-
t i a l equations with basal area, stocking
density, and volume (biomass) of a forest stand
changing with respect to time. Because these
relationships are functions of the size of the
average tree, the models contain parameters
derived from the expected growth of trees. The
even- aged, mono-speci es character of the simu-
lated forests allows the assumption that mathe-
matical functions for the expected response of
an average or typical tree are sufficient to
express these re1 ati onships among volume, stock-
ing, and basal area. These models work best if
the trees tend to be the same size, which helps
t o explain the use of these models in the more
genetically optimized, short-rotation, crop-like
Pi nus plantations. The underlying assumptions
of these models 1imit their applications to
even-aged stands, and the development of mixed-
aged models using t h i s approach i s difficult.
Unlike the spati a1 mono-species models we dis-
cussed previously, these models can, in some
cases, be solved analytically and, in all cases,
require only a moderate amount of computer time.
Mixed-age, Nonspatial Tree Models
These models simulate ecological succession
in naturally regenerated forests. Their empha-
s i s i s on birth/death processes affecting indi-
vidual trees, and the importance of tree growth
and form i s greatly deemphasized. They are not
particularly complex (i.e., birth and death of
trees might be treated as simple stochastic
processes; rep1 acement of trees as a first-order
Markov process), b u t frequently i t i s the stated
objective of the authors to attempt to capture
the salient aspects of succession with a minimal
model representation. In this objective, the
models are actually explorations into the con-
sequences of theories and assumptions on the
nature of ecological succession based on the
attributes of the species involved (Gleason
1926, Drury and Nesbit 1973).
The models can provide considerable insight
into patterns of ecosystem dynamics and can be
solved analytically without resorting to digital
computation. An example of t h i s modeling
approach (Noble and Slatyer 1978) uses the vital
attributes of species t o determine the expected
patterns of community successions generated by
competition among the species. Vital attributes
considered are the modes that a species uses t o
persist at a s i t e , the modes for establishment,
the avail abi 1i t y of a method or persistence
(e.g., seeds, vegetative sprouts) at different
l i f e stages of the plants (propagule, juvenile,
mature, extinct), and longevity of individuals.
Using these species attributes, they construct
schematic diagrams of changes that can be com-
pared with observational data from a given area.
Gap Models
Gap models simulate year-to-year changes in
diameters of each tree on a plot of known area.
These models do not account for the exact loca-
tion of each tree b u t use tree diameters t o
determine tree height and then use simulated
leaf area profiles to devise competition rela-
tionships due to shading. These models are
spatial in the vertical b u t not the horizontal
dimension. This simp1 i f ication greatly reduces
the cost of running these models and also elimi-
nates the consideration of complex spatial pat-
terns of trees, should t h i s be important in a
given application. The vertical gap models are
probably best used in studies of successional
dynamics of natural forests considered over long
time spans. Gap models have a1 so been the f i r s t
detailed succession simulators applied to a i r
pollution effects research.
Current Model Applications
Most models built s t r i c t l y for forestry use
are usually intended as applications in a
restricted set of specified circumstances.
 ORNL-OWG 7 8 -6488RAR
Given the great specificity of the models, they
s t i 11 simul ate cornnerci a1 ly important forest
YELLOW POPLAR ( I 1 BLACK OAK (I)
types, and i t is unfortunate that they have yet
to be used in any pollution effects studies.
Several of the succession models presented in
v
table 1 have been used in evaluating environ-
mental impacts on naturally occurring forests.
Botkin (1973, 1977) considered the effects of
CO2 enrichment on plant growth and subsequent
effects on forest dynamics. He found t h a t an
a r b i t r a r i l y assumed percentage change in rate of
photosynthate production at the individual plant
level in CO2-enriched atmospheres was not WHITE OAK ( R l BLACK CHERRY ( S )
manifested directly as a change in forest
growth. Other effects such as plant competition
and shading tended to 1ower the magnitude of the
system response. McLaughl i n and others (1978)
and West and other (1980) performed model
experiments on chronic a i r pollution stress
expressed as a change in growth rates of
poll uti on-sensi t i ve trees. They noted that the

response of growth over the long term and in
natural forests might vary in direction as well
as in magnitude from what one might predict from
1aboratory or greenhouse studies. Kickert ( t h i s
symposium) and Kercher ( t h i s symposium) have
also used these gap models of western forests to
investigate 1ong-term pollutant effects. All of
these studies identify a common problem; namely,
in natural forests where trees vary in spacing,
size, and competitive responses, one cannot
extrapol ate directly from 1aboratory studies t o
f i e l d conditions. Forest succession models can
provide and have provided a necessary adjunct t o
1abor atory-based assessments of environmental
effects. We will provide a detailed example of
such an application in the following section.
Gap Model Application
As used in the following example, the model
(the FORET model, Shugart and West 1977) con-
siders 33 forest tree species native to the
southern Appal achi an region and simul ates growth
of individual trees on a circular 1112-ha plot.
The growth of each tree on a plot i s incremented
yearly as a function of (1) total annual growing
degree days (5.6OC base), ( 2 ) the total leaf
area of t a l l e r trees on the plot, ( 3 ) total num-
ber of trees on the plot, and (4) the size of
the tree. A typical simulation i s illustrated
in figure 2.
The selection of a species for the plot and
subsequent i n i t i a t i o n and growth of the tree are
based on si lvicul tural characteristics of each
species. These characteristics include:
(1) s i t e requirements for germination,
( 2 ) pal atabi 1i t y of seedlings for browsers,
( 3 ) sprouting potential, (4) shade tolerance,
( 5 ) germi nati on and growth temperature require-
ments, ( 6 ) inherent growth potenti a1 ,
( 7 ) longevity, and (8) sensitivity to crowding
stress ( f i g . 2). The i n i t i a l trees established
on a plot with bare soil are those having shade-
intolerant growth requirements and germination
a f f i n i t i e s for mineral soil. As the simulation
p
TOTAL BIOMASS
OTHER SPECIES
100
50 'Y
. '& ---
0
200 400
TIME (yr)
Figure 2--Species and stand dynamics of a forest
with and without continuous exposure t o a i r
pollution stress (- unaffected; -----
affected).
proceeds, trees that have the a b i l i t y t o
germinate in leaf l i t t e r and grow under shaded
conditions are selected by the model. Leaf
l i t t e r is assumed to have accumulated to a level
commensurate with the total tree biomass for the
plot. The amount of shade cast by each tree i s
a function of leaf area of the tree and i s cal-
cul ated allometrical ly from i t s diameter by
totaling the leaf area of all t a l l e r trees on
the plot. Under optimal conditions, tree growth
i s assumed to occur at a rate that will produce
an individual of maximum recorded size ( d b h ) f o r
that species during the period of maximum
recorded age and i s based on a curci linear func-
tion that grows a tree t o two-thirds i t s maximum
dbh at one-half i t s age. Modifications reducing
this optimal growth are imposed on each tree by
some additive combination of shading and crowd-
ing from other trees on the plot and the sto-
chastic variation from optimum climate. Optimum
climate i s defined as the means of the minimum
and maximum growing degree-days within an indi-
vidual species range. Death i s a stochastic
process with the probabi 1i t y of dying inversely
related to the yearly growth increment. Total
stand density characteristics are calculated
from dbh. Ingrowth occurs by germination of
seeds and sprouting, and simulation may be ini-
tiated either from a bare plot or an existing
stand of a predetermined composition and
structure.
Validation of the FORET model was accom-
pl ished by simul ati ng a deciduous forest stand
with and without American chestnut as a viable
species (Shugart and West 1977). Simulations
with chestnut removed produced forests of simi-
1ar composition to the contemporary, post-
chestnut blight forest. With chestnut included,
the model produced a forest similar (Spearman
rank correlation - r = 0.83, see Siege1 1956) in
composition to the re1 ati vely undistrubed south-
ern Appalachian forest which existed around 1890
to 1910. All simulations were typically
repeated for a large number of plots ( 2loo),
and interpretations were based on average bi o-
mass of individual species and the forest stand
determined from the mu1 t i p l e runs.
By utilizing t h i s type model, we investigated
the results of the interaction of forest t r e e
competition and a i r pollution stress. In doing
t h i s , the f 011owing re1 evant questions concern-
ing the response of forests to a pollutant were
considered:
(1) What level of a i r pollution s t r e s s would
be required to significantly a1 t e r f o r e s t
growth and development?
( 2 ) How are s t r e s s effects integrated over
time?
( 3 ) How important i s competition i n moder-
ating or enhancing induced stresses on indi-
vidual species?
( 4 ) How are species responses integrated
into the response of f o r e s t systems?
Application of the model t o the study of the
effects of a i r pollution stress on growth and
development of eastern forests necessitated (1)
developing a r a t i onale f o r cl assifyi ng species
in terms of t h e i r relative s e n s i t i v i t y to t h i s
stress and ( 2 ) i ncorporati ng growth reductions
into the model which reflected species' sensi-
t i v i t y ranking and a range of impacts which
might be expected under f i e l d conditions.
Addressing the f i r s t task assumes that spe-
cies vary measurably in t h e i r growth responses
to chronic a i r pollution stress. Such a conclu-
sion i s intuitively obvious from a wealth of
data from controlled laboratory and f i e l d
studies where obvious differences in s e n s i t i v i t y
of foliage t o visible injury from a i r pollution
have been demonstrated. Data on relative sensi-
t i v i t y of f o r e s t trees to growth reduction from
chronic a i r pollution s t r e s s are limited, how-
ever. In t h i s application, we made the assump-
tion that trees most sensitive to f o l i a r injury
would also be most sensitive to growth inhibi-
tion. We group the 32 species into 3 sensitiv-
i t y classes (resistant, intermediate, and
sensitive), based on their relative s e n s i t i v i t y
to visible injury. The sensitivity classifica-
tion was based on 10 years of f i e l d survey data
of vegetation near a coal-fired e l e c t r i c plant
(McLaughlin and Lee 1974) and an extensive
sunmary of f i e l d and laboratory data on sus-
c e p t i b i l i t y of woody plants to SO2 and photo-
chemical oxidants reported by Davis and Wilhour
(1976). This classification then formed a
framework f o r addressing the second task,
determining appropriate levels of growth reduc-
tion to introduce into the modeled forest. For
eastern forests, t h i s task must also rely on the
rather 1imi ted data currently available from the
1i terature. However, one advantage of mathe-
matical models i s that a range of s t r e s s levels
may be simulated. While not providing exact
quantitative answers, such an approach does
permit one to bracket the range of likely
responses based on the best available data.
In the FORET approach, both the influence of
varying stress levels and the stage of f o r e s t
maturity at which stress was initiated were
examined. Results of a typical simulation a r e
presented in figure 2. Here, responses of
selected species are shown from a simulation in
which annual growth inhibitions of 20, 10, and 0
percent were imposed on seedlings in sensitive,
intermediate, and resistant sensitivity classes,
respectively. Increases in biomass of 4 major
species [yell ow popl ar (intermediate), white oak
( r e s i s t a n t ) , black oak (intermediate), and black
cherry (sensitive)] , the coll ecti ve "other" spe-
cies category, and total stand biomass were com-
pared with and without simulated a i r pollution
stress as the forest developed over time.
The results indicated that competition within
the forest stand may greatly modify responses
predicted from individual species' s e n s i t i v i t y
to stress. Both enhanced growth suppression
(black oak and black cherry) and reduced sup-
pressi on (ye11ow popl a r ) were demonstrated.
These responses were attributed to s h i f t s in the
competitive potenti a1 of these species induced
by differential stress applied within the f o r e s t
stand. An examination of total biomass of a l l
species indicated that suppression could be
greater than (as high as 20 percent) or less
than ( < 5 percent) that of the weighted average
suppression (7 percent) imposed in the simula-
tion.
Another useful capability inherent in sim-
ulation approaches i s that variations in stand
age and, relatedly, stand composition may be
introduced f o r the time of s t r e s s initiation.
In the FORET t e s t , stage of stand development
was also identified as an important modifier as
shown in figures 3 and 4. Yellow poplar, a
f ast-growing, shade-intolerant species which
showed growth stimulation when the seedling
forest was stressed ( i n i t i a t i o n time - year O),
 ORNL-DWG 7 8 - M 0 9 R
LIRIODENDRON TULIPIFERA
if0 1
40 % STRESS
BEGIN YEAR 0
----- BEGIN YEAR 50
......... BEGIN YEAR 400
0 50 400 450 200 250 309 350 400 450
YEARS
Figure 3--Response of ye1 1ow pop1 a r ( L i r i oden-
dron t u l i p i f e r a ) t o a 10% reduction in growth.
Growth reducing s t r e s s i s applied a t year 0,
year 50 or year 400.
showed growth reduction in the more mature
f o r e s t ( i n i t i a t i on time - year 50) where other
species compete more favorably in the closing
f o r e s t canopy. Black oak, on the other hand,
when stressed in the seedling f o r e s t showed a
g r e a t l y enhanced growth reduction. When. s t r e s s
was i n i t i a t e d a t year 50, however, the response
was q r e a t l y delayed until other more r e s i s t a n t
speci-es suih as Ghite oak began t o dominate (see
f i g . 2).
The e f f e c t s of d i f f e r e n t i a 1 levels of sen-
s i t i v i t y on growth and competition of f o r e s t
t r e e s which we have shown in f i g u r e 2 are sup-
ported by the f i e l d responses of deciduous t r e e s
measured by Brandt and Rhodes (1972, 1973). In
t h e i r studies of the e f f e c t s of 25 years of
limestone dust deposition on a deciduous f o r e s t ,
they found changes i n composition, with
increased dominance of ye1 1ow pop1 a r , white oak,
and red oak a t t h e s i t e of heavy dust accumula-
t i o n . Reduced l a t e r a l growth ( 2 18 percent) of
s e n s i t i v e species such as red maple, chestnut
oak, and red oak was accompanied by a 76 percent
increase i n l a t e r a l growth of yellow poplar a t
t h e t e s t s i t e near t h e limestone quarry (Brandt
and Rhodes 1973). Evidence of the amplification
of e f f e c t s of a b i o t i c s t r e s s by both i n t e r - and
intra-specific competition has also been
ORNL- DWG 7 8 - 4 9 6 0 8 R
QUERCUS VELUTINA
40 % STRESS
BEGIN YEAR 0
5
----- BEGIN YEAR 50
........ BEGIN YEAR 400
0
500 0 50 400 150 200 250 300 350 400 450 500
YEARS
Figure 4--Response of black oak (Quercus
velutina) t o a 10% reduction in growth. Growth
reducing s t r e s s i s applied a t year 0, year 50 or
year 400.
demonstrated by Fox and Caldwell (1978) in
studies with UV-B radiation. In s i t u a t i o n s of
severe mutualistic competition, some species
showed improved growth under the UV-B treatment,
a response a t t r i b u t e d t o improved competitive
status. Other examples of changes in p l a n t
competition under a i r pol 1ution s t r e s s were
reviewed by Guderian and Kuppers (1980) in t h e
preceding paper in t h i s session.
Validation of Forest Community Response t o S t r e s s
While the v a l i d i t y of model r e s u l t s may be
r e a d i l y checked against actual growth and devel-
opment patterns of "normal" f o r e s t s of a region,
evaluation of responses of disturbed f o r e s t s
becomes a much more d i f f i c u l t task. I t implies
developing a c a p a b i l i t y t o c l e a r l y distinguish
differences among measured values of parameters
of stand growth and composition and those which
would have occurred in t h e absence of pollutant
s t r e s s . Accomplishing t h i s necessitates e i t h e r
obtaining measurements on comparable stands over
a v a r i e t y of s t r e s s l e v e l s or documenting t h e
growth c h a r a c t e r i s t i c s of t h e stand in question
before the s t r e s s was i n i t i a t e d . In e i t h e r
case, the investigator i s faced with measuring
pollutant e f f e c t s in the f a c e of the wide vari-
e t y of b i o t i c and a b i o t i c variables control 1ing
growth of individual trees and forest
comnun i t i es.
Historicallyy documentation of forest
responses to rather high levels of gaseous pol-
lutants primarily SO2 and HFy from smelting
processes was f aci 1itated by the typical occur-
rence of we11 -defined gradients of stress with
distance from the sowce. Gordon and Gorham
(1963)Â for instance, were able to measure
i ncreased numbers of higher p1 ant species a1 ong
a 63-km gradient from the smelters at Sudburyy
Ontario. These changes followed a generalized
pattern of rep1 acement of more highly evolved
species of 1ater successional stages by the more
broadly adaptedy stress-tolerant genera1 i s t s
which Woodwell (1970) reported following point-
source radiation s t r e s s of a deciduous forest
comnun ity.
Present-day a i r pollution stress regimes can
generally be characterized as induced by gen-
e r a l l y lower levels of pollutants contributed by
mu1 t i p l e sources. High-level point sources have
been largely replaced by area sources where
1ocal topography and meteor01ogy combine t o
concentrate mu1 t i point eff 1uents. C1 assic
examples are the Los Angeles Basin in the West
and numerous industri a1 corri dors a1 ong river
valleys in the East. These areas provide good
possibilities for examining species and commu-
n i t y responses to chronic and occasionally acute
stress regimes.
Commun it.y-1 eve1 effects of oxi dants on
forests of the San Bernardino Mountains near Los
Angeles were described originally by Miller
(1973) and have formed a basis for a broadly
based study of a variety of ecosystem processes
at t h i s s i t e . Kickert and Gimnel (1980) used
these data in parameterizing a forest simulation
model to describe these changes. In the Easty
McCl enahen (1978) examined 7 deciduous forest
stands located along a gradient of chronic a i r
pollution stress on a 50-km portion of the
heavi 1y industri a1 ized Ohio River Valley.
Species richness evennessy and Shannon diver-
s i t y index were genera1 ly depressed for both
overstory and understory layers in the forest as
proximity to industrial a i r pollution sources
increased. Stem density in the overstory
decreasedy while lower s t r a t a showed increased
abundance of species along t h i s same gradient.
Shifts in re1 a t i ve species' importance were a1 so
noted.
Studies of the l a t t e r type provide very valu-
able data for describing the types of changes
that may x c u r under moderate pollution s t r e s s y
b u t are limited in their u t i l i t y for predicting
rates of change over time or at varying stress
levels. Information of t h i s type may be con-
tained in the chronology of t r e e growth at that
and other s i t e s however. Recent developments
in tree-ring analysis provide a potentially
powerful tool for analyzing both the rate and
direction of within-community changes.
Dendroecology i s a discipline of dendrochro-
no1 ogyy the science of dating annual growth
rings of woody plants ( F r i t t s 1971). I t can be
considered a companion tool with dendroclima-
tology to examine changes in tree growth in
re1 ation to local and regional environment. The
basic conceptsy applications, and limitations of
dendroecol ogy have been discussed by Fri t t s
(1971). In generaly i t r e l i e s on multivariate
s t a t i s t i c a l analysis t o identify principal
variables influencing tree growth. Resultant
equations are in themselves models of individual
tree growth over time. As a tool for studying
a i r pollution effects, dendroecology permits
separation of effects of tree age and local
cl imate from those induced by a i r pollution
(Nash and others 1975). Phillips and others
(1977ayb) have used t h i s approach to correlate
growth reductions in stands of lob1 011y and
white pine with production levels near an army
munitions plant. More relevant to the chal-
lenges of providing reliable predictions of
species and community-level changes i s the
potential u t i l i t y of t h i s technique for detect-
ing growth responses in our eastern regional
environment. Measurements of growth reductions
of white oak in apparent response to chronic
s t r e s s of t h i s type have been reported near
LaPorte, Indianay by Ashby and F r i t t s (1972).
In t h i s casey the decade during which anomalous
growth reductions occurred was associated with a
heavy incidence of smoke and haze in that region.
Documentation of pollutant histories in the
broader regional context represents a more
d i f f i c u l t task b u t one of great importance t o
efforts t o eventually develop a predictive
potential. A greatly expanded network of a i r
quality trends; howevery data for the past 40
yearsy during which emissions i n the Eastern
United States increased sharply, are lacking.
One potentially useful tool for obtaining his-
t o r i e s of exposure t o genera1 a i r pollution
s t r e s s i s heavy metal analysis in the individual
rings (Lepp 1977). This approach has been used
in Sweden (Symeonides 1979) t o construct his-
tories of heavy metal pollutiony a1 though Ti an
and Lepp (1975) caution that factors such as
radial transport and soil uptake must be f u l l y
understood to use t h i s technique accurately. In
the Swedish study, both lead and copper showed
l i t t l e lateral movement and were useful in con-
structing a decade-level history of metal pollu-
tion at the study s i t e . Recent developments
coupling x-ray emission spectroscopy (Val kovic
and other 1979) with growth-ring analysis show
promise for using a variety of trace elements
for historical analyses. As these techniques
are developed furthery they may provide useful
data for constructing historical indices of
regional-scale chronic stress.
The tools for validating or modifying f o r e s t
simulators as predictive tools appear t o be
either available now or close at hand. We feel
that dendroecol ogi ca1 approaches have tremendous
potential for unlocking a wide variety of
species/comunity/environment interactions which
will make t h i s task ultimately possible. Pro-
bably t h e g r e a t e s t value of t h e f o r e s t simula-
t o r s i s in predicting the consequences of s e t s
of I'most 1ogical I' assumptions regarding poll u-
tion e f f e c t s on t r e e s . Other assumed relation-
ships can be tested e a s i l y y and new information
may be added as i t i s developed (Kozlowski
1980). The mode1 i s merely a tool t o be used in
t h i s synthesis and refining process.
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Poster Summaries
Effects of Oxidant Air Pollutants on Pine Litter-fall and the Forest Floor
2
Rodney J. Arkley and Rudolph Glauser

Oxidant i n j u r y t o western yellow p i n e s (Pinus showed no t r e n d whatsoever.

ponderosa and P. Jeffrey!) i n t h e San Bernardino The response of t h e s e elements i s perhaps due
Mountains r e s u l t s i n n e e d l e i n j u r y followed by t o d e c l i n i n g c e l l w a l l t h i c k n e s s w i t h i t s calcium
i n c r e a s e d f a s c i c l e m o r t a l i t y and a b s c i s s i o n , p e c t a t e and an i n v e r s e d i l u t i o n e f f e c t on N , P,
decreased n e e d l e l e n g t h , branch m o r t a l i t y and and K.
f i n a l l y t r e e m o r t a l i t y . The d e g r e e of i n j u r y h a s The d a t a f o r t h e r e g r e s s i o n l i n e s shown i n
been e s t i m a t e d each y e a r f o r 6 y e a r s by a s c o r i n g Figure 2 a r e a s follows:
system based on o b s e r v a t i o n of a l l of t h e s e
f a c t o r s w i t h b i n o c u l a r s and combining them i n t o a
s i n g l e o x i d a n t i n j u r y s c o r e (01s). F o l i a g e
N(pct) -
,598
P ( p c t ) = .0707
-
-
.00255
.00034
QIS,
OIS,
r = -.264,
r = -.28,
n
n
=
=
187
185
d e n s i t y i s d i r e c t l y p r o p o r t i o n a l t o OIS a s K(pct) = .346 - .00331 OIS, r = -.37, n = 187
expected. The number of annual needle-whorls Ca(pct) = .303 + .00431 OIS, r = .58, n = 147
r e t a i n e d (W) i s r e l a t e d t o t h e s c o r e by W = 0.171
x OIS w i t h r = .75** and t h e grams of n e e d l e s p e r
twig by F = 2.71 x OIS w i t h r = .62*. Note t h a t
OIS d e c r e a s e s w i t h i n c r e a s i n g i n j u r y and a s c o r e
of 0 i n d i c a t e s a dead t r e e . Scores g r e a t e r t h a n 0 MEAN 1
35 i n d i c a t e no obvious i n j u r y .

EFFECT ON NEEDLE-FALL
The oven-dry weight of n e e d l e s c o l l e c t e d on
0.209 m2 s c r e e n s p l a c e d under p i n e s of v a r y i n g
I
I
- RANGE
0
*
"
n

OIS i s shown i n F i g u r e 1. The average annual

n e e d l e f a l l i n c r e a s e s from 1 3 1 gm/m2 under
h e a l t h y t r e e s t o 357 gm/m2 w i t h OIS of 9 t o 1 4 , ^-\+I
- \
5
/
and d e c r e a s e s a s t h e t r e e n e a r s d e a t h ,
The weight p e r n e e d l e f a s c i c l e i n t h e l i t t e r -
I \ - Â
N
.2S'Si
1 I ' -2
-
Â
f a l l decreases progressively with increasing
i n j u r y a s shown a l s o i n F i g u r e 1. (Sig. < .001). ./ b .20
The i n c r e a s e d l i t t e r - f a l l (170 p e r c e n t ) can 1 Ã

b e expected t o i n c r e a s e t h e t h i c k n e s s of l o o s e I
NEEDLE SIZE
- .IS
Z

d r y l i t t e r on t h e f o r e s t f l o o r w i t h consequent
i n c r e a s e d f i r e hazard and d e c r e a s e d s e e d l i n g
germination.
Oxidont Injury Score
P l a n t N u t r i e n t Content Dead 4 Heolthy
L i t t e r - f a l l samples were analyzed t o determine Fig. 1. E f f e c t of o x i d a n t i n j u r y on p i n e n e e d l e
t h e e f f e c t of o x i d a n t i n j u r y on t h e p l a n t n u t r i e n t f a l l and n e e d l e s i z e .
of t h e l i t t e r . The r e s u l t s a r e shown i n F i g u r e
2. The s c a t t e r of p o i n t s ( n o t shown) r e p r e s e n t e d
by t h e r e g r e s s i o n l i n e s i s wide, b u t 187 samples
were analyzed and t h e t r e n d s i n d i c a t e d a r e
c l e a r l y r e a l , s i n c e they a r e highly s i g n i f i c a n t
(P = < .001). Magnesium was a l s o analyzed b u t

p r e s e n t a t t h e Symposium on E f f e c t s of A i r
P o l l u t a n t s on Mediterranean and Temperate F o r e s t
Ecosystems, June 22-27, 1980, R i v e r s i d e , 3
I 1 I I 1 1
C a l i f o r n i a , U.S.A. z 01
10 20 30 40 50
2 ~ o i Morphologist
l and S t a f f Research A s s o c i a t e , Oxidant Injury Score
r e s p e c t i v e l y , Dept. of S o i l s and P l a n t N u t r i t i o n ,
Dead < Healthy
U n i v e r s i t y of C a l i f o r n i a , Berkeley, C a l i f . Fig. 2. Needle-fall c o n t e n t of p l a n t n u t r i e n t s .
 Regional Air Pollution Impacts on Forest Growth
Thomas V. Armentano, O r i e L. Loucks, and Wayne T. w i l l i a m s 2
Recent s t u d i e s underway i n t h e Ohio R i v e r Basin
Energy Study (Loucks 1980) have shown t h a t c h r o n i c
a i r p o l l u t i o n l e v e l s may b e reducing r e g i o n a l
growth over much of e a s t e r n North America. During
June through August, 1977, monitoring s t a t i o n s
r e p r e s e n t a t i v e of f o r e s t a r e a s recorded h o u r l y
maxima > 0.10 ppm on 1 4 t o 27 p e r c e n t of t h e days,
and maxima > 0.05 ppm on 70 t o 93 p e r c e n t of t h e
days.
A survey of e a s t e r n w h i t e p i n e (Pinus s t r o b u s
L.) s t a n d s i n r u r a l and urban l o c a t i o n s throughout
c e n t r a l and s o u t h e r n I n d i a n a showed widespread ozone
damage symptoms: c h l o r o t i c m o t t l i n g , c h l o r o t i c
dwarfing and premature n e e d l e senescence, on a
s c a l e from moderate t o s e v e r e . Sycamore ( P l a n t a n u s
o c c i d e n t a l i s L . ) , s i l v e r m a p l e (& saccharinum L . ) ,
yellow p o p l a r (Lireodendron t u l i p i f e r a L . ) , and t h e
b l a c k oak group (Quercus s p p . ) a l s o a r e somewhat
s e n s i t i v e . The 37.3 x l o 6 a c r e s of f o r e s t s i n t h e
Ohio Basin y i e l d about 40 f t 3 / a c r e 1 of wood an-
n u a l l y , b u t a n e s t i m a t e d 25 p e r c e n t of t h e f o r e s t
c o n s i s t s of 0 3 - s e n s i t i v e s p e c i e s , i n d i c a t i n g a
y i e l d r e d u c t i o n from o x i d a n t e f f e c t s (and i n t e r a c -
t i o n s w i t h o t h e r g a s e s and pathogens), r a n g i n g
from 3 t o 6 p e r c e n t a n n u a l l y (Table 1 ) .
Table I--Total l o s s i n wood p r o d u c t i o n i n Ohio
River Basin f o r e s t s e s t i m a t e d t o r e s u l t from d i r e c t
and i n d i r e c t a i r p o l l u t a n t e f f e c t s upon f o r e s t
growth and m o r t a l i t y r a t e s . T o t a l normal wood
y i e l d f o r t h e r e g i o n i n 1970 was 1 . 5 x 10' f t 3 .
Data e x p r e s s e d i n m i l l i o n s of c u b i c f e e t .
Wood Annual
Production Mortality
Loss 1 Loss T o t a l Loss
I
( p e t . ) 1 ( l o b f t 3 ) ( P C ~ . 1)( 1 0 b f t 3 ) (pet.)
Independent s t u d i e s i n d i c a t e 03 l e v e l s w i l l in-
c r e a s e i n t h e Ohio River Basin o v e r t h e n e x t 20
y e a r s , depending on i n c r e a s e s i n u t i l i t y NOx emis-
s i o n s . A c o n s e r v a t i v e energy development s c e n a r i o
' p r e s e n t e d a t t h e Symposium on E f f e c t s of A i r
P o l l u t a n t s on Mediterranean and Temperate F o r e s t
Ecosystems, J u n e 22-27, 1980, R i v e r s i d e ,
C a l i f o r n i a , U.S.A.
r e s e a r c h S c i e n t i s t , S c i e n c e D i r e c t o r , and
Research S c i e n t i s t , r e s p e c t i v e l y , The I n s t i t u t e of
Ecology (TIE), I n d i a n a p o l i s , I n d i a n a .
s u g g e s t s t h a t O 3 l e v e l s w i l l c o n t i n u e upward a t
l e a s t through 1985. A t t h e c u r r e n t r a t e of 03 i n -
c r e a s e , 0.02 ppm p e r decade, wood l o s s e s could
r e a c h 166 x l o 6 f t 3 , t w i c e t h e c u r r e n t e s t i m a t e d
l o s s e s (Table 1 ) .
A p a t t e r n of i n c r e a s i n g t r e e m o r t a l i t y i n t h e
r e l a t i v e l y young e a s t e r n f o r e s t s (30 p e r c e n t mor-
t a l i t y i n c r e a s e i n t h e N o r t h e a s t , and 10 p e r c e n t
i n t h e South from 1962 t o 1970) h a s been r e p o r t e d
d u r i n g a p e r i o d of s i g n i f i c a n t a i r p o l l u t i o n in-
c r e a s e s i n t h e N o r t h e a s t (U.S. F o r e s t S e r v i c e 1978).
T h i s m o r t a l i t y may b e a t t r i b u t a b l e , a t l e a s t i n
p a r t , t o d e g r a d a t i o n of a i r q u a l i t y . This hypothe-
sis i s supported by n e a r l y c o n s t a n t m o r t a l i t y i n
w e s t e r n f o r e s t s where a i r p o l l u t i o n i s g e n e r a l l y
low, d e s p i t e o v e r m a t u r i t y i n t h e s e f o r e s t s . I f mor-
t a l i t y l o s s e s i n t h e Ohio Basin were i n t e r m e d i a t e
between t h o s e of t h e Northeast and South from 1962
t o 1970, t h e l o s s of wood would b e 4.08 f t 3 / a c r e /
y r l . Proportionately greater losses a r e indicated
f o r 1985 and 2000 (Table 1 ) .
Other s t u d i e s underway i n d i c a t e t h e s e f o r e s t
l o s s e s could b e s i g n i f i c a n t f o r t h e g l o b a l C02
b a l a n c e (Armentano and R a l s t o n 1980). Because of
a f a v o r a b l e s t a n d age d i s t r i b u t i o n brought on by
p a s t h a r v e s t p a t t e r n s , temperate zone f o r e s t s now
s t o r e around 10' t o n s of carbon a n n u a l l y i n long-
l i v e d t r e e components, 20 p e r c e n t a s much a s t h e
carbon r e l e a s e d from f o s s i l - f u e l combustion. T h i s
s t o r a g e could c o n t i n u e f o r t h e n e x t two decades, b u t
i n c r e a s e d h a r v e s t , f o r e s t m a t u r a t i o n , and a i r p o l l u -
t i o n e f f e c t s can reduce carbon s e q u e s t e r i n g and wood
p r o d u c t i o n r a t e s . Thus, only management of f o r e s t s
focused on a b a l a n c e between wood p r o d u c t i o n and
wood accumulation can p r o v i d e optimum economic and
ecological benefits. I f present a i r pollution
t r e n d s c o n t i n u e , and i f r e l a t i o n s h i p s t o f o r e s t
growth suggested i n t h i s paper a r e s u b s t a n t i a t e d by
f u r t h e r r e s e a r c h , t h e long-term p r o d u c t i v i t y of f o r -
e s t s w i l l b e t h r e a t e n e d i n s e v e r a l r e g i o n s of t h e
United S t a t e s .
LITERATURE CITED
Armentano, T.V., and C.W. R a l s t o n .
1980. The r o l e of temperate zone f o r e s t s i n t h e
g l o b a l carbon c y c l e . Can. J. For. Res. 10:53-60.
Loucks, O.L. (ed.).
1980. Crop and F o r e s t Losses Due t o Current and
P r o j e c t e d Emissions from C o a l - f i r e d Power P l a n t s
i n t h e Ohio River Basin. TIE Report i n review.
Indianapolis.
U.S. F o r e s t S e r v i c e .
1978. F o r e s t S t a t i s t i c s of t h e U.S. 1977. De-
partment of A g r i c u l t u r e .
 Canopy Analysis of Pollutant Injured Ponderosa Pine in the San Bernardino
National Forest l 3
M.C. Axelrod, P.I. Coyne, G.E. Bingham, J.R. Kercher, P.R. Miller, and R.C. ~ u n ~ ~

Nine trees were selected from a ponderosa pine

tween known points to produce a set of initial

(Pinus ponderosa Laws.! stand which was estab-

estimates. The algorithm first computes the

lished after a fire in the mid-1950's. These

cumulative needle surface area as a function of

trees were classified into three injury groups,

height, then a linearizing transformation is made.

