Horticultural Reviews, Volume 9

HORTICULTURAL REVIEWS
VOLUME 9
Horticultural Reviews is sponsored by the
American Society for Horticultural Science
Editorial Board, Volume 9

A. A. De Hertogh
D. N. Maynard
J. N. Moore
HORTICULTURAL REVIEWS
VOLUME 9
edited by
Jules Janick
Purdue University
An avi Book
Published by Van Nostrand Reinhold Company
New York
An AVI Book
(AVI is an imprint of Van Nostrand Reinhold Company Inc.)
Copyright 0 1987 by Van Nostrand Reinhold Company Inc.
ISBN-0-442-24377-4
ISSN-0163-7851
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Contents
...
Contributors Vlll
Dedication xi
1 Energy-Efficient Greenhouses
Gordon J. Monk and J. M . Molnar
I. Introduction 1
11. Structural Modifications 2
111. Alternative Energy Sources 14
IV. Environmental Management 32
V. Future Developments 38
VI. Summary 39
Literature Cited 41
2 Plant Growth Regulators in Rose Plants

Yoram Mor and Naftaly Zieslin
I. Introduction 53
11. Propagation 54
111. Storage of Rose Plants 58
IV. Plant Development 59
V. Flower Senescence 65
Literature Cited 66
3 Nutrition of Container-Grown Woody Nursery Crops

Robert D. Wright and Alexander X . Niemiera
I. Introduction 75
11. The Container System 77
111. Methods of Nutrient Application 78
IV. Nutrient Requirements for Growth 80
V. Soil Testing 88
VI. Tissue Analysis 90
V
vi CONTENTS
VII. Timing of Nutrient Applications 90

VIII. Conclusions 93
Literature Cited 95
4 Elemental Status of Pine Bark-Based Potting Media

R . J . Ogden, l? A. Pokorny, H . A . Mills, and M . G . Dunavent
I. Introduction 103
11. Common Chemical Characteristics of Soil
Organic Matter and Softwood Bark 104
111. Cation Exchange Capacity 105
IV. Nitrogen 108
V. Phosphorus 112
VI. Potassium 113
VII. Soil Reaction, Lime, and Calcium 114
VIII. Magnesium 117
IX. Micronutrients 119
Literature Cited 124
5 Iron Deficiency Chlorosis

Ronald l? Korcak
I. Introduction 133
11. Soil Iron 137
111. Iron Uptake 144
IV. Iron 'Ikanslocation 153
V. Measures of Plant Iron S t a t u s 154
VI. Bicarbonate-Induced Chlorosis 158
VII. Iron Chlorosis: Horticultural Occurrences 161
VIII. Use of Chelates 169
IX. Conclusion 171
6 Ginseng: Industry, Botany, and Culture

J. T A. Proctor and W. G . Bailey
I. Introduction 188
11. Industry 188
111. Botany 195
IV. Culture 206
V. Concluding Remarks 229
7 The Honey Bee Pollination Component of Horticultural

Crop Production Systems
G . DeGrandi-Hoffman
I. Pollination in Agroecosystems 237
11. Pollination and the Production of Horticultural Crops 243
111. Mechanisms for Cross-Pollination 256
CONTENTS vii
IV. Concluding Remarks 259

8 Tissue Culture of Temperate Fruit and Nut n e e s

J a m e s l? H u t c h i n s o n and Richard H . Z i m m e r m a n
I. Introduction 273
11. Micropropagation 274
111. Virus Elimination 318
IV. Genetic Improvement 318
V. Germplasm Conservation 324
VI. Commercial Application 325
VII. Conclusions 327
9 Summer Pruning of Apple and Peach n e e s

Richard I? Marini and J o h n A . B a r d e n
I. Introduction 351
11. Peach 353
111. Apple 360
IV. Summary 367
10 Orchard Floor Vegetation Management

E. J . H a g u e and G . H . Neilsen
I. Introduction 377
11. Major Orchard Floor Management Systems 379
111. Effects on the n e e and Crop 382
IV. Effects on the Soil 405
Subject Index 43 1
Cumulative Subject Index 433
Cumulative Contributor Index 444
Contributors
W. G. BAILEY. Department of Geography, Simon Fraser University,

Burnaby, British Columbia, Canada V5A 1S6
JOHN A. BARDEN. Department of Horticulture, Virginia Polytech-
nic Institute and State University, Blacksburg, VA 24061
G. DEGRANDI-HOFFMAN. Carl Hayden Bee Research Center, U.S.
Department of Agriculture, Agricultural Research Service, 2000 E.
Allen Road, lhcson, AZ 85719
M. G. DUNAVENT. Department of Horticulture, University of Geor-
gia, Athens, GA 30602
E. J. HOGUE. Agriculture Canada, Research Station, Summerland,
BC, Canada VOH 1ZO
JAMES F. HUTCHINSON. Department of Agriculture and Rural Affairs,
Horticultural Research Institute, Knoxfield, P.O. Box 174, Ferntree
Gully, 3156 Victoria, Australia
RONALD F. KORCAK. Fruit Laboratory, U.S. Department of Agricul-
ture, Agricultural Research Service, Agricultural Research Center,
Beltsville, MD 20705
R. PAUL LARSEN. Extension Service, Utah State University, Logan
UT 84322
RICHARD P. MARINI. Department of Horticulture, Virginia Poly-
technic Institute and State University, Blacksburg, VA 24061
H. A. MILLS. Department of Horticulture, University of Georgia,
Athens, GA 30602
J. M. MOLNAR. Agriculture Canada Research Station, 6947 No. 7
Highway, P.O. Box 1000, Agassiz, BC, VOM 1AO Canada
GORDON J. MONK. Western Bio-%ch Engineering Ltd., 6576 Wellington
Avenue, West Vancouver, BC, Canada V7W 2H9
YORAM MOR. Department of Floriculture, Extension Services, Min-
istry of Agriculture, Hakirya, Tel Aviv 61070, Israel
viii
CONTRIBUTORS ix
G. H. NEILSEN. Agriculture Canada, Research Station, Summerland,

BC, Canada VOH 1ZO
ALEXANDER X. NIEMIERA.' Department of Horticulture, Virginia
Polytechnic Institute and State University, Blacksburg, VA 24061
R. J. OGDEN. Department of Horticulture, University of Georgia,
Athens, GA 30602
F. A. POKORNY. Department of Horticulture, University of Georgia,
Athens, GA 30602
J. T. A. PROCTOR. Department of Horticultural Science, University
of Guelph, Guelph, Ontario, Canada N1G 2W1
ROBERT D. WRIGHT. Department of Horticulture, Virginia Poly-
technic Institute and State University, Blacksburg, VA 24061
NAFTALY ZIESLIN.' Department of Ornamental Horticulture, The
Hebrew University of Jerusalem, Rehovot 76100, Israel.
RICHARD H. ZIMMERMAN. Fruit Laboratory, Room 105, Building
004, U.S. Department of Agriculture, Agricultural Research Service,
Agricultural Research Center-West, Beltsville, MD 20705
'Present address: Division of Agriculture, Arizona State University, Tempe, Ari-

zona 85287.
2Temporary address: Department of Horticultural Science, University of Guelph,
Guelph, Ontario, Canada N1G 2W1.
Edward L. Proebsting, Jr.
Dedication
“He was a gentleman on whom I built an absolute trust.” This line from
Shakespeare’sMucbeth sums up my assessment of Edward L. Proebsting,
Jr., to whom this volume is proudly dedicated.
Several years ago, the chairman of the Washington %e Fruit Research
Commission told me “this Commission will give Ed Proebsting whatever
he asks.” This comment did not mean that Ed was a favored pet of the
Fiesearch Commission, or that it was so flush with funds that it could
respond openly to all requests for research support. Quite the contrary,
the Washington n e e Fruit Research Commission is composed of hard-
nosed fruit growers who carefully and critically dole out research sup-
port, sometimes as though they were their own personal funds (which is
true to a certain extent), and sometimes as if fearing that such funds
might slip out to a wastrel heading for a Saturday night binge.
The reason for this liberal support of Ed Proebsting is obvious to
anyone who has known him for any period of time. He is a remarkably
productive horticulturist in research; in service to the Washington fruit
industry; and in open-handed assistance to his fellow horticulturists
everywhere. The commissioners know that funds invested with Ed
Proebsting would be prudently and wisely used and that there is a
reasonable probability of high return for dollars invested. In essence, he
instills confidence with his peers and with the fruit-growing fraternity by
continuous and conscientious attention to sound scientific principles in
answering the production problems of fruit growers. Even though he is
exceptionally close to growers and his feet have pressed the soil of
hundreds of Washington orchards, he never compromises either research
quality or time required to develop the type of data and answers that will
stand the test of time.
I t might be supposed, with considerable supporting evidence, that
Ed’s horticultural abilities and gentle nature are in part hereditary. His
xi
xii DEDICATION
father, Edward Proebsting, Sr., was one of those early giants of pomology
a t the University of California-Davis and his son, William H., is a
horticulturist a t Oregon State University. Ed Jr. was nourished and
educated in the enriched environment of UC-Davis, receiving a BS in
pomology in 1948. This was followed by three years at Michigan State
University, where he received a Ph.D. in horticulture under the tutelage
of A. L. Kenworthy, with a thesis entitled Cherry Growth and Develop-
ment as Influenced by Solar Radiation and Intensity of Nutrition. This
study proved to be a stepping stone to nearly a lifetime of studying
environmental effects of fruit growth, production, and fruit quality.
Except for a two-year naval tour during Korean War, Ed has spent 35
highly productive pars at Washington State University’s Irrigated Research
Center a t Prosser, Washington.
Early on in Prosser he learned that winter freeze injury was a recurrent
problem in the Washington fruit belt, particularly for stone fruits. Thus,
his research was aimed a t understanding and solving cold problems. He
developed a freezing system to simulate orchard conditions most closely
in order to study fruit bud hardiness. Using branches and buds of ‘Elberta’
peach trees, his major course of inquiry was aimed a t determining fruit
bud hardiness and morphological development from the first fall freezes
through spring bloom. In a state where orchard heating is widely prac-
ticed, his precise measurements of bud hardiness a t different morphologi-
cal stages was highly important in a scientific as well as economic sense.
His T-50 scales (temperature required to kill 50% of the fruit buds) were
widely adapted in Washington and other areas.
He also studied cold hardiness as influenced by nitrogen levels, cover
crops, irrigation practices, and other cultural and environmental factors.
Most of his studies covered many years under varying conditions. The
results were sometimes quite contrary to common orchard assumptions
and even published research data. For example, he showed conclusively
that fruit bud survival of high nitrogen trees was better than from
medium or low nitrogen trees, and this relationship was evident through-
out winter from mid-December to mid-March. In another 5-year study of
relationships between nitrogen levels and canning quality of ‘Elberta’
peaches, canning quality was improved by high nitrogen fertilization.
High nitrogen peaches were most closely associated with better flavor (as
preferred by taste panels), with finer texture, firmer flesh, less astrin-
gency, and lower tartness.
During the 1950s and 1960s, nearly every volume of the ASHS Pro-
ceedings had a research paper by E. L. Proebsting, Jr. The discussion
sections of his papers were interesting and scholarly. A 1958 Proceedings
article, for example, discussed an issue of major importance to all horti-
cultural researchers, that is, the dilemma of evaluating “constancy,” or
DEDICATION xiii
lack thereof, when trying to study vegetative efficiency of a fruit tree

under varying conditions of cropping and environmental stress. Even
through much of Proebsting’s early research was conducted with the
‘Elberta’ peach, his experimental methods and careful collection and
evaluation of the data were broadly applicable to other fruit crops.
Through the years, he also branched out into numerous studies with
cherries, plums, pears, and apples. The commercial use of phenoxy-based
growth regulators resulted in 20-50% increased yields of ‘Italian’plums in
Washington. This was due to Ed’s research and his assistance in develop-
ing grower practices. His research with daminozide and gibberellins on
sweet cherries was quickly translated into commercially usable practices.
In recent years, he has been deeply involved with other horticulturists
and plant pathologists in ice nucleation studies, using nucleation active
bacteria to reduce freeze injury of the target plants.
It is a long way from Prosser, Washington, to Sapparo, Japan, or
Ithaca, New York, where Ed has spent considerable time both visiting
and on sabbatical leave, but his quiet, helpful, concerned presence has
been of enormous benefit to young researchers there and elsewhere.
Wherever he has trod, horticultural researchers and the fruit industries in
general have benefitted by his presence.
Dr. Proebsting has been an active member of the American Society for
Horticultural Science for nearly 40 years and has held various Society
off ices, including chairmanships of five committees and associate editor
(twice).He was elected an ASHS Fellow in 1969, and was President of the
Society in 1983-1984. He received the Gourley Award in Pomology (1955),
the Woodbury Award ( 1958),and the National Food Processor’sAssocia-
tion Award (1979). He has received many honors and awards in his
adopted state of Washington, including being named the 1984 Cherry
King by the Washington Cherry Industry.
Every day is like a new adventure to Ed Proebsting. He never tires of
research orchards, or laboratories, and he is always a t hand with a
welcome smile and helpful hand to workers, technicians, colleagues,
visiting fruit growers, and fellow horticulturists. Ed Proebsting’s horti-
cultural philosophy was well stated in his 1984 Presidential address, “Life
in the Slow Lane,” wherein he indicated, “being a horticulturalist ain’t
necessarily a bed of roses.”
He went on to say that compared to the affluence of some professional
fields, we may be living our professional lives in the slow lane, but as
practicing horticultural scientists, “we have chosen a career that is both
rewarding and exciting. We are rewarded by the knowledge that our
efforts are contributing something of value to society. We have the
excitement that comes from discovery or from leading changes in the way
things are done. At the same time, we have the pleasure of working
xiv DEDICATION
directly with beauty in the fruits, vegetables, flowers, and landscapes of

our professions, and with the high-quality people who are attracted by
horticultural endeavors, either as professionals or as amateurs, and even
though we may not necessarily receive the monetary support or public
recognition of other professional groups, life in the slow lane need not
be dull.”
Those of us who have known or associated with E. L. Proebsting, Jr.,
know that “life in the slow lane” has been neither dull nor unproductive
for him. Einstein once said “only a life lived for others is a life worth-
while.” In nearly 35 years of friendship with Ed Proebsting, I would
suggest he nobly fits that definition.
R. Paul Larsen
Utah State University
Logan, Utah
Horticultural Reviews, Volume 9
Edited by Jules Janick
Copyright © 1987 Van Nostrand Reinhold Company Inc.
1
Energy-Efficient Greenhouses
Gordon J. Monk*
Western Bio-%ch Engineering Ltd.
West Vancouver, British Columbia, Canada V7W 2H9
J. M . Molnar*
Agriculture Canada Research Station
Agassiz, British Columbia, Canada VOM 1AO
I. Introduction 1
11. Structural Modifications 2
A . Coverings 2
B. Insulation 8
C . Thermal Screens 11
111. Alternative Energy Sources 14
A . Solar 14
B . WasteHeat 25
C . Geothermal 30
IV. Environmental Management 32
A . Root Zone Heating 32
B . Alternative Heating Methods 34
C . Environmental Control 35
V. Future Developments 38
VI. Summary 39
VII. Literature Cited 41
I. INTRODUCTION
In temperate climates, controlled-environment horticulture has become

an important sector for flower and bedding plant production and a small
but significant component of off-season production of vegetables, partic-
ularly tomato, cucumber, and lettuce. Until recently, however, the rapid
escalation of fossil fuel costs that followed the oil crisis of 1973 seriously
challenged the viability of this energy-intensive industry. Although world
oil prices did tumble suddenly in late 1985, many experts predict that
they will soon rise, eventually surpassing previous record high levels.
*The authors wish to thank their research assistants Bruce Ewert and Tony Wakelin
for their extensive help during the research review. Peggy Watson contributed gener-
ously by locating the references and Jennifer Lockyer organized the literature citations.
1
2 GORDON J. MONK AND J. M. MOLNAR
In response, researchers and commercial greenhouse operators were

forced to develop and test new energy-conserving structures and crop
management techniques. The use of alternative energy sources has also
been investigated to reduce the industry’s dependence on fossil fuels.
During this time, many significant changes were made to greenhouse
designs and their operations.
White (1979) described the heat transfer characteristics of green-
houses and introduced methods that were considered to make these
structures energy efficient. This chapter surveys the results of research
that has taken place over the last decade. The review focuses on new
energy sources, conservation techniques, heat distribution, and some
aspects of environmental control. I t concludes with a prediction of how
the knowledge gained will influence the development of greenhouse pro-
duction systems over the next ten years.
11. STRUCTURAL MODIFICATIONS
A. Coverings
The single most researched aspect of greenhouse design has been
identifying materials superior to glass for covering the structure, but
glass today remains the standard by which all other glazing materials
are measured.
Depending on the quality, glass can provide direct light transmittance
of 84-97%, which is greater than any substitute material. I t costs about
$6.50/m2, which is cheaper than most of the alternatives except fiber-
glass and double polyethylene film.
A conventional glasshouse is constructed from clear glass panes 3 mm
thick, 50 x 90 cm, supported by aluminum glazing bars on a galvanized
steel frame. The glass laps should be sealed with a silicon-based sealant.
Badger and Poole (1979)claimed the associated reduction in infiltration
losses can reduce energy consumption by 5-40%. Savings depend on
construction materials, the condition of the structure, and exposure to
prevailing winds. The cost will vary between $3.20 and $5.20/m2 of floor
area (1980 US.funds) and will pay for itself in about 18 months in a
climate similar to that of Pennsylvania (White and Aldrich 1980). The
disadvantage of lap-sealedglasshouses is the time and difficulty associated
with replacing broken panes. Fkduced infiltration also makes COa enrich-
ment more economical, but it increases humidity levels.
In recent years, the so-calledVenlo-type structure has become popular.
Developed in Holland, this design uses larger panes, 4 mm thick, 73 X 165
cm. Since fewer glazing bars are spaced further apart, the framework
1. ENERGY-EFFICIENT GREENHOUSES 3
blocks out less sunlight, resulting in higher plant canopy light levels.
These are also fewer glass laps through which to lose energy in unsealed
houses. However, the low ridges on narrow houses increase shading from
other structural components. The added shadows from ridges and gut-
ters tend to counterbalance the benefits from fewer glazing bars.
Unfortunately, glass offers very little resistance to conductive heat
transfer back out of the greenhouse. When glass forms a barrier between
relatively acquiescent masses of warm and cool air, its overall heat
transfer resistance (R-value)is 0.15 m2 OC/W. Wind and interior air cir-
culation reduce the thickness of the boundary layers of air and create
turbulence within them, accelerating the convective heat transfer. If the
glass laps are not sealed, the overall heat losses can increase by as much
as 10070,depending on the wind velocity.
These heat losses can be significantly reduced if an additional layer of
glazing is placed adjacent to the glass a t a minimum separation distance
of 10 mm. The sealed layers of glazing form a narrow space of trapped air.
The air space supplies considerable resistance to heat loss because the
slow process of convective heat transfer must now occur through the air
space itself and across two more laminar air boundary layers. Further-
more, the seal prevents external air movements from accelerating the
internal convective heat transfer processes.
Since the trapped air space is mainly responsible for increasing the
overall R-value, the type of glazing material forming the seal is not
critical. Accordingly, research to find suitable alteratives to single-layer
glass has focused primarily on double-layer coverings rather than on
other single-layer materials. Agrifilm 88 and low-emissivity glass are
examples of exceptions to this generalization.
The process of identifying new glazing materials begins in the labora-
tory. lksts are undertaken to determine R-values, transmittance of solar
and long-wavelength radiation, strength, degradation due to ultraviolet
( UV) radiation absorption, and corrosive susceptibility to pesticides and
other greenhouse chemicals. Computer simulation programs can then be
used to predict plant canopy light levels, energy savings, and economic
benefits. If results are promising, in situ experimentation is carried out
with a prototype structure over a period of several years.
Bond et al. (1977)presented solar, long-wavelength, and photosynthet-
ically active radiation ( PAR) transmittance for nine common materials
suitable for greenhouse coverings, as well as for 81 two-layer combina-
tions of these materials. The variation in solar energy transmission with
angle of incidence between the surface normal and solar direct-beam radi-
ation was also reported. Godbey et al. ( 1979)also reported the percentage
of transmitted direct-beam solar radiation that is diffuse radiation be-
low the cover materials. The nine materials were polyethylene (4-mil,
4 GORDON J. MONK AND 1. M. MOLNAR
UV-resistant ), flat fiberglass (25-mil), flat fiberglass (premium-grade,

40-mil1, two types of corrugated fiberglass (40-mil), polyester (5-mil),
glass (double-strength, 3-mil), polycarbonate ( 1.5-mil),and polyvinylfhoride
(3-mil)(note: 1mil = 0.001 in. = 0.00254 cm). Glass had a long-wavelength
or thermal radiation transmittance of 3%, the lowest of the p u p . Conversely,
single-layer polyethylene had by far the highest long-wavelength trans-
mittance a t 80%. The 40-mil fiberglass was the best diffuser, transmitting
89% of incident radiation as diffuse light. The polyester and polycarbonate
diffused only 23% of incoming light. However, if glazing thickness was
taken into account, glass scattered less light proportionally, diffusing
only 24% of the incoming radiation.
The heat loss from glazing materials was compared by Hellickson
( 1978). He found that the increase in heat loss from corrugated fiberglass
as compared to flat fiberglass was proportional to increased surface area
over projected area. Therefore, unless the extra strength supplied by the
corrugations is required, flat fiberglass is preferable. Single-layer 6-mil
polyethylene also transmitted 37% more heat than flat fiberglass.
Waaijenberg ( 1984a)summarized the results of research on the strength
and durability of plastic and glass products including their bending
stiffness and impact strength. In the case of double-glazed panels formed
from glass panes of different thicknesses, the thickest pane should always
be on the outside of the greenhouse cladding. Low temperatures and low
moisture contents tended to increase the bending stiffness of the plastic
materials. In order to investigate th’e UV stability, specimens of the
plastic panels were studied by means of accelerated aging tests in a Xenon
1200 apparatus. Acrylic panels did not manifest any deterioration of the
properties examined even after the maximum exposure time of 4500 hr,
which corresponds to about 15 years of outside exposure. With PVC
panels, the impact strength was found to have clearly deteriorated after
1500 hr of irradiation. Polycarbonate panels revealed a definite decrease
in impact strength between 1500 and 3000 hr of exposure. Nevertheless,
the impact strength remained sufficiently high for the material to con-
tinue functioning as a greenhouse glazing.
The spectral sensitivity of coverings was examined in detail by
McNaughton et al. (1981).The PAR transmittance by new glass was 90%,
higher than any other glazing tested. Thin 0.125- and 0.250-mm film
plastics such as polyethylene (AH1 or Garnite Greenhouse Film), poly-
ethylene copolymer ( Agphane 101), ethylene vinyl-acetate copolymer
(Permaflex),or polyvinyl chloride (Nylex)all transmitted about 88-9070
of PAR. llansmission was lower with reinforced ethylene vinyl-acetate
film (80% Graphlon) and with colored films (65-75%, green Graphlon,
green Permaflex 1. The rigid acrylic ( Acrylflute ) and fiberglass-reinforced
(Durolite f ) materials transmitted 76-8270, while rigid PVC (Novaroof)

transmitted 63-7390, depending on type.
Long-wave radiation transmission in these tests was 0 for glass and
less than 5% for rigid fiberglass (Durolite f ), rigid PVC (Novaroof),and
the double-walled acrylic material (Acrylflute).Thick (0.3-0.4 mm) PVC
film (Nylex) transmitted 10-1270, while polyethylene and EVA films
transmitted 40-6970.
Gustavsson et al. (1977)reported that double EVA film will provide
overall energy savings of 35-400/0,compared to a conventional single-span
glasshouse. The equivalent figure for German double acrylic (Skegdop-
pelplatte) was 40-50%. Royle ( 1984) reported that double-glass glazing
offered similar energy savings but reduced light transmission by 10%as
compared with single glass a t Somerset, England. There, the wet climate
produces low light levels, and Royle repeated the old rule of thumb that a
170loss of light equals a 1%loss of production. Because of this and a 20%
higher installation cost, double glass was not recommended for any
light-sensitive crops.
In Ontario, Jewett and Papadopoulos (1984)experimented with thin-
wall double PVC, which reduced heat loss by 22% and light levels by 18%
compared to a single-layerglass compartment. Nevertheless,they obtained
comparable seasonal tomato yields, even though early yields were delayed.
The reaction of cut flower and pot plant crops to reduced light intensities
and higher humidities under double glazing depends on the crop and the
glazing material according to Steinbuch and van de Vooren (1984). Be-
gonias and saintpaulia had a cultivation period one week shorter under
double glass but the cultivation period of chrysanthemum was longer.
Anthurium and ‘Sonia’rose showed increased production under double
glass but the quality of the roses suffered, especially in winter when thin
stems and more flowerless shoots were observed. In climates where natural
irradiation levels are high, light losses due to double glazing are obviously
less important.
The most popular rigid plastic glazings have been structured sheets of
double acrylic and double polycarbonate. Hinton-Mead ( 1982) achieved
41% energy savings with 22 mm acrylicand 34% with 10mm polycarbonate.
However, tomato yields and fruit quality under the polycarbonate were
reduced compared to acrylic. Norman (1983) and Goldsberry (1984)
found that polycarbonate is also susceptible to yellowing due to UV
degradation. The associated reduction in light transmission was meas-
ured by Landgren and Nilsson (1982)and found to be 0.8-1.070 annually.
Norman also reported that although acrylic produced higher humidity
levels and a slight tomato yield loss when compared to Venlo glass, it was
judged superior overall due to acceptable light transmission and high
energy savings. Goldsberry found that fiberglass-reinforcedplastic ( FRP )

laminated with Tedlar was another material that is resistant to yellowing.
The latest development in German technology (Anon. 1985a) has pro-
duced 32-mm double acrylic panels that are strong enough to eliminate
the need for structural glazing bars.
An inexpensive form of double glazing is inflated polyethylene film.
Goldsberry et al. (1982)compared double polyethylene with 6-mm dou-
ble polycarbonate, 140-g Tedlar-coated FRP panels and double-layer
Tedlar film (Nexglaze).A double polyethylene quonset required 9% more
fuel than the Nexglaze house but 1 and 34% less fuel than identical
polycarbonate and FRP houses, respectively. New 6-mil double polyeth-
ylene was also tested against new inflated 4-mil PVC, 2-year-olddouble-
layer 4-mil Tedlar film, and a single layer of 140-g Tedlar-coated FRP by
Ferare and Goldsberry (1984). When compared to the FRP panels, the
polyethylene and PVC reduced seasonal fuel consumption by about 40%,
while the Tedlar saved 35%. However, the PVC was not suitable for cold
winter conditions and condensate removal between the layers of Tedlar
was required for maximum transmission of solar radiation. Simpkins et
al. ( 1984)reported that high-density double polyethylene only transmit-
ted 62-64% of the available PAR, while O’Flaherty and Grant (1984)
found the mean PAR transmission to be 60.5%. Consequently, despite
their low cost, some double polyethylene films are obviously not suitable
for light-sensitive crops unless ambient irradiation levels are very high.
Recently, new clear polyethylene films have been developed that transmit
significantly more PAR. ‘Monsanto 703’ and ‘Tufflite 111’ have PAR
transmittances of 76.9 and 77.6%,respectively,according to tests conducted
a t the University of Illinois (Sherry 1986).
Inflated double polyethylene has been used to cover glasshouses dur-
ing the winter months. Snyder and Bauerle ( 1981)measured heat savings
of 40-60% annually with this application over four winters. Relative
humidity increased by only 2% on average but tomato-crop yields were
10.4% lower during spring and 15.4% during fall when compared to those
grown under single glass. The reductions were likely due to lower light
levels. Poole (1979)determined that irradiance reaching the crop would
drop by 15-20%. In spite of this, he calculated a one year payback period
attributable to 50%annual fuel savings and pointed out that this was the
least cost-highest return method of renovating an old greenhouse in poor
repair. These results were confirmed by Ingratta (1978)who measured an
18% decrease in light levels and a 40% reduction in fuel consumption
while determining a payback period of 12 months.
Polyethylene film can also be applied in single layer beneath the glass.
Vardiashvili et al. (1981)stretched film throughout the interior of a 200
m2 house, 0.06 m from the glass. The experiment was modified and
repeated by extending polyethylene across the house at gutter height,

thereby creating a large air buffer zone. Heat losses through the horizon-
tal buffer were 27-2970less than across the 0.06 m interlayer. O’Flaherty
and Grant (1984) lined a glasshouse with PVC panels. Although they
measured a 7.6% reduction in PAR compared to an unlined house, initial
tomato production was not affected. Starkey (1985)used 3-mil Melinex
as a secondary glazing to glass with an unspecified air gap thickness
and reported annual fuel savings of 30%.
The lack of outside air penetration into double-layer film greenhouses
reduces heat loss by infiltration but increases humidity levels. In cold
climates, condensation droplets form on the inside surface of the inner
glazing after nightfall. This results in relatively high rates of latent heat
transfer to the outside air via the layers of polyethylene and enclosed
airspace and causes problems with plant health. According to Silveston
et al. (1980)condensation contributes at least 15%of the total heat loss
in double polyethylene houses and as much as 20% when mylar is the
covering material. Since ventilation transfers heat directly to the outside
heat sink, the researchers used mechanical dehumidifiers to eliminate
latent heat loss but the costs were not justified on the basis of energy
savings alone. Commercially available surface-active agents ( surfactants)
were then used to promote film-wise condensation-a very thin, even
layer of water. Reducing the droplets to film decreases heat losses and in-
creases light transmittance. Heat losses were reduced by 5%but this was
insufficient to compensate for the additional cost. This is contrary to the
report of Bryenton et al. (1983),who discussed the application of ethox-
ylated fatty alcohols to glass, plastic, and fiberglass to encourage film-wise
condensation. In this case, a payback period of 1-2 years was projected.
Goldsberry ( 1979)reviewed the influences of wind speed on heat losses
and cited reports (Koths 1976; Morris 1959; Oldroyd 1975; Sheard 1974;
Whittle and Lawrence 1960)stating that windspeeds of 25-30 km/hr will
double fuel consumption in greenhouses. In exposed areas a windbreak
can be constructed to block winds from the prevailing direction. Morris
( 1960)hypothesized that a windbreak can account for 5-10% annual fuel
savings. Morris also reported on research conducted in Great Britain
indicating that 20% less fuel was required in a greenhouse given 40%
shelter by surrounding houses. The durability of various windbreak ma-
terials has been assessed by O’Flaherty (1977).White and Aldrich (1980)
cautioned that proper design of a windbreak is critical since poorly
engineered windbreaks may create eddy currents, which can actually
increase flow. They offered guidelines on siting a windbreak during their
discussion of a variety of energy-saving techniques.
Despite all of the research completed to date, single-layer glass is still
the most widely used glazing material in Europe and Canada because of
its unsurpassed light transmittance. In the United States, double-film