[(I) slight, (2) moderate, and (3) severe injury]

Linear interpolation is carried out on this new

in accordance with a scoring devised by P.R.

curve. The transformation is then inverted and

Miller. During the summers of 1978 and 1979, a

first differences taken, resulting in a reproduc-

detailed inventory was made of the canopy on each

tion of the original data and a new set of esti-

tree. The lengths of all main stem internodes

mates for the missing data. The new estimates are

were measured along with the number of primary

. used as initial values each iteration. A weighted

branches radiating from each of these internodes.

average of the needle conditions for each primary

At roughly every other main stem internode, sev-

branch was also computed. These averages are then

eral of the primary branches received a detailed

combined to yield an index of the condition of the

inventory. The length of the primary branch and

needles on the whole tree.

the number of internodes were recorded. This in-

ventory was carried on through to the secondary,

In table 1A, we see the total leaf area index

tertiary, and quarternary branch levels. The

for each needle age class, with the trees grouped

compass angle of each of the inventoried primary

into injury classifications. We can see the leaf

branches was also recorded. Whenever needles were

area indices decrease across injury classes with-

found on an internode, the following information

in age groups. There is a pronounced decrease

was recorded: (1) needle age, (2) needle condi-

in the retained leaf area with needle age even in

tion on a scale of 0-4, (3) the number of fasci-

the slightly injured group; the severely injured

cles, (4) the average needle length, (5) the

group has essentially no needle area except in the

average needle chord width, and (6) the length of

1 year age category. Table 1B shows the (weighted)

the internode bearing the needles. Note that

average needle condition for the inventoried trees

while not all the primary branches at a selected

and presents further evidence that ozone injury is

main stem internode were inventoried, the ones

dose accumulative with young needles being less

selected did receive a complete inventory.

affected by ozone than older needles. Since the

current year needles were still growing at the

A preliminary analysis of the 1978 canopy data

time the initial inventory was taken, they were

has been completed, characterizing the distribu-

inventoried separately the following year. This

tion of needle surface area for each tree. The

data is still being processed and is not yet

total needle surface area for each inventoried

available, but is expected to add proportionately

primary branch was computed separately for each

to the numbers presented here. The trend in leaf

needle age. In order to determine the distribu-

area decline indicates the competitive disadvan-

tion of needle surface area for a whole tree, we

tage of an injured tree.

estimated the needle area at the main stem inter-

nodes where no data was taken. The curve repre-

Table I--Needle properties by needle age for nine

senting needle area as a function of height is

trees under pollution stress.^-

approximately bell-shaped. We have developed an

algorithm designed to yield estimates of the

missing points and a smooth curve. The algorithm

(A) Leaf Area Index

is iterative and uses linear interpolation be-

Injury Class 1 Year 2 Year 3 Year

Slight 9.53 4.13 0.70

presented at the Symposium on Effects of Air

Moderate 5.23 0.73 0.00

Pollutants on Mediterranean and Temperate Forest

Severe 3.55 0.03 0.00

Ecosystems, June 22-27, 1980, Riverside,

(B) Average Needle condition2 of Canopy

California, U.S.A.

Slight 0.96 1.31 1.28

~lectricalEngineer, Lawrence Livermore Natl.

Moderate 2.18 2.42 NA

Laboratory (LLNL), Livermore, Calif.; Plant Phys-

Severe 2.63 NA NA

iologist, Southern Plains Range Research Station,

uSDA/SEA/AR, Woodward, Okla.; Environmental Scien-

tists, LLNL; Research Plant Pathologist, Pacific

" ~ o e snot include data for current year nee-

Southwest For. and Range Exp. Stn., Riverside,

dles. Averages for 3 trees in each injury class.

Calif.; and Environmental Scientist, LLNL.

~ e e d l econdition scale: 0 = green; 2 =
work performed under the auspices of the U.S.
chlorotic mottle. 4 = uniform yellow with
necrosis.

Department of Energy by the Lawrence Livermore

Natl. Laboratory under contract number W-7405-

ENG-48.

Photosynthesis and Stomatal Behavior in Ponderosa Pine Subjected to Oxidant Stress:

Water Stress Response 112

Gail E. Bingham and Patrick I. coyne3

Light response curves for net and/or gross

qX. When predawn ifiv was in the range from -3 to

photosynthesis, stomatal conductance, and needle

-5 bars, Cs (and Pn) decreased only slightly dur-

xylem potential of ponderosa pine (Pinus ponderosa

ing the normal daily decrease in ifiv due to

Laws.) subjected to water and/or ozone stress

changes in the diurnal course of irradiance.

were studied in the field and laboratory. In the

Minimum ^iy under these conditions seldom exceeded
field, measurements were made on a stand of pon-
-14 bars. However, when $x was forced below -15

derosa pine in the San Bernardino National Forest,

to -17 bars (by severing the fascicle from the

which has experienced long-term oxidant fumiga-

branch) a sharp decrease in Cs occurred, with

tions from the south coast air basin since their

complete stomatal closure (Cs<O.O1 cm/s) in the

establishment following fire in the mid-1950's.

range of -36 to -40 bars. When predawn

These trees were stratified for comparative stud-

started in the -5 to -8 bar range, however, a

ies into three groups (slight injury, moderate

bilevel relationship between Cg and ^iy was ob-
injury, and severe injury) having similar oxidant
served, with Cmax reaching an initial full light

symptoms, on the basis of the scoring system

value, followed at some later time by a step

of P.R. Miller.
decrease to a lower value. Conductance and Pn

remained at this significantly lower level

Controlled studies using ten healthy, uniform,

throughout most of the day, with an accelerated

Oregon-grown, six-year-old saplings growing in

closing trend toward the late afternoon. Late

55 1 containers were conducted to elucidate

afternoon values were usually only 60 to 70 per-

specific field responses. These trees were ap-

cent of their light corrected morning opening

proximately 2 m tall, and were randomly allocated

value.

to positions in two naturally-lighted, mylar-

covered, air and humidity conditioned exposure

Care had to be taken when interpreting pre-dawn

chambers. One chamber was supplied charcoal

$x measurements in the field. It was often ob-

filtered air and the other with air containing

served that at 'a predawn ifiv of -3 to -7 bars, the
0.01 ppm ozone for six hours/day during the
stomata would partially open in the predawn hours,

midday period.
resulting in Cs values approaching one-tenth of

their full light value. Under these conditions,

Measurements were made at regular intervals

sufficient transpiration occurred to make predawn

from May through October at the forest site and

qX very sensitive to wind speed. Differences be-

during the fumigation and water stress cycles in

tween predawn $x on calm and windy mornings as

the laboratory. During the laboratory study, net

large as 3 bars were observed.

photosynthesis (Pn) and stomatal conductance (Cs)

measurements were made at constant humidity and

The second factor depended solely on the pre-

temperature using the LLNL developed minicuvette

dawn xylem potential and controlled the maximum

system, with only the fascicle being measured and

stomatal conductance (Cmax) observed during the

a few surrounding needles exposed to light. The

diurnal cycle. This relationship could not be

rest of the tree was surrounded with heavy black

adequately examined in the forest due to untimely

cloth from before dawn until after light response

late season rains during the three years that

curves had been measured on three fascicles.

field studies were conducted, and was investigated

using potted trees. As predawn $x decreased below

The relationship between needle xylem poten-

-7 bars in control trees and about -10 bars in

tial and maximum stomatal conductance (Cmax)

fumigated trees, a drastic reduction in Cmax and

was not single valued, and depended on predawn

Pn was observed. In trees kept in filtered air,

Cmax was decreased from 0.36 to 0.036 cm/s by the

decrease of predawn $x from -5 to -15 bars. Since

"presented at the Symposium on Effects of Air

the majority of the tree was kept in the dark dur-

Pollutants on Mediterranean and Temperate Forest

Ecosystems, June 22-27, 1980, Riverside,

ing the measurement, the relationship between Cs

California, U.S.A.
and $x at QX values greater than can normally be

observed in nature (due to root and xylem resis-

tance) were examined. Conductance and Pn of fumi-

'work performed under the auspices of the U. S.

gated trees in this region was not significantly

Dept. of Energy by the Lawrence Livennore National

higher than that observed at potentials associated

Laboratory under contract number W-7405-ENG-48.

with full illumination. Reductions in GaX
of 25,

31, and 44 percent were measured for slight, mod-

3~nvironmentalScientist, Lawrence Livermore erate, and severely injured needles. Net photo-

National Laboratory, Livermore, Calif.; Plant

synthesis for the same needle injury classes was

Physiologist, Southern Plains Range Research

reduced by 38, 54, and 69 percent from the 12.9

Station, USDA/SEA/AR, Woodward, Okla.

mgco2/dir^-h rate measured in trees which had not

been exposed to ozone.

 Oxidant Impact on Ponderosa and Jeffrey Pine Foliage Decomposition1

2

J. N. Bruhn, J. R. Parmeter, Jr., and F. W. Cobb, Jr.

Litter decomposition was studied on four plots

Because not all fungi recorded in incubation

located along the oxidant dosage gradient in the

studies were identified, they are individually

San Bernardino Mountains (SBNF). Precipitation

referred to as categories rather than species.

decreased with decreasing oxidant dosage along

Live SNF foliage yielded approximately twice as

this gradient, while forest floor-level solar

many fungus categories (60) as foliage from any

radiation and temperature ir .eased. Ponderosa

other site. The two BEF plots harbored a few more

(Pinus ponderosa Dougl.) a1.d Jeffrey pine (Pinus

categories (29 and 32) than did HV (27), the

Jeffrey! Grev. & Balf.) dominated the most and healthiest SBNF site. HV, in turn, provided more

least severely impacted sites, respectively.

categories than any other SBNF site (15 to 20).

Nylon mesh envelopes of fresh-fallen litter were

Similar effects were not evident in forest floor

exchanged among healthy and sick trees on the

litter. Oxidants apparently affect live needle

four study sites. Decomposition was measured as

microflora in two ways. By reducing needle lon-

percents overall and nutrient (N, P, K. Ca and Mg)

gevity, internal microflora development is prema-

weight loss.
turely truncated. It also seems likely that oxi-

dants further reduce the variety of fungi coloniz-

During the first two years of decomposition,

ing live pine foliage by eliminating susceptible

its rate at each site was directly related to

species. Insofar as community functional pro-

precipitation and oxidant dose. While site mois-

perties are stabilized by a combination of species

ture apparently dominates litter decomposition,

adaptability and community diversity, reduction of

evidence suggests that oxidant injury to live

live foliage microfloral diversity by oxidants is

needles is directly related to the rate at which

viewed as weakening the functional stability of

they decompose. One year weight loss of ponderosa

these communities. The significance of such weak-

pine litter was negatively correlated (P .05) with

ening is unknown.

the oxidant injury scores (O.I.S.)(Miller 1973)

of litter source trees. One year weight loss by

In a growth chamber experiment, propylene oxide-

litter of both species was negatively correlated

sterilized pine needles were incubated on moist

(P .05) with the O.I.S. of litter destination

forest floor organic matter from either Che SNF or

trees.
one of the SBNF sites. No meaningful differences

in weight loss developed between treatments over

Live needle internal microflora may be initi-

22 weeks, showing that oxidants to date had not

ated in the foliar bud and can be thought of as

significantly impaired the abilities of study-site

pioneers in a succession of microorganisms re-

microflora to cause litter weight loss. In a sec-

sponsible for litter decomposition. Microbial

cond growth chamber experiment, propylene oxide-

populations involved in foliage decomposition

sterilized pine needles were incubated on a uniform

were studied via incubation of surface-sterilized

moist forest floor organic matter mixture in either

live and litter needles on water agar. Eight

filtered air or filtered air enriched with 20pphm

trees from each of two central Sierra Nevada sites,

ozone 8 hours daily for 14 weeks. The weight loss

Stanislaus National Forest (SNF) and Blodgett

difference between treatments did not reach signi-

Experimental Forest (BEF), were included for com-

ficance ( .05 P .01). Any effect in the field
parison with the 15 SBNF study trees. Both taxo-
would be slight and probably limited to surface

nomic richness and population density increased

litter during moderate to severe oxidant episodes.

with needle age. Both parameters increased simi-

larly with age on all four SBNF sites. However,

Acknowledgments: This project was funded in

both parameters increased faster on the somewhat

part with federal funds from the Environmental

less oxidant-impacted BEF, and both parameters

Protection Agency (E. P. A.) under contract number

developed fastest on the least oxidant-impacted

68-03-0273.

site (SNF).

LITERATURE CITED

presented at the Symposium on Effects of Air

Pollutants on Mediterranean and Temperate Forest

Miller, P. R.

Ecosystems, June 22-27, 1980, Riverside,

1973. Oxidant-induced community change in a

California, U.S.A.
mixed conifer forest. In Air pollution damage

to vegetation. p. 101-117. Advances in

2~nstructorof Forest Pathology, Michigan

Chemistry Series, Number 122. American

Technological University, Houghton, Mich.; Pro-

Chemical Society.

fessor and Associate Professor of Plant Pathology,

respectively, University of California, Berkeley,

Calif.

 Integrated Lake-Watershed Acidification Study

Carl W. Chen and Robert A. olds stein'

OBJECTIVES
series and in parallel will produce or consume

acids and will release chemicals that shift the pH

The Integrated Lake-Watershed Acidification

and other chemical equilibrium.

Study (ILWAS) was designed to determine the eco-

logical effects of acid rain under natural condi-

tions (EPRI, 1979, Goldstein et al., 1980). Since

the most widely reported effect of acid rain has

1-
-
Meteorology
Air duality
Rain Quality
been the acidification of lake water leading to

elimination of fish, it is of interest to learn

how and why the ecosystem becomes acidified by

- Throughfall Chemistry
acid rain.

Three forested watersheds (Panther, Woods, and

-Hydraulics
Sagamore) within 15 km of each other in the
-Geochemistry
Adirondack Park region of New York were selected
- Microbial Processes (& Lab)
for investigation. Each watershed has different
Hydrology
configurations and characteristics. Principle
-Aquatic Chemistry
hypothesis of the study is that these differences
Hydrology
may lend to different pH dynamics, i-e., Panther

Lake alkaline, Woods Lake acidic, and Sagamore

Lake in-between.
-Aquatic Chemistry
This research will produce a comprehensive data

- Paleoecology
Hydrology
base for the three watersheds covering a period of

almost 4 years, a series of interpretive reports,

and a calibrated and verified mathematical model.

Figure 1--Field Program Components

The wealth of understanding gained and the mathe-

matical model developed will be readily applicable

Field surveys are being conducted to character-

to other lake basins.

ize the properties of the basic ecosystem compart-

ments in each watershed. At selected locations,

measurements are made for ambient air quality, the

APPROACH
quantity and quality of waters that move through

the system from tree top to lake outlet (see fig.

ILWAS couples field investigation with theoret-

1). Data are collected monthly, weekly, synoptic,

ical modeling. The interactions between the model

and once only, depending on the parameters and

and the field research are practiced in an inter-

their temporal variability. The field program

active manner, each influencing and strengthening

began in 1977 and will be completed in 1981.

the other. Other research findings on mechanisms

and rates of acidification processes are inte-

grated into the model formulations.

MODELING

The model organizes the data into an integrated

FIELD PROGRAM
theoretical framework (Goldstein et al., 1980).

The model also serves as a vehicle to check the

The watershed ecosystem is envisioned to com-

consistency of theory and data from rainfall quan-

prise a cascade of basic compartments: atmosphere,

tity to lake outlet quality. Eventually, the

canopy, snowpack, catchment, soil layers, bogs,

model may provide scientific answers to such

stream segment and lake. These are the compart-

management questions as: What will and will not

ments that the acid rain must pass through before

happen if a certain air quality standard is im-

it reaches lake outlet. As it passes through each

posed, and if the acidity of precipitation is in-

compartment, biogeochemical processes acting in

creased or decreased.

presented at the Symposium on Effects of Air

The model flowchart shows the computation se-

Pollutants on Mediterranean and Temperate Forest

quence (fig. 2 ) . The model calculates dry depo-
Ecosystems, June 22-27, 1980, Riverside,
sition as a function of ambient air quality and

California, U.S.A.
simulates the quantity and quality of water in

throughfall, soil horizons, bogs, stream, lake,

2~ice-~resident,
Tetra Tech, Inc. , Lafayette, and lake outlet (Chen et al., 1978). All impor-

California; Program Manager, Electric Power

tant acidification processes are included in the

Research Institute, Palo Alto, California

model.


Ambient air quality
Basin properties
Dry deposition
Exudate
Calculate
Throughfall chemistry
Snow accumulation
Snowmelt
Ion-teaching
.
Calculate
Soil moisture
Vertical infiltration
Lateral outflow
Organic decomposition
+
Nutrient uptake
.Weathering
Cation exchange
Equilibration (At-sys, etc.)
Quantity and quality
of H20 through stream
Figure 2--Model Flowchart
The model uses a network of compartments to
account for spatial variability of ecosystem. It
updates meteorological conditions daily and am-
bient air quality weekly. The calculations are
performed on a daily time step to simulate the
dynamic responses of ecosystem throughout years.
Graphic outputs are provided to facilitate inter-
pretation.
PRELIMINARY FINDINGS
Data indicate that the H+ deposition rates are
seasonal and are approximately the same for all
three watersheds (Johannes and Altwicker, 1980).
The seasonal pattern of H+ deposition seems to
follow that of SO4 deposition (Johannes, 1980).
The deposition rates of various ions in acid rain
as measured at seven ILWAS stations correlate
well with those measured at the nearby MAP3S
stations. This is significant because it suggests
that acid rain data from regional monitoring sta-
tions may be used to perform preliminary calcula-
tion of acid rain effect for a new site.
The pH at Panther Lake inlet is normally 7.3 to
7.5 throughout the year. The pH at the outlet is
similar except during the snowmelt periods. Dur-
ing that period, pH drops to as low as 5.0. Some-
thing must have happened in the lake. Alterna-
tively, it was argued that the inlet was only a
small spring, not representative of all inflows.
The pH profiles measured in the lake show that
only surface water is acidified during the period
of snowmelt (Hendrey et al., 1980). What is the
source of H+ ions that acidify the lake surface?
To resolve the puzzle, the model was used to
help trace the source of water at the outlet. The
model was first calibrated to the Panther Lake
basin (Chen and Goldstein, 1980). After that,
precipitation falling directly on lake surface is
set to zero. This allows estimation of the con-
tributions of this input to the total observed
outflow. Approximately two-fifths of the peak
flow can be accounted for by the direct precipita-
tion to the lake surface. During that period,
lake water is inversely stratified with respect
to temperature. Direct precipitation which has a
pH of 3.8 to 4.2 quite possibly is deposited right
on the surface to acidify the lake surface water
(pH 5.0). Another possible explanation is surface
runoff resulting from snowmelt which has a high
acidity (Galloway et al., 1980).
The significance of the capability to manipu-
late the model to examine the effect of a single
process on the integrated response of the eco-
system should not be overlooked.
LITERATURE CITED
Chen, C.W., S.A. Gherini, and R.A. Goldstein.
1978. Modeling the Lake Acidification Process.
In Ecological Effects of Acid Precipitation,
-
Report of a Workshop, Galloway, United King-
dom, September 4-8, 1978, ed. M.G. Wood,
Electric Power Research Institute, Palo Alto,
CA, U.S.A., EA-79-6-LD.
Chen, C.W., and R.A. Goldstein.
1980. "Techniques for Assessing Ecosystem Im-
pacts of Air Pollutants." & Proc. Inter-
national Conference Air Pollutants and their
Effect on the Terrestrial Ecosystem [May 10-
17, 1980, Banff, Alberta, Canada].
EPRI Technical Work Statement for the Integrated
Lake-Watershed Acidification Study RP 1109.
1979. Electric Power Research Institute,
Palo Alto, California.
Galloway, J.N., C.L. Schofield, G.R. Hendrey,
and A.J. Johannes.
1980. "Sources of Acidity in Three Lakes Acid-
ified During Snowmelt." In Proc. of Inter-
national Conference on the Ecological Impact
of Acid Precipitation, [March 11-14, 1980,
Sandefjord, Norway].
Goldstein, R.A., C.W. Chen, S.A. Gherini, and
cipitation." Proc. of International Con-

J.D. Dean.
ference on the Ecological Impact of Acid

1980. A Framework for the Integrated Lake-

Precipitation, [March 11-14, 1980,

Watershed Acidification Study. & Proc. of Sandefjord, Norway] .

International Conference on the Ecological

Impact of Acid Precipitation, [March 11-14,

Johannes, A.H., and E.R. Altwicker.

1980, Sandefjord, Norway].

1980. "Atmosphere Imputs to Three Adirondack

Lake Watersheds." 3 Proc. of International

Hendrey, G.R., J.H. Galloway, and C.L. Schofield.

Conference on the Ecological Impact of Acid

1980. "Temporal and Spatial Trends in the Chem-

Precipitation, [March 11-14, 1980,

istry on the Ecological Impact of Acid Pre-

.
Sandefjord, Norway]
 Photosynthesis and Stomatal Response to Light and Temperature in Ponderosa Pine