greenhouses are dominant. Efforts are now underway to improve the
properties of glass. Pierce ( 1982)outlined the development of a molecular,
metal-covered glass, which admits over 90% of the available solar energy
but reflects 85% of the long-wave heat energy. White (1984)mentioned a
new glass trade named Solatex, which has a low iron content. The lower
iron content increases light transmission for double-strength float glass
from about 84 to 91%. Special surface treatments are also available to
reduce light reflection, permitting total transmission to reach 97%.
Emissivity is a measure of the degree to which a surface emits heat as
radiation. Hortiplus glass has been designed to have a low emissivity in
order to reduce long-wave radiation losses. Its exterior surface is covered
with a tin oxide film 0.42 pm thick, causing the emissivity factor to be
reduced to 0.3 as compared to 0.9 in ordinary clear glass (Glaverbell977,
1980). Its light transmittance is also 9% lower than ordinary glass.
However, Benoit et al. (1984)found that the effects of reduced luminosity
were compensated by modifying the environmental control. Compared to
an ordinary glasshouse, ventilators in the Hortiplus house were opened
56% farther to even out transpiration in the ‘Dombito’ tomato crop.
Relative air humidity was 1%lower on average. Heating pipe tempera-
tures were lowered an average of 6.75OC to reduce convection, which
theoretically decreased transpiration, thereby improving the supply of
nutrients and water. Air temperatures averaged 0.2OC higher and produc-
tion was maintained while decreasing overall energy consumption by 25%
during a February-May period.
B. Insulation
Unfortunately, common construction materials that effectively resist
the transfer of heat also block the transmission of light. In temperate
climates, low levels of solar radiation usually occur in winter when cold
temperatures maximize heat losses. With conventionally shaped green-
houses this means that any application of permanent insulation decreases
interior light levels, thereby reducing production of most crops. Buitelaar
et al. ( 1984)studied the effects of four insulation materials installed a t the
north and south sidewalls on flowering rates and production of tomato in
the Netherlands. Three of the materials reduced light transmittance
while the fourth was opaque. Not surprisingly, increased losses in produc-
tion were recorded as light transmission was decreased. In relative terms,
these losses increased as the level of ambient irradiation increased.
Therefore, the greatest percentage declines in production occurred in
summer. No significant differences in flowering rate were found with all of
the materials. However, a slightly earlier floweringwas recorded adjacent
to the north wall, as compared to the south wall. This may be explained
by a somewhat higher temperature a t the north wall.
Challa and Schapendonk (1984) found that there is no general rule
describing quantitatively the relation between light reduction and yield
since it is affected by the ambient light level, which varies seasonally and
by the stage of the crop. ?tyo principally different situations were
distinguished: young widely spaced plants and older plants growing in a
closed canopy. With young plants, a decrease in growth rate is less than
proportional to a reduction in light levels, which are reflected primarily by
a delay in the start of the production phase. With older plants, production
declines are proportional to a reduction in light except when ambient
irradiation levels are very low, in which case the production losses are
more than proportional to the light reductions.
In order to clarify the situation, Abdallah and Staley (1979) studied
the transmission of light and heat through each surface of a glasshouse
by means of computer simulation. The influences of aspect ratio, roof
slope, and orientation were determined by varying each parameter. They
found that in latitudes greater than 45', gable greenhouses oriented
east-west transmit more light in winter and less light in summer than
identical structures oriented north-south. Consequently, winter heating
loads and summer cooling loads could both be reduced by adopting an
east-west alignment.
Abdallah and Staley also calculated that only about 5% of the annual
incoming radiation is transmitted by the north-facing sidewall of a struc-
ture aligned east-west. By insulating this surface, heat losses would be
reduced by approximately 16%, depending on the aspect ratio and di-
mensions of the greenhouse. Interior light levels would be decreased on
summer days, especially during the early morning and late evening.
Nevertheless, Abdallah and Staley concluded that the fuel savings would
more than offset any crop reductions.
Lawand et al. also employed computer optimization (1975)to design a
greenhouse oriented east-west with a unique profile for northern lati-
tudes. The north-facing roof was eliminated and replaced by an insulated
sidewall inclined inwards a t an angle of 65' to the horizontal. By increas-
ing the slope of the south-facing roof to 35' to the horizontal, improved
light transmission of the low winter sun was achieved. The interior
surface of the north wall was coated with reflective material to redirect the
light down onto the crops. Energy savings of 40% were reported but crop
production data were unavailable.
Thomas ( 1978)used winter weather data from the Kew Meteorological
Office in England to determine the optimum angle for a specularly
reflecting insulated north wall located at a latitude of 51'. In this case,
diffuse radiation was taken into consideration to determine that an angle
of 75O to the horizontal would achieve the best results. In Virginia (37ON
latitude) Hartz and Lewis (1982)used a north wall sloped a t 60’ to the
horizontal to improve overall light transmission from 72 to 86% during
the period October-March. Energy savings of 14% were recorded but
these researchers cautioned that insulated north walls, by their nature,
can be installed only in single-span structures.
Bailey (1984) described a design developed in Japan, consisting of a
lean-to greenhouse in which the vertical north wall was covered with
mirrors. The mirrors were mounted on horizontal tubes, which were ro-
tated to ensure that the incoming sunlight was always reflected onto the
greenhouse floor. Under sunny conditions, average solar radiation inten-
sity on the greenhouse floor was 25% higher than outside, resulting in a
25% increase in tomato production.
In warmer climates near the equator where snow load is not a problem
and sunny, dry conditionsprevail, G m l i and Blackwell ( 1982)mmmended
that the roof slope of a gable structure be reduced to approximately 5%.
This method insulates the greenhouse in the sense that total surface area
of large ranges can be reduced by about 10%.
Besides the north wall, gables and sidewalls can be insulated with
opaque materials up to plant height. Badger and Poole (1979)estimated
that the additional savings could be as much as 10%. Langefeld (1983)
described the installation of transparent 2.5-cm Thermax panels on the
walls, which also provided a 10% fuel saving and a payback period of 6
months. Waaijenberg ( 1984b)concluded that the optimum sidewall insu-
lation consisted of double glass used in conjunction with a Bflon-FEP
film. Direct and diffuse light transmittance through this covering were 78
and 6870,respectively.
A much more sophisticated system of “dynamicinsulation”was developed
by Short and Shah ( 1981).Portable polystyrene pellets were pneumatically
conveyed diurnally to and from a 13-cmair space between the coverings of
a double-polyethylene greenhouse. The experimental system provided
nighttime heat load reductions of 90%, which enabled a soil-heating
system to maintain temperatures without any auxiliary heaters (Elwellet
al. 1983).Enshayan and Short ( 1985)studied the pertinent flow variables
of pneumatic conveying using a zipper tube roof distribution system
(Fig. 1.1).The zipper tube system worked well for experimental purposes
but would not be commercially feasible, since the zippers jammed with
polystyrene and were difficult to repair. The investigators concluded that
a new design is required that incorporates a pellet flow controller to
prevent plugging and improve removal of pellets from the roof.
Another dynamic insulation system was reported by Groh (1976). I t
utilized a liquid-based foam insulation material, which was injected
Fan
Fig. 1.1. Layout of the polystyrene pellet dynamic insulation system. (From Enshayan
and Short 1985.)
between two layers of polyethylene in the greenhouse roof at night and

pumped out in the morning. A test unit was constructed with permanent-
ly insulated walls to isolate the roof heat losses, and overall savings of
47% were recorded in comparison to an unfoamed double-polyethylene
house. The foam filled the length of the 14 m unit in about 16 minutes
but the researchers were unable to maintain adequate foam levels in the
roof consistently.
C. Thermal Screens
For several years, researchers have been assessing the feasibility of
installing retractable internal insulating covers for nighttime deploy-
ment (White 1979). This form of dynamic insulation can reduce heat
losses in five ways: ( 1 ) The screen resists conductive heat transfer and ( 2 )
may, depending on the material, block radiant heat losses to the sky
above. ( 3 ) Convective heat transfer from the inside air to the glazing
decreases because the internal air mass is divided in two, thereby restricting
internal air movement. ( 4 )Latent heat loss due to condensation is reduced
or eliminated since water vapour cannot move freely to the colder glazing
surfaces. ( 5 ) Finally, the heat transfer surface area is reduced if thermal
screens are deployed gutter to gutter.
In recent years, experimentation has concentrated on finding light-
weight, durable materials with high R-values that bundle into compact
12 GORDON J. MONK AND. J. M. MOLNAR
spaces. Hay and Goldsberry ( 1978)conducted a two-yeartest with single-

layer polyethylene that provided a 21-32% reduction in nighttime fuel
consumption. They concluded that polyethylene can provide minimum
seasonal energy savings of 20%. Bailey ( 1977)compared the effectiveness
of aluminized polyester, polyester containing aluminum powder, black
polyethylene, clear polyethylene, and a cellular polyethylene. During
deployment he obtained decreases in thermal transmittance of 43,28,32,
33, and 34, respectively, a t zero wind speed, though the type of aluminized
polyester used was not sufficiently durable for commercial use. Butters
( 1980)described the structure and mechanism of a thermal screen system
designed for Venlo ranges that initially yielded nighttime savings of 28%.
At the time of its introduction, Foylon had the highest R-value of any
available material. Goldsberry and llistan (1977)installed it in a fiber-
glass house with gaps a t truss intersections and gable ends, leaving 5% of
the enclosed area unlined. An overall reduction of 23% was obtained
during December-February, leading the researcher to speculate that
savings of 24% would be obtained in a totally enclosed block. However,
Van Every and Brundrett ( 1980)determined theoreticallyand experimentally
that sealing the cracks around thermal curtains to eliminate chimney
effects is of paramount importance. In fact, the configuration of the
cracks resulted in dramatic increases in heat transfer that actually led to
fuel demands approaching those for an unscreened house. Brundrett
(1981) later controlled leakage with end shelves and inflatable collars
with a modified system that paid for itself in 2-3 years.
A variety of other curtain materials and system configurations have
been tested. Aldrich and White ( 1979)found that heavy materials such as
foamed plastic and folding rigid board provided the greatest savings
among the thermal screens they tested. Daytime storage and transport
methods between open and closed positions were the most serious prob-
lems to solve.
In sunny climates it is advantageous if the screen can be used for
partial shading during daytime (Garzoli and Blackwell 1981). An inflated
polyethylene tube system can also provide partial shading during day-
time when it is deflated. At night, the tubes provide an almost perfect
seal, and Short and Huizing (1982)have predicted nighttime savings of
50% with no condensation or icing problems.
Albright (1983)recommended the use of double-layer screens, which
have demonstrated savings of 86% in winter and almost 100% in summer
for a payback period estimated at 2-5 years. Shapiro et al. ( 1979)described
a sewn double-layer vinyl-coated material containing a 2.0-cm layer of
continuous-filament polyester garment insulation that reduced energy
consumption by 45%. Staley et al. (1986) applied net present values
1, ENERGY-EFFICIENT GREENHOUSES 13
( N P V )economic analysis to nighttime heat load reduction data for three

thermal screen systems made from black terylene LS 100and a reinforced
aluminum foil-polyethylene laminate material. This resulted in a predic-
tion that thermal curtains capable of providing nighttime savings of
4340% will be a profitable investment under most circumstances unless
an unusually inexpensive source of fuel is available or high installation
costs are involved.
Thermal screen systems incorporating louvers have been developed
(Butler et al. 1984) that reduce heat loss by 46%. Unfortunately, the
louvers reduced light entering the greenhouse by 22% compared to a
control house, producing legginess in lupines and bedding plants. Lawson
(1984)reported on a “multiroll” thermal screen that bundles compactly
and fits inside the greenhouse wall. He also mentions a black polyester
material coated with aluminum, which retracts from a 3.2-m bay into a
space of 1 cm and provides savings of 70%.
Some greenhouse structures are not compatible with moving screens.
In Holland, fixed screens have been developed for these houses, which
have slits that can be opened to prevent the buildup of humidity. Van
Holsteijn and de Vogel ( 1984)compared humidity levels under the so-called
moisture-slit-screens with an uncovered control. When the slits were
open, humidity levels were only 1-3%higher, but with the screen completely
closed the difference was 1-1270, depending on the weather. Moisture-slit-
screens reduce heat losses by about 35% according to Van Holsteijn and
Schoenmakers ( 1984).In one commercial operation they observed slower
tomato plant development and fruit setting. At a second operation plants
had serious leaf shedding and delayed blooming, but this was thought to
have been caused by a temporarily high atmospheric COZ level. However,
Welles et al. ( 1983)concluded that the use of moisture-slit-screens did not
result in overall production loss based on observances of tomato crops a t
three commercial operations.
After initial reluctance, the commercial industry has now adopted the
use of thermal screens on a widespread basis. The growers had been
concerned about reduced light levels due to shading but most of the new
systems have rack and pinion drive, which provides good seals and
excellent bundling capacities.
Tkaditionally,thermal screens have been closed a t night and opened in
the morning. The advent of computer controls may change this practice.
Seginer and Albright ( 1979)theorized that the best time to close or open
a screen occurs when the potential gain from photosynthesis while open
equals the potential benefit of energy conservation when closed. This
strategy is currently being tested in Holland a t the Vleuten Experimen-
tal Station.
111. ALTERNATIVE ENERGY SOURCES
A. Solar
In his review article, White (1979)stated: “One of the most promising
long-range partial solutions to high greenhouse energy costs is in the
ultilization of solar energy for nighttime heating.” At the time many
other researchers shared White’s outlook, and subsequently a great deal
of solar heating research was undertaken. However, White also correctly
predicted that the researchers would find that “the biggest problem was
not in the technology of solar collection nor its application to the green-
house but in its cost effectiveness.” Other difficulties, associated with
declines in crop productivity, have been encountered in some instances
(Airhart 1984).
In most temperate climates there is sufficient solar radiation during
the months of March through October to create a buildup of surplus heat
in the greenhouse. The technology required to capture and store the
excess energy for nighttime heating is relatively simple and significant
savings can be achieved. However, many other methods of conserving
energy have proven to be as effective a t a lower cost. This is not to say that
solar energy research has been wasteful, producing nothing but failures.
On the contrary, some successful solar systems have been developed and
much has been learned. If another escalation in fossil fuel costs occurs,
many systems that are not cost effective today could be put into commer-
cial practice in the future. I t is comforting to note that while the supply of
fossil fuels is not inexhaustible, solar energy will be available for as long
as there are greenhouses.
Excellent review articles have been written that introduce the basic
principles of radiant energy transfer from the sun and the options that are
available for harnessing the available heat. Grace and Li (1974)concen-
trated on aspects of thermal energy storage. Aldrich (1980) described
how solar radiation strikes the greenhouse surfaces, is transmitted by the
glazing material, and is absorbed by the various interior surfaces, resulting
in passive heating. An outline of the types of systems that can collect and
store the excess heat has been prepared by Baird and Waters (1979).
Solar heating systems can be divided into two broad categories: pas-
sive and active. Passive systems can be defined as methods of capture and
utilization that require no internal energy input. Active systems require
electrical or mechanical energy inputs to facilitate energy capture. Gener-
ally speaking, passive systems are less expensive but, in turn, their
effectiveness is limited and dependent on climate to a greater degree.
Albright et al. (1981) discussed some of the main features of a pas-
sive system being designed for commercial use. These included aug-
mented thermal mass, expanded daily temperature ranges, and a highly

insulated thermal screen. In another system, water tanks were used for
heat storage in conjunction with movable foam insulation panels
(Kusianovich 1978).
McGowan and Black ( 1980)placed metal drums filled with water along
the north and south sidewalls of a greenhouse in lknnessee. Interior
temperatures of 12OC were maintained with outside temperatures of
-2OC, and the passive system provided 77% of the heat load from No-
vember to March. A similar system using a water bag solar collector is
technically and economically practical according to Murakami et al.
(1984). Mercier (1981) reported that in France, water drums, rock, and
miscellaneous storages were used in a conventional greenhouse to supply
69% of the energy requirement.
Active solar heating systems usually have separate solar collectors and
thermal storages, linked by some type of circulating fluid. Von Zabeltitz
and Damrath (1984)reviewed three options for the greenhouse collector
configuration. The collectors can be external to the greenhouse, inte-
grated into the greenhouse structure such as a liquid filter flowing
between a double roof glazing, or the greenhouse itself can function as a
collector. However, Van Die ( 1980)noted that solar heating will be used by
the greenhouse industry only if it is an integral component of the build-
ing design, since external collectors require additional land and funds.
During the design of an active solar heating system, transient and
steady-state thermal dynamic computer simulations are often required.
Zornig et al. (1977) demonstrated the necessity for a dynamic thermal
analysis for various design parameters. These involve changing tempera-
tures and insulation outside and inside the greenhouse as well as a t the
solar collectors, and changing temperatures and thermal lag in the heat
storages. Procedures for calculating the solar radiation capture by green-
houses using dimensionless factors have been determined by Abdallah
and Staley (1983). Radiation configuration factors were determined for
greenhouse lengths of 10-100 m, widths of 5-15 m, and roof slopes of 15O,
20°, and 25O. Among other things, the researchers concluded that the
extent of direct diffuse radiation loss increases as the effective albedo of
the plant canopy increases and is more dependent on the slope of a roof
rather than its width or length.
Lau et al. ( 1984) verified theoretical total transmission factor. values
with actual data from shed-type and conventional glasshouses (Thble
1.1).Tho different concepts were employed in treating the sky-diffuse
radiation on a tilted surface, namely, the isotropic model and the aniso-
tropic model (Klucher, 1979). In the isotropic model the intensity of
sky-diffuse radiation is assumed uniform over the sky dome. The aniso-
tropic model approximates partly cloudy sky conditions and may vary
Table 1.1. Total Tkansmission Factor: Short-Term Simulation Results

vs. Actual Data"
Solar shed Control house
Week Experimental Simulatedb Experimental Simulatedb
Oct 8-14 1.58 1.62 (1.73) 1.11 1.18 (1.26)

NOV12-28 1.78 1.52 (1.65) 1.37 1.23 (1.33)
Dec 17-23 2.44 2.99 (3.15) 1.78 1.95 (2.08)
J a n 14-20 2.29 2.22 (2.35) 1.61 1.48 (1.59)
Feb 11-17 1.81 1.67 (1.78) 1.30 1.24 (1.32)
Mar 10-17 1.70 1.58 (1.71) 1.17 1.15 (1.25)
Apr 8-14 1.20 1.26 (1.36) 1.02 1.07 (1.15)
May 13-19 1.06 1.14 (1.22) 0.96 1.05 (1.12)
J u n 10-16 1.04 1.13 (1.18) 0.92 1.00 (1.05)
Jul 15-21 1.07 1.16 (1.20) 0.90 1.01 (1.04)
Aug 12-18 1.29 1.21 (1.28) 1.00 1.05 (1.11)
Sept 16-22 1.32 1.48 (1.57) 1.04 1.10 (1.17)
=From Lau et al. ( 1984).

bNumbers in parentheses indicate results using the anisotropic diffuse radiation
model (Klucher, 1979).
from clear skies on one extreme to entirely cloudy skies on the other. The
results suggest that the isotropic radiation model more accurately pre-
dicts diffuse radiation on an inclined plane when beam radiation dominates,
as it did when the data were collected.
A computer model was written by Chandra and Willets (1980) to
predict the thermal behavior of a 6.7 x 12.2 m fiberglass quonset green-
house attached to a 3 x 1.9 X 13.4 m external crushed granite rock storage
with equivalent spherical particle diameter and porosity of 1.91 cm and
4690, respectively. Comparisons of predicted and measured greenhouse
air temperature and relative humidity were made for two days. Supple-
mental heating on the first day was almost negligible, whereas considera-
ble supplemental heating was required on the second day. The mean
deviations of the predicted and measured values of temperature and
relative humidity were 0.7OC and 2.590,respectively.
Lau and Staley (1985)are developing a generalized simulation model
appropriate for designers and engineers who must explore and evaluate
different solar greenhouse design alternatives. The simplified design
methods being used are an extension of the f-chart method for active solar
space and water heating systems (Klein et al. 1976) or solar load ratio
(SLR)method for passive systems (Balcomband Hedstrom 1977).RRsults
to date have been verified by actual data and suggest that the monthly
solar heating fraction may be correlated with a dimensionless variable in
the form of a logarithmic function,
1, ENERGY-EFFICIENT GREENHOUSES 17
Rotz et al. (1979) undertook a computer simulation study of energy

requirements in conventional, insulated, and solar-heated greenhouses
and insulated greenhouses with solar heating. The modeled greenhouses
were multispan glass and double-polyethylene structures with a floor
area of 4000 m2. Insulating techniques included thermal screens and an
insulating, double acrylic cover for the glass structure. The solar heating
systems studied used either well-insulated or uninsulated external water
collectors, and an insulated external air collector or an internal air collec-
tion system. The individual energy savings (lhble 1.2) varied from 1361
(15%) to 5309 GJ (58%)of heat energy for the glasshouse. The system
that provided the greatest fuel saving was a combination of a heavy
thermal screen and insulated external air collector system. When used
with either the acrylic or polyethylene greenhouse, the savings were 8299
GJ (90%)of heat energy.
McCormick (1977) created a similar computer simulation model that
partially determined installed costs and life cycle operational costs to
produce an economic analysis and optimization study. The model ana-
lyzed the cost of heating a 26 X 30 m glass greenhouse in Ohio using an
external water collector. The results were compared with the cost of oil
heating at several different fuel escalation rates (Fig. 1.2). The effect of
insulating the roof and north wall was also included in the study. The
optimum solar heating system size occurred with a 465 m2 collector
supplying about 55% of the energy used by the greenhouse. It is interest-
ing to note that insulating the greenhouse decreased the collector utiliza-
tion, driving up the cost of the delivered solar heat.
Active solar heating systems are classified according to the type of
material being used for the thermal energy storages. The thermal stor-
ages usually contain sensible heat, which is transferred in response to a
temperature differential. Phase change material can be used to store
latent heat. This is the energy transferred a t constant temperature when-
ever a material changes from a solid, liquid, or gaseous state. In green-
houses, most sensible-heat storages are large masses of either water or
rocks. Sokhansanj et al. (1982)developed testing and evaluation proce-
dures that can be used to select the best storage system for individual
applications. The decision is influenced by the type of solar collector
under consideration, the greenhouse structure, and the location of the
greenhouse.
Van de Velde et al. ( 1982)experimented with a unique collector-distribution
system consisting of flat, black PVC tubes, which covered 35% of the
ground surface. During daytime, the tubes absorbed solar energy and
heated water that was stored in a basin. At night the pipes radiated the
heat back into the greenhouse. Connellan ( 1984)found that a warm-water
floor storage system a t 25OC provided satisfactory greenhouse tempera-
tures. Performance during autumn months was lower than expected
18 GORDON J. MONK AND ,J. M. MOLNAR
Table 1.2. Computer-Simulated Energy Use in Heating a 4000-m2 Commercial

Greenhouse with Fuel Conservation Systems
Glasshouse Double polyethylene

house
Fuel Energy Energy

conservation requirement SavingsC requirement Savings'
system (GJ) (%) (GJ) (70)
Conventional heat
Oil-fired boiler + unit 9090 6908
heaters
Insulation
Double acrylic cover" 5636 38 - -
Thin thermal blanketb 5381 41 4177 40
Heavy thermal blanketb 3781 58 2715 61
Double acrylic cover and 3686 59 - -
thin thermal blanket
Double acrylic cover and 2516 72 -
heavy thermal blanket
Solar heat
Uninsulated water
collector 699 1 23 4939 29
Insulated water collector 5573 39 3636 47
Insulated air collector 5307 42 3506 49
Internal greenhouse
collector 7729 15 5559 19
Insulation and solar heat

Thin thermal blanket
and uninsulated
water collector 2596 62
Heavy thermal blanket
and insulated air
collector - - 950 86
Double acrylic cover,
heavy thermal
blanket, and insulated
air collector 791 91 -
"Glass is replaced by double acrylic material.

*Thin thermal blanket, R 1 1, heavy thermal blanket, R II 10.
'Compared to conventional glass greenhouse.
though, because the polyethylene film collector did not achieve the
desired temperatures. However, a low-cost polyethylene flat-plate collec-
tor was able to provide significant heat collection in Arizona (Mears and
Baird 1976). The collector supplied a plastic water bag beneath a green-
22
/
18
-
z
3
14 - \
\
\
5 -
/
0 /'' $~s"~~~!ihouse)
15%
5. 12-
fA
h
p 10-
6- 5%
Fuel oil
0 Differential fuel
escalation rate
(Fuel inflation)
2000 4000 6000 8000 Collector area ( 1 1 2 )
0 02 04 06 08 10
Percentage of total energy supplied
Fig. 1.2. Comparison of energy costs for solar and no. 2 oil heating for a glass
greenhouse in Ohio. (From McCormick 1977.)
house bench. Above the bag an insulated arch was used to form a plenum
through which interior air was circulated. Montero and Short ( 1984)also
obtained efficient performance from a plastic collector. I t supplied a
warm-floor heating system that maintained a night temperature of 15.5OC
in an insulated greenhouse in Spain. Overnight low temperatures in the
floor-reservoir-heated solar greenhouses can be predicted within 1OC
using an equation developed by Mahoney et al. (1980). Low-cost solar
collectors can also be made with concrete and their performance will be
comparable to other inexpensive collectors according to Bartok and
Aldrich (1984). Damrath (1982) installed a collector in a greenhouse,
which also served as a heat exchanger for nighttime heating.
Dale et al. ( 1977)designed a solar system that used groundwater in soil
a t a depth of 2-3 m to store heat from an insulated collector. The system's
efficiency was very low, unfortunately, due to rapid heat loss to the
surroundings (Dale et al. (1980). A system incorporating an external,
insulated tank for warm water storage was tested by Baird and Mears
(1976),but it was not economically feasible. Royle (1981a)has described
a system of mud storage being developed for a solar greenhouse on Prince
Edward Island. In Israel, a plastic greenhouse was built over an artificial

pond (Anon. 1984a).Heat was transferred to and from the greenhouse by
spraying water in one corner of the interior through a shower system. The
shower absorbed solar energy during the day, decreasing temperature
and increasing humidity. On cool nights, the shower helped maintain
suitable crop temperatures.
Solar ponds are shallow masses of water exposed directly to the sun’s
rays. As the pond heats up, temperature gradients develop, causing a
mixing action to take place that reduces surface temperatures. For this
reason, salt is usually added to solar ponds in order to produce density
gradients that conteract the effects of the temperature gradient, thereby
preventing mixing from taking place. A feasibility study of linking solar
ponds to greenhouses by means of insulated pipes was completed by
Carnegie et al. ( 1982). The dynamic thermal behavior of a solar pond was
modeled by Shah et al. (1981a) to study the overall performance of a
greenhouse heating system. During testing, a brine-source electric heat
pump was incorporated into the heat extraction system. Initial results
indicated that the combined system improved the effectiveness of both
the heat pump and the solar pond (Shah et al. 198lb). Fynn (1981)
reported that from mid-December to March the average system perform-
ance factor was 1.98. For ten days in January the pond supplied virtually
all of the heat required. Computer modeling by Carnegie et al. (1984)
determined that in the warm climate of California a solar pond could
supply 77% of the annual heat load a t a favorable cost for a double-
polyethylene greenhouse equipped with a single-layer thermal screen
made of polyethylene with a reflective coating on one side.
Fiesearchers a t Rutgers University developed a solar greenhouse with
external polyethylene collectors and warm-water storage beneath a porous
concrete floor. At night, the warm water flowed down simple polyethylene
heat exchangers and returned to the tank via the floor (Roberts et al.
1976).Mears et al. ( 1978)reported that over a September-May test period
solar energy provided 53% of the heat required for a payback period of 4.5
years. A 0.54 ha commercial demonstration project including insulating
curtains was built to study the system on a large scale (Cipolletti et al.
1981a). Fuel consumption was reduced significantly, with the collector
field contributing more than 25% of the heating load (Cipolletti et al.
198lb).Overall savings of 45% were achieved during the first year, 73% for
the next full season, and 68% during the following spring.
Ingratta and Blom (1980a) installed the Rutgers system in southern
Ontario. The solar collectors provided an efficiency of only 7% for a
September-June period and 40-45% in July and August, when little
nighttime heating was required (Ingratta and Blom 1980b). The energy
consumption (natural gas plus electricity) in the solar greenhouse was
less than a control house. However, the cost was greater because the solar
house used more electrical energy to run the collector circulation pump
(Jackson 1983).
Wilson et al. ( 1977)mathematically modeled greenhouse thermal behav-
ior and described many advantages of using small-sized crushed rock or
gravel for thermal storage. Milburn and Aldrich (1979) designed an
interior collection system to pull air from a greenhouse ridge into rock
storages beneath the floor. On a typical day the collectors were exposed to
306 MJ of solar radiation, and the system was able to collect and store 132
MJ of the total, using 11MJ of electricity in the process. This represents a
system operating efficiency of 43.2% and a coefficient of performance
(COP)of 12, which means that 12 MJ of heat were collected for every 1MJ
expended to run a motor. In Florida, Baird and Waters (1979)stretched
shade cloth across the attic of the greenhouse and sealed off the plant
growth area with clear polyethylene. Air was circulated from the attic to
rock storages under plant benches, resulting in substantial energy sav-
ings and improved crop performance.
Thermosiphoning solar collectors were built into the lower half of the
south sidewall of a greenhouse with a steeply sloping north roof. Sewer
rock was used for thermal storage, and Riemers (1979) reported that
inside temperatures of 4O-27OC were maintained when outside tempera-
tures were below -18OC. Click and Pile (1980)supplied rock storages in a
double-polyethylenehouse in Tbnnessee with a flat-plate collector a t the
insulated north wall. The system was self-sufficientfrom late February to
late November. An external rock storage that provided fall savings of 31%
was tested by Willets et al. (1980). Umarov et al. (1981)used a pebble
storage to contain 24-2670 of the solar radiation entering the effective
greenhouse area. This resulted in overall savings of 35-55%.
Ebeling and Kranzler ( 1982)connected a solar hop dryer to an existing
fiberglass greenhouse and retrofitted underbench rock storages. Initial
performance indicated annual savings of 3770,yielding a payback period
of 10 years for Cyprus. Rheinlaender and Photiades (1984)found that a
water solar system provided greater energy savings than the rock storage
system they tested.
Staley et al. (1981) described the design and construction of a glass
shed-type rock storage greenhouse with adjustable low-cost black cloth
collectors hanging in front of an insulated north wall. The solar shed used
29% less energy than a control house during a September-April period,
resulting in an annual savings estimate of 40% (Staley et al. 1982a).
Collector efficiencies ranged from 32 to 52% and tomato-crop yields were
not significantly different in the solar shed and control (Staley et al.
198213).
The researchers also studied the performance of a modified double-
polyethylene shed-type rock storage greenhouse that was not equipped

with any collectors (Staley et al. 1982~).In terms of 1982 Canadian
dollars, the capital cost of the solar heating system was $9.65/m2.Annual
gas savings in the commercial tomato operation amounted to $2.98/m2
compared to a similar operation nearby, and $6.28/m2 compared to a
neighboring cucumber operation with a double-polyethylene roof and
fiberglass sidewalls.
Soil can also be used to store sensible heat if channels are provided for
circulating air through the medium. The storage design and its capacity
are dependent on several factors, including the surface area of the chan-
nels, the air flow rate, and the moisture content of the soil. The pressure
drop in channel accumulators, heat transfer coefficients,and local drag in
relation to Reynolds number were investigated by Kim et al. (1980).
Magnussen ( 1982)studied the use of soil for seasonal heating by focusing
on the interrelationships between greenhouse area, storage volume, and
system performance. Puri (1981) undertook an economic evaluation of
moist ground soil for storing energy for greenhouses and for drying corn.
System life cycle evaluations of costs indicated that both pipe diameter
and insulation thickness are fairly insensitive parameters compared to
collector and storage costs. For example, a nominal storage capacity of
two days can be supplied by a storage volume of 0.043 m3per square meter
of floor area with a collector area of 0.14 m2per square meter of floor area.
This solar heating system can supply 17% of the annual greenhouse
heating demand. A solar system with a collector area of 1.14 m2 per
square meter of floor area can supply 30% of the heating requirement but
is no longer economical. A south wall air fluid solar collector has been
used to heat crop soil, resulting in energy savings of 45% and normal
‘‘Ilopic’ tomato production (Dale et al. 1984). Despite this, production
with ‘Dombito’tomato was about 50% below normal and it appeared that
circulating interior air through the soil might eliminate the need for the
solar collector.
Using the greenhouse itself as a solar collector, surplus daytime heat
can be stored and released a t night by circulating air through a network
of buried pipes. Design guidelines have been recommended by Sasaki and
Itagi ( 1979),who suggested reversing the air flow direction in the pipes at
night to maximize the temperature of the recovered heat. Monk et al.
( 1983)incorporated this feature in two earth thermal storage (ETS)heat
exchange systems using motorized dampers. One system is installed in a
glass gable house with supplementary heating (Fig. 1.3)while the second
system heats a double-polyethylenequonset without backup heaters. Air
temperatures in the pipes and the resulting fluctuations in soil tempera-
ture were measured by Staley et al. (1983). Longitudinal temperature
stratification was not significant. Mean air temperatures differed by only
Inlet dUC1 Charginp Inlet ducl Oischarginy

Outlet duct Dischaigmy Outletd~cl Charplnp
I I
Fig. 1.3. Glass earth thermal storage (ETS).(From Molnar et al. 1984.)
lo-2OC at each end of the storage. Therefore, the cost of facilitating air
flow reversal was not justified on the basis of the increased discharge
air temperature.
Molnar et al. (1984) compared the system component operating effi-
ciencies in the glass ETS house with those in the shed-type solar green-
house mentioned earlier. Generally, the solar shed heating system oper-
ated 4% more efficiently than the ETS system, which provided total
annual energy savings of approximately 20% (Molnar et al. 1983). How-
ever, Arcus Consulting Ltd. (1985)combined the results with capital cost
data and maintenance cost forecasts and found that the ETS system is
more cost effective than the solar shed. The ETS system provided posi-
tive NPV with natural gas heating and interest rates as high as 1570,
whereas the solar shed required an interest rate of 5% or more expensive
oil heating to be cost effective. Another advantage of the ETS system is
that it can easily be retrofitted into any conventional greenhouse that
does not have a concrete floor.
Certain benefits can be derived by using phase change material to
provide latent heat storage but stability problems have plagued experi-
ments over the long term (nkakura and Nishina 1981).Calcium chloride
hexahydrate has a high level of fusion and appears to be one of the most
promising materials in terms of cost (Kern and Aldrich 1979).Cadier and
Jaffrin (1981) used this material in stacked bags under a greenhouse
floor; 6070 of the captured heat was used in the phase change thermal
processes while the remainder raised the temperature of the surround-
ing ground.
When sunlight falls on plants, about 1.5% of the energy is used for
photosynthesis and 40-70% is used to drive their transpiration processes.
One hectare of tomatoes will transpire more than 467,700 liters of water
annually, using about 13.1 trillion joules of solar energy (Anon. 1980a).
Butterworth and Morgan (1981),Jewett et al. (1984)and other researchers
are currently studying the effectiveness of using heat pumps to recover
the latent heat of vaporization of the water while dehumidifying greenhouses.
Wind energy is another form of natural energy that is being evaluated for
greenhouse heating using 6.4-m diameter turbine windmills ( O’Flaherty
and Maher 1981).A severe storm caused serious damage to the system
because of the large turbine diameter. After damaged parts were replaced
with strengthened ones, reasonably satisfactory levels of energy were
obtained. Krause ( 1984a)described fiberglass rotor windmills that gener-
ate electricity used for warming water to 16O-2loC for heating cyclamen
crops. The optimum windspeed is 10 m/s when the mill produces 55 kW.
Surplus heat is stored in a 5-km length of Alkathene pipe buried in the
shape of a coil at a depth of 15 m.
The decomposition of manure and sawdust has long been used to heat
cold frames. This technique has been applied in a highly insulated
greenhouse in Rnnessee that has a requirement of 50% less heat than a
similar double-polyethylenegreenhouse (lbuliatos 1983).A bin filled with
sawdust supplies winter heating amounting to 35% of the annual require-
ment. In spring the sawdust is mixed with sand and pine bark for potting
soil and in fall the bin is refilled to continue the cycle.
The effects of radically altering the structural configuration of a green-
house in order to enhance solar energy capture have also been studied. A
hemispherical solar greenhouse was shown to be 17% more efficient a t
intercepting light and required 10% less energy as a result according to
Begin et al. (1984).Brown et al. (1979)extended the growing season by 3
months in an unheated double-glazed aquaculture greenhouse by build-
ing an insulated north wall with a parabolic configuration. Winter light
was reflected into an insulated 18,000-liter aquaculture pond with a
surface area of 16.7m2. Petrescu et al. (1981)also used parabolic modular
collectors for greenhouse heating. The system supplies 1-25 kW of heat in
the form of steam a t ;0Oo-15O0C and 15 bars. An east-west aligned
gothic arch-type greenhouse with an insulated north sidewall has been
used in combination with wet-earth storage for a hydroponic aquaculture
greenhouse (Van Toever et al. 1982).The capital cost of this structure will
be amortized in 6 years a t interest rates of 16% based on reduced operat-
ing costs and revenues from the sale of fish and vegetables.
Solar energy can also be used to cool greenhouses using absorption
chillers. Stickford et al. ( 1984) reported on a commercial solar greenhouse
range in Saudi Arabia that is completely energy self-sufficient. All elec-
trical requirementswere supplied by an array of photovoltaic cells. Dumont
and Cachard ( 1984)described a solar greenhouse designed to provide sea
water distillation as well. Daily production of fresh water in excess of 2