Exposed to Long-Term Oxidant Stress1

P a t r i c k I.Coyne and G a i l E. ~ i n ~ h a m *

Seasonal courses (May t o October 1977) o f about 20° i n May and 25OC i n J u l y . L i g h t r e s p i -
gross p h o t o s y n t h e s i s ( c a l c u l a t e d from CO2 up- r a t i o n ( e s t i m a t e d as Pgross-Pnet) was h i g h e s t i n
t a k e ) and stomatal conductance were c h a r a c t e r i z e d h e a l t h y younq needles and increased w i t h tempera-
as a f u n c t i o n o f l i g h t and gross and n e t photo- t u r e from 5° t o To t and then l e v e l e d o f f between
s y n t h e s i s were c h a r a c t e r i z e d as a f u n c t i o n o f Topt and 35OC. ~l though l i g h t r e s p i r a t i o n was
temperature (May and J u l y 1978) i n a s t a n d o f inversely related t o oxidant i n j u r y , the r a t i o o f
ponderosa p i n e (Pinus ponderosa m.)
Bernardino N a t i o n a l Forest. The CO2 d i f f u s i o n
i n t h e San Pnet/P ss tended t o decrease w i t h o x i d a n t injury.
~ ~ ~ a r e onx itd a? n~t s t r e s s e d t r e e s n o t o n l y had
pathway was p a r t i t i o n e d i n t o i t s stomatal and reduced r a t e s o f CO2 f i x a t i o n , b u t r e t a i n e d a
r e s i d u a l (mesophyll , c a r b o x y l a t i o n , e x c i t a t i o n ) smal l e r p r o p o r t i o n o f a s s i m i l a t e d carbon a f t e r
r e s i s t a n c e components f o r c o n d i t i o n s o f l i g h t r e s p i r a t i o n l o s s e s . Summary data f o r s e l e c t
s a t u r a t i o n and 20 C. These t r e e s have e x p e r i - parameters a r e compared i n Table 1.
enced l o n g t e r m o x i d a n t f u m i g a t i o n s from C a l i f o r -
n i a ' s South Coast A i r Basin s i n c e t h e i r e s t a b l i s h Table 1--Comparison o f s e l e c t parameters normal-
ment f o l l o w i n g f i r e i n t h e mid-1950's. Nine trees i z e d b y d i v i d i n g each mean b y t h e maxi-
were s t r a t i f i e d f o r comparative s t u d i e s i n t o t h r e e mum mean v a l u e i n each column.
chronic i n j u r y classes ( I - s l i g h t i n j u r y , I 1 -
moderate, I 1 1 - severe) h a v i n g s i m i l a r o x i d a n t I n j u r y Needle 1977 Means J u l y 1978
i n j u r y symptoms based on t h e s c o r i n g system o f Class Age Pmax ] Cmax lr;/r1 Pgross-Pnet
P. R. M i l l e r .
I 0 1.00 0.89 1.00 1.00
Maximum o r l i g h t s a t u r a t e d gross photosyn- 1 0.61 0.84 0.80 0.60
t h e t i c r a t e s (Pmax) and photochemical conversion 2 0.32 0.62 0.59 0.35
e f f i c i e n c i e s ( d P / d I ) I=o were h i g h e s t i n t h e c u r - II 0 0.92 0.92 0.82 0.95
r e n t needles and decreased w i t h i n c r e a s i n g nee- 1 0.53 0.75 0.71 0.51
d l e age and w i t h season. D i f f e r e n c e s among nee- 2 0.17 0.38 0.40 0.18
d l e age c l a s s e s w i t h i n an i n j u r y c l a s s d i v e r g e d III 0 '0.79 1.00 0.59 0.94
as t h e season progressed i n d i c a t i n g an a c c e l e r - 1 0.35 0.57 0.45 0.35
a t i o n o f senescence b y ozone. Maximum stomatal Max. value1 8.30 0.24 0.29 2.46
conductances (Cmax) and stomatal s e n s i t i v i t y t o
i n c r e a s i n g 1 i g h t (dC/dI) I=o d u r i n g opening f o l - ^ p a r meters d e f i n e i n t e x t . U n i t s : P, mg CO2
lowed a s i m i l a r t r e n d t o Pmax and dP/dI except g -7t-1-1; C, cm s-y ; r, s cm-1; age i n years.
i n t h e c u r r e n t needles i n which Cmax and dC/dI The d i f f e r e n t i a l response i n p h o t o s y n t h e s i s
were h i g h e s t i n t h e s e v e r e l y i n j u r e d t r e e s . T h i s and stomatal conductance among these f i e l d - g r o w n
suggests a p o s s i b l e f a c t o r c o n t r i b u t i n g t o d i f - ponderosa p i n e t r e e s growing i n a common e n v i r o n -
f e r e n t i a l ozone s e n s i t i v i t y i n t h i s stand. The ment i n d i c a t e d t h e presence o f e c o t y p i c v a r i a t i o n
r a t i o o f t h e stomatal r e s i s t a n c e f o r CO2 ( r l ) t o i n ozone s e n s i t i v i t y . The d i f f e r e n c e s among
t h e t o t a l r e s i s t a n c e (r', s t o m a t a l + r e s i d u a l ) i n j u r y classes were m a n i f e s t as an a c c e l e r a t i o n
decreased w i t h o x i d a n t i n j u r y , i n c r e a s i n g ' n e e d l e o f t h e normal d e c l i n e i n CO2 f i x a t i o n and stomatal
age, and season suggesting t h a t l o s s o f photo- conductance a s s o c i a t e d w i t h needle aging. P a r t i c -
s y n t h e t i c c a p a c i t y r e s u l t e d more from 1 i m i t a t i o n s
u l a r l y e v i d e n t were t h e premature senescence and
a t t h e c h l o r o p l a s t s than from r e s i s t a n c e t o CO2
a b s c i s s i o n o f needles o c c u r r i n g a t about t h e t i m e
d i f f u s i o n t h r o u g h t h e stomata.
gross CO2 uptake dropped t o 10 p e r c e n t o f t h e
Temperature optima (Top ) f o r photosynthesis p o t e n t i a l f o r c l a s s I c u r r e n t needles w i t h o u t
were s i m i l a r i n a l l i n j u r y c l a s s e s and averaged f o l i a r i n j u r y symptoms. T h i s occurred a t i n t e -
g r a t e d i n c i d e n t ozone doses as low as 450 ppm-h
i n s e v e r e l y i n j u r e d t r e e s ( c l a s s 111) o r as h i g h
'presented a t t h e Symposium on E f f e c t s o f as 800 ppm-h i n s l i g h t l y i n j u r e d t r e e s ( c l a s s I ) .
A i r P o l l u t a n t s on Mediterranean and Temperate
F o r e s t Ecosystems, June 22-27, 1980, R i v e r s i d e , A probable s c e n e r i o f o r o x i d a n t e f f e c t s can
C a l i f o r n i a , U.S.A. be described. As f o l i a r i n j u r y symptoms increase,
p h o t o s y n t h e t i c c a p a c i t y and n e t carbon accumula-
2 ~ 1 a n tP h y s i o l o g i s t , Southern P l a i n s Range t i o n p e r u n i t l e a f mass o r area, mass and area
Research S t a t i o n , USDA/SEA/AR, Woodward, Okla.; p e r needle, needle mass p e r u n i t area, and number
and Environmental S c i e n t i s t , Lawrence Livermore o f needle w h o r l s r e t a i n e d p e r t r e e d e c l i n e .
L a b o r a t o r y , Livermore, Cal i f . These f a c t o r s c o n t r i b u t e t o t h e steady l o s s o f
^ h i s work was supported by a U.S. Dept. o f t r e e v i g o r , weakening them t o t h e p o i n t o f v u l -
Energy c o n t r a c t (W-7405-ENG-48) w i t h Lawrence n e r a b i l i t y t o pathogenic organisms such as r o o t
Livermore L a b o r a t o r y . r o t t i n g f u n g i and b a r k b e e t l e s .
 The Effect of Air Pollution on Western Larch as Detected by Tree-Ring Analysis1
C a r l A . Fox and Thomas H. Nash 1112
The number of p u b l i c a t i o n s i n r e c e n t
y e a r s d e a l i n g w i t h t h e e f f e c t s of a i r
p o l l u t i o n on p l a n t s h a s been voluminous.
A v a s t m a j o r i t y of t h e s e have d e a l t w i t h
t h e e f f e c t s of s h o r t - t e r m ( h o u r s o r d a y s )
f u m i g a t i o n s on p l a n t r e s p o n s e . I n con-
t r a s t t o these studies, there is rela-
t i v e l y l i t t l e i n f o r m a t i o n on t h e l o n g -
term, c u m u l a t i v e e f f e c t s o f a i r p o l l u t i o n
o v e r decades. To examine t h e s e long-term
e f f e c t s , a dendroecological (tree-ring)
a n a l y s i s was employed t o a s s e s s t h e growth
r e s p o n s e o f w e s t e r n l a r c h ( ----- Larix
o-c c i d e n t a l i s N u t t . ) t o s u l f u r d i o x i d e .
- n e g a t i v e r e s i d u a l s o c c u r r i n g when s u l f u r
U t i l i z a t i o n of d e n d r o e c o l o g i c a l t e c h n i -
q u e s p r o v i d e d a means f o r q u a n t i t a t i v e l y
r e m o v i n g t h e e f f e c t o f c l i m a t e on t h e
growth response of l a r c h and, t h u s ,
p e r m i t t e d a n e x a m i n a t i o n of t h e r e s i d u a l
growth response i n terms of l o c a l s i t e
f a c t o r s , s p e c i f i c a l l y s u l f u r dioxide.
F i v e w e s t e r n l a r c h s t u d y s i t e s were
l o c a t e d i n t h e Columbia R i v e r V a l l e y n e a r
t h e lead-zinc smelter a t T r a i l , B r i t i s h
Columbia. This particular location
r e p r e s e n t s a unique study a r e a i n t h a t t h e
s u l f u r d i o x i d e g r a d i e n t has been w e l l
documented, b o t h o v e r s p a c e and t i m e ,
s i n c e s m e l t i n g a c t i v i t y began i n 1896.
S u l f u r e m i s s i o n s from t h e s m e l t e r
i n c r e a s e d i n t h e e a r l y 1 9 0 0 1 s , reached a
maximum i n 1930, and r e s u l t e d i n a con-
commitant d e c r e a s e i n a n n u a l t r e e growth.
A f t e r 1930, s u l f u r emissions decreased
d r a m a t i c a l l y w i t h t h e i m p l e m e n t a t i o n of
p o l l u t i o n abatement measures. However,
t h e growth r e s p o n s e of w e s t e r n l a r c h t o
t h e d e c r e a s e d s u l f u r e m i s s i o n s was n o t
immediate, and r e f l e c t e d t h e low f r e q u e n c y
n a t u r e of t h e v a r i a n c e o f t h e s u l f u r
d i o x i d e e f f e c t on t r e e growth.
p r e s e n t e d a t t h e Symposium on E f f e c t s o f
A i r P o l l u t a n t s on M e d i t e r r a n e a n and
Temperate F o r e s t Ecosystems, June 2 2 - 2 7 ,
1980, R i v e r s i d e , C a l i f o r n i a , U.S.A.
^ ~ e s e a r c hS c i e n t i s t , S o u t h e r n C a l i f o r n i a
Edison Company, Rosemead, C a l i f o r n i a ;
A s s o c i a t e P r o f e s s o r o f Botany, Arizona
S t a t e U n i v e r s i t y , Tempe, Arizona.
C o r r e l a t i o n and r e g r e s s i o n a n a l y s e s
were u t i l i z e d t o d e v e l o p m u l t i v a r i a t e
models f o r t h e l a r c h s t u d y s i t e s sampled
a t varying d i s t a n c e s from t h e T r a i l
smelter. C l i m a t i c models developed f o r
t h e c o n t r o l s i t e ( t r e e - r i n g ) chronology
were a p p l i e d t o s i t e c h r o n o l o g i e s l o c a t e d
within the s u l f u r dioxide affected area t o
remove t h e e f f e c t s o f c l i m a t e on t r e e
growth and examine t h e r e s i d u a l r e s p o n s e
of t h e system. The p a t t e r n of t h e r e s i -
d u a l s c l o s e l y resembled t h e s u l f u r emis-
s i o n s from t h e s m e l t e r with t h e most
e m i s s i o n s were g r e a t e s t .
F u r t h e r r e g r e s s i o n modeling i d e n t i -
f i e d t h e r e l a t i v e importance of s u l f u r
e m i s s i o n s , p r i o r y e a r ' s growth, tempera-
t u r e , and p r e c i p i t a t i o n t o a n n u a l f r e e
growth. I n those s i t e s closest t o the
s m e l t e r , s u l f u r emissions accounted f o r
t h e g r e a t e s t p r o p o r t i o n of t h e v a r i a n c e
c a l i b r a t e d b y t h e r e g r e s s i o n models. As
d i s t a n c e from t h e s m e l t e r i n c r e a s e d , t h e
variance a t t r i b u t a b l e t o s u l f u r emissions
d e c r e a s e d i n each s i t e model. I n a l l of
t h e s i t e models, t e m p e r a t u r e , p a r t i c u l a r l y
summer t e m p e r a t u r e , a p p e a r e d t o b e a
primary l i m i t i n g c l i m a t i c f a c t o r . Prior
growth a l s o accounted f o r c o n s i d e r a b l e
v a r i a n c e i n t h e models w i t h p r e c i p i t a t i o n
v a r i a b l e s a p p e a r i n g t o be o f l e s s e r
importance i n e x p l a i n i n g t h e v a r i a n c e of
the s i t e chronologies.
The r e s u l t s o f t h i s s t u d y demon-
s t r a t e t h e a p p l i c a b i l i t y of t r e e - r i n g
a n a l y s i s i n i d e n t i f y i n g and q u a n t i f y i n g
t h e long-term e f f e c t s o f a i r p o l l u t i o n on
f o r e s t communities. It a l s o p r o v i d e s a
b a s i s f o r examining t h e i n t e r r e l a t i o n s h i p s
between a i r p o l l u t i o n , c l i m a t e , and t r e e
growth.
 Acid Rain: Threshold of Leaf Damage in Eight Species from a Forest Succession1
2
B.L. Haines, M. S t e f a n i , and F. Hendrix

Eight p l a n t s p e c i e s were s u b j e c t e d t o a r t i - The volume weighted average r a i n f a l l pH f o r

f i c i a l a c i d r a i n s of pH 2.5, 2.0, 1.5, 1.0, and Coweeta i s 4.6 w i t h o b s e r v a t i o n s ranging from
0.5 i n o r d e r t o determine t h e t h r e s h o l d f o r and 3.2 t o 5.9. Because t h e pH s c a l e i s l o g a r i t h m i c
symptoms of damage. I n a previous study a r t i - w i t h a d e c r e a s e i n 1 pH u i t corresponding t o a
f i c i a l a c i d r a i n s of pH 5.5, 4.5, 3.5, and 2.5
?
10-fold i n c r e a s e i n t h e H c o n c e n t r a t i o n , i t i s
f a i l e d t o produce symptoms of damage. The pre- apparent t h a t a 100-fold i n c r e a s e in
t h e volume
s e n t s t u d y was designed t o extend t h e pH range. weighted average c o n c e n t r a t i o n of H a t Coweeta
The p l a n t s were E r e c h t i t e s , Robinia, Pinus, would change t h e pH from 4.6 t o 2.6 which i s
Quercus, Carya, Liriodendron, &, and Cornus n e a r t h e t h r e s h o l d of damage. However, t h e
from t h e F o r e s t s e r v i c e ' s owee eta ~ ~ d r o l o ~ i c - extremes may be f a r more c r i t i c a l than t h e vol-
Laboratory n e a r F r a n k l i n , North Carolina. Du- ume weighted average. With t h e lowest pH v a l u e
p l i c a t e 0.01 m l drops of each of t h e 5 simulated recorded f o r Coweeta being pH 3.2, merely a 10-
a c i d r a i n s o l u t i o n s were a p p l i e d t o a s i n g l e f o l d i n c r e a s e i n a c i d i t y t o pH 2.2 i n a s i n g l e
mature l e a f on each of two p l a n t s of seven spe- s p r i n g o r summer storm seems l i k e l y t o b r i n g
c i e s . The e n t i r e experiment was performed twice. damage o r d e a t h t o mature l e a v e s of t h e flowering
I n Pinus n e e d l e t i p s were immersed i n t h e solu- p l a n t s a t Coweeta.
t i o n s . The s i z e s of n e c r o t i c s p o t s a r e shown i n
PLANT SPECIES MD LIFE FORM pH OF DROPLETS APPLIED
Fig. 1. Droplets of pH 2.0 produced brown nec-
0.5 1.0 1.5 2.0 2.5
r o t i c s p o t s on a l l s p e c i e s except Pinus w h i l e
d r o p l e t s of pH 1.0 produced necroses on l e a v e s of
a l l s p e c i e s examined. Pinus n e e d l e s were dam-
aged a t pH v a l u e s between 1.0 and 0.5. Damage was
confined t o younger n e e d l e s and was evidenced by
HERBACEOUS WED (N 6) -
ERECHTIKS HlEMClFOLIA(L) RAF.

browning and c o l l a p s e . For angiosperms t h e s i z e s

of n e c r o t i c s p o t s i n c r e a s e d w i t h decreasing pH.
PUUS STROBUS L.
This s u g g e s t s t h a t t h e l e a f has some b u f f e r i n g
TREE ( N - 8 )
c a p a c i t y which was p r o g r e s s i v e l y overcome by
i n c r e a s i n g d r o p l e t a c i d i t y . Comparison of r e -
s u l t s w i t h l i t e r a t u r e s u g g e s t s t h a t developing
l e a v e s a r e more e a s i l y damaged than a r e mature
l e a v e s used i n t h i s study. No s u c c e s s i o n a l
t r e n d s i n s u s c e p t i b i l i t y were observed.
-
ROBIHIA PSEUDO-ACACIA La
SHRUB TO TREE ( N 6 )

p r e s e n t e d a t t h e Symposium on E f f e c t s of A i r
P o l l u t a n t s on Mediterranean and Temperate F o r e s t
Ecosystems, June 22-27, 1980, R i v e r s i d e ,
C a l i f o r n i a , U.S.A.

' ~ s s i s t a n t P r o f e s s o r of Botany, Department

of Botany, U n i v e r s i t y of Georgia, Athens, GA
ACER RUBRUH L.
30602; Graduate Research A s s i s t a n t i n P l a n t
Pathology, and Prof. of P l a n t Pathology, Dept. THE ( N = 0

of P l a n t Pathology and Genetics, U n i v e r s i t y of

Georgia, Athens, Georgia 30602. Research
supported by U.S. EPA Grant R80530510 and NSF
g r a n t s DEB 7619930, DEB 7705324A01 and DEB
7904537 t o t h e U n i v e r s i t y of Georgia.

Figure 1. Average diameter i n mm of n e c r o t i c

s p o t s on l e a v e s s u b j e c t e d t o d r o p l e t s of 5 pH
levels. (*) Spot diameters n o t measured on
needles, s e e t e x t f o r d e s c r i p t i o n of damage.
 Prioritization of Research on Air Quality Related Resources of the National Parks1
E. A. Howard, L. If. Boyd, W. M. Brock, D. W. Morse, S. S. Shin and K. L. steel-sabo2

The National Park Service (NPS) has the legal

that are susceptible to the effects of air pollu-

responsibility and authority to preserve and pro-

tion. The availability of current research

tect the resources of lands under its jurisdiction.

information or literature on susceptibility must

Air pollution has been recognized as a significant

be similarly considered in establishing research

threat to the parks and the 1977 Amendments to the

priorities. Time constraints on recovering infor-

Clean Air Act provided the Service the authority

mation or research results on air quality effects

and responsibility to protect resources that are

on natural ecosystems has prompted the development

air quality related. The NPS Air Quality Office

of a quick access annotated bibliography that uses

intends to conduct research programs to determine

a codin8 system based specifically on natural and

air pollution effects, t o emphasize the use of

cultural resources of NPS units. The bibliography

sensitive receptors to detect effects, as well as

has been used on several occasions to provide

to ensure that scientific evidence is readily a-

lists of references in support of litigation and

vailable when NPS must make determinations of

testimony for hearings.

adverse effects. These efforts are subject to

time and funding limitations and therefore guide-

Setting priorities for research on air quality

lines on setting research priorities are being

effects is altered by other influences and con-

developed.
straints (fig. 1). Funding and time limitations

must be incorporated into the prioritization. In-

The first step in establishing cause and effect

sufficient funding may cause postponement of the

relationships is to identify the pollutants that

highest priority research until these requirements

may.affect park resources (fig. 1). Pollutants

can be integrated into the budget cycle. Time

from existing or planned sources may be identified

constraints also have considerable influence on

by monitoring or estimated from permit require-

priority setting. Substantive scientific data on

ments. If sources are not known, pollutants must be

air quality effects cannot be produced within the

identified through some early warning process or

short time schedules of hearings or litigation.

on the basis of the potential development of re-

The timing of the budget cycle creates difficulty

gional natural resources. Existing sources of air

in obtaining sustained funding for long-term

pollutants that may affect the natural or cultural

effects research. Political requirements at the

resources of park units must be ranked according

Washington level may override the regional pri-

to pollutant type, concentration, frequency, and

oritization. The setting of priorities must be

duration of exposure of the resource. The rankine

continually updated in conjunction with any change

must also incorporate the estimated lead time be-

imposed by these influences or constraints.

fore new pollutant sources begin operation. The

identification and ranking of pollutant sources is

necessary to economize the search for potential

effects.

POLLUTANT SOURCES

The identified pollutants are then used as a

RANKED ACCORDING TO

basis of comparison between the list of resources

IMMINENCE AND DAMAGE

present on a park unit and the literature or cur-

POTENTIAL

rent research information on resource susceptibility

(fig. 1). The presence or absence of information

on park resources must be incorporated into the

LISTS OF RESOURCES LITERATURE AND

setting of priorities. If these data are unavail-

FOR PARKS OF THE RESEARCH INFOFNA-

able, obtaining information on air quality related

REGION TION ON RESOURCE

resources may be the most pressing need. If the

SUSCEPTIBILITY

resource inventories are available, then the in-

formation from these inventories is used in the

comparison between resources present and those

OTHER INFLUENCES

OR CONSTRAINTS

presented at the Symposium on Effects of Air

Pollutants on Mediterranean and Temperate Forest

PRIORITIZE

Ecosystems, June 22-27, 1980, Riverside,

RESEARCH

California, U.S.A.
NEEDS

2~hysicalScientist and Environmental Protec-

tion Specialists, Denver Service Center, National

Figure I--Considerations in setting research

Park Service, U.S. Department of Interior, Denver,

priorities for air quality related resources.

Colorado.

 Estimation of Adverse Effects of Air Pollution on Danish Forests1

2
I b Johnsen

E f f e c t s o f a i r p o l l u t i o n on Danish f o r e s t s do Areas of woodland in Denmark r e l a t e d t o s p e c i e s .

p r o b a b l y o n l y o c c u r under c o n d i t i o n s , where SO2, 1976. - ( Numbers in 1000 ha. ) -
NO,, a n d / o r 0 3 a r e p r e s e n t t o g e t h e r , t h u s r e s u l t - Fagus
- silvatica 75 Res .
i n g in s y n e r g i s t i c a c t i o n on t h e l e a v e s . The Quercus r o b u r 25 Res.
maximum observed monthly a v e r a g e urban immission Fraxinus e x c e l s i o r 9 Sens.
o f NOx and SO2 a r e w i t h i n t h e r a n g e f o r syner- Acer psuedoplatanus 5 Res.
g i s t i c action hereof. I n r u r a l regions only the Other deciduous spp. 24
most s e n s i t i v e s p e c i e s may b e a f f e c t e d by combi- Deciduous t o t a l 138
n a t i o n s of S02, NOx and 03. Oxidants, and 0 3 i n
p a r t i c u l a r , p r o b a b l y p l a y a s t r o n g e r r o l e in Picea abies Sens.
r u r a l a r e a s s i t u a t e d in t h e o u t e r p e r i p h e r y o f Picea s i t c h e n s i s 173 sens.
c i t i e s . Here l e v e l s exceeding t h e v a l u e s a t Abies spp . 27 Sens.
which e f f e c t s on r a t h e r s e n s i t i v e s p e c i e s occa- P i n u s mugo 30 Res.
s i o n a l l y o c c u r d u r i n g summer time. Unstable Other c o n i f e r o u s spp. 46
s u p e r a d i a b a t i c c o n d i t i o n s combined w i t h h i g h Coniferous t o t a l 276
i n s o l a t i o n l e a d t o h i g h 0 3 formation r a t e s , and
h i g h ground l e v e l S O ~ / N O l~e v e l s a r e observed Woodland t o t a l 4 14
when h i g h s t a c k e m i s s i o n s a r e t r a n s p o r t e d t o t h e
ground r e l a t i v e l y c l o s e t o t h e s o u r c e .
Immission l e v e l s and t h r e s h o l d v a l u e s
Fluorides a r e only o f importance in v e r y re- -
F
s t r i c t e d a r e a s around brick factories, f e r t i l - O3
izer industries etc. , and o f minor s i g n i f i c a n c e -3
(ugm , d i u r n a l means)
i n r e l a t i o n t o Danish forestry.
Urban 40-100. 50-100 25-50 0.5-5
Rural 10-40 5-20 25-200 0.2-0.5
Heavy m e t a l s and hydrogen i o n s r e s u l t i n
i n d i r e c t e f f e c t s a s a d v e r s e e f f e c t s on t h e n u t r i - Single 250 500 100 1
e n t s t a t u s of t h e s o i l and t h e s o i l m i c r o b i a l 100 50
p r o c e s s e s . The map shows a r e a s of Denmark Comb. 50 50
(shaded a r e a s ) where t h e s o i l i s b e l i e v e d t o b e 100 50
most v u l n e r a b l e t o a c i d p r e c i p i t a t i o n ; i t i s
r e a s o n a b l e t o b e l i e v e t h a t t h e s e a r e a s a r e co-
i n c i d e n t w i t h t h o s e e x p e c t e d t o b e most a f f e c t e d
by i n c r e a s i n g heavy m e t a l l e v e l s in t o p s o i l due
t o a t m o s p h e r i c f a l l o u t . The most v u l n e r a b l e
s o i l s a r e i n t e r m e d i a t e between t h e v e r y pod-
z o l i s e d s o i l s o f t h e a l l u v i a l p l a i n s (Western
J u t l a n d ) and brown e a r t h s / c l a y i s h s o i l s w i t h high
b u f f e r c a p a c i t y , and are c h a r a c t e r i z e d by t h e i r
h i g h c o n t e n t o f moraine sand.

p r e s e n t e d a t t h e Symposium on E f f e c t s of A i r
P o l l u t a n t s on Mediterranean and Temperate
F o r e s t Ecosystems, June 22-27, 1980, R i v e r s i d e ,
C a l i f o r n i a , U.S.A.

A s s o c i a t e P r o f e s s o r o f P l a n t Ecology,
U n i v e r s i t y of Copenhagen, Denmark.

F i g u r e I--Areas of Denmark (shaded a r e a s )

dominated by moraine sand d e p o s i t s .
 Changes in Southern Wisconsin White Pine Stands Related to Air Pollution Sensitivity1
David F. ~ a r n o s k y ~
F i f t e e n e a s t e r n white pine (Pinus strobus L. )
sample p l o t s c o n s i s t i n g of a t o t a l of 1523 t r e e s
were e s t a b l i s h e d within a 1 3 km r a d i u s of t h e
coal-burning 1054 MW Columbia Generating S t a t i o n ,
located 40 km north of Madison, Wisconsin. The
sample p l o t s were e s t a b l i s h e d i n 1971 and observed
f r e q u e n t l y during t h e growing season f o r 4 years
and then annually a t t h e end of each growing season
f o r t h e next 5 years. These p l o t s consisted of
p l a n t a t i o n s with t r e e s having uniform ages within
each p l a n t a t i o n but with t r e e s ranging i n age from
1 0 t o 40 years old across t h e 15 p l o t s . Baseline
study during t h e 4 years 'before t h e p l a n t began
operation i n 1975 showed t h a t some 1 0 percent of
t h e white pine t r e e s were s e n s i t i v e i n some degree
t o ambient a i r p o l l u t i o n a s determined by t h e
presence of needle t i p h r n and/or c h l o r o t i c dwarf
symptoms i n one o r more years. The s e n s i t i v e
t r e e s occurred randomly i n t h e p l o t s .
The most common type of a i r p o l l u t i o n symptom
found i n t h e 'baseline study was tipburn c o n s i s t i n g
of reddish brown dieback (0.1 t o 3 cm i n l e n g t h )
on f i r s t - y e a r needle t i p s . The s e v e r i t y of symp-
toms v a r i e d from t r e e t o t r e e and from year t o
year. The most severely a f f e c t e d t r e e s had
stunted t o p s , s h o r t needles, poor needle r e t e n t i o n ,
and were c h a r a c t e r i s t i c of an a i r pollution-induced
syndrome c a l l e d t h e c h l o r o t i c dwarf disease.
However, c h l o r o t i c mottling of new needles, com-
mon t o t h e c h l o r o t i c dwarf disease., did not occur.
Tipburn symptom development u s u a l l y began during
t h e e a r l y summer when t h e new needles were elon-
gating. Thus, t r e e s 'began showing symptoms i n
e a r l y t o mid June, and symptoms developed through-
out t h e growing season.
Continuous a i r monitoring f o r s u l f u r dioxide
('SO2) and ozone t o 3 ) , 'begun i n 1973 and continued
t o t h e present time, revealed t h e common occur-
rence of SO2 and O-; concentrations i n t h e range
of 0.4 t o 5.0 pphm f o r 1 t o 3 hours during t h e
summer months. Maximum one-hour averages recorded
1 during t h e study were 11 pphm SO2 and 1 3 pphm 03.
These l e v e l s , while low i n terms of a i r q u a l i t y
standards, have "been shown t o be within t h e range
of concentrations of t h e s e p o l l u t a n t s reported t o
i n j u r e g e n e t i c a l l y s e n s i t i v e e a s t e r n white pine
t r e e s i n c o n t r o l l e d fumigations.
p r e s e n t e d a t t h e Symposium on E f f e c t s of A i r
P o l l u t a n t s on Mediterranean and Temperate Forest
Ecosystems, June 22-27, 2980, Riverside,
C a l i f o r n i a , U.S.A.
o ore st
G e n e t i c i s t , New York Botanical
Garden Cary Arboretum, Millbrook, N.Y.
During t h e 5 years since t h e power p l a n t began
operation, t h e r e have been no d e t e c t a b l e d i f f e r -
ences i n t h e occurrence o r s e v e r i t y of symptoms.
With only a few exceptions, t h e same t r e e s have
shown symptoms a f t e r t h e p l a n t began operation a s
before it s t a r t e d . However, m o r t a l i t y has reduced
intermediate genotypes (those i n j u r e d i n 1 o r 2
years of t h e 9 year study) by 4.7 percent and t h e
s e n s i t i v e genotypes (those i n j u r e d i n 3 o r more
y e a r s ) by 10.4 percent (Table 1 ) . I n comparison,
only 2.0 percent of t h e t o l e r a n t t r e e s have died
during t h e course of t h e study. Most of t h e mor-
t a l i t y of t h e s e n s i t i v e genotypes was apparently
due t o t h e i r being slower growing and t h u s unable
t o compete f o r l i g h t , water, and n u t r i e n t s with
t h e i r neighboring t r e e s . As a r e s u l t , t h e white
pine stands studied a r e undergoing a gradual t r a n -
s i t i o n towards having a i r - p o l l u t i o n t o l e r a n t geno-
types dominate.
Acknowledgments: This study was supported i n
p a r t by Grant R803971, Environmental Protection
Agency, and by t h e Wisconsin Pi-iblic Service
Corp., t h e Madison Gas and E l e c t r i c Co., t h e W i s -
consin Power and Light Co., and t h e College of
A g r i c u l t u r a l and L i f e Sciences, University of
Wisconsin.
Table I ~ M o r t a l i t yover a 9-year period of e a s t -
ern white pine t r e e s d i f f e r i n g i n a i r p o l l u t i o n
sensitivity.
Air
Pollution
sensitivity1 If Trees Died Mortality
Tolerant
Intermediate
Sensitive
Totals :
l ~ h es e n s i t i v i t y rankings were based on t h e
following: t o l e r a n t t r e e s did not have a i r -
pollution-induced t i p b u r n symptoms during t h e
course of t h e study; intermediate t r e e s had
symptoms f o r 1 o r 2 y e a r s ; and s e n s i t i v e t r e e s
had symptoms f o r 3 o r more years.
 Effect of Os and Os + NO2 on Growth of Tree Seedlings1
Lance W. ~ r e s s ~

Seedlings of 10 eastern forest tree species were

Table 1. Height growth and dry weight expressed as

exposed to 0.05, 0.10, or 0.15 pprn 0 and seedlings

percent of control for seedlings of 10 tree species

of 7 tree species were exposed to 0.10 pprn 0 and/or

exposed to 0.05, 0.10, or 0.15 pprn 0 for 6 hr/day

3 3

0.10 pprn NO in 6 hr/day exposures for 28 consecu-

for 28 consecutive days.

2
tive days. The exposures were performed when the

Percent of control

seedlings were 2-4wk-old in indoor exposure

chambers of the CSTR design.

Species
The Environmental Protection Agency has deter-

Loblolly Pine '100a looa lOOa

mined that the threshold for significant growth

(Pinus taeda L.) 82 b 85ab 90ab

effects due to extended 0 exposures (7 hr daily

3
avg. for 2 no) for sensitxve vegetation is between

73 b 79 b 72 bc
59 c 74 b 64 c
0.06 and 0.10 ppm. Three species in this study ex-

Pitch Pine lOOa lOOa lOOa

hibited a threshold for significant suppressions in

(P.- rigida Mill. ) 96ab 94a 84ab

that range, while three other species exhibited a

87 b 83a 77ab
threshold for significant effects between 0.10 and

74 c 78a 68 b
0.15 pprn (Table 1). However, two species exhibited

Virginia Pine lOOa lOOa lOOa

a threshold for significant growth suppressions at

(L virginiana 95a 98a 120a

or below 0.05 ppm. Those two species, loblolly pine

Mill. ) 89a 97a 93a

and American sycamore, are probably the most im-

86a 87a 86a

portant of the species tested to the forest industry.

Sweetgum lOOa lOOa l60a

The significant effects at 0.05 pprn 0 were not

3
(Liauidambar 109a 9lab 88ab
accompanied by foliar injury.

styraciflua L.) 71 b 76ab 65 bc

In previous studies there have been indications
55 b 60 b 52 c
that low concentrations of NO might be stimulatory
Sycamore lOOa lOOa lOOa
2
to plant growth or alleviate 0 phytotoxicity.
(Platanus 104a 84a 57 b
3
Similar indications were noted for some of the tree
occidentalis L.) 73 b 43 b 27 b
species in this study. Two species (white ash and
79 b 36 b 19 b
green ash) exhibited greater growth in the NO Green Ash lOOa lOOa lOOa
2

treatment than the control, and the only significant

(Lpennsylvanica 98ab 86ab 86a
interactive effects were significantly less than
Marsh. ) 76 bc 7lab 75a
additive (sweetgum and white ash).
70 c 64 b 75a
White Ash lOOa 100 b lOOab
Stimulations of growth at low 0 concentrations

3 {Fraxinus 112a 125a 11la

have been noted in the past, and some species ex-

americana L.) 91a 92 b 87 b

hibited growth stimulations in this study (Table 1).

However, apparent growth stimulations appear to be

85a 83 b 81 b
Willow Oak lOOa lOOa 1 OOa
dependent in part on the plant species and the

(Quercus phellos 99a 99a 94a

parameter being evaluated.

L. ) 96a 92a 83a

The relationship that these data have to field
81 b 89a 83a
conditions is not clear. This study demonstrates
Sugar Maple 100 b lOOa 1OOab
the potential for adverse effects at pollutant con-
(& saccharum 95 b 88a 161a
centrations below the current NAAQS, but future
L.) 108a 90a 116ab
research will have to determine whether such effects
88 c 58b 69 b
can be detected in the field.
Yellow Poplar 100 b 100 b 1OOa
(Liriodendron 160a 147a 133a
tulipifera L.1 108 b 11lab 96a
presented at the Symposium on Effects of Air

Pollutants on Mediterranean and Temperate Forest

Ecosystems, June 22-27, 1980, Riverside,

California, U.S.A.
Values for each column/species followed by the

same letter are not different at p = 0.05 according

to the ~uncan'sNew Multiple Range Test.

'~esearch Associate of Plant Pathology, Vir-

ginia Tech., Blacksburg, Va. Currently Assistant

Ecologist, Argonne National Laboratory, Argonne, 111.

 Impact of Oxidant Air Pollution on Ponderosa and Jeffrey Pine Cone Production1

Robert F. ~ u c k ~

Cone p r o d u c t i o n by ponderosa and J e f f r e y p i n e PONDEROSA PINE

100
t r e e s 10 cm o r more i n dbh was a s s e s s e d on 19
p l o t s l o c a t e d i n t h e San Bernardino Mountains o f 80
s o u t h e r n C a l i f o r n i a . These p l o t s were e s t a b - z

-
l i s h e d along a gradient of oxidant a i r p o l l u t i o n . 60
Each t r e e was r a t e d a n n u a l l y u s i n g an o x i d a n t a i r w
p o l l u t i o n index based upon t h e number o f y e a r 3 40
c l a s s e s o f n e e d l e s r e t a i n e d , t h e i r c h l o r o t i c con-
d i t i o n , t h e i r l e n g t h and t h e amount o f branch aw 0a 2o
m o r t a l i t y p r e s e n t . T h i s r a t i n g was made i n d e -
p e n d e n t l y i n t h e upper and lower crown f o r each
Or z
CÂ¥ O 0-1 2 3 4 5
t r e e . Other t r e e c h a r a c t e r i s t i c s , e . g . , age,
dbh, h e i g h t , were a l s o measured once d u r i n g t h e JEFFREY PINE
c o u r s e o f t h e 6 y e a r s t u d y . Cones were v i s u a l l y
counted w i t h i n t h e crown o f each t r e e each ¤ 80
September and October. M
CK
0
*
a
60
Although s e v e r e o x i d a n t a i r p o l l u t i o n i n j u r y a -I
was a s s o c i a t e d w i t h reduced cone p r o d u c t i o n , t h e 0 U 40
most i m p o r t a n t c o r r e l a t e w i t h cone p r o d u c t i o n 0"

Q_ 20
was crown c l a s s , i . e . , t h e p o s i t i o n o f a t r e e ' s
crown r e l a t i v e t o t h o s e o f i t s n e i g h b o r s . Dom-
inant t r e e s bore t h e g r e a t e s t proportion of 0 1 2 3 4 5
cones. I n ponderosa p i n e dominant t r e e s com-
p r i s e d 32 p e r c e n t o f t h o s e p r e s e n t on t h e 19 100
p l o t s b u t b o r e 80 p e r c e n t o f t h e cones. When t h e 8 o PONDEROSA PINE
dominant and codominant crown c l a s s e s were com-
bined t h e y comprised 58 p e r c e n t o f t h e ponderosa
p i n e s b u t b o r e 96 p e r c e n t o f t h e cones. J e f f r e y
p i n e showed a similar p a t t e r n . I n b o t h J e f f r e y
and ponderosa p i n e , cone p r o d u c t i o n i n c r e a s e d
s i g n i f i c a n t l y w i t h age (Ho:b=O: ponderosa p i n e ;
dominant: 7 = -92.33 + 1 . 8 6 (Age), r2 = 0.811**;
codominant: 7 = -74.48 + 1.077 (Age), r2 =
0.4216*: J e f f r e y p i n e ; dominant: 7 = 4.74 + 0.434
(Age), r2 = 0.765**; codominant: 7 = -8.28. +
0.243 (Age), r2 = 0.7456**). 100
JEFFREY PINE
Three p a t t e r n s were observed: (1) I n dominant
J e f f r e y p i n e s 130 y e a r s o r o l d e r fewer s e v e r e l y
i n j u r e d t r e e s b o r e cones t h a n u n i n j u r e d ones
( l b F ( 4 , 25) = 1 1 . 9 8 , p < 0.05) ; (2) i n dominant
ponderosa p i n e s 130 y e a r s o r o l d e r s e v e r e l y i n -
j u r e d t r e e s b o r e fewer c o n e s / t r e e t h a n u n i n j u r e d
ones (2a x2 = 21.07, p < 0.005) ; and (3) i n
f4l
b o t h dominant and codominant J e f f r e y and ponder-
osa pines severely i n j u r e d t r e e s bore s i g n i f i -
c a n t l y fewer cone c r o p s t h a n u n i n j u r e d ones dur- SLIGHT SEVERE
i n g t h e 6 y e a r s o f t h e s t u d y ( x 2 ' s v a l u e s < 0.01 SMOG INDEX
i n 5 c a s e s , < 0.05 i n 1 c a s e ) .
- F i g . 1. P r o p o r t i o n o f t r e e s producing cones
' p r e s e n t e d a t t h e Symposium on E f f e c t s of A i r i n a given smog c l a s s . ( a ) Ponderosa p i n e ; (b)
P o l l u t a n t s on M e d i t e r r a n e a n and Temperate F o r e s t J e f f r e y pine.
Ecosystems, J u n e 22-27, 1980, R i v e r s i d e ,
C a l i f o r n i a , U.S.A. F i g . 2. P r o p o r t i o n o f c o n e s / t r e e borne by
a s s o c i a t e P r o f e s s o r o f Entomology. D i v i s i o n t r e e belonging t o a given smog c l a s s . ( a ) Ponder-
of Biological Control, University of California, o s a p i n e ; (b) J e f f r e y p i n e .
Riverside, C a l i f .
 Lichens as Air Quality Monitors1
P.D. Lulman
.
R J . Fessende
S.A. McKinnon
The Athabasca t a r s a n d s , l o c a t e d i n a remote
region i n n o r t h e a s t e r n A l b e r t a , i n t h e b o r e a l
f o r e s t r e g i o n o f Canada, c o n t a i n v a s t r e s e r v e s
(approx. 600 b i l l i o n b a r r e l s ) o f bitumen. A t t h e
p r e s e n t time t h e r e a r e o n l y 2 companies o p e r a t i n g
i n t h e r e g i o n , producing s y n t h e t i c crude o i l from
t h i s bitumen, Suncor I n c . and Syncrude Canada Ltd.
Suncor, which began production i n 1967, e m i t s
approximately 200-300 tonnes o f SO2 p e r day.
Syncr-ide, which began production i n 1978 and
which i s l o c a t e d o n l y 10 km from Suncor, h a s been
e m i t t i n g about 50-100 tonnes p e r day. I n t h e
n e x t 10-20 y e a r s t h e r e i s a good p o s s i b i l i t y t h a t
many more companies w i l l begin o p e r a t i n g i n t h e
r e g i o n a s w e l l . T h i s h a s l e a d t o a concern about
t h e e f f e c t s o f changes i n a i r q u a l i t y on t h e
vegetation of the region. Since lichens a r e
known t o be more s e n s i t i v e t o a i r p o l l u t a n t s ,
p a r t i c u l a r l y SO2, t h a n most h i g h e r v e g e t a t i o n , i t
was decided t o monitor changes i n l i c h e n growth
a s an e a r l y warning o f t h e impact o f p o l l u t a n t s
on a l l components o f t h e v e g e t a t i o n .
I n 1976, Syncrude Canada Ltd. i n s t a l l e d a n e t -
work o f 56 permanent p l o t s i n a r a d i a t i n g p a t t e r n
c e n t e r e d on t h e 2 o p e r a t i o n s and a t d i s t a n c e s o f
up t o 47 km. Within each p l o t 20, 200 cm perma-
-
n e n t a u a d r a t s c o n t a i n i n a t h a l l i o f Parmelia s u l -
cats o r Hypogymnia physodes, 2 abundant bark-
-
l i c h e n s , were e s t a b l i s h e d and photographed. Most
o f t h e & a d r a t s were e s t a b l i s h e d o n w h i t e s p r u c e
P i c e a e l a u c a . a 1thouah a s m a l l e r number w i r e
(--
-
white birch (Betula papyrifera).
- -
e s t a b l i s h e d on' balsam f i r (Abies balsamea) and
? photographs
were p r o j e c t e d an surface areas of selected
l i c h e n t h a l l i were measured. I n 1979, a l l o f t h e
q u a d r a t s i n 12 s e l e c t e d p l o t s were rephotographed
and t h e photographs analyzed a s b e f o r e . The ob-
j e c t i v e s o f t h i s p a r t i a l resurvey were ( a ) t o
measure changes i n t h e s u r f a c e a r e a o f t h a l l i o f
P. s u l c a t a , t h e dominant l i c h e n s p e c i e s , i n t h e
3 y e a r p e r i o d 1976-1979, ( b ) t o t e s t t h e s i g n i f i -
cance o f t h e s e changes a s a f u n c t i o n o f d i s t a n c e
P r e s e n t e d a t t h e Symposium on E f f e c t s of A i r
P o l l u t a n t s on Mediterranean and Temperate F o r e s t
Ecosystems, June 22-27, 1980, R i v e r s i d e ,
C a l i f o r n i a , U.S.A.
~ n v i r o n m e n t a l i s t , Calgary Power L t d . ,
Calgary, A l b e r t a , Canada, formerly, Research
A s s o c i a t e , LGL L t d . , Environmental Research
A s s o c i a t e s , Edmonton, A l b e r t a , Canada; Head,
T e r r e s t r i a l Environment S e c t i o n and Research
A s s i s t a n t , Environmental A f f a i r s Dept., Syncrude
Canada L t d . , Edmonton, A l b e r t a , Canada.
9
and/or d i r e c t i o n from t h e emission s o u r c e s , and
( c ) t o assess the p r a c t i c a b i l i t y of the l i c h e n
network a s a l o n g term monitoring system f o r
d e t e c t i n g a i r q u a l i t y impacts on v e g e t a t i o n .
There was a s i g n i f i c a n t i n c r e a s e i n p r o j e c t e d
thallus surface area i n 6 plots, a significant
d e c r e a s e i n 2 p l o t s , and no s i g n i f i c a n t change i n
4 p l o t s . I n most c a s e s t h e n e t change i n l i c h e n
s u r f a c e a r e a d u r i n g t h e 3 y e a r p e r i o d was l e s s
than 10%. However, w i t h i n - p l o t v a r i a b i l i t y was
very l a r g e w i t h c o e f f i c i e n t s o f v a r i a t i o n common-
l y between 300-400%. The changes i n p r o j e c t e d
t h a l l u s s u r f a c e a r e a were s i g n i f i c a n t l y b u t
weakly, r e l a t e d t o d i s t a n c e ; t h e t h a l l i c l o s e s t
t o t h e emission s o u r c e s i n c r e a s i n g more i n a r e a
than t h o s e a t a d i s t a n c e , where i n f a c t t h e r e was
a n e t d e c r e a s e i n s u r f a c e a r e a on t h e average.-
F i e l d o b s e r v a t i o n s i n d i c a t e d t h a t t h e r e was no
r e l a t i o n s h i p between l i c h e n c o l o r and/or v i g o r
and d i s t a n c e and/or d i r e c t i o n from t h e emission
s o u r c e . It was e v i d e n t i n a comparison o f t h e
1976 and 1 9 7 9 p a i r e d photographs t h a t t h e r e was
a s i g n i f i c a n t l o s s o f whole t h a l l i and p o r t i o n s
o f t h a l l i from most o f t h e q u a d r a t s . It was n o t
unusual f o r 10-20% o f t h e t h a l l i p r e s e n t i n 1976
t o be m i s s i n g i n 1979. T h i s appeared t o be p r i -
m a r i l y r e l a t e d t o t h e n a t u r a l p r o c e s s o f bark
e x f o l i a t i o n , although animal a c t i v i t y and i n s e c t
g r a z i n g a r e a l s o suspected c a u s e s .
T h i s technique i s capable o f d e t e c t i n g s i g n i -
f i c a n t changes i n t h e s u r f a c e a r e a o f P. s u l c a t a
t h a l l i o v e r a 3 y e a r p e r i o d , however, i t r e q u i r e s
c a r e f u l s t a n d a r d i z a t i o n and a l a r g e number o f
samples due t o t h e l a r g e w i t h i n - p l o t v a r i a b i l i t y
i n t h i s c h a r a c t e r i s t i c . The h i g h v a r i a b i l i t y i s
r e l a t e d t o t h e r a t h e r non-uniform arowth o f t h e
P. s u l c a t a t h a l l u s a s w e l l a s t h e v a r i e t y o f
n a t u r a l processes causing l o s s e s of portions of
t h e t h a l l u s . Perhaps o t h e r a t t r i b u t e s o f t h e
t h a l l u s would make b e t t e r i n d i c e s o f growth ( o r
l a c k o f growth), however, none were i n v e s t i g a t e d .
The e x t e n s i v e l o s s e s o f whole t h a l l i r a i s e ques-
t i o n s about t h e permanence o f t h i s system and
hence i t s v a l u e a s a l o n g term monitoring system.
Because t h e predominant reason f o r t h a l l u s l o s s
i s bark e x f o l i a t i o n , t h i s problem i s very much
r e l a t e d t o t r e e s p e c i e s . Spruce a r e p a r t i c u l a r l y
bad i n t h i s r e g a r d . There was very l i t t l e t h a l l -
u s l o s s from b i r c h o r f i r . Notwithstanding t h e
above d i f f i c u l t i e s , i t was concluded t h a t t h e r e
was no a i r q u a l i t y caused damage t o P. s u l c a t a
between 1976 and 1979 a s i n f e r r e d from t h e rela-
t i o n s h i p s between t h e n e t change i n t h a l l i s u r -
f a c e a r e a and d i s t a n c e and/or d i r e c t i o n from t h e
e m i s s i o n s o u r c e s . T h i s conclusion i s supported
by f i e l d o b s e r v a t i o n s of l i c h e n c o l o r and v i g-o r .
 Acid Precipitation in California and Some Ecological Effects1

John G. McColl and Mary K. Firestone

2

Wet and d r y p r e c i p i t a t i o n s were monitored on Table 1 --Mean i o n i c concentrations o f wet p r e c i p i -

an event b a s i s i n 1978-9 a t Berkeley and San Jose t a t i o n d u r i n g the study p e r i o d i n 1978-9 (peqI1).
( p o l l u t i o n source areas i n t h e San Francisco Bay
area) , Davis and Par1 i e r ( i n t h e c e n t r a l a g r i c u l -
t u r a l and rangelands) , Chal lenqe ( l o w e r S i e r r a n
f o r e s t ) , and Tahoe C i t y (on t h e shore o f Lake
Tahoe) . Concentrations o f t h i r t e e n i o n i c species
H' (Lab.) 22.1 6.8 10.9 13.0 38.0 7.9 6.3 14.6
and s p e c i f i c conductance and volume were measured
(Table 1 ) . ~ a + 36.9 11.8 15.3 12.8 33.1 14.5 15.1 22.0
A c i d r a i n (pH < 5.6) was common a t a l l e i g h t K+ 1.5 0.6 1.4 2.1 1.5 1.1 1.1 1.4
s i t e s . Mean pH o f storms v a r i e d from 4.24 a t San ~a2+ 6.0 4.3 8.0 10.9 12.6 3.3 5.6 4.6
Jose t o 5.20 a t Davis, and t h e l o w e s t pH o f any kJ2+ 9.6 1.7 3.6 7.0 9.7 3.8 5.7 5.3
storm was 3.71 a t San Jose. The primary cause o f ~ e3+ 0.3 0.0 0.6 0.4 0.8 0.2 0.4 0.4
t h e a c i d i t y was probably t h e a i r p o l l u t a n t NOxy ~ n2+ 0.1 0.0 0.1 0.6 3.1 0.1 0.1 0.1
f o l l o w i n g i t s d i s s o l u t i o n i n wet p r e c i p i t a t i o n . cu2+ 0.1 0.1 0.1 0.2 0.3 0.1 0.2 0.7
NO? was t h e anion most c l o s e l y c o r r e l a t e d w i t h H', zn2+ 0.3 0.1 0.2 0.2 0.6 0.1 0.1 0.2
and NO3 g e n e r a l l y occurred i n g r e a t e r concen- NH; 8.0 4.1 40.0 11.9 19.1 9.7 35.5 12.1
t r a t i o n than SO$-.
T o t a l d r y d e p o s i t i o n s o f chemical c o n s t i t u e n t s NO; 13.7 6.7 43.4 19.9 16.4 11.1 22.6 16.4
between rainstorms were o f t h e same o r d e r as t o t a l Cl- 40.1 3.7 11.2 7.7 38.9 14.2 14.0 -23.1
SO-: 10.2 13.3 13.8 8.6 10.0 6.2 19.0 11 .7
wet d e p o s i t i o n s d u r i n g storms. Dry d e p o s i t i o n
d u r i n q summer would g r e a t l y increase t h e amounts
recorded i n t h i s study which was conducted i n t h e Cond .
wet season o n l y . However, more research i s needed (pmho/cm) 13.9 .3.9 17.4 10.6 16.2 6.4 9.6 10.7
i n procedures f o r q u a n t i f y i n g d r y atmospheric pH (Lab.) 4.7 5.2 4.7 4.9 4.4 5.1 5.2 4.8
deposition. Vol. (cm) 52.7 78.0 19.7 110.7 21.0 64.9 39.7 62.0
' A1 though NOS c o n c e n t r a t i o n (pg/1) and a c i d i t y
'BE Berkeley, TC Tahoe C i t y , KE Kearny ( f i e l d
(H" c o n c e n t r a t i o n , p g / l ) of wet p r e c i p i t a t i o n were
g r e a t e s t i n p o l l u t i o n source areas, t o t a l depo- s t a t i o n a t par1 ier) , CH Challenge, SJ San Jose, HO
s i t i o n ( k g l h a ) o f NO3 and H+ were g r e a t e s t i n t h e Hop1and, DA Davis , and NA Napa .
non-urban r e c e p t o r areas o f Napa and Chal lenqe;
t h i s was l a r g e l y a f u n c t i o n o f t h e g r e a t e r p r e c i -
p i t a t i o n volumes a t these two s i t e s (Table 1 ) . I n t h e i n o r g a n i c s o i l -chemical s t u d i e s , s o i l s
c o v e r i n g a wide range o f p a r e n t m a t e r i a l s and age
Thus e c o l o g i c a l e f f e c t s may be expected i n t h e
coast ranges and S i e r r a s w i t h i n t h e g e n e r a l east- are b e i n a t r e a t e d w i t h a c i d i n p u t s , and t h e subse-
to-west "wash-out f a n " o f wet p r e c i p i t a t i o n , as quent l e a c h i n g p a t t e r n s o f i o n s ( i n c l u d i n g ~ l + 3 ,
w e l l as w i t h i n p o l l ution-source areas. H+, Nat, K+, Mg++, Ca^) are b e i n g determined.
E f f e c t s o f these a c i d i n p u t s t o C a l i f o r n i a The b i o l o g i c a l i n v e s t i g a t i o n s i n c l u d e assess-
s o i l s are c u r r e n t l y b e i n g assessed. These inves- ment o f p o s s i b l e e f f e c t s on t h e s o i l - p l a n t
t i g a t i o n s on s o i l e f f e c t s i n c l u d e research on both n u t r i e n t system. The f i r s t 10-week p o t - t r i a l ,
u s i n g a Yolo s e r i e s s o i l and qrowina b a r l e y and
t h e i n o r g a n i c phase o f s o i l and on t h e o r q a n i c l -
b i o l o q i c a l components. c l o v e r , has j u s t been completed. Prel i m i n a r y
r e s u l t s i n d i c a t e marked qrowth increases i n t r e a t -
ments where t h e p l a n t s were sprayed w i t h s o l u t i o n s
o f pH 2.0 and 3.0. This r e s u l t i s a t t r i b u t e d t o
p r e s e n t e d a t t h e Symposium on Effects of A i r g r e a t e r a d d i t i o n s o f SO^- and NO?, and/or t o
P o l l u t a n t s on Mediterranean and Temperate F o r e s t increased a v a i l a b i l it y of s o i l n u t r i e n t s i n these
Ecosystems, June 22-27, 1980, R i v e r s i d e , lower-pH treatments. However, s p o t t i n g o r leaves
C a l i f o r n i a , U.S.A. by a c i d d r o p l e t s a l s o occurred i n t h e pH 2.0
treatments. N i t r i f i c a t i o n , deni t r i f i c a t i o n and
~ s s o c i a t eProfessor of S o i l Science and n i t r o g e n f i x a t i o n a c t i v i t i e s are b e i n g measured i n
Associate S p e c i a l i s t i n S o i l Microbiology, b o t h t h e rhizosphere and non-rhi zosphere s o i l .
r e s p e c t i v e l y , U n i v e r s i t y o f C a l i f o r n i a , Berkeley,
C a l i f o r n i a , U.S.A.
 Leaf Litter Decomposition in the Vicinity of a Zinc Refinery

Emmissions from a large zinc refinery in north-

It was found that the rate of decomposition

ern Ontario have been monitored for 10 years and

of the leaf litter was lower at sites nearest to

an accumulation of zinc, copper, cadmium, arsenic

the refinery and that contaminated foliage de-

and lead in the soil and vegetation surrounding

composed more slowly than control foliage at all

the refinery complex has been documented. It was

sites. The microarthropod population was domi-

considered that decomposition of leaf litter as a

nated by mites and springtails. The number of

part of the nutrient cycling process might be a

microarthropods was generally lower at the sites

convenient method to monitor the initial impact

nearest to the refinery and was also lower in the

of these metals on the environment. This inves-

litter baqs containing the contaminated foliage.

tigation was undertaken in the fall of 1977. Ny-

The numbers of microarthropods appeared to fluc-

lon mesh bags containing 10 q (oven-dry weight)

tuate with moisture content of the litter. It

of trembling aspen (Populus tremuloides) were set

was found that the concentrations of zinc, copper,

out at several locations around the refinery.

cadmium, lead, arsenic, iron, sulphur and sele-

Two sources of aspen foliage were utilized inclu-

nium in the leaf litter increased with time at

ding foliage from a control location and conta-

the sites nearest to the zinc refinery but only a

minated foliage collected near the refinery. The

slight increase was noted at the more distant

exposure locations for the litter bags included

sites.

three sites in close proximity to the refinery,

-
(Plot A, B, C ) one site 2.2 km from the refinery
(Plot D) and one at a control location (80 km

southwest of the refinery) (Plot E) .

'
z
I-
60"
PLOT NO A
CONTAMINATED LITTER ..*-*.
m CONTROL LITTER -
The litter bags were allowed to over-winter

g
0
45-

and triplicate bags of each litter type were col-

a 30-
lected in May, June and August of 1978 and in

I-
May, July and August in 1979. Microarthropods
z
were extracted from the bags in modified Tulgren

funnels in the laboratory. The litter was then

;
u

15-
n

dried to determine the leaf weight loss and pro-

cessed for chemical analysis.

NOV MAY JUN AUG MAY JUL AUG

1977 1978 1979

Figure 1. Pattern of decomposition of two types

of leaf litter at two sites.

presented at the Symposium on Effects of Air

Pollutants on Mediterranean and Temperate Forest

Ecosystems, June 22-27, 1980, Riverside,

California, U.S.A.

plant Pathologist, Ministry of the Environment,

199 Larch Street, Sudbury, Ontario, P3E 5P9.

NOV MAY JUN AUG MAY JUL AUG

1977 1978 1979

Figure 2. Zinc accumulation is typical of metal

accumulation in leaf litter.