liters per square meter of floor area was observed.
B. Waste Heat
Vast amounts of waste heat are available everywhere in the industrial-
ized world. In Canada, for instance, 82 sources of reject heat capable of
heating over 1000ha were identified (Van Die and Le Blanc 1983).Usually
the heat is in the form of warm water that has been used to condense
steam or cool machinery or manufactured products. As far as the green-
house industry is concerned this source of low-cost energy poses two
main problems. More often than not, the source is a considerable distance
away from the greenhouse site. Second, the heat is usually available only
a t low temperatures in the 25O-35’c range, although O’Flaherty and
Maher (1979)have described a process by which power plant condenser
water can be delivered a t 90°C. The scheme is based on the use of steam
bled from the low-pressurestage of one of the plant’s turbines. This steam
is passed to a heat exchanger in which its heat is transferred to the
greenhouse heating water. The technique involves the sacrifice of a small
proportion of the plant’s electrical output.
Energy can be recovered from low-temperature water by increasing
flow rates through a distribution network of pipes that have a large
surface area to facilitate heat transfer. Sometimes it is cheaper or more
efficient to use a heat pump to elevate the water temperature before it
enters a conventional distribution system.
A 500-m2double-polyethylene greenhouse was heated for two years
using four water-to-airheat pumps with reject heat as the source (Rotz et
al. 1981). The heat pumps were operated a t a lower cost than with
conventional natural gas heating. The most critical parameter in the
economic analysis was the life of the heat pump equipment. If the life
dropped from 20 to 10 years, due to compressor failure, for example, the
system would no longer compete with natural gas.
The difficulty of transporting waste heat is not so easily solved. A
large-diameter, insulated pipeline is required to carry the water over long
distances. For example, in Romania, a pipeline carries water from a power
plant to an 80-ha operation 6 km away (Van der Horst 1972).Insulation 18
cm thick ensures that the temperature loss during transmission never
exceeds 1OC. Since transmission lines are very expensive, a large area of
greenhouse must be connected in order to make a project feasible. In
countries such as Holland and Denmark, this is not necessarily a prob-
lem because their greenhouse industries are highly concentrated. How-
ever, in North America, the dispersed markets and widespread availability
of land in many areas have resulted in isolated greenhouse development.
26 GORDON 1. MONK AND 1. M. MOLNAR
Despite this, many waste heat feasibility studies have been undertaken
and low-temperature heating experiments are ongoing. Some projects
have been completed and both the utilities supplying the heat and the
operators involved report they are satisfied with the mutual benefits they
have derived.
Gillham (1974) reviewed the Romanian system to see if thermal dis-
charge water in Ontario could be used by the greenhouse industry there.
As a result, specific recommendations were established that provide a
framework for development of waste heat utilization. Meekhof (1977)
conducted a similar study applicable to Michigan and concluded that
waste heat utilization will be a least-cost alternative to a utility only if the
consumers own the system or pay rent.
The best sources of waste heat are nuclear power plants, chemical
plants, and refineries according to a report by Resource Management
Consultants Ltd. (1979). Thermal power plants, pulp and paper mills,
and steel mills were identified as being less attractive sources based on
reliability of supply, characteristics of waste heat recovery systems, and
the associated costs. The report also predicted that waste heat green-
houses will become significantly competitive only after the year 2000.
Schaupmeyer ( 1981) described the various systems available to recover
reject heat including subsoil piping, fin tube heat exchangers, direct
contact exchangers, medium transfer, shell heating, and surface heating.
Friday (1982a) in an independent economic assessment of waste heat
greenhouses concluded that significant economies would be achieved if
the reject heat delivery system is sized to handle only a portion of the
maximum greenhouse heat load; otherwise, the capital costs associated
with the delivery piping would be excessive. Olszewski ( 1977)studied a
bimodel system incorporating evaporative pads to heat a greenhouse in
winter and cool it in summer. Operating profits occurred as long as the
thermal effluent remained above 26.7OC.
Schisler and Bakker-Arkema ( 1975) evaluated a mathematical model
applicable to soil warming achieved by circulating waste heat in a buried
pipe grid. Crop yield models were developed for pea bean, soybean,
tomato, and sweet corn. The parameters used in these models were found
to be adequate for calculating a least-cost present-value criterion designed
to evaluate and select agricultural uses of waste heat.
Computer simulation was used to design a 1.1-hagreenhouse utilizing
24OC water from a power plant (Manning et al. 1980;Manning and Mears
1981). The greenhouse featured a flooded floor system that dissipated
heat from an underground rock-water heat storage up through a porous
concrete floor. Additional heat exchange area was supplied by heat
exchangers that were constructed by draping plastic over a distribution
pipe that circulated the warm water. A prototype was built using thermal
screens to reduce heat loads by 30% or more (Manning et al. 1981).The

results showed that an evaporative cooling system can be used to cool
thermal effluent in summer as well as inside air temperatures. Mears et al.
(1982)observed that when the warm water from the plant was at normal
temperature (unspecified),the reject heat could provide all of the energy
required. I t was also found that the large thermal mass of the flooded
floor was capable of meeting the heating requirements for many hours
without heat input.
Roberts et al. (1980) compared the effectiveness of the flooded floor
system with another system that circulated effluent through polyethyl-
ene pipes embedded on 15-cmcenters in porous concrete. Mats, normally
used in skating rinks to form ice, were then placed on the floor. Overall, it
was found that the dry-floor system was cheaper to install but delivered
50% less heat (Roberts 1983). Tbrkewitsch and Brundrett (1978)evalu-
ated the suitability of the flooded-floor system for Canadian conditions
using computer simulation. They also examined conventional fin tube
multirow heat exchangers and a system that circulated greenhouse air
between sealed troughs of warm water beneath the greenhouse floor. The
capital cost of the flooded floor system was lowest but the heat transfer
rate was insufficient by about 70%. Heat exchangers offered no storage
capacity and had high operational costs due to the fan power required.
The third system was judged to be the most attractive, particularly
because of its storage capacity, which allowed for 2-4 days of waste heat
supply interruption. Nevertheless, heat exchangers are being used to
distribute heat from power station cooling water a t 27O-35OC to an 8-ha
greenhouse range in England (Royle 1981b).
Madewall et al. ( 1975) described a 0.2-ha experimental greenhouse
complex in Alabama designed to compare heat exchange systems using
power plant condenser water. Because of the relatively low water tempera-
ture available, initial tests were conducted using simple contact heat
exchangers. Sensible heat was transferred from warm water flowing over
aspen fiber pads to recirculated air a t saturation. Based on the results,
Carter and Pile (1982)reported that condenser cooling water a t a mini-
mum 21OC can be used by direct-contact heat exchangers for tomato
production when ambient temperatures are as low as -11.1OC. If con-
denser cooling water temperatures fell below 2loC, supplementary heat
was necessary for optimum production. Tomato yields ranged from 5.5 to
6.5 kg per plant in the zones heated with waste heat, compared with 6.9 to
7.5 kg per plant in a conventionally heated zone.
A reject heating system in Minnesota incorporates underground tub-
ing and a forced-air system (Widmer 1979).The design criteria included
the use of 29.5OC thermal effluent, a minimum outside air temperature of
-34.5OC, and a tolerable double-polyethylenegreenhouse temperature of
10°C. Subsequent winter lows were 10% lower than average and the
greenhouses were heated with minimal difficulty. Ashley ( 1979) consid-
ered the economics of the operation and found that the total capital and
operating costs associated with waste heat delivery were $3200 per
hectare (1978 U.S. funds). At that time, the system was also providing
savings of $2000 per hectare based on no. 2 fuel oil costs of $0.12 per liter.
Ashley predicted that fossil fuel costs would rise in comparison to waste
heat costs and found the overall economics to be satisfactory at that time.
Drakes (1980)compared water contact heat exchangers with fin tube
radiators. Heat losses from houses with evaporative heating were 25-40%
higher than with the fin tube radiators. Thermal effluent can also be
circulated through plastic “Q mats,” which are placed in the floor in the
growing area (Gallagher 1982).Plants can be grown between the mats or
through holes fabricated in the mats. With a waste water temperature of
3OoC and an outside air temperature of -4OoC, the air temperature in
double-polyethylene quonsets could be maintained at 10°C. The mass
transfer unit in the ‘Tilacell”system for heat exchange (Shawet al. 1976)
proved to be an efficient heat exchanger, but the resulting high humidities
caused mechanical failures and horticultural problems. Furthermore, the
system cost 300-800% more than conventional systems in terms of capi-
tal and operating expenses, which could not be justified by the savings
achieved.
One of the simplest and least expensive methods of extracting heat
from thermal effluent is to stream the water over the exterior surface of a
greenhouse. The system has an advantage in that greenhouse humidity
remains constant because the interior air does not contact the water. If
the water is cooler than the greenhouse but warmer than the outside air,
the heat transfer between the water and the greenhouse is negative, but
the surface flow can still be beneficial since the water acts as an insulator
(Walker et al. 1981). In extremely cold climates, external fogging and
ice and snow build-up may adversely effect light transmission ( Schaup-
meyer 1982).Under these conditions, the surface flow itself would likely
be impeded.
Walker et al. (1981) found that the water flow rate is optimized by
balancing the amount of energy required to pump the water to the ridge of
the greenhouse against the heating benefit derived from the water. Equa-
tions were developed that describe the optimum flow rate. Experiments
conducted with a small greenhouse in Illinois by Walker ( 1978)indicated
that 3OoC water with a flow rate of 0.094 liters per square meter of
greenhouse area was sufficient to heat the greenhouse to 15OC when the
outside temperature was -6OC. Computer simulation has been used to
compare the heating and cooling requirements of conventional and surface-
heated greenhouses maintained a t 18'C daytime and 16'C nighttime

(Lazarus et al. 1981). Results indicated that a surface-heated greenhouse
requires 55.9% less energy when thermal effluent temperatures vary
between 13' and 25'C. If effluent temperatures were raised 3'C, energy
savings of 72.7% were predicted. A double-layer polyethylene-covered
greenhouse was also studied (Walker et al. 1982). Thermal effluent a t
8'-2OoC provided savings of 22.5%, while water a t 18'-3OoC provided
savings of 80.4%. White (1985)reported that effluent a t 10'C does not
freeze when outside temperatures drop to -29'C as long as the surface
flow over a greenhouse is not interrupted.
The feasibility of using waste heat sources other than power plants for
greenhouse heating has also been investigated. Iverson et al. (1978)
recommended that a heavy-water plant in Nova Scotia be used to develop
a viable greenhouse industry. In New Brunswick, utilization of reject
heat generated by the regasification of Algerian natural gas has been
considered (McBean et al. 1979). Dalton (1982) described a hot-water
recovery system supplied by an ethylene plant. The thermal effluent
arrives a t the 0.8-ha greenhouse range a t a temperature of 40.5'C. Two
525-kW boilers maintain the temperature and supply a conventional
overhead pipe distribution system. In Saskatchewan, Maginnes and
Green (1983)found that 32'C water from an oil refinery was sufficient to
heat a double-glazed greenhouse if a reflective thermal screen was used to
reduce night time heat 10ss.
Friday (198213)examined the use of exhaust gases from pipeline com-
pressor stations in the United States as a source of high-temperature
( 6Oo-77'C) thermal effluent. This concept is being tested in Canada (Ho
Lem 1979). %NO 2500-kW reciprocating engines were selected for waste
heat recovery because of their nearly continuous operation on an annual
basis. Back pressure on the engine limits maximum heat recovery to 50%.
The 370'C exhaust from a compressor station jet engine has been mixed
with outside air and blown between the layers of polyethylene on a
greenhouse roof (Anon. 1981a).The system was predicted to pay for itself
within two years. Condensation has been eliminated and COZ from the jet
exhaust passes into the growing area until levels of 5000-6000 ppm are
reached, while nitrous oxides are filtered out by the polyethylene.
In Scotland a distillery supplies thermal effluent from condensers a t
5Oo-7O0C to a 0.2-ha range (Sheard 1980).The water is pumped directly
into a conventional piped system. A distillery in Ireland also supplies hot
water to a 0.7-ha range (Dixon et al. 1982).However, the effluent tempera-
ture fluctuates from 20' to 80'C. A 55,000-liter hot-water storage tank
and calorifier system are used to maintain an input water temperature of
70'C.
Waste heat can be obtained from nonindustrial sources. In West Germany,

methane gas from an 8-ha refuse dump is used to generate 58 kWh of
electricity (Krause 1984b). The reject heat is being used economically to
heat a small pot plant nursery. Heating costs at a 40,000-m2greenhouse
operation outside Hamburg have been reduced by two-thirds overall by
using condenser water from one of the city’s garbage incinerators (Anon.
1983).Waste heat from another of the city’s garbage incinerators has cut
heating costs in half compared to oil (Krause 1984~). Using garbage
incinerators to supply waste heat to greenhouses makes sense because
incinerators and greenhouses are both usually located close to cities.
Biomass burning is another method for greenhouse heating ( McDonnell
1985). A system in California combusts peach pits from a local canner
to produce 293 kW. The initial investment was returned in less than
one year.
C. Geothermal
Geothermal heat has been successfully extracted from groundwater,
hotsprings, soil, and deep-mine air for greenhouse heating. Hot-water
reservoirs ranging in temperatures from 52O to 93OC are widely distri-
buted across the western United States and Canada (Icerman, 1983).
Depths vary along with flow rates. Some sources supply water sufficient
for a 23,000-m2greenhouse. Unfortunately, these sources are usually in
very remote areas, unsuitable for the establishment of greenhouse opera-
tions. Furthermore, geothermal waters can be corrosive or contain dis-
solved minerals that make them unsuitable for heat distribution systems.
Nevertheless, geothermal fluids are being used to heat greenhouses in
California (Boron, 1979)and Israel (Anon. 1984b).
Pasternak and Rappeport (1982) outlined the use of soil warming,
direct-contact systems, forced-convection systems, and water curtain
systems for distributing geothermal heat. In Utah, a 120-mwell has been
tapped that produces 39OC water a t a rate of 3800 liters/min (Anon.
1981b). The greenhouse operators grow roses and estimate that the
investment will be paid off in 5 years.
Groundwater at 2OoC has been circulated through pipes in root zones
under winter propagation structures (Regulski 1983). Soil temperatures
of 10°C were maintained when outside temperatures dropped to -8OC,
with energy savings of 40-5070.Greenhouse heating usually requires
warm temperatures, which means that heat must be extracted by means
of a heat pump. However, heat pumps have been used in France to heat
groundwater to only 2OoC (Lawson 1985). The 2OoC water adequately
heated a tomato crop when it was circulated through a water mattress or
small-diameter pipes lying on the surface of the soil. Energy savings of

60% were reported.
Groundwater can be pumped directly into an evaporator heat exchanger
unless there is a danger of freezing in the external loop (Lawson 1981a).
In that case, the heat is extracted from the groundwater by means of a
closed loop containing an antifreeze solution. Lawson ( 1981b) reported
that this type of extraction system removes heat from soil at a location
where, below a depth of 1 m, the temperature remains constant a t 8OC.
Holes 10 m deep were drilled in line with one another. Loops of nylon
tubing with refrigerant running through them were placed in the holes
and connected in series. I t was estimated that for a 15 x 37 m greenhouse
cropped with tomato and originally heated by a conventional boiler
system, the payback period would be 5 years. Fuel input would be halved
compared with oil heating, and this benefit would continue long after the
installation was paid off.
Spieser (1983)described a commercial tomato operation that partially
heats a 9 X 71 m double-polyethylenegreenhouse with three heat pumps
that use 15OC groundwater as a heat source. Each heat pump uses 2
liters/sec of water, reducing its temperature by 4OC before returning it to
the aquifer. The heat pumps move air at a rate of 2300 literdsec each,
raising its temperature from 2OoC to 31'C. Costs for heating ranged
between 41 and 71%of heating with natural gas, resulting in positive cost
effectiveness predictions.
In Ontario, water-to-water heat pumps are being used to maintain soil
temperatures in a greenhouse a t 15OC for the production of Alstroemeria
(Anon. 1985b). By using three 5-ton heat pumps, 1020 m2of flower beds
are heated and cooled through ground loops.
Another source of geothermal energy is air in thermal equilibrium with
temperatures underground. A particular study is being devoted to air
from deep mines, where temperatures remain constant (13O-16OC)and
oxidation provides an increase in COz. Duncan and Walker (1981)stated
that in the United States such a greenhouse requires 8.4 times less energy
than conventional heating. Air from a deep coal mine in Kentucky was
used to ventilate a greenhouse a t a rate of one-half to one air exchange per
minute (Walker et al. 1976). Greenhouse temperatures were 2.8O-1.6'C
lower, respectively, than ideal set-point. Buxton et al. (1979) cautioned
that mine air ventilated greenhouses may have relative humidities near
100%constantly during the winter. Therefore, these greenhouses may be
suitable for horticultural production from mid-February to November
only (Buxton et al. 1977). Carbon dioxide levels reached 2500 ppm and
toxic gases were not detected. However, in other locations, toxic gases
could be a problem.
IV. ENVIRONMENTAL MANAGEMENT
A. Root Zone Heating