 Effects of Chronic Air Pollution Stress on Allocation of Photosynthate by White Pine
S. B. McLaughlin, R. K. McConathy, and D. Duvick2
A1 location of ^c-photosynthate by in s i t u
branches of nine field-grown white pine trees was
studied t o determine whether distribution patterns
differed between trees with apparent differences
in s e n s i t i v i t y to a i r pollution stress. Three
trees were selected in each of three sensitivity
classes which were differentiated on the basis of
needle length, mottling, and duration of reten-
tion. Previous studies (Mann e t a1. 1980) indi-
cated that photosynthetic potential of foliage
from trees in these three classes was not sig-
nificantly different. Growth ring analysis of
increment cores indicated that average annual
increment of intermediate and sensitive trees was
98% and 47%, respectively, of that attained by
the tolerant trees (7.8 mm y r - l ) over the past
18 years (Fig. 1 ). Sensitive trees showed a
marked decline in annual growth during the past
10 years.
Foliage was labelled with ^COz four times
during the growing season (June, July, August
and November). Of the paired branches from each
tree labelled on each date, one was harvested
a f t e r seven days and the remainder in November
a t the end of the growing season. Photosynthate
allocation patterns were compared by determining
levels of f o l i a r retention and allocation to
nearby branches. In June when elongating needles
were approximately 50%of t h e i r final length, ^ C
movement patterns in needles of three age classes
(Fig. 2 ) indicated that contribution of ^C-
photosynthate by old needles t o new needle growth
was occurring. This process was most rapid in
tolerant trees which retained needles from two
prior years and l e a s t significant in sensi tjve
trees. Lower levels of incorporation of photo-
synthate into f o l i a r tissues occurred subse-
quently (X = 35% in June, 27% in July, and 5% in
August a f t e r seven days). There were no d i s t i n c t
differences in f o l i a r retention of 1 4 C between
the three s e n s i t i v i t y classes which could be
associated w i t h the distribution of high levels
of ozone (1h avg > 0.08 ppm) near the study
area. ~ i g nfii c a n t 2-year-old needle retention
on tolerant trees did not extend beyond July.
Presented a t the Symposium on Effects of Air
Pollutants on Mediterranean and Temperate Forest
Ecosystems, June 22-27, 1980, Riverside,
California, U.S.A.
%search Staff llember, Research Associate,
and Technician, respectively, Terrestrial Ecology
Section, Environmental Sciences Division, Oak
Ridge National Laboratory, Oak Ridge, Tennessee,
U.S.A.
Generally higher levels of transfer of 1 4 c from
foliacje into branches were noted in the tolerant
trees throughout the growin season. Higher
levels of trans1 ocation of ^C-photosynthate out
of 1-year-01 d need1 es than current-year needles
were also found for a l l sensitivity classes.
Enhanced current needle elongation (+25%sensi-
tive, +lo%intermediate, and +3% tolerant) com-
pared to the previous year was associated with
lower than average ozone levels and higher than
average rainfall during the growing season. Data
support the hypothesis that growth limitations
in sensitive trees are a function of stress-
induced reductions in photosynthate availability
which result from reduced needle length (a func-
tion of decreased availability of carbohydrates
for needle growth) and premature needle senes-
cence. The resulting chronic decline of sensitive
trees i s likely a result of interactions between
a i r pollution stress and s i t e factors which may
include secondary belowground pathogens.
Acknowledgments: Research sponsored by the
U. S. Environmental Protection Agency under Inter-
agency AgreementNo. 40-740-78 and the U.S.
Department of Energy under Contract W-7405-eng-26
with Union Carbide Corporation.
Literature Cited
Mann, L. K., S. B. McLaughlin, and D. S. Shr i ner.
1980. Seasonal physiological responses of white
pine under chronic a i r pollution stress. Env.
Exp. Bot. (in press).
240 1
1960'62 'â‚ '66 '68 '70 '72 '74 '76 '78 1980 0 24 168
YEAR TIRE SINCE LABtLLIHt (m)
F i g u r e I ( l e f t ) - - A v e r a g e annual d i a m e t e r g r o w t h
o f 3 t r e e s i n each o f 3 s e n s i t i v i t y c l a s s e s .
F i g u r e 2 ( r i g h t ) - - D i s t r i b u t i o n o f ^C p h o t o s y n t h a t e
i n f o l i a g e 0, I , and 7 days a f t e r ^cog u p t a k e i n
June.
 Effects of SO2 and Ozone on Photosynthesis and Leaf Growth in Hybrid Poplar
Reginald D. Noble
and
Keith F. ~ e n s e n 2
P l a n t s i n t h e n a t u r a l environment a r e o f t e n si-
multaneously exposed t o a combination of many atmos-
pheric p o l l u t a n t s . Thus i t i s of i n t e r e s t t o as-
c e r t a i n t h e n a t u r e of i n t e r a c t i o n of t h e s e pollu-
t a n t s on p l a n t development and metabolism. Two pol-
l u t a n t s commonly encountered i n t h e atmosphere which
a r e p a r t i c u l a r l y t o x i c t o p l a n t s and would appear
t o o f f e r p o t e n t i a l f o r i n t e r a c t i o n a r e SO2 and 03.
METHODS
P l a n t s of hybrid p o p l a r #207 (Populus d e l t o i d e s
Bartr. X P. t r i c h o c a r p a Torr. & Gray) were grown
from c u t t i n g s under greenhouse c o n d i t i o n s . S i x
week o l d p l a n t s were fumigated i n c o n t r o l l e d envi-
ronment chambers f o r 12 hours p e r day f o r 24 con-
s e c u t i v e days. Four s e t s of p l a n t s were used a s
follows : S e t 1-Controls (no fumigation) ; S e t 2-Fu-
migated with SO2 a t 0.5 ppm; S e t 3-Fumigated with
03 a t 0.25 ppm; S e t 4-Fumigated with SO2 a t 0.5
pprn p l u s O3 a t 0.25 ppm. A t t h e end of t h e 24-day
fumigation p e r i o d , number of l e a v e s p e r p l a n t , l e a f
f r e s h and d r y weight and l e a f a r e a were determined.
During t h e l a t e r s t a g e s of t h e fumigation, photosyn-
t h e s i s measurements were determined f o r s i n g l e a t -
tached l e a v e s from p l a n t s fumigated with SO2 and
those fumigated with s02+03. Photosynthetic mea-
surements were made a t ambient C02 l e v e l s (300 pprn
C02) and a t 1000 pprn C02.
RESULTS
P l a n t s fumigated with SO2 showed l i t t l e v i s i b l e
evidence of i n j u r y while those fumigated with O3 o r
a combination of t h e two began t o develop t i n y ne-
c r o t i c s p o t s a f t e r 5 t o 7 days of exposure. Soon
a f t e r lower l e a v e s began t o a b s c i s e and by 14 days
e x t e n s i v e i n j u r y was apparent on l e a v e s below t h e
7 t h node from t h e apex. Both O3 and S02+03 t r e a t -
ments caused a reduction i n t h e number of l e a v e s p e r
p l a n t ; however, t h e r a t e of l e a f formation was n o t
a f f e c t e d f o r t h e f o u r groups. The number of l e a v e s
p e r p l a n t was lower i n t h e O3 treatment group than
i n t h e S02+03 group, suggesting an a n t a g o n i s t i c
_L
Presented a t t h e Symposium on E f f e c t s of A i r
P o l l u t a n t s on Mediterranean and Temperate F o r e s t
Ecosystems, June 22-27, 1980, Riverside,
C a l i f o r n i a , U.S.A.
p r o f e s s o r of B i o l o g i c a l Sciences, Bowling Green
S t a t e U n i v e r s i t y , Bowling Green, Ohio; Research
S c i e n t i s t , U.S. Department of A g r i c u l t u r e , F o r e s t
S e r v i c e , Delaware, Ohio.
r e l a t i o n s h i p . Fumigation tended t o cause a decrease
i n l e a f a r e a , l e a f dry weight and l e a f f r e s h weight.
This was i n c r e a s i n g l y t r u e a s t h e d i s t a n c e from t h e
p l a n t apex increased. P o l l u t a n t i n t e r a c t i o n eval-
u a t i o n s revealed an a n t a g o n i s t i c r e l a t i o n s h i p be-
tween t h e responses t o O3 and SO2. Ozone which r e -
tarded growth most had l i t t l e o r no e f f e c t on a r e a ,
dry o r f r e s h weight of l e a v e s above node 5; however,
t h e s e parameters were reduced by 25 t o 40 p e r c e n t
i n l e a v e s fumigated with O3 a t node 9. These and
r e l a t e d observations demonstrate t h a t O3 r e t a r d s
growth of leaves i n t h i s p l a n t only during t h e l a t -
e r s t a g e s of t h e i r development. Ozone fumigated
l e a v e s with v i s i b l e i n j u r y comparable t o t h o s e fu-
migated with both SO2 and O3 contained l e s s d r y
m a t t e r per u n i t a r e a .
Photosynthetic measurements on c o n t r o l s a s w e l l
a s p l a n t s fumigated with SO2 and S02+03 revealed
l i t t l e d i f f e r e n c e i n r a t e of C02 a s s i m i l a t i o n a t
300 pprn COT Even i n l e a v e s of S02+03 fumigated
p l a n t s where v i s i b l e i n j u r y was e x t e n s i v e t h e
photosynthetic r a t e was reduced by only 10 t o 20
p e r c e n t . When t h e C02 c o n c e n t r a t i o n was e l e v a t e d
t o 1000 pprn C02 photosynthetic r a t e s f o r c o n t r o l s ,
SO2 fumigated and S02+03 fumigated l e a v e s (with no
v i s i b l e i n j u r y ) were e l e v a t e d three-fold t o approx-
imately 35 mg C02 dm'-hr-l. I n l e a v e s i n which
i n j u r y was apparent t h e photosynthetic r a t e i n c r e a s -
e s were u s u a l l y l e s s than two-fold, t o a l e v e l of
approximately 1 8 mg C02 dm-2hr-1 (whether t h e in-
jury was severe or barely discernable).
DISCUSSION
Ozone, and O3 i n combination with SO2 h a s t e n
senescence and promote r a t e of l e a f drop t o t h e
e x t e n t t h a t c o n t r o l p l a n t s have approximately 50
p e r c e n t more l e a v e s than fumigated ones. Fumiga-
t i o n with SO2 causes no such e f f e c t . Analysis of
a t t a c h e d l e a v e s f o r treatment e f f e c t s c l e a r l y shows
an O3 e f f e c t b u t l i t t l e o r no SO2 e f f e c t . I n com-
bination these pollutants a t t h i s concentration
i n t e r a c t i n an a n t a g o n i s t i c manner i n r e l a t i o n t o
growth parameters.
Measurements of e f f e c t s of fumigation on photo-
s y n t h e s i s i n d i c a t e t h a t l e a f photosynthesis i s n o t
d r a s t i c a l l y reduced even i n t h e presence of con-
s i d e r a b l e i n j u r y . I t appears t h a t i n j u r y tends
t o reduce l e a f p o t e n t i a l t o respond t o enhance-
ment c o n d i t i o n s more s o than t h e a b i l i t y t o f i x
C02 under c o n d i t i o n s where C02 may be l i m i t i n g .
 Behavior of Airborne Fluorides in Soils'
Janina Polomski, Hannes Flfihler, and P e t e r
Accumulation of airborne f l u o r i d e s i n s o i l s i s
s a i d t o be a s i d e e f f e c t of minor importance i f it
i s compared w i t h t h e p h y t o t o x i c i t y of man-made
gaseous f l u o r i d e . When s o i l f l u o r i d e was consid-
ered i n t h e p a s t , F-uptake of p l a n t s , l e a f i n j u r y
o r y i e l d reduction was of prime concern. Not much
i s known about t h e f a t e of F-pollutants a f t e r
being deposited onto t h e s o i l s u r f a c e o r incor-
porated i n t h e p l a n t l i t t e r l a y e r . Our study aims
a t providing experimental t o o l s and d a t a f o r a
b e t t e r understanding of long-termed e f f e c t s . The
o b j e c t i v e i s t o p i n down t h e most s i g n i f i c a n t ,
c o n t r o l l i n g mechanisms of t h e F-mobility i n s o i l s .
F i e l d experiments: I n t h e v i c i n i t y of a
30000 t Aluminum smelter, b u i l t 1908, we ob-
served t h a t F a c t u a l l y accumulates i n the s o i l
(Table). The s o i l s sampled a t various d i s -
tances d i f f e r i n many r e s p e c t b u t a r e a l l
calcareous. The F-contents do depend upon
d i s t a n c e from t h e emission source.
distance from total F watersoluble F dissolved F F-content of
Al-smelter 1 %-year old
(km) 1 (pg F per gram soil)
(pg F per ml
soil solution)
pine needles
( P P ~
In s o i l s , f l u o r i d e c o e x i s t s i n many d i f f e r e n t
forms which makes t h e t o t a l F-content t o be a
hardly i n t e r p r e t a b l e s o i l c h a r a c t e r i s t i c s .
I n a pine stand, l o c a t e d 1.8 km from t h e
smelter t h e magnitude of t h e annual F-input and
leaching l o s s e s were determined (kg F per y and
ha) : 1-3 kg within dead p l a n t m a t e r i a l , 6-10 kg
atmospheric washout, 40-80 kg adsorption and
sedimentation. The leaching l o s s e s o u t of t h e
r o o t zone were estimated t o be some 80-100 kg.
Under pine canopies t h e F-contents i n t h e
s u r f a c e l a y e r (humus) a r e s y s t e m a t i c a l l y higher
than i n t h e s u b s o i l (mineral horizon) whereas
i n u n c u l t i v a t e d open land under g r a s s cover
f l u o r i d e accumulates i n t h e s u b s o i l .
p r e s e n t e d a t t h e Symposium on E f f e c t s of A i r
P o l l u t a n t s on Mediterranean and Temperate Forest
Ecosystems, June 22-27, 1980, Riverside,
C a l i f o r n i a , U.S.A.
~ g r o n o m i s t ,s o i l p h y s i c i s t and chemist,
r e s p e c t i v e l y , Swiss F o r e s t r y Research
I n s t i t u t e , Binnensdorf, Switzerland.
laserâ
Laboratory experiments : What c o n t r o l s the
f l u o r i d e t r a n s p o r t i n s o i l s ? I n case of a non-
calcareous s o i l (pH 5.2) the amount of f l u o r i d e
adsorbed within a mixture of s o i l and NaF-
s o l u t i o n being e q u i l i b r a t e d on a shaker
(3340 \lg ~ / g s) o i l equals approximately t h e
F-adsorption i n t h e course of a p e r c o l a t i o n
through a s o i l s l a b (Fig.)
Breakthrough-Curves of S i b Clay Soil with and without CaCO,
Cumulated Outflow [Number of Pore Voluminal
The c l o s e agreement between t h e two experi-
ments i n d i c a t e s t h a t adsorption and p r e c i p i t a -
t i o n a r e f a s t r e a c t i o n s a s compared with t h e
v e l o c i t y of t h e p e r c o l a t i n g s o i l s o l u t i o n .
In calcareous s o i l s , however, the exchange
k i n e t i c s between t h e l i q u i d and s o l i d phase
c o n t r o l s t h e e x t e n t of accumulation and
depletion. The exchange proceeds too slow
t o keep up with t h e pace of t h e moving water.
This explains t h e amazingly high F-mobility
observed i n case of t h e F-contaminated
calcareous s o i l s .
I n t h e course of t h e p e r c o l a t i o n experi-
ments t h e l e a c h a t e s from t h e s o i l columns
contain appreciable amounts of s o l u b i l i z e d
organic matter and aluminum. For a given
s o i l type t h e leaching l o s s e s depend i n t h e
f i r s t p l a c e upon t h e F-content of t h e i n f i l -
t r a t i n g s o l u t i o n and f u r t h e r upon s o i l type
and t h e F-compounds used t o make up t h e
p e r c o l a t i n g s o l u t i o n . Under f i e l d conditions
with 8-10 ppm F i n the s o i l s o l u t i o n such
phenomena may a l s o occur. A s i g n i f i c a n t amount
of s o l u b l e organic m a t e r i a l can be l o s t which
may r e p r e s e n t an energy s u b s t r a t e shortage
f o r t h e microorganisms, and furthermore t h e
Al-concentrations possibly reach phytotoxic
l e v e l s . A t t h i s p o i n t t h e experimental evi-
dence r a t h e r backs up questions than pro-
vides d e f i n i t e answers.
 Multiple Pollutant Fumigations Under Near Ambient Environmental Conditions
Using a Linear Gradient Technique1
P.B. Reich, R.G. Amundson, and J.P. L a s s o i e
I n v e s t i g a t i o n s of t h e impact of atmospheric
p o l l u t a n t s on p l a n t s normally i n v o l v e a r t i f i c i a l -
l y r a i s i n g t h e c o n c e n t r a t i o n of c e r t a i n g a s e s a-
round s t u d y p l a n t s . Such experiments t y p i c a l l y
u t i l i z e e n c l o s u r e s which g r e a t l y modify o t h e r
environmental f a c t o r s . Hence, t h e s e experiments
may n o t a c c u r a t e l y t e s t p l a n t responses under
ambient c o n d i t i o n s . The development of open-top,
f i e l d f u m i g a t i o n chambers (Mandl and o t h e r s 1973)
and a new l i n e a r g r a d i e n t exposure system (Shinn
and o t h e r s 1977) have been a t t e m p t s t o s o l v e
t h i s problem.
I n 1979 we c o n s t r u c t e d a modified l i n e a r gra-
d i e n t system and 6 open-top chambers. The gra-
d i e n t system enabled t h e simultaneous exposure of
numerous s t u d y p l a n t s t o a c o n c e n t r a t i o n g r a d i e n t
of SO2 and 03. S i n c e t h e system does n o t use
e n c l o s u r e s , p l a n t s a r e e a s i l y accessed and ex-
p e r i e n c e near-ambient environmental c o n d i t i o n s .
A p l o t of soybeans (Glycine s. v a r . Hark) i n
I t h a c a , NY was used i n a n a t t e m p t t o answer t h e
following questions.
1 ) Can t h e g r a d i e n t system be used t o expose
p l a n t s i n t h e f i e l d t o c o n t r o l l e d l e v e l s of a i r
p o l l u t a n t s without s i g n i f i c a n t l y a l t e r i n g t h e i r
environments?
2) Is p l a n t growth s i g n i f i c a n t l y d i f f e r e n t
i n open-top chambers v e r s u s i n a l i n e a r g r a d i e n t
system?
3) What r e d u c t i o n s i n soybean seed y i e l d w i l l
r e s u l t from exposure t o moderate l e v e l s of SO2
and 03 d u r i n g t h e seed m a t u r a t i o n p e r i o d ?
During t h e seed m a t u r a t i o n p e r i o d , t h e p l a n t s
were exposed t o SO2 and O3 f o r about 55 h o v e r a
20-day p e r i o d . On a l l o c c a s i o n s , a l i n e a r con-
c e n t r a t i o n g r a d i e n t was c r e a t e d by t h e system.
Mean c o n c e n t r a t i o n s of SO2 and O3 a t t h e "high"
end of t h e g r a d i e n t were 0.16 and 0.06 ppm, r e -
s p e c t i v e l y , w h i l e ambient c o n c e n t r a t i o n s of b o t h
p o l l u t a n t s were about 0.02 ppm. Maximum h o u r l y
means a t t h e "high" end of t h e g r a d i e n t were0.50
and 0.17 ppm S O 2 a n d 03, r e s p e c t i v e l y . By moni-
t o r i n g t h e p l o t i n a g r i d d e s i g n , we were a b l e
t o account f o r t h e e f f e c t s of wind on p o l l u t a n t
levels. I n t h e open-top chambers, p l a n t s were
p r e s e n t e d a t t h e Symposium on E f f e c t s of A i r
P o l l u t a n t s on Mediterranean and Temperate F o r e s t
Ecosystems, June 22-27, 1980, R i v e r s i d e ,
C a l i f o r n i a , U.S.A.
~ r a d u a t es t u d e n t of F o r e s t S c i e n c e , C o r n e l l
U n i v e r s i t y , I t h a c a , NY; P o s t D o c t o r a l A s s o c i a t e ,
Boyce Thompson I n s t i t u t e , I t h a c a , NY; and A s s i s -
t a n t P r o f e s s o r of F o r e s t S c i e n c e , C o r n e l l
U n i v e r s i t y , I t h a c a , NY.
2
exposed t o 0.30, 0.10, and 0.02 (ambient) ppm SO2
f o r 72 h , over a 30-day p e r i o d , d u r i n g t h e pod-
f i l l i n g stage.
Comparisons of t r e a t m e n t s r e v e a l e d t h a t t h e
g r a d i e n t fumigations reduced t o t a l bean y i e l d p e r
p l a n t and d r y mass p e r bean by a s much a s 39 and
14 p e r c e n t , r e s p e c t i v e l y . I n t h e open-top cham-
b e r s , t o t a l bean y i e l d p e r p l a n t and d r y mass p e r
bean were reduced by a s much a s 1 0 and 7 p e r c e n t ,
r e s p e c t i v e l y . It i s of i n t e r e s t t o n o t e t h a t
b o t h t h e "high" and "medium" (mean SO2-0.09 ppm,
mean 03-0.04 ppm) t r e a t m e n t s i n t h e g r a d i e n t sys-
tem caused much g r e a t e r r e d u c t i o n s i n y i e l d t h a n
d i d t h e 0.30 ppm SO2 t r e a t m e n t i n t h e open-top
chamber. I n i n t e r p r e t i n g t h e s e d a t a , one must
remember t h a t t h e p l a n t s i n t h e g r a d i e n t system
did e x p e r i e n c e short-term peaks much h i g h e r t h a n
t h e i r mean c o n c e n t r a t i o n exposures.
I n any c a s e , 03, a l t h o u g h p r e s e n t a t moderate-
l y low l e v e l s , was ( a l o n e o r s y n e r g i s t i c a l l y w i t h
SO2) probably r e s p o n s i b l e f o r t h e g r e a t e s t r e -
d u c t i o n s i n seed s i z e and y i e l d .
I n open-topchambers, one can a c c u r a t e l y con-
t r o l pollutant levels. In contrast, the l i n e a r
g r a d i e n t system a l l o w s f u m i g a t i o n s which v a r y
w i t h time and wind, a s under ambient c o n d i t i o n s .
The l i n e a r g r a d i e n t system proved t o b e a n e f -
f e c t i v e means of fumigating p l a n t s w i t h m u l t i p l e
p o l l u t a n t s under ambient environmental condi-
t i o n s . Another m o d i f i c a t i o n of t h e g r a d i e n t sys-
tem i s c u r r e n t l y underway s o a s t o a l l o w f o r
a n a l y s i s of t h e i n t e r a c t i o n s between SO2 and 03,
and t h e e f f e c t s of such on v a r i o u s hardwood t r e e
seedlings.
T h i s s t u d y was supported i n p a r t by t h e North-
e a s t e r n F o r e s t Experiment S t a t i o n , F o r e s t S e r v i c e ,
U.S. Department of A g r i c u l t u r e , through t h e Con-
s o r t i u m f o r Environmental F o r e s t r y S t u d i e s (For-
e s t S e r v i c e Grant 23-169) and by t h e C o r n e l l
U n i v e r s i t y A g r i c u l t u r a l Experiment S t a t i o n
(McIntire-Stennis P r o j e c t NY(C)-147556).
LITERATURE CITED
Nandl, R.H., L.H. Weinstein, D.C. McCune, and
M. Keveny .
1973. A c y l i n d r i c a l , open-top chamber f o r t h e
exposure of p l a n t s t o a i r p o l l u t a n t s i n t h e
f i e l d . J. Environ. Qual. 2:371-376.
Shinn, J . H . , B.R. Clegg, and I1.L. S t u a r t .
1977. A l i n e a r - g r a d i e n t chamber f o r exposing
f i e l d p l a n t s t o c o n s t a n t l e v e l s of a i r
p o l l u t a n t s . DOE-Preprint UCKL-80411.
 Changes in Plant Communities with Distance from an SO2 Source

Paul R. scale2

Investigations were carried out to assess the

occurs despite their own greater exposure to S O .

effects on plant community composition of sulphur

The low-bush blueberry (Vaccinium angustifoliumL

dioxide (SO2) emitted from an iron sintering

Ait.) for example, becomes very abundant at dis-

plant in Wawa, Ontario, Canada. Diversity indices

tances of 20 to 30 tan yet shows considerable SO2

and the ordination techniques of Correspondence

damage in that area. The first substantial change

Analysis (C.A.) and Principal Component Analysis

in species abundance occurs at a distance of 40 km

(P.C.A.) were used to discern community level

where beaked hazel (Corylus cornuta Marsh.) becomes

changes. Of special interest were the subtle

very abundant before a very rapid decline. This

changes in areas noted to be only slightly affected

undoubtedly contributes to the drop in the abund-

by SO2 emissions. Over 60 upland birch stands were

ance of the large leaved aster (Aster macrophyllus

accessed along a north-easterly transect away from

L.) at that distance.
the source at distances of 12 to 55 km. Species

specific data was collected on the tree, shrub and

Dealing collectively with all species of a com-

ground flora.
munity to discern changes caused by a pollutant

is, for theoretical reasons and from experience, a

Changes in species composition are dramatic as

more satisfactory approach than changes in indivi-

the 30 year SO2 source is approached. The total

dual species abundance. Diversity indices have

percent cover of the dominant, predominantly boreal,

been used extensively for this purpose. However,

ground flora species are shown in Figure 1. They

the data from Wawa indicates that although diver-

reflect the complex pattern of species abundance

sity indices can deal with spatial/structural

which occurs along the transect. Characteristic is

changes in the community, they do not adequately

the transitory increase in abundance which occurs

reflect the subtler changes in species composition.

in most species. What is particularly evident is

Ordination techniques, such as C.A. and P.C.A., are

the ability of many species to take advantage of

better suited for this purpose. For example, the

the increasing damage caused first to the tree

Shannon-Weiner Diversity index has a transitory

canopy and subsequently to the shrub layer. This

increase between 20 and 30 km before declining at

distances below 20 km. In contrast, the first

axis of C.A. using ground flora data shows a simple,

linear change from 12 to 33 km (Figure 2). C.A.

is most effective in reflecting overall trends

whereas P.C.A. is particularly useful in discerning

differences in stands of very similar composition.

On the basis of P.C.A. using shrub density data,

the separation of sites in intermediate areas from

controls can be achieved. The boundary between the

two groups in Figure 2 corresponds well with the

line designated in 1973 and 1974 as separating areas

which do and do not show visible leaf damage.

Figure 1--Changes in ground flora species abundance

with distance using a running average of percent

cover: Am, Aster macrophyllus; An, Aralia

nudicaulis; Df, Deschampsia flexuosa; Dl, Diervilla

lonicera; Me, Maianthemum canadense; Va, Vaccinium

angustifolium.

presented at the Symposium on Effects of Air

Pollutants on Mediterranean and Temperate Forest

Ecosystems, June 22-27, 1980, Riverside,

California, U.S.A.

Figure 2~Distancevs Axis 1 of ground flora

~ r a d u a t eStudent, Institute for Environmental

(percent cover) and shrub density (number of

Studies, University of Toronto, Toronto,

stems under 5 cm diameter at breast height)

Ontario, Canada.
data using C.A. and P.C.A., respectively.

 Lichens as Ecological Indicators of Photochemical Oxidant Air Pollution1

Lorene L. S i g a l and Thomas H. Nash 111~

From t h e l a t e 1 9 6 0 ' s t o t h e p r e s e n t , both t h e o c c u r r i n g i n t h e f i e l d . I n j u r y was documented a s a

c o n c e n t r a t i o n and dose of photochemical oxidant a i r
p o l l u t a n t s (ozone and peroxyacetyl n i t r a t e o r PAN)
have continued t o i n c r e a s e i n t h e mountainous a r e a s
surrounding t h e Los Angeles Basin. Since 1972, a
m u l t i - d i s c i p l i n a r y team of e c o l o g i s t s has monitored
and analyzed t h e e c o l o g i c a l consequences of t h i s
p o l l u t i o n i n t h e c o n i f e r f o r e s t ecosystem of t h e
San Bernardino Mountains. Our l i c h e n study p a r a l -
l e l s and expands t h e parameters of t h e aforemen-
t i o n e d s t u d y and a l l o w s a comparison of t h e r e l -
a t i v e s e n s i t i v i t y of l i c h e n s v e r s u s higher p l a n t s
t o oxidants.
H i s t o r i c a l l y , l i c h e n s have been demonstrated t o
be s e n s i t i v e i n d i c a t o r s of a i r p o l l u t a n t s such a s
s u l f u r d i o x i d e . More r e c e n t l y , t h e r e i s s t r o n g
evidence t h a t l i c h e n s a r e a l s o s e n s i t i v e t o hy-
drogen f l u o r i d e and heavy m e t a l s . The p r e s e n t
s t u d y documents f o r t h e f i r s t time t h a t l i c h e n s a r e
a l s o s e n s i t i v e t o ozone and peroxyacetyl n i t r a t e .
F i e l d i n v e s t i g a t i o n s were conducted i n f o u r
mountain ranges surrounding t h e Los Angeles Basin.
A f i f t h s i t e i n Cuyamaca Rancho S t a t e Park, e a s t of
San Diego, was chosen a s t h e c o n t r o l a r e a . R e s u l t s
of sampling l i c h e n s on Quercus k e l l o g g i i Newb. and
c o n i f e r s showed o v e r a l l s p e c i e s r i c h n e s s decreased
by 38 p e r c e n t i n t h e h i g h l y impacted San Bernardino
Mountains. Cover v a l u e s f o r s p e c i e s on Q. kellog-
decreased by 16 p e r c e n t ; f o r l i c h e n s on con-
i f e r s t h e d e c r e a s e was 78 p e r c e n t . The l a t t e r de-
c r e a s e i s l a r g e due t o t h e f a c t t h a t t h e "sen-
s i t i v e " f r u t i c o s e l i c h e n s p e c i e s a r e found mainly
on c o n i f e r s . There was an i n v e r s e r e l a t i o n s h i p
between t h e cover of l i c h e n s a t b r e a s t h e i g h t on
c o n i f e r s and t h e oxidant dose e s t i m a t e s a t s i t e s i n
t h e San Bernardino Mountains ( f i g . 1 ) . A t h r e s h o l d
i s seen a t 180 ppm-hrs above which t h e l i c h e n s a r e
almost a b s e n t .
s i g n i f i c a n t r e d u c t i o n i n g r o s s photosynthesis.
Parmelia s u l c a t a Tayl. e x h i b i t e d g r e a t e r s e n s i t i v -
i t y t o ozone and PAN than Hypogymnia enteromorpha
(Ach.) Nyl. These r e s u l t s were c o n s i s t e n t w i t h
f i e l d observations.
A s a r e s u l t of c o l l e c t i o n s made by H. E . Hasse
and t h e subsequent p u b l i c a t i o n of h i s "Lichen
F l o r a of Southern C a l i f o r n i a " i n 1913, t h e p r e s e n t
d i s t r i b u t i o n and v i t a l i t y of a number of l i c h e n s
was compared t o t h e i r d i s t r i b u t i o n a t t h e t u r n of
t h e century. There was a 50 p e r c e n t d e c r e a s e i n
species richness. I n a d d i t i o n , marked morpho-
l o g i c a l d e t e r i o r a t i o n was s e e n i n t h e t h a l l i of
e x i s t i n g f o l o s e s p e c i e s such a s Hypogymnia e n t e r -
omorpha. Forty-two p e r c e n t of t h e t h a l l i c o l l e c t e d
i n t h e San Bernardinos were bleached and 44 p e r c e n t
were convoluted i n c o n t r a s t t o no bleaching o r con-
v o l u t i o n i n c o l l e c t i o n s made i n t h e Cuymacas.
T h a l l u s dimensions were decreased by approximately
50 percent i n t h e San Bernardino c o l l e c t i o n s .
The importance of l i c h e n s t u d i e s l i e s i n t h e i r
s e n s i t i v i t y t o a i r p o l l u t i o n and t h e i r p o t e n t i a l
u s e a s b i o i n d - i c a t o r s . By monitoring t h e abundance
and d i s t r i b u t i o n of s e n s i t i v e l i c h e n s , i t i s pos-
s i b l e t o e s t i n a t e t h e magnitude of a i r p o l l u t i o n .
u s e of l i c h e n s i n t h i s m a t t e r may be f e a s i b l e n o t
only a s c u r r e n t l y a p p l i e d i n n o r t h e r n Europe w i t h
s u l f u r d i o x i d e a i r p o l l u t i o n , b u t a l s o over l a r g e
geographical a r e a s impacted by photochemical
oxidant a i r p o l l u t i o n .
Acknowledgment: This s t u d y was supported by
National Science Foundation Grant DEB-7610244 t o
D r . Thomas H. Nash 111, Dept. of Botany/Micro-
biology, Arizona S t a t e U n i v e r s i t y , Tempe, Arizona
85281.
-f 300 I i r
ORNL- DWG 8 0 - 7556 ESD
r 1

Fumigation s t u d i e s i n t h e l a b o r a t o r y provided a
d e f i n i t i v e way of t e s t i n g t h e response of f i e l d
s e n s i t i v e and f i e l d t o l e r a n t s p e c i e s t o c o n t r o l l e d Y
-
-CAMP PAIVIKA
-
p 220
c o n c e n t r a t i o n s of ozone and PAN s i m i l a r t o those w
'n
w -
in 180 BARTON FLATS
n
0
SKY FOREST ,GREEN VALLEY CREEK
140 - -
p r e s e n t e d a t t h e Symposium on E f f e c t s of A i r
P o l l u t a n t s on Mediterranean and Temperate F o r e s t
DEERLICK
^
Ecosystems, June 22-27, 1980, R i v e r s i d e , 0 100
1
O f 2 3 4 5 6 7
C a l i f o r n i a , U.S.A. 1. COVER OF LETHARIA VULPINA AND
HYPOGYMNIA ENTEROMORPHA
2
Research A s s o c i a t e , Oak Ridge National Lab- F i g u r e 1--Percent l i n e a r cover a t b r e a s t h e i g h t of
o r a t o r y , Environmental Sciences Division, P.O. Box L e t h a r i a v u l p i n a (1.) Hue and Hypop-ymnia
X, Oak Ridge, Tenn. 37830. Operated by Union enteromorpha (Ach.) Nyl. a s a f u n c t i o n of t h e
Carbide Corporation under c o n t r a c t W-7405-eng-26 oxidant dose e s t i m a t e s (ppm - h r s . = t h e mean
w i t h t h e U.S. Department of Energy; and A s s o c i a t e oxidant c o n c e n t r a t i o n i n ppm m u l t i p l i e d by t h e
P r o f e s s o r of Ecology and Lichenology, Arizona time of exposure) a t s i t e s i n t h e San Bernardino
S t a t e U n i v e r s i t y , Tempe, Ariz. 85281. Mountains.
 The Effects of Air Pollutants on Forest Ecosystems in S.R. Slovenia1
Marjan Solar
INTRODUCTORY DATA
S l o v e n i a w i t h i t s s u r f a c e o f 20,251 km2 repre-
sents 8% o f t h e t e r r i t o r y o f Yugoslavia (255,804
km2). I t i s s i t u a t e d i n t h e extreme northwestern
p a r t o f t h e s t a t e between t h e Alps, t h e Pannonian
p l a i n , t h e A d r i a t i c sea and t h e D i n a r i c o r o g r a p h i c
system. H a l f o f t h e s u r f a c e i s covered by f o r e s t s
(1,000,000 ha), 2/3 o f which a r e p r i v a t e l y owned.
The p o r t i o n o f c o n i f e r s amounts t o 56%, t h e average
wood mass i s n e a r l y 200 m3/ ha. The t o t a l increment
per year i s 4,000,000 m3, t h e c u t t i n g 3,000,000 m3.
Two-thirds o f t h e wood h a r v e s t possess a t e c h n i c a l
value. The p o r t i o n o f f o r e s t r y w i t h i n t h e n a t i o n a l
income i s 4% and i n d i c a t e s w e l l t h e l e v e l o f t h e
i n d u s t r i a l development o f Slovenia.
S l o v e n i a i s d i v i d e d up i n t o 15 f o r e s t management
areas and each o f those i n t o t h e s o c i a l l y and p r i -
v a t e l y owned s e c t o r s . The p r i n c i p l e s o f management
a r e u n i f i e d . The b a s i c p r i n c i p l e i s t o manage as
much p r o n a t u r a l l y as p o s s i b l e t o secure t h e perma-
nency o f y i e l d and f u n c t i o n s o f t h e f o r e s t . The f o r -
e s t s e r v i c e e n j o y s i n some places a 200-year
tradition.
The study o f e f f e c t s o f a i r p o l l u t a n t s on t h e
f o r e s t s goes back t o t h e year 1926, and systematic
i n v e s t i g a t i o n s were s t a r t e d i n 1969.
OBJECTIVES OF INVESTIGATION
We w i s h t o determine t h e parameters, i n t e n s i t y
and p e r s p e c t i v e s o f t h e e f f e c t s o f a i r p o l l u t i o n on
t h e f o r e s t s i n Slovenia, w i t h t h e aim t o secure
r e l i a b l e f o u n d a t i o n s f o r t h e p l a n n i n g o f t h e indus-
t r i a l development r u n n i n g p a r a l l e l l y w i t h t h e p r e -
servation o f the multifunctional forest, f o r the
r e g u l a t i o n o f i n d e m n i t i e s and t h e e l a b o r a t i o n o f
c o r r e c t n o r m a t i v e concerning t h e maximum t o l e r a b l e
concentrations valuable f o r t h i s s p e c i f i c f o r e s t
area.
PRELIMINARY FOUNDATIONS REQUIRED
Ecology (geology, s o i 1 s , phytocoenology , c l imate)
as bases f o r t h e d e t e r m i n a t i o n o f f o r e s t ecosystems
n o t s t r e s s e d by emission, f o r e s t i n v e n t o r y as a ba-
s i s f o r t h e d e t e r m i n a t i o n o f t h e normal management
P r e s e n t e d a t t h e Symposium on E f f e c t s o f A i r
P o l l u t a n t s on Mediterranean and Temperate F o r e s t
Ecosystems, June 22-27, 1980, R i v e r s i d e ,
C a l i f o r n i a , U.S.A.
~ r a d u a t ef o r e s t engineer, h i g h e r research
c o o p e r a t o r o f t h e I n s t i t u t e f o r t h e F o r e s t and
Wood Economy a t t h e B i o t e c h n i c a 1 F a c u l t y i n
L j u b l j a n a , Yugoslavia.
s i t u a t i o n o f t h e f o r e s t s and a r e g i s t e r o f e m i t t e r s .
METHODS OF ASSESSING THE EFFECTS
OF POLLUTED AIR ON FORESTS
Symptomatics, chemical analyses, increment a n a l -
yses, b i o i n d i c a t o r s , presence o f d i f f e r e n t l y aged
needles, presence o f secondary p e s t s , i n f r a c o l o r -
aerophotographs.
SPECIFIC FEATURES OF THE TERRITORY OF SLOVENIA
Accentuated ~ o n f i g u r a t i o n ~ i n d u s t royf t e n l o c a t e d
i n narrow v a l l e y s , temperature i n v e r s i o n , extremely
h i g h emission values (shocks), h i g h p r o p o r t i o n o f
s u s c e p t i b l e c o n i f e r s , e r o d i b l e s o i l s , predominancy
o f SO2 and HF.
RESULTS OF INVESTIGATION
The t o t a l s u r f a c e o f v i s i b l y damaged f o r e s t s i n
S.R. Slovenia amounts t o 22,000 ha i . e . 2.2% o f t h e
t o t a l f o r e s t area ( s i t u a t i o n i n 1977). The i n t e r n a l
d i v i s i o n o f damaged f o r e s t s i s t h e f o l l o w i n g :
Group A-the narrower emission area- t h e f o r e s t des-
t r o y e d and e x i s t e n s i a l l y threatened, o r s u r f a c e s
under t h e impact o f c r i t i c a l emission c o n d i t i o n s
e n t a i l i n g t h e t o t a l d e s t r u c t i o n o f a11 f o r e s t
f u n c t i o n s - d e s t r u c t i o n o f t h e f o r e s t ecosystem.
Surface 4,000 ha.
Group B-the w i d e r emission a r e a - f o r e s t under p e r -
manent emission impact, y e t t h e e x i s t e n c e o f t h e
f o r e s t i s n o t threatened, i t s f u n c t i o n s a r e however
reduced, o r surfaces under t o heavy emission con-
d i t i o n s t o a l l o w a normal t h r i v i n g o f t h e f o r e s t .
The emission causes an i n s t a b i l i t y o f f o r e s t eco-
systems. Surface 16,000 ha.
Group C-Periodic emission a r e a - p e r i o d i c occurrence o f
damages, p o s s i b l y a c u t e ones, b u t between i n d i v i d -
u a l emission i n f l u e n c e s i n every case l o n g e r normal
s i t u a t i o n s a r e i n t e r c a l a t e d d u r i n g which t h e f o r -
e s t can recover. Surface 2,000 ha.
Based on t h e chemical analyses o f Spruce needles
as t o t h e increased c o n t e n t o f sulphur and f l u o r i d e ,
t h e s u r f a c e o f f o r e s t s w i t h hidden i n j u r i e s i s es-
t i m a t e d t o 25,000 ha. Survey and d e t a i l e d maps o f
i n j u r e d f o r e s t have been made f o r a11 b i g g e r emis-
s i o n s areas. On t h e b a s i s o f r e l a t i v e r e s i s t a n c e
c a p a c i t y o f f o r e s t t r e e s and o f t h e i r p r o p o r t i o n a l
presence i n t h e f o r e s t v e g e t a t i o n communities a
categorization o f f o r e s t s w i t h respect t o t h e i r
s u s c e p t i b i l i t y t o a i r p o l l u t i o n has been c a r r i e d o u t .
The process o f a f u r t h e r improvement o f t h e
emission damages i n t h e f o r e s t s observed d u r i n g t h e
l a s t t h r e e years i s n o t l i k e l y t o proceed because
o f t h e increase o f consumption o f c o a l possessing
h i g h c o n t e n t s o f combustible s u l p h u r .
 CONCLUSIONS
On t h e b a s i s o f i n v e s t i g a t i o n s c a r r i e d o u t i n
t h e t o t a l f o r e s t area o f Slovenia i n f l u e n c e d by
emission, o f knowledge concerning t h e ecology o f
t h i s t e r r i t o r y and t h e p a r t i c u l a r i t i e s o f e f f e c t s
o f t h e p o l l u t e d a i r , we a r e a b l e t o p r e d i c t exact-
l y t h e f u t u r e o f a c e r t a i n f o r e s t under c e r t a i n
emission i n f l u e n c e .
The s t a n d p o i n t o f f o r e s t e r s i s always t h e f o l -
lowing: Wherever t h e f o r e s t has been destroyed
o r i s on t h e way o f d e s t r u c t i o n , t h e a i r i s 10-
c a l l y oolluted t o a r e l a t i v e l y c r i t i c a l extent.
The a i r p o l l u t i o n i s t o o h i g h a l s o i n places
where t h e f o r e s t i s hampered i n i t s development.
Appealing t o t h e normatives f i x e d by law and
concerning t h e maximum emission values allowed
means n o t h i n g b u t t h a t t h e normatives a r e s e t
t o o h i g h and t h a t t h e f o r e s t e r s a r e bound t o
lower them.
The R e s o l u t i o n issued on t h e xth ~ e e t i no f~ t h e
IUFRO Group 2.09-Air P o l l u t i o n , L j u b l j a n a 1978 i s
proposing t h e normatives i n s u r i n g t h e normal
t h r i v i n g o f a11 k i n d o f f o r e s t s on average and
extreme s i t e s . The values f o r SOi a r e f o l l o w i n g
( i n pg/m3/air) : 50 as y e a r l y average, 100 as 24 h
average, 150 as & h value, and f o r HF ( i n
ug/m3/air) 0.3 as y e a r l y average and 0.9 as & h
value. For extreme s i t e s t h e normatives a r e t w i c e
as sharp.
I f we compare t h e normatives f i x e d by Resolu-
t i o n t o those f i x e d by law i n d i f f e r e n t c o u n t r i e s
we r e a l i s e t h e l a s t ones a r e much t o o h i g h .
 Population Differences in Response to Sulfur Dioxide: a Physiological Analysis1
G. E. Taylor, Jr. and D. T. Tingey
The environment e x e r t s a profound i n f l u e n c e on
a p l a n t ' s a b i l i t y t o s u r v i v e and reproduce, and
consequently v e g e t a t i o n i n a g i v e n a r e a w i l l
p o s s e s s a s e t of morphological and p h y s i o l o g i c a l
t r a i t s t h a t enhance f i t n e s s . A t t h e l e v e l of
populations, these site-specific a t t r i b u t e s a r i s e
through e i t h e r a m o d i f i c a t i o n of t h e phenotype
(phenotypic p l a s t i c i t y ) o r a change i n t h e gene
pool (ecogenetic adaptation). Both r e s p o n s e s a r e
common s t r a t e g i e s among n a t u r a l p o p u l a t i o n s exper-
i e n c i n g d i s p a r a t e c l i m a t i c , e d a p h i c and b i o t i c
stresses. It is h y p o t h e s i z e d t h a t e l e v a t e d l e v e l s
of a t m o s p h e r i c p o l l u t a n t s have e l i c i t e d i n a com-
p a r a b l e manner a v a r i e t y of t r a i t s t h a t enhance
s u r v i v a l and r e p r o d u c t i o n i n n a t i v e p l a n t s inhab-
i t i n g pollution-stressed areas.
One example of e c o g e n e t i c a d a p t a t i o n i n
r e s p o n s e t o a i r p o l l u t i o n i s t h e e v o l u t i o n of
s u l f u r d i o x i d e r e s i s t a n c e w i t h i n p o p u l a t i o n s of
Geranium c a r o l i n i a n u m , a n herbaceous w i n t e r a n n u a l
common i n d i s t u r b e d h a b i t a t s i n t h e S o u t h e a s t e r n
United S t a t e s . I n comparison w i t h t h e i r counter-
p a r t s from p o l l u t i o n - f r e e r e g i o n s , p o p u l a t i o n s
sampled from a r e a s e x p e r i e n c i n g v a r i a b l e SO2
s t r e s s f o r 3 1 y e a r s were c o n s i s t e n t l y more r e s i s -
t a n t t o SO2 under c o n t r o l l e d exposure c o n d i t i o n s .
This i n f r a s p e c i f i c v a r i a t i o n is g e n e t i c a l l y deter-
mined and q u a n t i t a t i v e l y c o n t r o l l e d . P l a n t s of
c o n t r a s t i n g SO2 r e s i s t a n c e were used t o invest!+
g a t e t h e p h y s i o l o g i c a l b a s i s of t h i s a d a p t a t i o n .
I n d i v i d u a l p l a n t s were p l a c e d i n a whole-plant
gaseous exchange system i n which c o n c u r r e n t s t e a d y
s t a t e measures of l e a f r e s i s t a n c e t o w a t e r vapor
e f f l u x and SO2 i n f l u x were monitored i n t h e d a r k
and l i g h t a t p o l l u t a n t c o n c e n t r a t i o n s o f 0.4, 0.6,
and 0.8 u l 1 - l . For r e s i s t a n t and s e n s i t i v e
p l a n t s a t each c o n c e n t r a t i o n , e s t i m a t e s of t o t a l
SO2 f l u x (pg c m 2 h r l ) a s a f u n c t i o n of l e a f
r e s i s t a n c e t o Hz0 e f f l u x were modeled u s i n g l i n e a r
r e g r e s s i o n t e c h n i q u e s . From e s t i m a t i o n p r o c e d u r e s ,
t o t a l f l u x was p a r t i t i o n e d i n t o l e a f s u r f a c e and
internal fractions.
T o t a l SO2 f l u x v a r i e d a s a f u n c t i o n of l e a f
r e s i s t a n c e and d i d n o t d i f f e r among r e s i s t a n t and
s e n s i t i v e p l a n t s a t 0.4 and 0.6 u l l 1 SOz. Con-
v e r s e l y , a t t h e h i g h e s t c o n c e n t r a t i o n t o t a l SO2
f l u x was n o t t h e same f o r t h e 2 p l a n t groups a s a
consequence of d i s p a r a t e s l o p e parameters. Irre-
s p e c t i v e of c o n c e n t r a t i o n t h e l e a f s u r f a c e and
i n t e r i o r were major s i n k s f o r SO2 (Fig. 1 ) . Each
f r a c t i o n increased l i n e a r l y with concentration,
' p r e s e n t e d a t t h e Symposium on E f f e c t s of A i r
P o l l u t a n t s on M e d i t e r r a n e a n and Temperate F o r e s t
Ecosystems, J u n e 22-27, 1980, R i v e r s i d e ,
C a l i f o r n i a , U.S.A.
2 ~ o s t d o c t o r a lA s s o c i a t e and P l a n t P h y s i o l o g i s t ,
T e r r e s t r i a l D i v i s i o n , U.S. Environmental Protec-
t i o n Agency-Corvallis Environmental Research
Laboratory, C o r v a l l i s , Oregon, U.S.A.
2
however t h e r a t i o o f i n t e r n a l t o t o t a l f l u x de-
c r e a s e d s t e a d i l y . The a b s o l u t e v a l u e s f o r i n t e r -
n a l SO2 f l u x were s t r i k i n g l y s i m i l a r f o r b o t h
p l a n t groups a t each c o n c e n t r a t i o n . T h e r e f o r e , i n
s p i t e of t h e v a r i a t i o n i n t o t a l SO2 f l u x t o t h e
p l a n t , t h e r a t e of SO2 absorbed i n t o t h e l e a f
i n t e r i o r was e q u i v a l e n t f o r r e s i s t a n t and s e n s i -
t i v e plants. Since l e a f r e s i s t a n c e t o water vapor
e f f l u x i s t h e same f o r a l l p l a n t s i n b o t h ambient
and S02-polluted atmospheres, o v e r t p l a n t - t o - p l a n t
d i f f e r e n c e s i n t h e i r r e s p o n s e t o SO2 r e f l e c t
d i s p a r a t e i n t e r n a l biochemical p r o c e s s e s a f f e c t i n g
pollutafit t o x i c i t y , perturbation or c e l l u l a r
repair.
The f a c t t h a t p l a n t d i f f e r e n c e s i n r e s p o n s e t o
SO2 a r e g e n e t i c a l l y c o n t r o l l e d and n o t a r e s u l t of
p o l l u t a n t exclusion is relevant t o an understanding
of t h e r a p i d e v o l u t i o n of p o p u l a t i o n r e s i s t a n c e i n
-
G. carolinianum. T h i s s p e c i e s i s a n a n n u a l t h a t
t h r i v e s f o r s i x months a s a w i n t e r r o s e t t e d u r i n g
which new l e a f growth i s minimal and y e t t r a n s -
p i r a t i o n remains a c t i v e . This h a b i t coupled w i t h
t h e p l a n t ' s i n a b i l i t y t o t r a c k and e x c l u d e e l e -
v a t e d l e v e l s of SO2 may p r e d i s p o s e t h i s s p e c i e s
t o accumulate more SO2 d e r i v a t i v e s t h a n o t h e r
co-occurring s p e c i e s t h a t a r e e i t h e r more ephemeral
o r c a p a b l e of a v o i d i n g SO2 u p t a k e increasing
stomata1 resistance. These s p e c i e s ' a t t r i b u t e s
may e x p l a i n t h e c o n s i s t e n t f i e l d o b s e r v a t i o n s t h a t
show c. carolinianum t o b e a s e n s i t i v e b i o l o g i c a l
i n d i c a t o r of e l e v a t e d SO2 l e v e l s .
Acknowledgments: Research sponsored by t h e
N a t i o n a l Academy of Sciences-National Research
Council through a P o s t d o c t o r a l Research A s s o c i a t e -
s h i p t o t h e s e n i o r author. S e n i o r a u t h o r acknow-
l e d g e s s u p p o r t f o r manuscript p r e p a r a t i o n under
C o n t r a c t W-7405-eng-26 by t h e Department of Energy
w i t h Union Carbide Corporation, a t Oak Ridge
N a t i o n a l Laboratory, Oak Ridge, Tennessee.
6.0 , t r 1
-- RESISTANT PLANTS SENSITIVE PLANTS
SULFUR DIOXIDE CONCENTRATION ( p l I"')
F i g u r e I--Absolute v a l u e s f o r t o t a l , s u r f a c e and
i n t e r n a l l e a f f l u x of SO2 a s a f u n c t i o n of
p o l l u t a n t c o n c e n t r a t i o n i n r e s i s t a n t and' s e n s i t i v e
plants.
 Ozone Injury to Pines in the Southern Sierra Nevada of California1
D e t l e v R. Vogler and John Pronos
The F o r e s t S e r v i c e began e v a l u a t i n g t h e impact
o f ozone on S i e r r a Nevada f o r e s t s i n 1974. Eval-
u a t i o n e f f o r t s were expanded and i n t e n s i f i e d i n
1977, and s i n c e then t h e primary methods o f
assessment have included, ( 1 ) m o n i t o r i n g ambient
ozone levels, and (2) r a t i n g i n j u r y t o p i n e f o l -
iage i n permanent t r e n d p l o t s .
A i r p o l l u t i o n i n j u r y t o commercial p i n e stands
located between 4000 and 8000 f e e t e l e v a t i o n
occurs q u i t e u n i f o r m l y along most o f t h e southern
S i e r r a Nevada f r o n t range adjacent t o t h e San
Joaquin V a l l e y and up i n t o t h e major r i v e r d r a i n -
ages. Symptoms a r e n o t confined t o l o c a l i z e d
areas downwind from major p o l l u t a n t sources.
Rather, ozone i s c a r r i e d down t h e e n t i r e San
Joaqu i n A i r Basin and eastward i n t o t h e S i e r r a .
Metropo I i t a n areas c o n t r i b u t i n g t o t h e ozone dose
inc I ude Stockton, Modesto, Merced, Fresno,
Visa1 i a B a k e r s f i e l d , and perhaps even Sacramento
and t h e San Francisco Bay Area.
OZONE MONITORING
F i v e f o r e s t e d l o c a t i o n s i n t h e southern S i e r r a
were moni+ored f o r season-long ozone dosages
between 1977 and 1979. One s i t e -- Whitaker
Forest -- was monitored c o n t i n u o u s l y d u r i n g t h i s
3-year period. The s i t e s ranged i n e l e v a t i o n
from 5400 f e e t t o 7540 f e e t , and a l l were a t
l e a s t 50 m i l e s from suspected m e t r o p o l i t a n sources
of a i r pollution.
Ozone l e v e l s a t each s i t e exceeded both t h e
Federal (0.12 ppm) and S t a t e (0.10 ppm) Standards
each year they were recorded. Based on t h e num-
b e r o f hours exceeding t h e Standards, 1977 was
t h e w o r s t year f o r ozone, w h i l e l e v e l s d e c l i n e d
s t e a d i l y i n 1978 and 1979. During t h e summer o f
each year d a i l y peak ozone values commonly ranged
between 0.10 ppm and 0.14 ppm. These values can
be compared t o those i n t h e San Bernardino Moun-
t a i n s o f southern C a l i f o r n i a , where maximum d a i l y
ozone l e v e l s f r e q u e n t l y range from 0.20 t o 0.33
ppm, and where i n j u r y t o p i n e f o r e s t s i s corres-
pondingly more severe.
TREND PLOTS
Presented a t t h e Symposium on E f f e c t s o f A i r
P o l l u t a n t s on Mediterranean and Temperate Forest
Ecosystems, June 22-27, 1980, Riverside,
C a l i f o r n i a , U.S.A.
'plant P a t h o l o q i s t s , P a c i f i c Southwest Region,
F o r e s t Service, Ui s . ~e partment of A g r i c u ture,
San Francisco, Ca I i f o r nia.
2
Trend p l o t data were co I l e c t e d f o r t h r e e con-
s e c u t i v e years beginning i n 1977. I n 1977 and
1978 ozone i n j u r y r a t i n g s of p l o t s s c a t t e r e d
throughout a sampl i n g area o f over 1 m i I l i o n acres
ranged between no i n j u r y and moderate i n j u r y . A l -
though t h e m a j o r i t y o f 1978 p l o t s showed more o x i -
dant symptoms t h a n 1977 p l o t s , t h e r e was no sta-
t i s t i c a l l y s i g n i f i c a n t t r e n d evident. The change
between 1977 and 1979, however, was more dramatic.
Nineteen o f 27 p l o t s evaluated i n 1979 showed
a d d i t i o n a l i n j u r y o v e r 1977 levels, and seven o f
these p l o t d i f f e r e n c e s were s t a t i s t i c a l l y s i g n i f i -
c a n t (P = 0.05). I n 1979, f o r t h e f i r s t time,
several p l o t s f e l I i n t o t h e severe i n j u r y category.
The t r e e s i n these p l o t s averaged 2 years o f
needle r e t e n t i o n w i t h ozone symptoms on Z-year-
ol'd needles.
T h i s obvious increase i n i n j u r y was n o t expec-
t e d because measured ozone l e v e l s i n 1979 were
g e n e r a l l y lower than i n e i t h e r 1978 o r 1977.
Considerable p h y s i o l o g i c a l s t r e s s from a 2-year
drought (1976-1977) c o u l d account f o r p a r t o f t h e
observed increase i n f o l i a r i n j u r y . One v i s i b l e
response o f pines t o t h e drought was a dramatic
decrease i n needle r e t e n t i o n , which tended t o
produce more severe ozone i n j u r y r a t i n g s . F u t u r e
recovery o f t r e e s from drought stress, i n t h e form
o f increased needle r e t e n t i o n , may r e s u l t i n a
r e l a t i v e r e d u c t i o n o f measureable a i r p o l l u t i o n
i n j u r y d u r i n g t h e n e x t few years.
CONCLUSIONS
The o v e r a l l l e v e l o f forest-wide i n j u r y i n t h e
south'ern S i e r r a Nevada can be termed s l ight, w i t h
some l o c a l p o p u l a t i o n s o f s u s c e p t i b l e pines show-
ing moderate o r worse i n j u r y . Unexpectedly h i g h
amounts o f i n j u r y detected i n c e r t a i n areas i n
1979 placed some p l o t s i n t o t h e severe category.
I t i s p r e s e n t l y u n c l e a r whether t h i s i s a perma-
nent t r e n d of increasing impact, o r j u s t a short-
term r e s u l t o f drought stress.
With o n l y 3 years o f m o n i t o r i n g data, t r e n d s
i n annual ozone dose a r e d i f f i c u l t t o assess.
Ozone l e v e l s seemed t o decrease s l i g h t l y each year
between 1977 and 1979, even though v i s i b l e i n j u r y
t o pines increased. Measured v a r i a t i o n s i n sea-
sonal ozone dose may be associated more w i t h
weather p a t t e r n s than w i t h decreased pol I u t a n t s
a t t h e source. Maximum d a i l y ozone values i n t h e
S i e r r a s t i l l remain about 1/2 o r less of those
o c c u r r i n g i n southern C a l i f o r n i a . As ozone con-
t i n u e s t o be t r a n s p o r t e d i n t o t h e western slopes
o f t h e S i e r r a Nevada, s e n s i t i v e pines w i l I con-
t i n u e t o show a d d i t i o n a l i n j u r y and w i l I s l o w l y
decl lne.
 Modifications of Chemical Contents of Precipitation
by Passage through Oak Forests1

George T. Weaver and Jon D. ones^

Bulk precipitation monitored within oak forests

creased during winter. In the spring of 1980

in the Ozark Physiographic Province in Illinois

maximum bulk precipitation pH was 6.6 but passage

was strongly acidic (pH<4.5) throughout most of

through the canopy caused a decrease in pH as

. 1978. Exceptions occurred during late summer and great as 0.8 units. Soluble organic compounds,

early autumn when pH rose to 5.8. This pattern

apparently leached from some oaks, were present

with similar pH values reoccurred during the au-

in sufficient quantities to impart a dark stain

tumn and winter of 1979-80 and differs from sea-

to canopy drip during this period and may be as-

sonal trends usually reported from the northeast-

sociated with the reversal in H exchange in the

ern United States where periods of greatest and

canopy.

least acidity are summer and winter, respectively.

During the 1979-80 period, the highest concentra-

The concentrations of ions in soil water fonn-

tions of ~ 0 occurred
~ ~ durin~
-
autumn, and on an
ed three distinct patterns relative to concentra-

equivalence basis, exceeded H concentrations by

tions in bulk precipitation and canopy drip. Hy-

a factor of 10. Concentrations of ~ a ~ , g and,

drogen ion concentration was decreased markedly

K+ were also high during autumn. It is hypothe-

by passage through these ecosystems and the quan-

sized that considerable neutralization of strong

tity remaining in soil water was only about 10

acids occurred in the atmosphere due to the pre-

percent as great as in bulk preci itation. The

sence of airborne dust which normally exists dur-

concentrations of four ions -,-:oS m 3 - , ca2+-
ing autumn.
and W2+ - increased as water passed through these
ecosystems. The levels of these ions in soil wa-

Additional neutralization occurred as precipi-

ter were as great as 37 times and 9 times the

tation passed through the forest canopy, especial-

levels in bulk precipitation and canopy drip, re-

ly during autumnal leaf senescence. At this time

spectively, although major differences occurred

canopy drip pH exceeded bulk precipitation among seasons and between ions. The contents of

up to 0.7 units. The concentrations of Ca2qH , by

K+ and ~ 0 in ~
bulk-precipitation

also were in-

M g 2 ,,'K 804 and ~ 0 in~canopy

~ drip
- increased

creased by up to 25 times by passage through the

markedly compared to bulk precipitation, particu-

forest canopy. However, concentrations of these

larly during autumn. During the winter, canopy

particular ions either decreased or remained simi-

drip pH decreased to values as low as 3.8 and dif-

lar to concentrations in canopy drip after passage

fered little from bulk precipitation pH. In 1978,

through the soil.

68 percent of the precipitation reaching the for-

est floor as canopy drip was acidic (pH<5.6). The

Between October 1979 and April 1980, ca2+ and

removal of nutrient ions from the canopy also de-

sod2- were the predominant ions (equivalence basis)

associated with meteorologic and hydrologic proc-

esses in these ecosystems. The importance of Ca2+

presented at the Symposium on Effects of Air

in these ecosystems was anticipated since it is

Pollutants on Mediterranean and Temperate Forest

selectively accumulated by some species of oaks in

Ecosystems, June 22-27, 1980, Riverside,

these forests. It is also apparent that elevated

California, U.S.A.
levels of 'H are being deposited in association

with in these ecosystems, but the impacts

~ s s o c i a t eProfessor of Forestry, Southern

remain undetermined.

Illinois University, Carbondale, 111.; Forestry

Aide, Baruch Science Institute, Georgetown, S.C.

 Seasonal Variation of Inorganic and Organic Sulfur in Coniferous Needles

Intensified by SO2 Pollution '
2
Karl Friedrich Wentzel und SUnther Gasch

-
Method:

Needles front 50 - 70 year old Norway spruce from 1. gardens and

parks of Wiesbaden town (slight damage), and 2. Wiesbaden forests
(no v i s i b l e injury) were used as bioindicators f o r $0.-uptake.
I n octobre we picked 05., 1.5 and 25. year old needles, i n m y
we picked 1, 2 and 3 year old needles fro@the tops of the sam
trees. Inorganic and organic sulfur was determined according t o
JSger und Steubing (1970).

-
Results:

1. Air pollution measuremnt 1976 - 1979. Datas i n microgram fig. 1 :Map of Wiesoaden/FRG
SO per cbw air. 11 = annual average, 12 = 95 percentile of
2
30 win. values:
Zone II 1 2

Industrial Area 120 - 140 400 - 500

Town 7 0 - 90 200-300
Forests <50 < 120

2. Total sulfur content increases with needle age as shown i n

Fig. 2. I n town the S contents d the needles are 100 - 200 ppm
higher than those froa the forests.
500. - Wiesbaden Town
3. The inorganic fraction of a l l needle samples surmounts the
-- - - - Wiesbaden Forest
300
0.5 1.0 1.5 2.0 25 3D Needle4 e
organic fraction. The difference i s greater i n autumn than i n
Fig. 2 :Sulfur content ( p p of dr. wt.) i n spruce needles from
spring and more apparent i n town than i n the forests. Wiesbaden town (means of 39 tree samples) and forests (mans of
16 tree samples).
4. The v a l i d i t y of using coniferous needles as indicator of a i r
pollution effects i s best when octobre datas of inorganic S are
Oclobre- deltas
used. . . . . -
May deltas

5. Organic sulfur content decreases during the vegetation period 1 A Decrease of inoro. S

while the inorganic fraction increases. During winter time the

opposite occurs (Fig. 3). From t h i s i t i s suggested that i n
spring t i n e a part of the organic sulfur d m n d i s covered by
remvinq the inorganic amount, which p a r t i a l l y i s of air-borne
origin.

1 Presented a t t h e I n t e r n a t i o n a l Symposium on
E f f e c t s o f A i r P o l l u t a n t s on Mediterranean and
Temperate Forese Ecosystems, June 22-27, 1980,
I
Wiesbaden Town 39 trees in casualorder
-
Riverside, C a l i f o r n i a , U.S.A.
Fig. 3 : Seasonal alteration of inorganic and organic S i n spruce
2 Forest E c o l o g i s t and chemist, r e s p e c t i v e l y , needles frw Wiesbaden town (mans of 6 semi-annual needle sets).
Hessische Landesanstalt f u r Umwelt, Wiesbaden,
West Germany
 Sulfur Dioxide and Oxidant Effects on Californian Coastal Sage Scrub1
W.E. Westman and K.P. p r e s t o n z
Two f i e l d s t u d i e s have r e l a t e d o x i d a n t and
s u l f u r dioxide p o l l u t i o n t o a deterioration i n
t h e s t r u c t u r e and f u n c t i o n o f s o u t h e r n C a l i f o r -
n i a ' s drought-deciduous shrublands ( c o a s t a l sage
scrub). I n a s t u d y o f 67 s i t e s (0.63ha) f r o m
San F r a n c i s c o t o E l R o s a r i o ( B a j a C a l i f o r n i a ) ,
d a t a were c o l l e c t e d on s p e c i e s c o v e r and 43
h a b i t a t v a r i a b l e s d e s c r i b i n g topography, s o i 1 ,
c l i m a t e , v e g e t a t i o n s t r u c t u r e , age s i n c e f i r e ,
g r a z i n g i n t e n s i t y , and a i r p o l l u t i o n o f s i t e s .
The v a r i a b l e which showed t h e h i g h e s t s i q n i f i -
c a n t c o r r e l a t i o n w i t h p e r c e n t f o l i a r cover o f
n a t i v e species was t h e mean annual o x i d a n t
c o n c e n t r a t i o n (r=-.58, P<.001). E l e v a t i o n and
mean maximum temperature o f t h e warmest month
a l s o showed h i g h l y s i g n i f i c a n t c o r r e l a t i o n s
(r=-.52). The p a r t i a l c o r r e l a t i o n c o e f f i c i e n t s
o f o x i d a n t s w i t h p e r c e n t cover remained h i g h
when c o v a r i a t i o n s w i t h e l e v a t i o n , mean maximum
temperature o f t h e warmest month ,and d i s t a n c e
f r o m t h e c o a s t were e x t r a c t e d ( r = - .41,- .35,- .42
respectively) . The i n t e r r e l a t i o n s h i p s o f these
v a r i a b l e s were i n v e s t i g a t e d f u r t h e r by means o f
p a t h a n a l y s i s . A p a t h model ( c h i - s q u a r e p r o b a b i -
l i t y = .87) r e l a t e d environmental f a c t o r s t o a
r e d u c t i o n i n t h e p e r c e n t c o v e r o f n a t i v e species.
The p a t h model suggested t h a t o t h e r f a c t o r s c o r -
r e 1 a t e d w i t h d e c l in i ng c o v e r were a c t i n g p r i m a r i -
1y t h r o u g h t h e i r i n f l u e n c e on o x i d a n t concentra-
t i o n i n p r e d i c t i n g t h e d e c l i n e i n cover o f n a t i v e
scrub species.
I n c r e a s i n g c o n c e n t r a t i o n s o f o x i d a n t s were
a l s o a s s o c i a t e d w i t h a d e c l i n e i n species r i c h -
ness (r=-.23; P<.05) and i n c r e a s e i n e q u i t a -
b i l i t y ( W h i t t a k e r ' s Ec i n d e x ; r=.24; P<.05).
Diversity i n f l o r i s t i c a l l y s i m i l a r s i t e s o f high
and low annual o x i d a n t c o n c e n t r a t i o n s was
compared. H i g h l y p o l 1u t e d s i t e s have fewer
species p e r abundance c l a s s and a l o w e r t o t a l
species r i c h n e s s ("7=18 v . T=29 a t l e s s p o l l u t e d
s i t e s ) . C o n c e n t r a t i o n of dominance i n c r e a s e s i n
t h e more p o l l u t e d s i t e s . M a j o r r e s u l t s have been
r e p o r t e d i n Wes tman ( 1979).
I n a second f i e l d study, t h e e f f e c t s o f s u l f u r
d i o x i d e emissions (up t o 0.13 ppm f o r a p e r i o d o f
25 y e a r s ) from an o i l r e f i n e r y were s t u d i e d n e a r
Santa Maria, on t h e r u r a l c e n t r a l c o a s t o f C a l i -
f o r n i a . Stands o f c o a s t a l sage s c r u b w i t h b l a c k
sage (Sal v i a me1 1if e r a ) downwind o f t h e r e f i n e r y
were compared w i t h stands i n r e l a t i v e l y p o l l u t i o n -
Â¥'Â¥presenta t t h e Symposium on E f f e c t s o f A i r
P o l l u t a n t s on Mediterranean and Temperate F o r e s t
Ecosystems, June 22-27, 1980, R i v e r s i d e ,
C a l i f o r n i a , U.S.A.
' ~ s s o c i a t e P r o f e s s o r o f Geography and graduate
student, r e s p e c t i v e l y , U n i v e r s i t y o f C a l i f o r n i a ,
Los Angeles, Cal i f o r n i a.
f r e e upwind s t t e s . I n j u r y t o i n d i v i d u a l S a l v i a
shrubs and changes i n community s t r u c t u r e and
f l o r i s t i c c o m p o s i t i o n were recorded. Stomata1
r e s i s t a n c e was found t o be s i g n i f i c a n t l y l o w e r
on t h e p o l l u t e d s i t e s w i t h a concomitant 35 p e r -
c e n t i n c r e a s e i n mean t r a n s p i r a t i o n r a t e s o f
S a l v i a . F l o w e r i n g c a p a c i t y o f S a l v i a , measured
by t h e number o f f l o w e r w h o r l s p e r f l o w e r s p i k e ,
was a l s o s i g n i f i c a n t l y reduced. S i g n i f i c a n t
decreases i n t h e h e i g h t / w i d t h r a t i o o f S a l v i a
were h y p o t h e s i z e d t o be caused by s u l f i t e -
mediated d e s t r u c t i o n o f i n d o l e a c e t i c a c i d (Yang
and Saleh, 1973) and p o s s i b l e i n h i b i t i o n o f
a p i c a l dominance. S i g n i f i c a n t r e d u c t i o n s i n
photos.ynthetica11.y a c t i v e t i s s u e o f S a l v i a
r e s u l t e d f r o m i n c r e a s e d d e f o l i a t i o n and r e d u c t i o n
i n l e a f s i z e a s s o c i a t e d w i t h SO2 s t r e s s . The
evidence i n d i c a t e s t h a t these f a c t o r s decreased
t h e s h r u b s ' a b i l i t y t o compete w i t h t h e more r-
s e l e c t e d annuals. As such, t h e number o f
species, p r i m a r i l y annuals, i n c r e a s e d c o n s i d e r -
a b l y i n t h e most o o l l u t e d s i t e s (X=27 v. 7=18.on
c o n t r o l s i t e s ) . These SO2 - a s s o c i a t e d changes
i n community s t r u c t u r e and f l o r i s t i c c o m p o s i t i o n
suggest t h a t r e t r o g r e s s i o n i s o c c u r r i n g , c a u s i n g
t h e 26-year o l d s t a n d t o resemble a 7-year o l d
p o s t - f i r e sera1 'stage. R e s u l t s a r e r e p o r t e d
more f u l l y i n P r e s t o n ( 1 980). L a b o r a t o r y s t u d i e s
a r e i n progress t o assess s e n s i t i v i t y o f sage
scrub species t o SO2 i n j u r y .
Preston, K.P.
1980. E f f e c t s o f s u l f u r d i o x i d e p o l l u t i o n on
c o a s t a l sage scrub. M.A. Thesis. Department
o f Geography, U n i v e r s i t y o f C a l i f o r n i a , Los
Angel es .
Westman, W.E.
1979. O x i d a n t e f f e c t s on C a l i f o r n i a n c o a s t a l
sage scrub. Science 205:1001-1003.
Whi t t a k e r , R.H.
1972. E v o l u t i o n and measurement o f species
d i v e r s i t y . Taxon 21 :213-251.
Yang, S.F. and S.A. Saleh.
1973. D e s t r u c t i o n o f i n d o l e - 3 - a c e t i c a c i d
d u r i n g t h e a e r o b i c o x i d a t i o n o f s u l fit e .
Phytochemistry 12:1463-1466.
 Miller, Paul R., technical coordinator.
I

1980. Proceedings of the symposium on effects of air pollutants on Mediterranean

and temperate forest ecosystems, June 22-27, 1980, Riverside, California, I

U.S.A. Gen. Tech. Rep. PSW-43,256 p. Pacific Southwest Forest and Range Exp.
1 Stn., Forest Sew., U.S. Dep. Agric., Berkeley, Calif. 1

I
I

I These proceedings papers and poster summaries discuss the influence of air pollution on
terrestrial and related aquatic ecosystems. They describe single species-single pollutant
I

relationships; interactions of producers, consumers, and decomposers under pollutant I

I stress; and the use of ecological systems models for interpreting and predicting pollutant

effects.
I
I Retrieval Terms: air pollution injury, acidic precipitation, pollutant stress, terrestrial I

habitats, species-pollutant relationship. mathematical models.