Root zone heating is accomplished by distributing heat on or under a
propagation bench, raised bench, or soil bed. This strategy reduces
heating costs for two basic reasons. The heating system does not have to
build up heat in the greenhouse peak so that it moves down toward the
thermostat and plant canopy. Instead, heat rises up through the crop
reducing the load on the system. Second, researchers have found that a
variety of crops produce normally in reduced air temperatures if their root
structures are embedded in warm soil. Lower set-point temperatures can
reduce heat loads significantly. Stefanczyk ( 1984)has discussed the pros
and cons of bottom heating while describing three commercially avail-
able heat distribution systems.
Shen and Mears (1977)used dynamic computer simulation to investi-
gate the feasibility of heating double-polyethylene greenhouses with
warm water on the soil. Results indicated adequate temperatures would
be achieved only if a thermal screen was used to release nighttime heat
load. The performance of such a system is influenced by pipe diameter
and spacing, floor depth, greenhouse air temperature, hot-water tempera-
ture, water flow rate, and plant canopy density. Puri (1982) generated
steady-state design curves that consider these variables for the determi-
nation of long-term thermal performance. Parker et al. (1981) chose to
create a dynamic soil model including temperature-induced vapor flow
and soil-water-potential-induced liquid flow. It allows for variable soil
thermal conductivity, hydraulic conductivity, thermal capacity, soil water
potential, water vapor diffusion coefficient, and water vapor density.
Steady-state, constant-property, soil-warming models for three pipe con-
figurations were also generated by Merbaum et al. (1983) and were
presented in dimensionless graphic form. A procedure for establishing
pipe depth, spacing, and water flow rate to fulfill root zone heating de-
mand is described.
The effect of thermal effluent heating on temperature, heat flow, and
moisture distribution in sand, a peat-vermiculite mixture, and Wooster
silt-loam was examined by Roller and Elwell ( 1980).Water inlet tempera-
tures of 25', 30°, 35O, and 40'C were used to provide a range of informa-
tion that would cover temperature levels normally encountered in power
plant condenser water. I t was determined that depending on soil type and
heating water temperature, up to 40% of the annual greenhouse heat load
can be supplied by root zone heating under Ohio conditions (Roller and
Elwell 19811. In addition, lettuce growth response was dramatically
improved when soil temperature was maintained above 17'C, even when
the minimum nighttime air temperature was lowered below normally

accepted values. This resulted in a 25% reduction in fuel consumption.
Canham (1976) reported on experiments with tomato crops showing
that a reduction in air temperature of 5.6'C can be compensated by
warming the soil to about 25'C. Energy savings of 48% were measured by
Ingratta ( 1978)when tomato root temperatures were maintained a t 24'C,
while night temperatures were reduced from 16.5' to 10'C. Gross dollar
returns per plant equalled or exceeded returns from the standard crop-
ping conditions. A t normal air temperatures, soil warming increased
total tomato yields by 40% in the spring and 8%in the fall (Gosselin and
Trudel 1984). Soil warming caused greater effects on tomato production
in fall when night air temperatures were reduced. Cucumber yields were
not significantly affected by soil warming when grown in fall but were
increased by 36% in spring. Hurewitz et al. (1984)found that root zone
heating of tomato seedlings increased growth, basipetal photosynthate
translocation, photosynthesis, and 32Puptake but did not compensate for
lower air temperature (16'C). Nontheless, Janes and Giacomelli ( 1983)
pointed out that soil heating during spring bedding plant propagation
speeds germination and seedling development, allowing later starting of
the crop. This eliminates two to three weeks of greenhouse operation
during the coldest time of the year.
The effects of root zone heating in crops of calceolaria were studied by
White and Biernbaum ( 1984). From December 1979 through March 1980
a control and a soil-warmed crop were exposed to night break lighting to
accelerate flowering. Root zone heating of 2Oo-22'C increased fresh and
dry shoot weights, improved development, and increased the number of
flowers. The treatment was less effective on a second set of crops grown
from March through June without lighting.
Zeroni et al. (1984)found that a decrease in night air temperature from
18' to 6 1'C was tolerated by 'Sonia' rose plants without loss of yield or
quality when roots were held a t the optimum temperature of 21 * 1'C.
Moss ( 1982)reported that cultivation of roses a t a root zone temperature
of 25'C with low night temperatures of 12' and 9'C represented a
considerable energy saving. However, the use of root warming a t a higher
night temperature of 18'C would be preferred because of higher yield and
more constant production, resulting in reduced energy consumption per
bloom produced.
l b o major types of commercial root zone heating systems have been
studied in the United Kingdom (Zondag and Brugger 1985). The first
system used 6-mm tubes made of EPDM, a synthetic rubber that is also
used to make spark plug wires, outdoor high-tension wires, and gaskets
to seal airplane windows. The EPDM tubing was spaced a t 4-cm centers
on benches and developed hot spots due to improper bleeding of the lines.
34 GORDON J. MONK AND 1. M. MOLNAR
In bedding plant trays, media temperature varied as much as 10°C,

depending on the location of the feed and return lines. This was elimi-
nated by covering the tubes with wet sand. If the sand had a chance to dry
out though, the hot spots reoccurred. The tubing could heat the entire
greenhouse space if water temperatures were raised but this had an
adverse effect on plant growth. A second system used 19-mm tubing on
30-cm centers. This system was cheaper to install and heat distribution
was satisfactory.
B. Alternative Heating Methods

Infrared ( IR)heating systems combust fuel at very high temperatures,
resulting in increased efficiencies. The hot exhaust gases are circulated
through overhead pipes. Radiant heat transfer raises the temperature of
crops and other interior surfaces directly rather than warming the air.
This would suggest that air temperatures can be lowered, resulting in
reduced heating requirements.
Several experiments have compared the energy consumption of IR
systems with conventional systems but the results have varied tremendously.
Knies et al. ( 1983)grew tulip and lily with the NOR-RAY-VACIR system
manufactured by Robert Gordon Appliance Inc. and the Vitotherm IR
system manufactured by Vito Technieken. In the latter system, insulat-
ing material with decreasing thickness and retarders are inserted in the
pipe. This measure evens out temperature distribution along sections
having a maximum length of 20 m. A low-overheadhot-water pipe system
was used as a control. The net energy consumption of the Vitotherm
system was 6%lower, while the NOR-RAY-VACsystem provided savings
of 13%. Both IR systems distributed radiation intensity poorly but
consistent air temperatures were realized. The conventional hot-water
boiler efficiency was 76%. Burning efficiencies of the NOR-RAY-VACand
Vitotherm (with flue gas condenser) systems were higher a t 86 and 8870,
respectively. Blom and Ingratta (1981)tested the CO-RAY-VAC IR sys-
tem manufactured by Robert Gordon Appliance Inc., with roses and
chrysanthemums. Rose stem productivity did not appear to be significantly
affected and energy savings of approximately 15-20% were recorded.
Chrysanthemums showed an 18% fuel saving and soil temperatures
remained normal.
The performance of IR heating in a glasshouse was compared to that of
conventionally heated glass and double-polyethylene houses by Heins
and Rotz (1980).Poinsettias, mums, and geraniums grew better in both
glasshouses than in the double-polyethylene houses. The IR heated
glasshouse used 30% less fuel than the glass control house, while the
double-polyethylene house used 40% less fuel than the glass control
house. The results were included in an economic study that evaluated 12
energy conservation methods including thermal screens and double acrylic
glazing (Rotz and Heins 1980).For plants that grow well under reduced
light conditions, the most economical system was a double-polyethylene
structure with unit heaters or inflated polyethylene over glass. For more
light sensitive crops, the most economical technique was IR heating in
a glasshouse.
Youngman ( 1983)installed CO-RAY-VACIR burner units in a 14,400-m2
range growing pot plants on benches. The IR heaters used 62% less fuel
than conventional gas-fired unit heaters with polyethylene convection
tubes. Electrical energy consumption was reduced by 70%. The combined
energy savings were sufficient to pay the entire cost of equipment and
installation in two years.
Heat pumps can be used to extract heat from outside air (Royle 19831.
In the case of the gas-engine-driven heat pumps studied, the efficiency
was 124%a t an outside temperature of -1OOC and 152%a t 18OC. It was
noted that the capital costs of each heat pump can usually be recovered in
five years. Heijna (1976)reported on the proper operation of condensing
heat exchangers designed to recover heat from the stack gases of natural-
gas-fired boilers. The dew point of the gases depends on the COZconcen-
tration and varies between 55' and 58OC. To obtain reasonable recovery
of the latent heat available, the temperature of the gases leaving the
exchanger should be considerably lower than 55OC. Lawand ( 1983)exam-
ined the possibility of using high-pressure sodium vapor lamps to heat
double-glazed and insulated greenhouses. The lighting systems were
installed with an electrical load of 125 W/m2, enabling them to provide
40-100'70 of annual heat load depending on the structure. However, the
high capital investment could not be justified on the basis of energy
savings alone. Tomato yields from traditional operations without lights
are about 20 kg/m2. If these figures can be doubled to 41 kg/m2 using
supplementary lighting, it will be economically feasible.
C. Environmental Control
Modern greenhouse operations are large and complex. Environmental
control equipment often includes such equipment as hot-water mixing
valves, lights, watering systems, nutrient injection systems, misters,
ridge ventilators, fans, and evaporative pads. In the past, this equipment
has been controlled by individual timers, humidistats, and thermostats.
Many of these devices can be installed in a greenhouse range leading to

complicated wiring connections. Furthermore, their dynamic character-
istics force the operator to contend with set-point drift, changing temper-
ature differentials, coarse scaling, and balance. In fact, if the operation of
the different types of environmental equipment is not integrated properly
they can start functioning a t odds with each other. This results in inac-
curate control and a lot of wasted energy.
Greenhouse control computers solve these problems by providing much
greater control capabilities in terms of capacity, speed, and flexibility.
Complex crop blueprints can be implemented efficiently through opti-
mum coordination of the environmental equipment. Bryenton et al.
(1983) reported that adoption of computer controls will provide energy
savings of 10-1570as a result. The associated payback periods for ranges
2300 m2 and larger will be from two to three years.
Several user-friendly computers are now on the market and many large
operations have installed them. lhntau ( 1982)described the development
of a greenhouse climate control system using a commercial microcomputer
based on the Motorola 6809 microprocessor. The programs use self-
tuning control algorithms, and a simplified mathematical model is in-
cluded to calculate the influence of outside weather conditions on the
inside climate.
live microprocessor-based systems have been designed to control the
collection, storage, and use of solar energy as well as conventional control
equipment. The system developed by Robinson and Kranzler (1982) is
built around a Motorola 6802 control processor unit. I t controls the
transfer of solar energy from a collector to a rock storage and into the
greenhouse by operating air circulation fans. Data acquisition is also
performed with the provision for remote monitoring and data transfer.
Van Zinderen Bakker et al. (1983)and Monk et al. (1985)have described
a commercial greenhouse computer system with user-accessible soft-
ware, which allows the control algorithms to be altered for control of
shed-type rock storage and earth thermal storage solar greenhouses. All
input/output signals are distributed via field modules located close to
the area to be controlled, thereby reducing wiring costs, maintenance,
and complexity. The computer has an expandable capacity totalling 512
analog input channels, 2048 digital inputs, and 256 digital outputs.
Temperatures in all environments have been maintained within half a
degree of each other. The results indicate that the computer will increase
efficiency of the solar systems by about 20% while reducing maintenance
and greatly decreasing manpower requirements.
Advanced control software makes it possible for researchers to experi-
ment with new heating and ventilating strategies designed to increase
crop production and maximize energy efficiency. For instance, air temper-
atures can be lowered for the latter portion of the night without affecting
the development of certain crops. Physiological studies suggest that
"split-night'' temperatures may be feasible since most nighttime growth
processes can be sufficiently completed during the first few hours of
darkness as long as temperatures are warm. Thorne and Jaynes (1977)
report that eight cultivars of chrysanthemums and lilies, as well as
marigolds and petunias, have been grown under a split night temperature
regime of 16OC until 11:OO P.M. and 7OC until 6:OO A.M. No reductions in
the size, appearance, or stage of development of any plants were observed
when compared to those grown under a traditional warm-night regime. In
February, energy savings of 6% were achieved due to the reduction in
temperature of 9OC over a 7-hr period each night. Fuel savings of about
20% will be achieved in Connecticut during the January-April period if
greenhouse temperatures are maintained a t 16OC for the first part of the
night and then dropped to 7OC for 8 hr (Anon. 1980b). Tomato plants
grown under the split-night regime grew slightly slower, but reached the
same size as plants grown under warm-night conditions. Fruit yield was
12% lower from plants subjected to the split-night temperatures but most
tomato plants in Connecticut are grown in greenhouses for spring
transplanting. Therefore, effects on yield of transplanted crops would
probably be less significant.
Seginer and Raviv (1984)described a method of estimating the most
economical constant nighttime temperature for a particular crop: it
requires growth, engineering, and economic data. Growth chamber data
on tomato seedling temperature sensitivity was used to predict that the
most economical night temperature is the mean of the physiologically
optimal night temperature and the outside temperature. Split-night tem-
peratures of 20' and 12OC and constant night temperatures of 20°, 16O,
and 12OC were tried with cucumber crops (Van de Vooren et al. 1978).
No temperature influence on the rate of production could be recognized in
this experiment, suggesting that the low night temperature of 12OC be
used for production. Kooistra (1984)has reported the results of several
other temperature experiments. Energy savings with cucumbers were
achieved when lower night temperatures were introduced after the cucum-
bers started the productive stage. Night temperatures for sweet peppers
could be reduced from the moment of planting if day temperatures were
increased. Large numbers of experiments with tomatoes did not produce
recommendations for altered temperatures but some reductions have
taken place in commercial practice.
O'Flaherty and Maher ( 1981) reported unacceptable heat losses when
humidistats were used to control humidity in double-polyethylenegreen-
houses with fan ventilation. This problem was overcome by using a

computer to place an upper limit on the number of minutes in any one
hour during which the fans could be used to control humidity.
V. FUTURE DEVELOPMENTS
Energy efficiency in greenhouses can be accomplished by ( a )increas-

ing production on a unit floor area basis or ( b) reducing energy consump-
tion. This chapter has not examined the challenge of obtaining higher
crop yields since recent research has concentrated on reducing heat loads,
finding alternative energy sources, and testing infrared heating methods.
However, the advent of computerized control and better knowledge of
crop blueprints is shifting the focus of research toward intensifying
indoor horticultural practices.
At the present time, control computers can only react to changes in
temperature, humidity, or light levels. Flexible control strategies can
only be adopted after we learn more about plant requirements and the
dynamics of growth processes. This will be done by computer simulation
using intricate growth and climate models. By understanding the influ-
ence of environmental factors on growth and development, the green-
house environment can be adjusted to compensate for climatic deviations.
This type of sophisticated climate control is essential for achieving
improved crop quality and yields.
Interest rates will vary over the foreseeable future but the return to low
rates of 5-770is unlikely. Therefore, different approaches must be taken to
energy conservation that do not require the operators to undertake large
capital investments. For example, breeding research should be expanded
to develop new crops or cultivars that can be grown a t lower tempera-
tures. Low-energy cultivation techniques must also be studied. This
should include hydroponic systems with root zone warming that do not
produce excessive humidity levels.
Alternative energy sources will be used more extensively whenever
possible, especially if fossil fuel costs rise dramatically again. However, it
is clear that problems of availability and climatic factors will continue to
limit the use of geothermal, waste heat, and solar energy. The develop-
ment of low-temperature crops could lead to major breakthroughs in this
regard. The efficiency of solar heating systems could be made more
efficient if control computers could predict weather conditions over the
short term. This may be achieved by developing relatively inexpensive
barometric pressure sensors that will supply information to complex
control algorithm software in conjunction with radiation and tempera-
ture data. Alternatively, the day may come when weather forecast agen-
cies will have central computers designed to feed forecast data to

environmental control computers. By anticipating the loads placed on
the internal environment, the control computer could adjust solar heat
storage conditions in advance to maximize system effectiveness. In the
meantime, more work is needed to make existing heating methods more
energy efficient.
Higher crop yields will also be attained by increasing crop density. In
Europe double and triple layer transport systems are being installed in
greenhouses producing pot plants. The effect of alternately exposing the
plants to artificial and natural light requires further study. The design of
the transport system must be improved as well to make them cheaper and
more reliable.
In the future, greenhouses will become more like modified growth
chambers. The use of double glazings will increase and supplementary
lighting will be used more extensively where sunlight poses limitations
on growth. Since heat loads are being reduced, lights will also contribute
more significantly to the heating requirements of the structure. Rational
operation of lights and thermal screens will be interfaced and screens will
be deployed whenever the insulating benefits exceed the costs associated
with operating the lamps.
Finally it is clear that individual growers must bear the responsibility
of adapting the results of research to the set of conditions imposed by
the nature of their own operations. This requires that the operators be-
come more familiar with the interrelationships between climate and
crop development.
VI. SUMMARY
The high cost of heating greenhouses has led to many developments

that have greatly improved the energy efficiency of this intensive form of
horticulture. Inflated double-polyethylene glazing is now being used
extensively in place of glass wherever climatic conditions and crop require-
ments permit. Glass continues to be a popular glazing material because
of its superior light-transmitting properties. However, in temperate cli-
mates its use is almost invariably accompanied by the operation of
thermal screens for reduced nighttime heat losses. Growers regard double-
layer inflated films as an attractive alternative to glass because of their
low cost and high R-values. The durability and light transmittance of
these materials are continually being improved. The use of more durable
and expensive double-acrylic materials is becoming more widespread in
cases where operators can afford the investment. However, reduced light
levels may justify the use of high-intensity growing lamps if electricity