I I

Synthesis Session

Opening Remarks and Summary of

Walter E. westman*

A b s t r a c t : I n t h e opening remarks by W. E. Westman, t h r e e ma-

j o r q u e s t i o n s were r a i s e d : ( 1 ) How can r e s i l i e n c e of ecosys-
tems t o a i r p o l l u t i o n damage be p r e d i c t e d ? Models c o n s t r u c t e d
from key physiognomic, p h y s i o l o g i c a l o r l i f e - h i s t o r y a t t r i b u t e s
of dominants w i t h i n a community c o u l d form t h e b a s i s Of a n a u t -
e c o l o g i c a l approach. Using a s y n e c o l o g i c a l approach, a t l e a s t
f o u r d i s t i n c t components can be recognized ( e l a s t i c i t y , ampli-
t u d e , h y s t e r e s i s , m a l l e a b i l i t y ) which r e f l e c t d i f f e r e n t a s p e c t s
of t h e r e c o v e r y p r o c e s s . These could be measured i n t h e f i e l d
o r d e r i v e d from modeling, u s i n g such community-level a t t r i -
b u t e s a s components of d i v e r s i t y , f o l i a r c o v e r , and s i m i l a r -
i t i e s i n composition. ( 2 ) What i n d i c a t o r s of e f f e c t s of a i r
p o l l u t i o n on ecosystem n u t r i e n t c y c l i n g a r e most r e l i a b l e ?
Observing changes i n t h e m i n e r a l composition of f r e s h l i t t e r -
f a l l d u r i n g peak l i t t e r f a l l p e r i o d s may be p r e f e r a b l e t o mea-
s u r i n g f o l i a r n u t r i e n t c o n c e n t r a t i o n s i n r e l a t i v e l y s m a l l sam-
p l e s of f o r e s t s p e c i e s . ( 3 ) What i s t h e r o l e of c u l t u r a l v a l u e s
i n a i r p o l l u t i o n r e s e a r c h ? What a r e t h e s o c i a l r e s p o n s i b i l i t i e s
of s c i e n t i s t s , and how c a n t h e y be d i s c h a r g e d ? Examples were
g i v e n of t h e r o l e of c u l t u r a l v a l u e s and p e r c e p t i o n s i n t h e con-
d u c t of a i r p o l l u t i o n r e s e a r c h and i n t e r p r e t a t i o n of i t s r e -
s u l t s . S c i e n t i s t s may a s s i s t decision-makers i n i n t e r p r e t i n g
t h e s i g n i f i c a n c e of r e s u l t s of a i r p o l l u t i o n e f f e c t s on f o r e s t s
by i l l u s t r a t i n g t h e e x t e r n a l c o s t s g e n e r a t e d i n t h e economy by
l o s s of ecosystem f u n c t i o n s , a s w e l l a s s t r u c t u r e , due t o a i r
p o l l u t i o n damage. I n t h e open d i s c u s s i o n t h a t f o l l o w e d , p a r t i -
c i p a n t s d i s c u s s e d t h e u s e of a i r p o l l u t i o n s i m u l a t i o n models
i n making d e c i s i o n s a b o u t land use. Modelers i n d i c a t e d t h a t r e -
p a r a m e t e r i z a t i o n of e x i s t i n g v e g e t a t i o n models t o l o c a l c o n d i -
t i o n s c o u l d p r o v i d e a n e f f i c i e n t means of a p p l y i n g e x i s t i n g mo-
d e l s t o l o c a l s i t i n g d e c i s i o n s . F i e l d b i o l o g i s t s and managers
e x p r e s s e d some r e s e r v a t i o n s a b o u t t h e l e v e l of p r e c i s i o n t o be
achieved from such a procedure. Some a t t r i b u t e s of t h e f o r e s t
ecosystem which a r e most i n d i c a t i v e of p o l l u t i o n s t r e s s , and
hence most u s e f u l l y i n c o r p o r a t e d i n such models, were d e t a i l e d ,
i n c l u d i n g physiognomic a t t r i b u t e s , v i s i b l e f o l i a r i n j u r y symp-
toms, wood growth r a t e s and l i c h e n composition. A t l e a s t n i n e
a r e a s i n which r e s e a r c h i s needed on t h e e f f e c t s of a i r p o l -
l u t a n t s on f o r e s t s were s u g g e s t e d .

p r e s e n t e d a t t h e Symposium on E f f e c t s of A i r OPENING REMARKS

P o l l u t a n t s on M e d i t e r r a n e a n and Temperate F o r e s t
Ecosystems, J u n e 22-27, 1980, R i v e r s i d e ,
C a l i f o r n i a , U.S.A.
^ ~ s s o c i a t eP r o f e s s o r of Geography, U n i v e r s i t y
of C a l i f o r n i a , Los Angeles, C a l i f o r n i a
The purpose of t h i s f i n a l d i s c u s s i o n s e s s i o n
i s t o i d e n t i f y ecosystem-level c o n c e p t s p e r t i -
n e n t t o t h e s t u d y of a i r p o l l u t a n t e f f e c t s of
f o r e s t s , and t o p r o v i d e a n o p p o r t u n i t y f o r d i s -
c u s s i o n between p a r t i c i p a n t s a t t h e Symposium
r e g a r d i n g key q u e s t i o n s a r i s i n g from t h e Sympo-
sium proceedings. To a s s i s t i n t h i s e f f o r t ,
Symposium s e s s i o n chairmen ( P a t r i c k Coyne, J o e
McBride, Samuel McLaughlin, J r . , Joseph Shinn,
William H. Smith, and David Tingey) w i l l s e r v e
a s a panel t o f i e l d d i s c u s s i o n q u e s t i o n s from
t h e audience. I have been asked t o open t h e
s e s s i o n w i t h some "provocative" q u e s t i o n s r e -
g a r d i n g ecosystem-level s t u d i e s of a i r p o l l u t i o n
e f f e c t s , a r i s i n g from t h e Symposium p r e s e n t a -
t i o n s and f i e l d t r i p s of t h e l a s t f o u r days. I
cannot g u a r a n t e e t h a t a n y t h i n g more t h a n my
g a r i s h r e d t i e w i l l be p r o v o c a t i v e , but I would
l i k e t o pose t h r e e q u e s t i o n s f o r your c o n s i d e r -
ation.
HOW CAN RESILIENCE O F ECOSYSTEMS TO A I R
POLLUTION DAMAGE BE PREDICTED?
The term " r e s i l i e n c e " h a s been used t o r e f e r
t o t h e degree, manner and pace of r e s t o r a t i o n
of i n i t i a l s t r u c t u r e and f u n c t i o n i n a n eco-
system a f t e r d i s t u r b a n c e (Westman, 1978). Most
of t h e papers i n t h i s Symposium have d i s c u s s e d
n o t r e s i l i e n c e , b u t what h a s been termed eco-
system " i n e r t i a " (Orians, 1975; Westman, 1978).
' I n e r t i a " i s t h e a b i l i t y of a n ecosystem t o r e -
s i s t displacement i n s t r u c t u r e o r f u n c t i o n when
s u b j e c t e d t o a d i s t u r b i n g f o r c e . I n t h e c a s e of
a i r p o l l u t i o n , i n e r t i a can be measured by d e t e r -
mining t h e minimum c o n c e n t r a t i o n of a p o l l u t a n t
a t which impact t o a n ecosystem o c c u r s . We have
c o n s i d e r a b l e information on t h e l e v e l s of c h r o n i c
or acute a i r pollution t h a t w i l l i n i t i a t e in-
jury t o i n d i v i d u a l s p e c i e s , and a l e s s ample
body of evidence r e g a r d i n g t h e l e v e l s of pol-
l u t a n t s n e c e s s a r y t o i n i t i a t e community-level
changes.
Ecosystem models, once e f f e c t i v e l y v a l i d a t e d ,
can p o t e n t i a l l y s e r v e a s t o o l s f o r t h e p r e d i c -
t i o n of r e s i l i e n c e . Since it is impractical t o
model a l l f e a t u r e s of a n ecosystem, however, mod-
e l e r s need t o know which a t t r i b u t e s of a s p e c i e s
make t h e organism most v u l n e r a b l e t o p o l l u t i o n
i n j u r y . Thus, t o be of maximum a s s i s t a n c e t o
modelers, e x p e r i m e n t a l i s t s need t o determine
which physiognomic a n d / o r p h y s i o l o g i c a l a t t r i -
b u t e s of a s p e c i e s ( o r s p e c i e s a s s o c i a t i o n ) a r e
most u s e f u l i n d i c a t o r s of t h e i n e r t i a and r e -
s i l i e n c e of s p e c i e s . Noble and S l a t y e r (1976)
and Grime (1979) have made p r o g r e s s r e c e n t l y
i n i d e n t i f y i n g a t t r i b u t e s of s p e c i e s which may
be u s e f u l i n p r e d i c t i n g t h e i r a b i l i t y t o r e -
c o l o n i z e burned o r c l e a r e d s i t e s .
We might be a b l e t o speed t h e b u i l d i n g of
e f f i c i e n t models of a i r p o l l u t i o n e f f e c t s on
ecosystems by examining s p e c i e s f o r t h o s e a t -
t r i b u t e s which a r e most v u l n e r a b l e t o p o l l u t i o n
damage, o r which enhance recovery f o l l o w i n g da-
mage. During t h i s symposium, we heard t h a t li-
chens and mosses tend t o be more v u l n e r a b l e
t o a i r p o l l u t a n t s because of t h e i r l a c k of waxy
c u t i c l e s . Beyond t h i s , it would be u s e f u l t o
know more a b o u t t h e r e l a t i v e e f f e c t s of such
l e a f a t t r i b u t e s a s mesophylly, s c l e r o p h y l l y
and succulence i n p r o v i d i n g r e s i s t a n c e t o ab-
s o r p t i o n of p o l l u t a n t s . What i s t h e p r e d i c t i v e
v a l u e of phenological a t t r i b u t e s , such a s ever-
greenness vs. deciduousness, o r l i f e - c y c l e
a t t r i b u t e s such a s annual vs. p e r e n n i a l r e p r o -
d u c t i v e c y c l e s ? What a r e t h e e f f e c t s of crown-
s p r o u t i n g a b i l i t y , o r l a c k t h e r e o f , on eco-
system recovery following a i r p o l l u t i o n damage?
These q u e s t i o n s form p a r t of what might be
c a l l e d the"autecologica1 approach" t o eco-
system r e s i l i e n c e , s i n c e they f o c u s on s p e c i e s
a t t r i b u t e s which, when i n c o r p o r a t e d i n a n eco-
system model, can be used t o s y n t h e s i z e key
p r o p e r t i e s of ecosystems.
It i s p o s s i b l e , however, t h a t p r o g r e s s may
be made more q u i c k l y by a t t e m p t i n g g e n e r a l i z a -
t i o n s of ecosystem r e s i l i e n c e by s t u d y i n g com-
munity-level processes of recovery i n p a r t i c -
u l a r biomes. I n o r d e r t o o r g a n i z e a s t u d y of
community-level response t o d i s t u r b a n c e it i s
u s e f u l t o recognize a t l e a s t f o u r d i s t i n c t
components of r e s i l i e n c e (Westman, 1978) :
The r a p i d i t y of r e s t o r a t i o n of a s t a b l e s t a t e
following d i s t u r b a n c e . To u s e t h e analogy of
a m e t a l c o i l , e l a s t i c i t y of t h e c o i l i s t h e
time r e q u i r e d t o s p r i n g back t o i n i t i a l s i z e
a f t e r stretching a c e r t a i n distance.
Amplitude
The zone from which t h e ecosystem w i l l r e -
t u r n t o a s t a b l e s t a t e . By analogy, amplitude
i s t h e d i s t a n c e beyond which a c o i l cannot be
s t r e t c h e d w i t h o u t being permanently deformed.
Hysteresis
The degree t o which t h e p a t h of r e s t o r a -
t i o n ( s u c c e s s i o n ) i s a n e x a c t r e v e r s a l of t h e
p a t h of d e g r a d a t i o n ( r e t r o g r e s s i o n ) . By ana-
logy, h y s t e r e s i s i s t h e degree t o which t h e
r e g i o n t e m p o r a r i l y occupied by a c o i l i n
s p r i n g i n g back d i f f e r s from t h e r e g i o n through
which t h e c o i l moved when i n i t i a l l y s t r e t c h e d .
Malleability
The d e g r e e t o which t h e s t a b l e s t a t e e s t a b -
l i s h e d a f t e r d i s t u r b a n c e d i f f e r s from t h e
o r i g i n a l steady s t a t e . Similarly, malleabil-
i t y i s t h e degree t o which a s t r e t c h e d c o i l
remains s t r e t c h e d a f t e r t h e deforming f o r c e i s
removed.
These components of r e s i l i e n c e a r e s u b j e c t
t o measurement (Westman, 1978). I n t h e c a s e of
t h e s t u d y of o x i d a n t e f f e c t s on p i n e s i n t h e
San Bernardino Mountains, i n e r t i a was d e t e r -
mined by observing t h e l e v e l s of o x i d a n t s a t
which damage t o p i n e s f i r s t appeared. Amp-
l i t u d e might be e s t a b l i s h e d by i d e n t i f y i n g t h e
sample p l o t s , among t h e s e v e r a l used, a t which
p i n e s a r e no longer r e p l a c i n g themselves ( i f
t h e s e a r e taken t o be climax s p e c i e s f o r t h e
r e g i o n ) , and determining t h e lowest mean and
peak concentrations of oxidants a t which t h i s
threshhold e f f e c t i s observed. E l a s t i c i t y could
be measured a s t h e time necessary f o r recovery
of such a s i t e once p o l l u t a n t s t r e s s i s r e -
moved. M a l l e a b i l i t y could be measured, using
a percentage s i m i l a r i t y index, by comparing t h e
community composition of t h e new steady s t a t e
which was e s t a b l i s h e d following p o l l u t i o n
s t r e s s t o t h e p r e - s t r e s s composition. I n ex-
amining h y s t e r e s i s , one would a s k whether t h e
f i r s t s p e c i e s t o disappear from t h e ecosystem
were t h e l a s t t o r e t u r n , using, f o r example, a
rank c o r r e l a t i o n c o e f f i c i e n t .
It i s obvious t h a t i n many s i t u a t i o n s r e -
covery cannot be observed because t h e chronic
s t r e s s continues ( a s i s t h e c a s e i n t h e San
Bernardino Mountains). F u r t h e r , t h e p o s t - s t r e s s
recovery period may be on t h e order of centur-
i e s , i n which c a s e ecosystem models using a u t e c -
o l o g i c a l approaches must be r e l i e d upon f o r
quicker p r e d i c t i o n s . Nevertheless, t h e r e a r e
s i t u a t i o n s i n which a p o l l u t i n g source has been
removed o r reduced, and recovery can be observ-
ed. I n t h e s e s i t u a t i o n s , compilation of t h e
components of r e s i l i e n c e f o r a p a r t i c u l a r p l a n t
formation may a i d us i n g e n e r a l i z i n g about t h e
impact of new p o l l u t i o n sources on a s y e t un-
impacted a r e a s of v e g e t a t i o n of s i m i l a r type.
Thus i n considering t h e p r e d i c t i o n of eco-
system r e s i l i e n c e , we may be wise t o focus both
on s e n s i t i v e a t t r i b u t e s of i n d i v i d u a l s p e c i e s
o r s p e c i e s a s s o c i a t i o n s , and of community-level
changes i n s p e c i e s r i c h n e s s , -composition, fo-
l i a r cover, e t c . which can form t h e bases f o r
observing community-level components of r e s i l -
ience.
WHAT INDICATORS OF EFFECTS OF A I R POLLUTION ON
ECOSYSTEM NUTRIENT CYCLING ARE MOST RELIABLE?
The n u t r i e n t budget of an e n t i r e ecosystem
r e p r e s e n t s one ecosystem l e v e l a t t r i b u t e t h a t
can r e v e a l much about growth-potential and func-
t i o n i n g a t t h e supraorganismal l e v e l . Deter-
mining t h e n u t r i e n t budget f o r even a small por-
t i o n of t h e landscape, however, i s a very c o s t -
l y and time-consuming process. Hence some
speakers (Bruce Wiersma and K. W. Brown, Allen
Legge) described a t t e m p t s t o determine e f f e c t s
of a i r p o l l u t a n t s on n u t r i e n t c y c l i n g by mea-
surement of c o n c e n t r a t i o n s of mineral elements
i n f o l i a g e , a s p o s s i b l e b i o i n d i c a t o r s of pol-
lution-induced ecosystem-level damage. Paul
Zinke described some of t h e many ecosystem com-
partments which must be considered i n concep-
t u a l i z i n g ecosystem n u t r i e n t c y c l i n g .
I would l i k e t o i s s u e a c a u t i o n a g a i n s t t h e
use of f o l i a r n u t r i e n t c o n c e n t r a t i o n s , i n t h e
absence of considerable c o n t e x t , f o r t h e study
of p o l l u t i o n s t r e s s . S o i l s c i e n t i s t s have long
recognized t h a t f o l i a r n u t r i e n t concentrations
a r e dependent on s o i l n u t r i e n t concentrations,
and have used t h e f o l i a g e a n a l y s e s a s i n d i c a t o r s
of "available" concentrations of t h e elements
i n t h e s o i l i n a number of instances. Thus i t
becomes very important f o r t h e a i r p o l l u t i o n
researcher t o characterize s o i l heterogeneities
i n h i s o r h e r study. Secondly, e c o l o g i s t s know
t h a t f o l i a r n u t r i e n t concentrations vary temp-
o r a l l y a s t h e l e a f passes from e a r l y growth
s t a g e s t o senescence and l e a f f a l l . The changes
a r e due t o t h e change i n t i s s u e and c e l l u l a r
component r a t i o s with age, t o t h e changing r a -
t i o of photosynthate t o mineral elements, and
t o withdrawal of more mobile n u t r i e n t s i n t o
stems before l e a f f a l l . Thus t h e h a r v e s t of
leaves a t d i f f e r e n t times of year makes nu-
t r i e n t analyses of these leaves i n a p p r o p r i a t e
f o r use a s samples from a s i n g l e population.
Furthermore, species d i f f e r i n t h e i r a b i l i t i e s
t o a s s i m i l a t e , r e t a i n o r accumulate t o luxury
l e v e l s , p a r t i c u l a r mineral elements. The a b i l -
i t y of some s p e c i e s t o accumulate c e r t a i n heavy
metals, f o r example, i s t h e b a s i s f o r biogeo-
chemical prospecting. Luxury accumulation of
potassium by many s p e c i e s i s w e l l known. Cal-
cium, being immobile, tends t o i n c r e a s e i n con-
c e n t r a t i o n i n leaves with age, but t h e i n i t i a l
a b i l i t y t o a s s i m i l a t e calcium d i f f e r s from spe-
c i e s t o s p e c i e s . Thus t h e a n a l y s i s of f o l i a r
n u t r i e n t concentrations without regard t o s p e c i e s
o r ecotype i s t o be avoided.
The m u l t i p l e axes of v a r i a t i o n presented by.
d i f f e r e n c e s i n s p e c i e s , mineral p r o p e r t i e s , time
of year and s o i l concentrations implies t h a t a
much more massive sampling program must be un-
dertaken t o observe meaningful t r e n d s from f o l i a r
analyses than has c h a r a c t e r i z e d some of t h e a i r
p o l l u t i o n s t u d i e s reported.
Short of a f u l l n u t r i e n t budget a n a l y s i s ,
changes i n t h e mineral composition of f r e s h
l i t t e r f a l l during peak l i t t e r f a l l periods may
provide a more s u i t a b l e i n d i c a t o r of n u t r i e n t
changes due t o p o l l u t i o n s t r e s s , a s t h i s com-
ponent is standardized i n time, and weighted t o
t h e f o l i a r biomass composition of t h e f o r e s t .
Even s o , l a r g e sample-sizes a r e needed, and
much c a u t i o n i n e x t r a p o l a t i o n of r e s u l t s w i l l
s t i l l be necessary.
WHAT IS THE ROLE OF CULTURAL VALUES I N
A I R POLLUTION RESEARCH?
WHAT ARE THE SOCIAL RESPONSIBILITIES OF
SCIENTISTS, AND HOW CAM THEY BE DISCHARGED?
As n a t u r a l s c i e n t i s t s , we tend t o r e l e g a t e
t h e s o c i a l context of our research t o o t h e r
segments of society. We do s o a t r i s k , however,
because t h e r e a r e both s o c i a l i s s u e s upon which
we a r e most q u a l i f i e d t o comment, and socio-
p o l i t i c a l f o r c e s which a f f e c t t h e choice and
conduct of our r e s e a r c h problems. I n t h e course
of t h e present Symposium, I was amused t o note
t h e v a r i a t i o n i n perception of t h e a i r q u a l i t y
i n Riverside during t h e period, by v a r i o u s p a r t i -
c i p a n t s . Some thought t h e smog l i g h t , o t h e r s
oppressively heavy. The smog c o n c e n t r a t i o n was
a constant; c u l t u r a l values were a t work i n
influencing perceptions. As a second example,
some speakers emphasized t h e r o l e of a c i d r a i n
a s a f e r t i l i z e r , w h i l e o t h e r s emphasized i t s
toxic properties. Of c o u r s e , whether any pol-
lutant w i l l exercise its toxic properties w i l l
depend upon r a t e and d u r a t i o n of a p p l i c a t i o n ,
and c o n c e n t r a t i o n , a s w e l l a s upon t h e physio-
l o g i c a l s t a t e of t h e r e c e p t o r organism and i t s
ecosystem. I n t h e c a s e of c h a r a c t e r i z i n g t h e
p r o p e r t i e s of a c i d r a i n , however, t h e a t t r i b u t e s
chosen f o r emphasis were chosen f o r r e a s o n s
having t o do w i t h t h e c u l t u r a l a t t i t u d e s and
v a l u e s of t h e s p e a k e r s , and n o t because of d i s -
agreement over e m p i r i c a l o b s e r v a t i o n s . A s a
t h i r d example, we heard from one F o r e s t S e r v i c e
representative during the f i e l d t r i p t h a t the
damaged p o r t i o n s of t h e San Bernardino National
F o r e s t were n o t e x p e r i e n c i n g d i f f i c u l t y i n r e -
p r o d u c t i o n . But, t h e E.P.A. sponsored, Univ-
e r s i t y of C a l i f o r n i a - F o r e s t S e r v i c e r e s e a r c h
team provided evidence t o t h e c o n t r a r y . Were
t h e s e two p a r t i e s d i s a g r e e i n g over e m p i r i c a l
o b s e r v a t i o n s , o r was t h e i r d i f f e r e n c e one of
c u l t u r a l v a l u e s and p e r c e p t i o n s a p p l i e d i n e v a l -
u a t i n g t h e s i g n i f i c a n c e of t h e same body of
d a t a ? A s a f o u r t h example, r e c a l l t h a t about
50% of t h e a d u l t American p o p u l a t i o n smokes,
thereby b r i n g i n g i n t o t h e i r lungs s e v e r a l times
t h e ambient l e v e l s of p a r t i c u l a t e s and t o x i c
gases p r e s e n t i n p o l l u t e d urban a i r . W i l l
t h e s e people p e r c e i v e t h e human h e a l t h hazards
of outdoor a i r i n t h e same way a s nonsmokers?
W i l l they a s s i g n t h e same weights t o t h e i r im-
portance?
A t t h e very l e a s t , s c i e n t i s t s have a r e -
s p o n s i b i l i t y t o d i f f e r e n t i a t e c l e a r l y between
e m p i r i c a l o b s e r v a t i o n s and normative ( v a l u e )
judgments. But does our r e s p o n s i b i l i t y t o
decision-makers and t h e p u b l i c s t o p t h e r e ?
Much h a s been s a i d r e g a r d i n g t h e a p p r o p r i a t e -
n e s s of s c i e n t i s t s i n o f f e r i n g v a l u e judgments
t o s o c i e t y , and I w i l l n o t e n t e r t h e d e b a t e
h e r e . I would, however, l i k e t o s u g g e s t a way
i n which s c i e n t i s t s can h e l p t o c l a r i f y i n
s o c i a l l y - m e a n i n g f u l terms t h e s o c i a l c o s t s .of
a i r p o l l u t i o n damage.
I b e l i e v e decision-makers would prof it from
a f u l l e r understanding of t h e e f f e c t s of a i r
p o l l u t i o n n o t only on t h e marketable a s p e c t s
of ecosystem s t r u c t u r e ( s t a n d i n g timber, c r o p
y i e l d , t o u r i s t r e v e n u e ) , b u t a l s o on t h o s e a s -
p e c t s of ecosystem f u n c t i o n i n g which c r e a t e h i d -
den c o s t s i n t h e market p l a c e . To t a k e a n ex-
ample from t h e San Bernardino N a t i o n a l F o r e s t
o x i d a n t s t u d y which I have p r e v i o u s l y d i s c u s s e d
(Westman, 1977), c o n s i d e r t h e d o l l a r c o s t s t o
s o c i e t y of t h e l o s s of t h e s o i l binding f u n c t i o n
from a i r p o l l u t i o n damage t o p i n e s i n t h i s f o r -
e s t . A s of 1972, 57% of t h e t r e e s over a 4000
h e c t a r e a r e a w i l l be r e p l a c e d by a r e t r o g r e s -
s i v e v e g e t a t i o n of f o r b s and g r a s s e s , and t h a t
e r o s i o n l o s s e s from t h e l a t t e r w i l l i n c r e a s e
i n t h e p r o p o r t i o n observed when c h a p a r r a l was
converted t o g r a s s l a n d by t h e U.S. F o r e s t Ser-
v i c e a t San Dimas i n t h e neighboring San G a b r i e l
Mountains, it is p o s s i b l e t o e s t i m a t e e r o s i o n
l o s s e s from t h e s t r e s s e d f o r e s t . A t c u r r e n t
r a t e s of cleanup of sediment from s t r e e t s , sew-
e r s and d e b r i s b a s i n s , t h e c o s t of damage from
l o s s of t h e s o i l binding f u n c t i o n of t h e San
Bernardino Mountain p i n e s is $27 m i l l i o n p e r
y e a r . These c o s t s a r e being absorbed c u r r e n t l y
by t h e g e n e r a l p u b l i c n o t only a s d i r e c t t a x
c o l l e c t i o n t o l o c a l governments c l e a n i n g up
t h e sediment, a s only a p o r t i o n of t h e sediment
i s being recovered i n t h i s way. The c o s t s a r e
being absorbed a l s o i n terms of l o s s e s t o f i s h -
e r i e s i n c o a s t a l w a t e r s where spawning a r e a s
a r e smothered by sediments, by p u b l i c works
a l l o c a t i o n s f o r new dams, by flood damage f o l -
lowing storms i n s i l t e d f l o o d channels. Rarely
i f ever a r e t h e s e c o s t s a t t r i b u t e d t o smog and
considered i n t h e c o s t - b e n e f i t a n a l y s i s of pro-
p o s a l s t o i n s t a l l emission c o n t r o l d e v i c e s .
F u r t h e r , s o i l binding i s only one of t h e func-
t i o n s destroyed through d e a t h of t h e p i n e s . Loss
of t h e f u n c t i o n s of n u t r i e n t c a p t u r e and r e -
tention, pollution absorption, c l i m a t i c reg-
u l a t i o n and energy f i x a t i o n a l l have t h e i r so-
c i a l c o s t s , c a p a b l e of a t l e a s t p a r t i a l enumer-
a t i o n and e v a l u a t i o n (Westman, 1978).
Complex a s t h i s process of s o c i a l c o s t i d e n t i -
f i c a t i o n is, I suggest t h a t i t i s a t o p i c i n
which s c i e n t i s t s can p l a y a l a r g e r r o l e t h a n we
have t o d a t e , and a t o p i c t o which, i t may be
argued, we have a r e s p o n s i b i l i t y t o contribute-.
SUMMARY OF AUDIENCE PANEL DISCUSSION
Use of a i r p o l l u t i o n s i m u l a t i o n models
i n d e c i s ion-making
Discussion ensued on t h e c u r r e n t a p p l i c a b i l -
i t y of computer models t o such immediate ques-
t i o n s a s how t o s i t e power p l a n t s t o minimize
damage t o v e g e t a t i o n . Those w i t h experience
i n b u i l d i n g models expressed c o n s i d e r a b l e con-
f i d e n c e t h a t t h e s e could be used, upon r e p a r a -
m e t e r i z a t i o n , t o h e l p r e s o l v e such q u e s t i o n s
i n a number of p a r t s o f t h e country. Experi-
mental b i o l o g i s t s and land managers expressed
concern t h a t t h e models were s t i l l t o o broadly
conceived t o provide a c c u r a t e e s t i m a t e s of d i f -
f e r e n c e s between s i t e s i n a s i n g l e v e g e t a t i o n
type. A number of t h e s p e c i f i c i s s u e s t h a t were
broached a s p a r t of t h i s d i s c u s s i o n a r e a s f o l -
lows :
T r a n s f e r a b i l i t y of models between v e g e t a t i o n
types
What l e v e l of r e s o l u t i o n of a model i s needed
i n o r d e r t o provide a c c u r a t e p r e d i c t i o n s of
p o l l u t i o n e f f e c t s on v e g e t a t i o n , of u s e t o
land managers? Are s i n g l e g e n e r a l models f o r
each biome s u f f i c i e n t , o r do we need a model f o r
each of 400 o r 500 American v e g e t a t i o n t y p e s ?
How should a model be c o n s t r u c t e d t o maximize
i t s t r a n s f e r a b i l i t y between r e g i o n s ? S e v e r a l
suggested responses were o f f e r e d :
-1. The work of e c o l o g i s t s , a i r p o l l u t i o n
r e s e a r c h s c i e n t i s t s and modelers could be
c o o r d i n a t e d i n a nationwide r e s e a r c h l a b o r a t o r y
consortium t o c o n s t r u c t working models of a i r
p o l l u t i o n e f f e c t s on v e g e t a t i o n f o r t h e major
r e g i o n a l p l a n t formations. Land managers con-
cerned w i t h p a r t i c u l a r s i t i n g o r f o r e s t r y ques-
t i o n s i n a r e g i o n of t h e c o u n t r y could submit
a r e q u e s t t o t h e consortium f o r a d a p t a t i o n of a
r e g i o n a l model t o h i s o r h e r p a r t i c u l a r prob-
lem. One advantage of t h i s approach i s t h a t
b i o l o g i s t s and modelers who a r e most f a m i l i a r
w i t h t h e assumptions i n t h e model would be
a v a i l a b l e t o o p e r a t e t h e model. R e p e t i t i o u s
r e s e a r c h e f f o r t s could be avoided, and exper-
i e n c e accumulated from v a r i o u s r e g i o n a l e f f o r t s .
2 . Models which i n c o r p o r a t e t h e m e c h a n i s t i c
b a s i s f o r a i r p o l l u t i o n e f f e c t s on v e g e t a t i o n
may be most e a s i l y adapted t o new v e g e t a t i o n
t y p e s w i t h accuracy. "Mechanistic" bases may
c o n s i s t of p h y s i o l o g i c a l models of t h e e f f e c t s
of p o l l u t a n t s , temperature and m o i s t u r e on
n u t r i e n t a s s i m i l a t i o n and p h o t o s y n t h e s i s , o r
t h e y may c o n s i s t of ecosystem-level models i n
which s e n s i t i v e a t t r i b u t e s of t h e ecosystem
( l i t t e r f a l l , evergreenness vs. deciduousness,
s p e c i e s l o n g e v i t y ) a r e modeled.
3 . The N a t i o n a l Power P l a n t Team of t h e United
S t a t e s F i s h and W i l d l i f e S e r v i c e h a s a v a i l a b l e
a power p l a n t s i t i n g model which, although con-
s i d e r i n g v a r i o u s c o n s t r a i n t s , does n o t i n c o r -
p o r a t e a model of a i r p o l l u t i o n e f f e c t s on vege-
t a t i o n i n d e t a i l . They have r e c e n t l y o b t a i n e d ,
however, a copy of Kercher's (Lawrence Liver-
more Laboratory) f o r e s t growth model, and p l a n
t o make t h i s a v a i l a b l e f o r p u b l i c use, and per-
haps u l t i m a t e l y l i n k i t t o t h e i r e x i s t i n g s i t i n g
model.
4. A f o r e s t growth model f o r a p a r t i c u l a r ve-
g e t a t i o n t y p e a p p e a r s t o r e q u i r e 1 - 2 person-
y e a r s of e f f o r t t o c o n s t r u c t from s c r a t c h . Re-
p a r a m e t e r i z a t i o n of a n e x i s t i n g model i s r e -
garded a s a l e s s c o s t l y and time-consuming way
t o provide a model f o r a new v e g e t a t i o n type.
An important l i m i t i n g f a c t o r t o t h i s e f f o r t
i s t h e p a u c i t y of f i e l d and l a b o r a t o r y fumi-
g a t i o n d a t a on e f f e c t s of a i r p o l l u t a n t s , a l o n e
and i n combination, on s p e c i e s , e s p e c i a l l y over
extended p e r i o d s . I n t h e absence of such f i e l d
d a t a , s e n s i t i v i t y a n a l y s i s of t h e computer
model may be used t o e s t a b l i s h t h e l i k e l y com-
ponents of t h e ecosystem which w i l l be most
a d v e r s e l y a f f e c t e d by a p a r t i c u l a r s t r e s s , and
t o produce q u a l i t a t i v e s c e n a r i o s of worst c a s e
events.
5. E x i s t i n g models f o r many v e g e t a t i o n t y p e s
do n o t i n c o r p o r a t e long-term c y c l i c a l e v e n t s
such a s f i r e s , seed c y c l e s , e t c . However, mo-
d e l s such a s t h o s e of Kercher do i n c o r p o r a t e
t h e s e e f f e c t s . The f o r e s t growth model a v a i l -
a b l e through Oak Ridge N a t i o n a l Laboratory i s
c o n s i d e r e d c a p a b l e of a p p l i c a t i o n t o a l l e a s t e r n
U. S. f o r e s t t y p e s except Southern p i n e f o r e s t s .
Ecosystem-level i n d i c a t o r s of p o l l u t i o n s t r e s s
What a t t r i b u t e s of s p e c i e s o r of ecosystems
can be used by f i e l d e c o l o g i s t s and modelers
a s s e n s i t i v e i n d i c a t o r s of l i k e l y p o l l u t i o n -
induced changes?
1. Physiognomic a t t r i b u t e s ( l e a f c u t i c l e t h i c k -
n e s s and c h e m i s t r y ) , phenologic a t t r i b u t e s
(age t o r e p r o d u c t i o n , d u r a t i o n of f o l i a g e ) ,
and l i f e h i s r o r y a t t r i b u t e s (annual vs. p e r -
e n n i a l n a t u r e , l o n g e v i t y of g e n e r a t i o n s ) a r e
s e e n a s community-level i n d i c a t o r s . L i t t e r -
f a l l may be a u s e f u l i n d i c a t o r of n u t r i e n t
budget processes.
2. Growth r i n g s (dendrochronology) may be
used a s a long-term r e c o r d of f o r e s t growth
responses. I n Pennsylvania, f o r example, t r e e
r i n g a n a l y s i s i n f o r e s t i n t h e v i c i n i t y of pow-
e r p l a n t s has been conducted.
3 . V i s i b l e i n j u r y symptoms, such a s t h e f o l i a r
i n j u r y index used by P a u l M i l l e r and co-work-
e r s , may be u s e f u l . The Tennessee Valley Auth-
o r i t y h a s recorded v i s u a l i n j u r y symptoms on
t r e e s surrounding a number of i t s power p l a n t s .
4. The use of p a r t i c u l a r l y s e n s i t i v e s p e c i e s ,
such a s l i c h e n s , was i l l u s t r a t e d e a r l i e r i n
t h e Symposium.
Use of models i n s e t t i n g a i r p o l l u t i o n s t a n d a r d s
Can a i r p o l l u t i o n models be used by agen-
c i e s concerned w i t h e s t a b l i s h i n g minimum con-
c e n t r a t i o n s of ambient exposure t h a t w i l l c a u s e
damage t o v e g e t a t i o n ?
1. To t h e e x t e n t t h a t s t a n d a r d s i n c o r p o r a t e
s o c i a l values a s well a s s c i e n t i f i c c r i t e r i a ,
a computer model of t h e type being d i s c u s s e d
cannot be used t o s e t a s t a n d a r d .
2. On t h e o t h e r hand, computer models have
been used t o determine l e v e l s of f o r e s t growth
r e d u c t i o n from p a r t i c u l a r l e v e l s of p o l l u t a n t s .
This information a l o n e , o r converted t o econom-
i c l o s s f i g u r e s , can be u s e f u l a s i n f o r m a t i o n
f o r c r i t e r i a documents used i n s t a n d a r d s e t t i n g .
3 . Models have been used t o c a l c u l a t e r a -
d i a t i o n dose t o people i n t h e v i c i n i t y of
n u c l e a r power p l a n t s . The I n d i a n P o i n t power
p l a n t was modified t o reduce damage of e f f l u -
e n t t o f i s h l i f e , based on information o b t a i n -
ed through a computer model.
4. Models of f o r e s t growth have i l l u s t r a t e d
t h a t even a 5-10% decrement i n t r e e growth
due t o a i r p o l l u t i o n can have s e v e r e long-
term e f f e c t s on f o r e s t growth and composition.
This is a p r e d i c t i o n t h a t would have been d i f -
f i c u l t t o make w i t h p r e c i s i o n i n t h e absence
of a computer model.
A Research Agenda
During t h e course of t h e s y n t h e s i s s e s s i o n
s e v e r a l suggestions were made regarding a r e a s
i n which important information i s p a r t i c u l a r l y
lacking. These r e s e a r c h t o p i c s a r e l i s t e d be-
low.
P a t t e r n s of Recovery i n Stressed Vegetation--
While many ecosystem s t u d i e s have documented
t h e ' i n e r t i a ' of t h e systems, fewer have ob-
served t h e recovery process. The emission
source a t T r a i l , B r i t i s h Columbia, i s an ex-
ample of a p o i n t source whose emissions de-
creased d r a m a t i c a l l y i n t h e l a t e 1930's.
S t u d i e s of subsequent recovery of damaged ve-
g e t a t i o n could t e l l us much about r e s i l i e n c e
of t h a t f o r e s t ecosystem.
P r e c i p i t a t i o n Chemistry--Much needs t o be
known about t h e chemical transformations i n
r a i n and snow a s they pass over t h e s u r f a c e s
of v e g e t a t i o n and s o i l , and how t h e s e i n t u r n
a f f e c t t h e f l u x of n u t r i e n t s i n t h e ecosystem.
Mechanistic Models For P a r t i c u l a t e ( i n -
cluding heavy m e t a l ) Pollutants--Models f o r
t h e i n j u r y t o p l a n t s from t o x i c gases a r e more
advanced than those f o r p a r t i c u l a t e s . How
do p a r t i c u l a t e s e n t e r p l a n t s and a t what r a t e s ?
How a r e t h e i r e f f e c t s r e g i s t e r e d ? How can
they be modeled?
Synergisms--Much i s unknown about t h e i n -
t e r a c t i v e e f f e c t s of s e v e r a l a i r p o l l u t a n t s
on i n d i v i d u a l s p e c i e s and on ecosystems. Fu-
migation s t u d i e s should i n c o r p o r a t e p o l l u t a n t
combinations a s w e l l a s t e s t the effects
of s i n g l e p o l l u t a n t s .
growth. How do t h e s e e f f e c t s i n t e r a c t with
ambient a i r p o l l u t i o n l e v e l s ? How w i l l t h e
a f f e c t of C02 on g l o b a l c l i m a t e f u r t h e r a f -
f e c t p o l l u t i o n response of f o r e s t s ?
Funding of Research on F o r e s t Growth Models
S p e c i f i c a l l y B u i l t t o Incorporate A i r Pol-
l u t i o n Effects--Most e x i s t i n g models sim-
u l a t e f o r e s t growth i n t h e absence of pollu-
t i o n e f f e c t s , and were funded by agencies o t h e r
than those concerned with a i r p o l l u t i o n .
REFERENCES
Grime, J. P.
1979. P l a n t s t r a t e g i e s and vegetation pro-
c e s s e s . 222 p. John Wiley and Sons,
New York.
Noble, I. R., and R. 0. S l a t y e r .
1976. P o s t - f i r e succession of p l a n t s i n
Mediterranean ecosystems. 2 Proc. Symp.
Environmental Consequences of F i r e and
Fuel Management i n Mediterranean Ecosys-
tems. H. A. Mooney and C. E. Conrad, eds
p. 27-36. F o r e s t Service, U. S. Dept.
Agric., Washington, D. C.
Orians, G. H.
1975. D i v e r s i t y , s t a b i l i t y and m a t u r i t y i n
n a t u r a l ecosystems. & I Unifying Concepts
i n Ecology. W. H. van Dobben and R. H.
Lowe-McConnel, eds. p. 64-65. Junk,
The Hague.
Westman, W. E.
1977. How much a r e Nature's s e r v i c e s worth?
Science 197: 960-964.