and energy costs are not prohibitive.
North wall and perimeter insulation is now standard in most locations.
However, its use is limited by light level considerations. Reflective sur-
faces can be put on inside surfaces including the floor to reduce loss of
light greatly.
Thermal screen installations are increasing as materials and drive
systems improve. Rack and pinion drive is essential for compact bundling
capability. Nighttime energy savings of 40-5090are generally reported.
However, in small-sized ranges, installation may not be economic on the
basis of energy savings alone. For this reason, there is increasing interest
in materials that will supply partial shade for increased crop productivity
while delivering savings in the area of 20-30%. Experiments concerning
the rational operation of screens on cloudy days are now being conducted.
A tremendous amount of research has been undertaken to find alterna-
tive sources of energy for heating greenhouses. Solar systems have been
developed that may become cost effective. Unfortunately their use is
constrained by climate conditions. Temperate climates generally do not
provide sufficient solar radiation during late fall, winter, and early spring,
when heating demands are highest. Attempts to find cost-effectivemeth-
ods for seasonal storage of solar energy have failed. In certain areas solar
heating systems will become more prevalent but it will take another large
escalation in fossil fuel prices or a limitation on supply before solar
heating becomes common.
The use of geothermal and waste heat energy for heating is also limited
by the availability of sources. Low source temperatures can be a problem
but heat pumps can be used to overcome this. Nevertheless, there are
locations where the greenhouse industry is sufficiently concentrated in
close proximity to reliable and cost-effective sources. Under these cir-
cumstances this method of heating has been an unqualified success.
As the greenhouse industry expands and rebuilds, geothermal and waste
heat will gradually become a more important source of energy. Reject heat
from the incineration of garbage may be the most promising new source.
Root zone heating is generally cost effective and beneficial for most
greenhouse crops. However, it does represent a substantial investment
and requires careful monitoring and control. I t is usually installed in the
growing medium such as soil or nutrient solution for maximum benefit
except for the United States, where it is commonly used beneath raised
benches and propagation benches. Lately there has been increased in-
terest in heating concrete floors, possibly in conjunction with ebb and
flood irrigation.
Alternative heating methods such as infrared and heat pumps have
produced mixed results. Capital costs and long-term reliability remain
questionable. In the meantime significant advancements have been made

on boiler design. Flue gas heat recovery units are used extensively with
natural gas heating, increasing fuel efficiency by 10-15% and giving pay
back periods of 1-2 years. COZenrichment can also be achieved by pump-
ing a portion of the flue gases from the heat recovery unit directly into
the greenhouse.
Computers have increased the accuracy of environmental control in
greenhouses by fine tuning and coordinating equipment operations,
resulting in energy savings of 10-1570. Split-night temperature regimes
can now be invoked with additional savings of 20% annually. The full
potential of computerized control has not yet been realized because
further experimentation with sophisticated control algorithms must be
carried out.
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Plant Growth Regulators

in Rose Plants*
Yoram Mor
Department of Floriculture, Extension Services,
Ministry of Agriculture,
Hakirya, R l Aviv 61070, Israel
Naftaly Zieslin
Department of Ornamental Horticulture,
The Hebrew University of Jerusalem,
Rehovot 76100, Israel
I. Introduction 53
11. Propagation 54
A. Propagation by Seeds 54
B . Propagation by Cuttings 55
C . Grafting and Budding 56
D. Root Regeneration 57
E . Micropropagation 57
111. Storage of Rose Plants 58
IV. Plant Development 59
A. Renewalshoots 59
B . Release of Lateral Buds from Inhibition and Flower
Bud Formation 60
C . Leaf and Flower Abscission 63
D. Flower Bud Development and Pigmentation 64
V. Flower Senescence 65
A . Ethylene 65
B . Abscisic Acid 66
C . Cytokinin 66
Literature Cited 66
I. INTRODUCTION
Garden and greenhouse roses, known in floricultural practice as Rosa

hybrida, are woody perennials without lateral bud dormancy (Zieslin and
Moe 1985)and are perpetually flowering,noninductiw plants. The sprouting
*Abbreviations used: ABA, abscisic acid; BA, benzylamino purine; IAA, indolacetic
acid; I B A , indolbutyric acid; GA, gibberellins; P B A , 6(benzylamino)-9 ( 2
tetrahydropyranyl) 9H purine; PGR, plant growth regulators; STS, silver thiosulfate;
TIBA, triiodo benzoic acid.
53
54 YORAM MOR AND NAFTALY ZIESLIN
of axillary buds is recurrent if apical dominance is removed and the

environmental conditions for growth are suitable. Due to the terminal
position of the flower, there is no antagonism between vegetative growth
and flowering. Potentially, each lateral shoot should bear a flower if no
abortion of atrophy of the flower bud occurs (Zieslin et al. 1973).
Roses are vegetatively propagated mainly by bud grafting on suitable
rootstocks or by cuttings (Post 1952). The rootstock is propagated by
cuttings or from seeds, depending on the species. Flower initiation and
development of garden and greenhouse roses are affected by factors like
root to shoot ratio, readiness of lateral buds to sprout, formation of basal
shoots (called renewal or structural shoots), growth rate of lateral shoots,
and sensitivity of flower buds to abortion (Zieslin et al. 1973). Many of
these factors are related to the genetic properties of rose cultivars and
rootstocks and correlate with changes in the content and activity of
endogenous plant hormones. Furthermore, some of these factors are
influenced by exogenous application of plant growth regulators a t the
proper stage of development.
The use of plant growth regulators (PGR)plays an important role in
horticulture. live books and a recent review have been published on this
subject (Larson 1985; Nickel1 1983; Weaver 1972)but only two citations
refer to roses. There are, however, various studies on the activity of plant
hormones in Rosa hybrida and the possible use of PGR in controlling the
physiological processes in rose plants. The objective of this review is to
present a comprehensive survey of the literature on the endogenous
changes of plant hormones and the uses and implications of PGR in rose
growth and development.
11. PROPAGATION
A. Propagation by Seeds
Seed propagation is used in breeding and in production of certain
rootstocks, e.g., R. canina and R. multiflora. Fruits (hips) are prone to
abscission. Application of various auxins prevents abscission of hips in
several rose species and promotes development of parthenocarpic fruits
(Prosser and Jackson 1959). Gibberellins (GA)were more effective than
auxins in preventing hip abscission in all species tested. GA, was the
most effective gibberellin to promote fruit set, while GA1 was the most
effective for promotion of hip growth (Jackson and Blundell 1964).The
fruit set was correlated with the endogenous levels of gibberellinlike,
growth-promotive substances in untreated hips of R. arvensis (Prosser
and Jackson 1959).GA promoted formation of fertile seeds (achenes)and
2. PLANT GROWTH REGULATORS IN ROSE PLANTS 55
seed germination, and GA application has been adopted by rose breeders

(Allen 1967). Rose achenes contain a growth inhibitor (Prosser and
Jackson 1959; Jackson and Prosser 1961; Jackson and Blundell 1965)
that was identified by Milborrow ( 1967) as abscisic acid (ABA).It was
suggested that the dormancy and germination of rose achenes are con-
trolled by a balance between ABA and growth promoters such as gibber-
ellins and cytokinins (Jackson and Blundelll963; Jackson 1968; Yu et al.
1975). However, Svejda and Poapst ( 1972)showed that the elimination of
the inhibitor from the achenes was not followed by simultaneous increase
in germination. Tillberg (1983) demonstrated that the decline in the
endogenous levels of ABA during stratification of rose achenes was not
adequate enough to promote germination. Tillberg et al. (1982a,b)suggested
that the decrease in ABA content in the achenes is only a prerequisite for
activation of growth promoters that regulate the release from dormancy
and germination. The promoters involved in the release of seeds from
dormancy were identified as cytokininlike substances, while GA and
indolacetic acid (IAA)were involved in the germination of rose achenes
(Tillberg et al. 1982a, b 1.
B. Propagation by Cuttings
The promotive effect of auxin on rooting of rose cuttings was reported
by Kirkpatrick (1940). Dipping the bases of leafy rose cuttings in IBA
solution of low concentration for 24 hr or using 1 mg/g of IBA as a talc
powder resulted in -100% rooting of many cultivars and species. The
efficacy of IBA on rooting of rose cuttings was confirmed and this
treatment was introduced into practice (Laurie and Stillings 1949).
IBA is more effective than IAA or NAA in promoting rooting of rose
cuttings. At concentrations of 2 g/liter, 25.4 roots per cutting were
formed after treatment with IBA, while only 7.2 roots were formed after
treatment with IAA or NAA of the same concentration (Moe 1973). It is
possible, however, that the optimal concentrations of NAA and IAA in
rooting of rose cuttings differ from those that were employed in the
experiments. It was shown (Grueber and Hanan 1981)that 5 mg/liter of
NAA had an effect similar to that obtained with 500 mg/liter of IBA, but
no differences between IBA and IAA were observed in rooting of R.
damascena cuttings (Khosh-Khui and Tafazoli 1979).When high concen-
trations of auxins were used in rooting of rose cuttings, negative results
were also reported in some experiments. It is possible that the difference
in response to auxin concentrations higher than 200 mg/liter may be due
to variations in the plant material such as species and cultivars, stage of
maturity, and environmental conditions, like storage and rooting temper-
atures prior to and during the application of the rooting compounds. A
56 YORAM M O R AND NAFTALY ZIESLIN
quick dip of cuttings in a solution of 750 mg/liter of IBA resulted in

increased dry weight of roots of R. canina over that of R. multiflora
(Azimi and Bisgrove 1975).Concentrations of IBA higher than 1.5 g/liter
increased the root dry weight of R. canina, but reduced root dry weight of
R. multiflora. A momentary dip in a solution of 4 g/liter of IBA increased
rooting of R. manetti by 22%. The same treatment had no effect on
rooting of R. indica or R. multiflora cuttings (Ohkawa 1980).Without an
auxin treatment there was no difference in rooting between stored and
unstored cuttings. However, pretreatment with NAA of cuttings stored
for 3 weeks a t 5 O C reduced the number of roots per cutting from 10 to 3.
Similar results were also obtained with cuttings treated with IAA
(Papandreou 1972). Promotion of rooting by the treatment with high
concentrations of IBA was obtained in rose cuttings from mature shoots
of R. indica but not in soft-wood cuttings (van de Pol and Breukelaar 1982).
Increasing the duration of the treatment with 200 mg/liter of IBA from
30 to 120 sec reduced the rooting of ‘Baccara’rose cuttings from 47 to 7%
and reduced the fresh weight of roots from 750 m to 450 mg per cutting
(Zieslin et al. 1971b). The results were less variable if lower concentra-
tions of auxin were utilized. When concentrations of 1-10 mg/liter of IBA
were used for 24 hr treatment, the rooting of more than 30 rose cultivars
was promoted without any negative symptoms (Kirkpatrick 1940). In
spite of the advantage of a long-term treatment with low concentrations
of auxins, which was also suggested in more recent studies (Grueber and
Hanan 1981, Hanan and Grueber 1984),a momentary dip of the bases of
leafy rose cuttings in a high concentration of auxins has become a
common practice. The use of high concentrations of auxins could be a
consequence of convenience and of the positive results that have been
obtained by this method in rooting of hardwood cuttings of other species
(Hartman and Kester 1975). In roses, however, it appears that longer
duration of the treatment with lower concentrations of auxins is more
beneficial than the quick dip of cuttings in a high concentration. No data
are available on the changes in the endogenous levels of hormones in rose
cuttings during rooting.
C. Grafting and Budding

There is no commercial use of PGR in grafting and budding of roses.
-
Application of BA to the bud (0.75% in lanolin paste) resulted in 100%
sprouting of the grafted buds and stimulated shoot growth (N. Zieslin
and Y. Mor, unpublished). In other experiments 2 g/liter of BA in lanolin
paste applied to the scion inhibited rather than stimulated the outgrowth
of the grafted buds (Fann et al. 1983). The lack of response to BA in this
experiment could be due to the fact that the BA was applied a t a distance
from the bud and not to the bud itself (Ohkawa 1984).
D. Root Regeneration
Root regeneration of rose plants after grafting and transplanting is
affected by auxin (Fuchs 1986). The number of new roots and the root
fresh weight increased significantly when roots of different rose cultivars
grafted on R. canina inermis were treated with various concentrations of
IBA or NAA. A similar response was obtained when 1g/liter of IBA was
applied to root segments of R. multiflora ‘Kanagawa’ (Ohkawa 1973).
E. Micropropagation
In uitro propagation has become a common practice in the cultivation
of roses, particularly in the propagation of flowering pot roses, where
grafting on suitable rootstocks is not required. The tissue culture of roses
does not differ substantially from that of other ornamental plants. The
methods of micropropagation of roses in uitro were recently reviewed
(Hughes 1981; Skirvin et al. 1986). Therefore, only the aspects concern-
ing the different effects of PGR in micropropagation are included in
this review.
The first shoot organogenesis from rose callus tissue was reported by
Hill (1967). A large number of leaf and petaloid primordia were formed
when 20 mg/liter of GA, were included in the growth medium containing
0.5 mg/liter of NAA and 0.2 mg/liter of kinetin.
Shoot and bud differentiation but not root differentiation were obtained
from callus tissue of ‘Super Star’roses only when 0.1 mg/liter of IBA and
5 mg/liter of kinetin were included in the growth medium (Jacobs et al.
1968). However, Khosh-Khui and Sink (1982a)failed in their attempts to
induce shoot formation from callus of R. manetti and R. hybrida cv.
‘Super Star’, using various growth media and different ratios of plant
hormones. The micropropagation of roses was advanced by using excised
shoot tips 3-4 to 20 mm in length (Jacobs et al. 1969, Skirvin and Chu
1979),apices of 0.6-1 mm (Elliot 1970)or nodule segments with a lateral
bud (Avramis et al. 1982, Davies 1980, Hasegawa 1979).
Jacobs et al. (1969) showed that leaf growth occurred only in the
absence of NAA and when 4-8 mg/liter of kinetin was used as a source of
cytokinin. On the other hand, root initiation was observed only in the
absence of kinetin and in the presence of 0.5-2.0 mg/liter NAA. Further
studies showed that BA and zeatin are prefered as cytokinins over the
kinetin (Elliot 1970; Hasegawa 1979). Hasegawa ( 1980) showed better
results using IAA in comparison with IBA and NAA, but NAA is still
widely used as a source of auxin. There are differences in the species and
cultivar response to various concentration of cytokinins and auxins in the
medium (Bressan et al. 1982, Davies 1980, Khosh-Khui and Sink 1982b).
Shoot multiplication was also improved by application of TIBA to the
main shoot in the culture tube (Bressan et al. 1982), although this
compound is rarely employed in tissue culture. Including GA in the
growth medium usually inhibits bud differentiation (Jacobs et al. 1970;
Elliot 1970). The promotive effect of GA on the development of shoot
primordia in rose tissue culture reported by Hill (1967)has remained an
exceptional case. However, recent results showed (D. de Vries and L.
Dubois, ITV, Wageningen, personal communication) that addition of
GA, to medium after autoclaving significantly promoted the growth of
plantlets in uitro.
Rooting is usually not possible in a growth medium enriched with
cytokinins (Elliot 1970). Thus, rooting of rose plantlets was obtained by
transfer into medium without hormones (Davies 1980, Skirvin and Chu
1979) or by transplanting the plantlets into a medium containing a low
concentration of auxin but without cytokinin (Hasegawa 1979, 1980;
Khosh-Khui and Sink 1982~). No significant difference was found between
IAA, IBA, and NAA in their effect on rooting (Hasegawa 1980). How-
ever, better rooting of the plantlets was reported when a combination of
two auxins-IAA and NAA or IBA and NAA-was used (Khosh-Khui
and Sink 1982~). An improvement in rooting of microcuttings of roses
was obtained by a dip of the plantlet base for a few hours in an aqueous
solution of 1 mM of IAA instead of a continuous culture on a medium
containing auxin (Collet 1985). Auxins and cytokinins enhanced ethyl-
ene evolution in airtight vessels containing callus tissue of rose (R.
hybrida L.). The increase of the ethylene levels had no adverse affect on
the growth of the callus (Wulster and Sacalis 1980).
111. STORAGE OF ROSE PLANTS
Young rose plants from nurseries can be stored in cold for several
months after lifting (Yerkes and Gardener 1935). Undesirable phenom-
ena that occur during storage include precocious sprouting of etiolated
lateral buds, shoot decay, dieback during and after storage, and the
development of various diseases (Marth 1943, Yerkes and Gardner 1935).
lleatment of stored rose plants with methyl or ethyl esters of NAA,
either by spraying or by evaporating a volatile formulation, was very
effective in preventing early sprouting (Marth 1943).There was no sprouting
in plants sprayed with 0.005% solution of a-naphthalene methyl or ethyl
acetate or fumigated for 16 hr a t 21OC with 10 mg/m3 of a volatile
formulation, and stored for 51 days at Oo-5OC. In untreated plants, 48