Long-term E f f e c t s of P o l l u t a n t Exposure-- Westman, W. E.

There i s a need f o r l a b o r a t o r y s t u d i e s of 1978. Measuring t h e i n e r t i a and r e s i l i e n c e
long-term exposures ( g r e a t e r than one month) of ecosystems. BioScience 28: 705-710.
of a i r p o l l u t a n t s t o species. I n t h e f i e l d ,
exposures t o f o r e s t s p e c i e s occur over many
y e a r s . It i s d i f f i c u l t t o understand long-
term e f f e c t s on reproduction and growth i n
t h e absence of chronic exposure s t u d i e s .

Monitoring Data For Ambient P o l l u t a n t Con-

centrations--More of t h e s e d a t a a r e needed
f o r a l l p a r t s of t h e United S t a t e s i f use
of computer models with r e a l i s t i c ambient a i r
c o n c e n t r a t i o n s i s t o be achieved. The same
d a t a a r e needed f o r i n t e r p r e t a t i o n of f i e l d
observations.

I n t e r a c t i o n of A i r P o l l u t i o n S t r e s s e s With
Management Practices--How does t h e e f f e c t of
an a i r p o l l u t a n t on a f o r e s t d i f f e r under d i f -
f e r e n t t h i n n i n g regimes? This question may
be examined i n t h e next phase of r e s e a r c h i n t h e
San Bernardino Mountain pine f o r e s t s .

Carbon Dioxide Enrichment i n t h e Atmosphere--

Rising ambient CO2 l e v e l s a r e a f f e c t i n g f o r e s t
 Integration: a Role for Adaptive
Environmental Assessment and
Management '
L
Nicholas C. Sonntag, Robert R. Everitt, Michael J. Staley

Abstract: Adaptive Environmental Assessment and Management

(AEAM) is a proven methodology for integration and analysis

of environmental research and management. Simulation

modelling workshops are an essential part of the process.

They act as a catalyst to focus analysis, stimulate

discussion and foster communication amongst managers,

planners and scientific disciplinarians.

Some of the philosophy and methodology are described

along with a case study example on the Alberta Oil Sands

Environmental Research Program (AOSERP). The emphasis is

on how AEAM can help foster integration and communication

of scientific information.

Over the last decade we have seen phenomenal

based, and inspite of a desire for interdisciplis-

growth of environmental science. It has spawned

ary coordination most large research programs have

a proliferation of public and private institutions

failed to integrate their results. One reason for

dedicated to environmental research and protection.

this is the lack of a proper forum for communica-

Environmental research and management have become

tion amongst scientific specialists; another is

highly skilled, influential, and more often than

the absence of a systematic framework for synthe-

not, respected professions. Governments have

sizing results. While some programs have been

created such well meaning provisions as the

successful at pulling the individual studies to-

National Environmental Policy Act (NEPA), the Clean

gether few have been able to make the results

Air Act and the Water Quality Act in the United

relevant to environmental management. Many argue

States, and the Environmental Assessment and

that this can be remedied by raising the quality

Review Process (EARP) in Canada.

of environmental research. Unfortunately there is

a dichotomy between environmental research and

These gallant efforts to research and protect

environmental management. Research science cannot

the environment have over the years generated

effectively guide managers because scientists do

voluminous amounts of information; information

not readily comprehend management concerns;

intended ultimately to provide knowledge to help

management cannot adequately direct research

mankind better manage the world's environment and

science since managers often do not have scienti-

resources.
fic knowledge or the breadth of understanding such

knowledge provides. .This inability of scientists

The challenge today is to analyze and make

and managers to interact effectively has been a

sease of the vast amount of data and knowledge.

major stumbling block in developing progressive

Unfortunately most research has been disciplinary

policies and attitudes towards the environment both

in the public and private sector.

presented at the Symposium on Effects of Air

Adaptive Environmental Assessment and Management

Pollutants on Mediterranean and Temperate Forest

(AEAM), well documented in (tolling (19781, has

Ecosystems, June 22-27, 1980, Riverside,

evolved over the last ten years into a proven

California.
methodology for integrating environmental research

while linking research and management. AEAM uses

"ESSA Environmental and Social Systems Analysts

computer simulation modelling in a unique and novel

Ltd., respectively, Vancouver, British Columbia.

way. Within a structured modelling workshop, the

task of constructing a simulation model acts as a

are not of equal importance and judgment is the

catalyst to focus analysis, stimulate discussion

toolof management not exhaustive research.

and foster communication amongst managers, plan-

ners, and scientific disciplinarians.

MODELLING WORKSHOPS: THE CORE OF AEAM

BIASES OF AEAM

A modelling workshop is a 3 to 5 day meeting of

Ecologist's understanding of ecosystem struc-

a group of scientists, planners, and managers in-

ture and behaviour have come from four basic

volved in the design and execution of an environ-

properties which determine how ecological systems

mental study. No papers are presented, there is

respond to change (Holling, 1978).

no keynote speaker, there are simply 3 to 5 days of

focussed activity on the problems at hand. The

1. The parts of an ecological system are con-

development of a computer simulation model of the

nected to each other in a selective way (this has

physical, biological, and social aspects of the

implications for what should be measured).

problem serves as the focus for the workshop.

2. Events are not uniform over space. (This

Participants in the workshop do need any

has implications for how intense impacts will be

knowledge of computers or modelling to contribute

and where they will occur.)

to the workshop and to gain from its results.

Workshop facilitators translate participant input

3. Sharp shifts in behaviour are natural for

into quantitative relationships that can be pro-

many ecosystems. (Traditional methods of monitor-

grammed into the simulation model. The facilita-

ing or assessment can misinterpret these and make

tors can be viewed as information translators for

them seem unexpected or perverse.)

it is the participants who conceptualize the model.

Therefore the model that evolves from a workshop

4. Variability, not constancy, is a feature of

is as much a product of the ideas and concerns of

ecological systems that contributes to their per-

people unfamiliar with modelling as it is a product

sistence and to their self monitoring and self

of those familiar with simulation techniques.'

correcting capabilities.

The obvious objective of a modelling workshop

Underlying each of these properties is the fact

is to build and run a computer simulation model of

that environmental systems are characterized by

the bio-physical system of interest. However, the

overwhelming uncertainty. Man's understanding of

resultant model is not an end in itself. Usually

the underlying biological, social and physical

its predictions are not very precise and it often

processes and interactions is minimal, and will

lacks obvious features of the actual system.

remain so for the forseeable future. The complex

Rather, the model is a focus for communication

and pervasive nature of environmental and social

promoting objectivity and honesty. Building the

issues guarantees our ignorance will always exceed

model forces the participants to formalize their

our knowledge.
understanding of the system components and inter-

actions. This facilitates easier evaluation of the

The many sources of error in environmental sys-

importance of interactions to the system and the

tems ensure that no matter how broad or deep an

workshop objectives. Often favoured factors turn

analysis inevitably something outside will in-

out to be irrelevant for predictions, therefore

fluence the results and violate the predictions.

requiring less future effort both in model develop-

The conclusion is that environmental and social

ment and data-gathering programs.

systems are fundamentally unpredictable. If you

accept this hypothesis then, how can environmental

As with many modelling studies, a workshop gen-

science produce effective research and management.

erated model confers the ability to test hypotheses,

research plans and different management policies

First and foremost we must never promote re-

without risk. However, that is where the similarity

search and analysis as the panacea for prediction

usually ends. Since a workshop model is designed

of the fate of society and its environment.

and built by all the participants its structure

Second, we must recognize that decisions are al-

and resultant dynamics are "transparent" to the

ways made under uncertainty, and ignorance.

user. The model is comprehensible. This inspires

Realizing this we must capitalize on methods that

trust and increases insight thereby promoting

help focus ideas and information, integrate con-

generalization of the model projections. It also

cepts, and guide decision making. Better decisions

facilitates easier evaluation of those factors left

will usually be made with a clear picture of both

out of the model.

knowledge and ignorance, and a broad appreciation

of the consequences of action.

The modelling workshop style prevents the build-

ing of a sophisticated state of the art simulation

The key tool of AEAM is the computer modelling

model. The workshop model is invariably simple in

workshops. These short intense meetings circum-

structure and inefficient in operation. Further by

vent the natural scientific tendency for reduction-

striving to simplify the problem the need for

ism and the eternal cry for more studies. Parti-

sophisticated tools such as complex implicit (or

cipants in these workshops are forced to recognize

explicit) finite difference methods for approximat-

that all components of natural resource systems

ing differential equations are usually avoided.

Although the precision of the model's predictions

a minimum of environmental damage. This policy

may suffer simplification does not deter from the

resulted in particular attention being paid by

objectives of participant involvement, communica-

regulatory agencies to the need for development of

tion and understanding. People are the key

an environmental research program for the Athabasca

components in the modelling workshop not the model.

Oil Sands region. Consequently, late in 1973,

This does not have to be the case after the work-

officials of Alberta Environment (Research Secre-

shop. Often the model structure established serves

tariat) and Environment Canada (Environmental

as an excellent guide for future modelling efforts

Management Service) separately produced internal

when more consideration can be given to the "art"

reports recommending a comprehensive environmental

of simulation modelling.
research program. Projections in each of the

reports favoured an environmental research program

By definition any simulation model is "wrong"

lasting 10 years, with total costs estimated in the

since it must be a simplification of reality.

range of $30 million to $40 million." (Smith

However, complex models do not necessarily make

1979) .
better predictions. As the number of variables

increases, so does the number of assumptions about

The Alberta Oil Sands Environmental Research

how they are related and the chances of making a

Program (AOSERP) began in April, 1975. Early

critically wrong assumption rises rapidly. There-

investigations were used primarily to establish

fore parsimony is the underlying workshop modelling

large data bases. The data bases were intended

theme. Interactions and relations should make sense

to facilitate the construction of models to aid in

when interpreted in terms of physics and biology.

predicting physical, chemical, biological, and

Logical consistency and clarity are stressed in

social impacts of Oil Sands development.

the building of the workshop model and go far in

maintaining its "transparency" to the participants.

In the fall of 1979, AOSERP sponsored an AEAM

modelling workshop. By this time the program had

The incorporation of modelling workshops in the

been organized into four systems: Air, Land,

AEAM process is designed to be iterative. By

Water and Human. The objectives of the workshop

locating them between sequential field programs

were: (1) to construct a simulation model that

the researchers and managers have the opportunity

would provide a mechanism of integration of the

to adapt research plans and management policies

plethora of AOSERP data and information; (2) to

in light of new insights emerging from the modell-

delineate the interrelationship between the sys-

ing workshop exercise. The AEAM process, and

tems (Air, Land, Water, Human) that are basic to

specifically modelling workshops, can be success-

a general understanding; (3) to identify and

fully implemented at any point in a study, right

evaluate data gaps and uncertainties about system

up to the end. The integration and coordination

function; and (4) to evaluate and recommend

aspects of modelling workshops provide a useful

approaches to environmental management in the oil

vehicle for communication among those responsible

sands areas, including mechanisms for technology

for preparing environmental overviews and assess-

transfer from AOSERP to Alberta Environment regu-

ments. This ensures they are pertinent, credible,

latory branches (Staley and others 1979). The

and address the questions being asked. Further

participants in the workshop included the director

the resultant model (after some refinement)

of AOSERP, the heads of the Air, Water, Land and

provides a very effective device for summarizing

Human systems, the chairman of the Research Secre-

and presenting the results of a study to policy

tariat of Alberta Environment, as well as numerous

makers, administrators and/or funding agencies.

government scientists and planners, and private

A workshop held in 1979 on the Alberta Oil Sands,

consultants.

is a good example of such an application.

Simulation Model

ASSESSMENT OF THE ENVIRONMENTAL EFFECTS

OF ALBERTA OIL SANDS DEVELOPMENT

During the bounding exercise, the workshop

decided to consider the entire AOSERP study area

Setting
which comprises approximately 2.86xlo4 square

kilometers of northeastern Alberta, Canada with the

'The development of the Athabasca Deposit, one

spatial resolution based on the area's 13 water

of several oil sands deposits in Alberta, has been

drainage units. To assess impacts over a meaning-

the subject of intense interest for several de-

ful period, a thirty year time horizon with a

cades. The Athabasca Deposit contains more than

yearly time step was used. Important system

600x10" barrels of bitumen reserves, and consti-

phenomena operating on a shorter time scale were

tutes about 88 percent of known oil sands in

represented implicitly within the one year step.

Alberta. Consequently its potential to augment

The simulation model was divided into four inter-

the oil supply of Canada has been a driving force

related submodels: human, physical transport,

in present development, and will continue to gen-

aquatic biology, and terrestrial.

erate pressure for further development . . .

The human submodel consisted of three components:

...The Government of Alberta has an established

policy of environmental legislation which allows
1. An industrial component that generated a

for the orderly development of resources with

number of different development scenarios.

 2. A population component t h a t estimated t h e
population based on background growth and develop-
ment r e l a t e d growth s c e n a r i o s .
3 . A land component t h a t c a l c u l a t e d t h e land
requirements f o r urban, i n d u s t r i a l and t r a n s p o r t a -
t i o n needs.
The t r a n s p o r t submodel was concerned with t h e
p h y s i c a l t r a n s p o r t of water, a i r , and a s s o c i a t e d
p o l l u t a n t s throughout t h e AOSERP a r e a . The water
component was a simple hydrological model of t h e
flow w i t h i n t h e 13 water drainage u n i t s . Relevant
water p o l l u t a n t s were s e l e c t e d a s water q u a l i t y
i n d i c a t o r s and t h e i r concentrations were c a l c u l a t -
ed f o r each water drainage u n i t .
The a q u a t i c biology submodel was concerned with
t h e impacts of commercial and r e c r e a t i o n a l f i s h e r -
men, instream flows, and instream p o l l u t a n t s on
four common f i s h s p e c i e s .
The t e r r e s t r i a l submodel was r e s p o n s i b l e f o r
r e p r e s e n t i n g t h e v e g e t a t i o n and w i l d l i f e dynamics.
The model concentrated on t h e economically impor-
t a n t s p e c i e s moose and beaver and t h e i r h a b i t a t .
RESULTS
A d e t a i l e d d e s c r i p t i o n of t h e simulation model
and i t s r e s u l t s a r e found i n S t a l e y e t a l .
(1979). However, t h e r e s u l t s of applying t h e AEAM
process a r e f a r more important than t h e o u t p u t o f
t h e model. Each submodel made e x p l i c i t a number
of d a t a and information gaps. A t t h e conclusion
of t h e workshop i t was apparent t h a t many of t h e
d a t a gaps revealed by t h e simulation modelling
e x e r c i s e could be f i l l e d by a p p r o p r i a t e reorgani-
z a t i o n of t h e AOSERP d a t a base. However, i t a l s o
became c l e a r t h a t a f u r t h e r conceptual understand-
ing of t h e environmental system under study was
needed before a d e t a i l e d a n a l y s i s of t h e e f f e c t s
of o i l sands development could be made. This has
important i m p l i c a t i o n s f o r f u t u r e r e s e a r c h s i n c e
t h e d a t a b a s e l i n e has been e s t a b l i s h e d . Further
r e s e a r c h should concentrate on understanding t h e
dynamics of t h e system. I n o t h e r words t h e focus
needs t o be on those t h i n g s t h a t cause v a r i a t i o n
and change i n t h e system, and not on t h e c u r r e n t
s t a t e of t h e system.
The simulation model i t s e l f provided a v e h i c l e
f o r i n t e g r a t i o n of f i v e years of environmental
r e s e a r c h . The process of b u i l d i n g t h e model i n t h e
workshop revealed a number of important r e l a t i o n -
s h i p s between t h e (Air, Water, Land, Human)
systems t h a t a r e b a s i c t o an o v e r a l l understand-
ing.
The a p p l i c a t i o n of t h e AEAM process t o AOSERP
i s n o t complete. The next phase which should
be completed i n t h e f a l l of 1980 w i l l concentrate
on developing t h e model a s a t o o l t o a i d environ-
mental management i n t h e o i l sands a r e a . The
many conceptual weaknesses and bad d a t a i n t h e
simulation model w i l l be remedied through a s e r i e s
of t e c h n i c a l meetings t o be held with t h e s t a f f of
each of t h e Human, Land, Water and A i r systems.
The r e s u l t s of t h e s e meetings w i l l guide model
refinement. The f i n a l model w i l l be put t o g e t h e r
i n an i n t e g r a t i o n workshop where t h e p a r t i c i p a n t s
of t h e o r i g i n a l workshop and t h e t e c h n i c a l meetings
w i l l i n t e r a c t i v e l y game with t h e model. Once t h e
model goes through t h i s " t r i a l by f i r e u before i t s
c r e a t o r s t h e model w i l l be used t o evaluate a number
of environmental management s t r a t e g i e s . This w i l l
be done i n an one day p o l i c y workshop focussed on
t h e evaluation of t h e model's p r o j e c t i o n s . I t i s
t h e r e s u l t s of t h i s l a t t e r s t a g e t h a t w i l l measure
t h e degree of success of t h i s a p p l i c a t i o n of AEAM.
CONCLUSIONS
The AEAM process r e p r e s e n t s t h e combined l e a r n -
ing of a number of i n t e r n a t i o n a l s c i e n t i s t s and
p r a c t i t i o n e r s and i s i n a s t a t e of "dynamic e q u i l i -
brium", c o n t i n u a l l y adapting i n l i g h t of new
experience. But t h e term "adaptivew s t r e s s e s a
more important lesson, t h e need f o r r e s e a r c h and
management t o be open t o change and t o be adapt
i n both s t y l e and content when new information
becomes a v a i l a b l e . While AEAM i t s e l f i s i n a
c o n t i n u a l s t a t e of change, i t s two underlying -
themes, expect t h e unexpected, and l e a r n t o p l a n
and p l a n t o l e a r n , never change.
U t i l i z i n g t h e modelling workshop provides many
b e n e f i t s t o t h e success of t h e AEAM e x e r c i s e . I t
f o r c e s p a r t i c i p a n t s t o focus on t h e r e l e v a n t i s s u e s ,
promotes i n t e r d i s c i p l i n a r y communication,
i d e n t i f i e s information needs, provides a framework
f o r evaluation of e x i s t i n g information and manage-
ment a c t i o n s , and i s a guide f o r environmental
p o l i c y design.
AEAM through i t s use of simulation i n t h e work-
shop s e t t i n g provides a mechanism f o r i n t e g r a t i n g
information and f a c i l i t a t i n g t h e a n a l y s i s of
impacts r a t h e r than massaging b a s e l i n e d a t a . I t
i s through thoughtful s y n t h e s i s , a n a l y s i s and
e f f e c t i v e communication of environmental information
t h a t environmental managers and r e s e a r c h e r s w i l l
make e f f e c t i v e use of t h e i r resources: time,
money and e x p e r t i s e .
LITERATURE CITED
Holling, C.S. ( e d i t o r ) . 1978. Adaptive Environ-
mental Assessment and Management. 377 p .
Wiley Inter-Science, Chichester.
Smith, S.B. ( e d i t o r ) . 1979. Alberta O i l Sands
Environmental Research Program Interim Report
covering period April 1975 t o November 1978.
Prepared by A.S. Mann, R . H . Hursey, R.T.
Seidner, and B. Kasinska-Banas. Edmonton,
Alberta.
S t a l e y , M . J . , and o t h e r s . 1979. Report of a s i m -
u l a t i o n modelling workshop on t h e environmental
e f f e c t s of Alberta O i l Sands development. 73 pp.