buds per plant sprouted during the storage. Wenty one days following
the transfer of the plants to greenhouse conditions there was no statisti-
cal difference in the number of new shoots between the treated and
controlled plants. The NAA-esters had also a positive effect on root
growth. More new roots developed, shortly after planting, in treated than
in control plants (Marth 1942). %ice as many flowers were produced in
treated plants grown outdoors in comparison to the control plants. Appli-
cation of a supraoptimal concentration of a volatile NAA-ester (300
mg/m3) resulted in a twofold increase in the number of flower petals. This
treatment also delayed for 2 weeks the development of Botrytis cinerea
fungus in stored plants.
Exogenous application of ethylene during cold storage caused an
increase in the number of sprouted lateral buds. Concentrations of ethyl-
ene higher than 3 ppm also promoted flower abortion (Widmer and
Swanson 1970). Exposure of various rose cultivars to 1000 ppm ethylene
during storage a t 5OC caused inhibition of bud sprouting and cane
dieback after replanting (Meadows and Richardson 1983).The deteriora-
tion of cold-stored rose plants after planting was attributed to evolution
of ethylene during storage (Widmer and Swanson 1970). However, in
neither of the studies cited above was ethylene evolution measured.
IV. PLANT DEVELOPMENT
A. Renewal Shoots
Formation of renewal shoots is indispensable for maintenance of gar-
den and greenhouse rose plants. These vigorous, juvenilelike shoots
growing from the basal parts of the plant are also known as structural
shoots or “bottom breaks” (Zieslin et al. 1973, Zieslin and Mor 1981b).
Outgrowth of these shoots is a complex phenomenon in which environ-
mental and endogenous factors are involved (Khayat and Zieslin 1982;
Zieslin and Khayat 1983). The sprouting of the quiescent buds, which
give rise to the renewal shoots, is prevented by accumulation of an
inhibitory complex in the basal part of the rose plant. One of the inhibi-
tors was identified as ABA (Zieslin and Khayat 1983).Following pruning
(removal of canopy), ABA was partially transformed by light into an
inactive form (Zieslin and Khayat 1983). Prior to the outgrowth of the
basal shoots, high cytokinin activity was found in the proliferating tissue
of the bud union (the “crown”). This cytokinin activity rapidly declined
following the sprouting of the basal buds (Zieslin and Khayat 1983).
llansport of cytokinins toward the upper sprouting buds has been dem-
onstrated in rose plants (Van Staden et al. 1981a, b; Van Staden 1982).
This upward transport of cytokinins may be influenced by a high concen-
tration of auxin in the young rose buds (Moe 1971).
Stimulation of renewal shoot formation in rose plants by application of
PGR began prior to the investigations of the role of the endogenous plant
hormones in this phenomenon. The inhibition of sprouting of the basal
buds was attributed to auxins. Therefore, in order to overcome the
inhibition, TIBA, an auxin transport inhibitor, was applied in lanolin
paste to the basal part of the rose plant with positive results (Asen and
Hamner 1953). This effect of TIBA, however, was not confirmed in later
experiments (Carpenter and Rodriguez 1971; Preis et al. 1972). On the
other hand, application of 100 mg/liter of NAA almost completely inhibited
sprouting of basal buds (Zieslin and Mor 1981b).
Tho plant growth regulating substances, cytokinin and ethylene, are
effective in promotion of the formation of renewal shoots. The cytokinins
BA and PBA were tested in roses (Parups 1971, Carpenter and Rodriguez
1971, Preis et al. 1972, Carpenter 1975, Faber and White 1977, Okhawa
1979). The most effective treatments were those applied directly to the
basal part of the plant, either in a lanolin paste (Carpenter and Rodriguez
1971, Parups 1971, Faber and White 1977)or as a foam spray (Carpenter
and Rodriguez 1971).The stimulation of the growth of renewal shoots by
cytokinins was enhanced by the removal of the canopy (pruning).When
cytokinin was applied directly to the basal buds, renewal shoots were also
formed without pruning, especially if a score was made above the bud,
while foliar application of cytokinin was ineffective (Preis et al. 1973,
Ohkawa 1979).The effect of cytokinins promoting the outgrowth of basal
buds in roses was nullified if GA was added to the solution (Parups 19711.
GAS itself was also ineffective (Preis et al. 1972). Extract of seaweed,
presumed to contain cytokinin, applied as a spray to the lower part of the
plant, promoted formation of renewal shoots in roses (Raviv 1985).
Exposure of rose plants to ethylene promoted sprouting of lateral buds
(Zimmerman et al. 1931). Ethylene simultaneously caused severe leaf
abscission. The introduction of ethephon, a liquid, ethylene-releasing
compound, enabled direct application of ethylene to the basal part of the
plant without affecting the foliage. Ethephon promoted sprouting of the
basal buds in roses and its effect was improved by scoring above the buds
(Zieslin et al. 1972, Anon. 1973). Both cytokinins and ethephon can be
successfully used in the commercial production of plants in rose nurseries.
B. Release of Lateral Buds from Inhibition and

Flower Bud Formation
The number of buds sprouting on the distal end of the rose branch
following decapitation is limited (Zieslin and Halevy 1976e, Zieslin et al.
1976). Usually no more than one or two of the upper buds sprout, but in
some cultivars there are three or four sprouting buds. Under certain
conditions even the uppermost bud does not sprout (Durkin 1965, Kohl
and Rundle 1974, Michalak and Mynett 1978, Ohkawa 1984).The inhibi-
tion of lateral buds along the rose shoots is greater in the basal than in the
upper part of the shoot. Extracts from different parts of ‘Baccara’ rose
shoots showed a basipetally increasing gradient of ABA content (Zieslin
et al. 1978).Spraying ‘Helen naubel’ rose plants with 100-200 ppm ABA
or immersing the plants in ABA solution of the same concentration,
delayed sprouting by 4 to 5 days, but did not affect growth of the shoots
(Cohen and Kelley 1974). Similar effect of ABA on ‘Bacarra’ roses was
also reported (Zieslin et al. 1978).
Exogenous cytokinins can overcome the lateral bud inhibition. Appli-
cation of cytokinins, either BA or PBA in lanolin paste, 0.125-0.570to the
stump, above the uppermost bud promoted its sprouting and the develop-
ment of the young shoot (Kohl and Rundle 1974, Ohkawa 1984).However,
lateral buds on soft, immature shoots did not respond to this treatment
(Kohl and Rundle 1974). The lack of response and even inhibition of the
buds was also noticed after application of cytokinins to softwood cut-
tings of miniature roses (L. A. M. Dubois, Institute for Horticultural
Plant Breeding, Wageningen, The Netherlands, personal communica-
tion). The lateral buds, in position 2 or 3 from the distal end of the
decapitated branch, sprout less readily, and those which do sprout very
often develop shoots with aborted flowers (Zieslin et al. 1973, Zieslin and
Halevy 1975). Application of 0.75% PBA in lanolin paste to the second
bud stimulated sprouting and decreased atrophy of the flowers on shoots
that developed from the treated buds (Zieslin et al. 1985). Prevention of
flower atrophy in shoots of ‘Marimba’rose was also obtained when young
shoots were sprayed with 50 mg/liter of BA soon after sprouting (Mor
and Halevy 1984).Spraying the whole shoot or plant with PBA caused a
twofold increase in the number of sprouting buds in ‘Baccara’ roses
(Zieslin and Halevy 1976d),and in ‘Jeanne la Joie’, a miniature climbing
rose (Richards and Wilkinson 1984). This cultivar also responded to BA
treatment (100 mg/liter) by increased number of flower buds in the
inflorescence. Promotion of sprouting of lateral buds by cytokinins is
influenced by light. Darkening of buds that were treated with cytokinins
reduced the promoting effect by over 50% (Preis et al. 1973).
Shoots that developed from the uppermost buds on the rose branch
had higher activity of endogenous plant hormones, including cytokinins,
than shoots that developed from the lower buds (Zieslin and Halevy
1976a).This activity was influenced by light intensity and other environ-
mental factors. Darkening of the young shoot that sprouted from the
uppermost bud on a decapitated branch resulted in a parallel decrease in
the transport of I4C-labeledcarbohydrates toward the darkened shoot.
Application of 50 mg/liter of BA to the darkened shoot restored the

translocation (Mor et al. 1981). However, when the young shoots were
tested for endogenous cytokinins, higher levels were found in the darkened
shoots than in the shoots in light (Van Staden et al. 1981b). This contra-
diction may indicate a difference in response between the endogenous,
naturally occurring cytokinins and exogenously applied synthetic cytokinin
in the dark. Possibly natural cytokinins are inactivated in the dark, while
the exogenously applied cytokinins are not. After decapitation, the endog-
enous levels of cytokinins are lower in the uppermost lateral bud than in
the bud beneath (Van Staden et al. 1981a).In the leaves subtending these
buds, the level of cytokinins was higher in the uppermost leaf than in the
leaf beneath. Van Staden ( 1982)has shown that in roses 14C-zeatinmoved
from the subtending leaf to the bud during the process of sprouting. On
day 6 after decapitation, the level of cytokinins increased in the upper-
most bud, which had already started sprouting, while in the subtending
leaf the level of endogenous cytokinins decreased (VanStaden et al. 1981a).
Gibberellins are also involved in the developmental processes of rose
shoots and flowers. The level of endogenous GA was much higher in the
flowering shoots than in the shoots with aborted flower buds (Zieslin and
Halevy 1976a). The levels of GA in the leaves of the uppermost shoot
were higher than in the leaves of the shoot beneath, which is more prone
to flower atrophy (Zieslin and Halevy 1976a). Under low light and low
temperature conditions, which increase the incidence of the flower atro-
phy is roses, a decrease in GA activity was found in the young shoots.
This decrease was more pronounced in the lower than in the uppermost
shoot on the same branch (Zieslin and Halevy 1976~). Application of GA
to young rose shoots of many different cultivars increased the number of
flowers, produced longer stems and caused earlier flowering (Van Onsem
and Haegeman 1962, Zieslin and Halevy 1976b, El Shafie 1978, El Shafie
et al. 1980). The stem length and fresh weight of rose flowers increased
after one application of 10-100 mg/liter of GA, (Mastalerz 1965).
Simultaneous or sequential application of BA and GAS to the lateral
buds of young rose shoots had an additive effect on the increase in dry
weight and translocation of labeled carbon to the shoots. This effect,
however, was accompanied by deterioration of the young tissue and was
detrimental to the shoot apex (Y. Mor, N. Zieslin, and J. Van Staden,
unpublished). The reason for this effect is not clear but could stem from a
disturbed hormonal balance in the rose tissue.
Promotion of flower formation in roses by chlormequat (CCC) was
found in pot roses (Bornemann and Muhe 1967, Mor et al. 1986) and in
greenhouse roses (Byrne et al. 1971, Zieslin et al. 1971, Mor et al. 1977).
However, an inhibition of growth and a reduction in number of flowers
was also reported (Moe 1970). The promotion of flowering was obtained
only when the whole plant was treated. Application of CCC to individual
branches on the plant had no effect on flowering (Zieslin and Halevy
1976b, d). I t is possible that CCC affects the partitioning of plant
metabolites in roses. After a soil drench application of CCC, more 14C
from 14C02supplied to a leaf was found in the lower young shoots (Y. Mor
and A. H. Halevy, unpublished results). Without CCC more labeled
carbohydrates were translocated to the uppermost shoot. The diversion
of assimilates toward the lower shoots may prevent flower bud atrophy in
these shoots. Awad et al. (1981)showed that CCC treatment promoted
the development of floral organs in rose apices. CCC is known as an
inhibitor of gibberellins, but the levels of endogenous GA in rose plants
were not reduced by application of CCC (Zieslin and Halevy 1976d).
The two developing uppermost shoots on a decapitated rose branch
produce relatively high levels of ethylene (Zieslin and Halevy 1976~).
After exposing the plants to low light conditions for 6 hr, the ethylene
emanation from the same shoots was reduced by 70%. After 2 days in low
light conditions, there was an increase in ethylene emanation from the
second shoot while the ethylene emanation from the uppermost shoot
remained low. On day 4, the ethylene emanation from the second shoot
reached the initial level. This increase in ethylene production was corre-
lated with the occurrence of the flower bud atrophy in the second bud
(Zieslin and Halevy 1976~). Spraying roses with 1000 ppm ethephon once
or with 100ppm twice caused an abundant atrophy of flower buds (Zieslin
and Halevy 1976b). The increase in atrophy was accompanied by symp-
toms similar to those described after exposure of roses to ethylene, i.e.,
leaf abscission, cessation of growth, and abnormally large number of
sprouting buds (Zimmerman et al. 1931, Piersol 1974). Promotion of
flower formation by low concentration of ethephon ( 10-30 ppm) was also
reported when outdoor-grown garden roses were sprayed with ethephon
(Hassan et al. 1976).
C. Leaf and Flower Abscission

The recurrent shoot growth of roses is accompanied by continuous leaf
abscission from the older parts of the plant (Zieslin and Mor 1981a).
Foliar application of 250 mg/liter of BA reduced the winter leaf abscission
of ‘ForeverYours’roses that is common in this cultivar (Byrneet al. 1971).
Application of 100 mg/liter of PBA reduced leaf abscission of ‘Pink
Margo Koster’pot roses by 10%during shipment in the dark (Halevy and
Kofranek 1976).The same treatment also prevented flower bud abscission,
which usually occurs during shipment of this cultivar.
Ethylene is known to induce defoliation in roses (Zimmerman et al.
1931).In a special field trial on defoliation of roses the plants were sprayed
with several ethylene-releasing compounds on four dates from June to

October (Lurssen 1982). The compounds used in the trial included 0.2%
solutions of 1-aminocyclopropane-1-carboxcylic acid ( ACC ), a precursor
of ethylene synthesis, its derivative SDF 1664, and two sulfonic deriva-
tives of chloroethane, HOL 1247 and HOL 1302. In June, none of the
compounds induced defoliation. From July on, ACC caused defoliation of
90-95%. SDF 1664, which released only small amounts of ethylene (5%of
that released by ACC), was almost as effective as ACC in August and
October treatments. HOL 1247 and HOL 1302, which released ethylene
fivefold and fourfold, respectively, more than ACC, were active only in
October treatments. It was inferred from this study that the action of
ethylene depends on the concentration of hormone receptors rather than
on concentration of the hormone, which determines the reaction of a
plant (Lurssen 1982).
D. Flower Bud Development and Pigmentation

Development of the flower bud in roses is very sensitive to adverse
environmental conditions. The involvement of hormonal balance in the
flower development associated with alteration of air temperature was
studied in ‘Baccara’roses (Zieslin et al. 1974,1979).At low night temper-
ature a high proportion of ‘Baccara’ flowers are malformed. This malfor-
mation is characterized by proliferation of the reproductive organs,
abnormally high number of petals, and a reduction in the ratio of length
to diameter of the flower bud ( Halevy and Zieslin 1969, Lindenbaum et al.
1975). The malformation was associated with a decrease in the endoge-
nous levels of free and bound gibberellins and a sharp rise in endogenous
cytokinins (Zieslin et al. 1979).The levels of endogenous cytokinins a t the
early stage of development of the flower bud were 15 times higher a t
22/13OC day/night temperature than the level found in flower buds of
plants grown at 28/18OC. The changes in the ratio of gibberellins to
cytokinins were positively correlated with the length to diameter ratio of
the flower bud. An injection of GA3 into the receptacles of the flowers
grown a t low night temperatures prevented the proliferation of the repro-
ductive organs and increased the length to diameter ratio of the bud.
However, the number of petals was not reduced by GA3 treatment. An
injection of cytokinin alone to the receptacle of the rose bud caused a
fourfold increase in the number of the malformed flowers (Zieslin et al.
1974, 1979). The antagonism between cytokinins and GA in roses has
already been mentioned in regard to the sprouting of basal buds ( Parups
1971).
Blackening of red rose petals in low temperature is caused by high
content of anthocyanin pigments as well as that of tannins (Zieslin and
Halevy 1969). GA, treatment caused accumulation of anthocyanins in
petals of ‘Baccara’roses. This effect was more pronounced a t low tempera-
tures (20/14'C, day/night) than in higher temperatures ( 3O/2O0C).The

synergistic effect of GA, and low temperature on the accumulation of
anthocyanins in rose petals was found also in uitro (Zieslin et al. 1974,
1977).Application of GA3 to young rose leaves promoted the retention of
leaf anthocyanins and delayed the visual appearance of chlorophyll
( Mastalerz 1965).Promotion of anthocyanins biosynthesis in rose petals
by cytokinin was also reported (Nakamura et al. 1980).
V. FLOWER SENESCENCE
The process of flower senescence in roses is similar to that described in

other flowers (Halevy and Mayak 1979, 1981; Mayak and Halevy 1980).
However, some of the phenomena associated with senescence in rose
flowers differ from other plants. For example, the petal curling (sleepi-
ness) response of carnation flowers to ethylene does not exist in roses.
Flower bud abscission is also uncommon in roses except for a few culti-
vars (Halevy and Kofranek 1976). The general aspects of rose flower
senescence have been comprehensively reviewed ( Halevy and Mayak
1979,1981),and therefore only those aspects concerning PGR are reviewed
in the following section.
A. Ethylene
One of the plant hormones that plays a major role in flower senescence
is ethylene (Halevy and Mayak 1981). Emanation of ethylene by rose
flowers was reported for the first time by Fisher ( 1950).The time course of
ethylene emanation by rose petals resembles that of carnation and other
flowers and is composed of three distinct phases: (1)a low steady state;
( 2 )an accelerated rise to a maximum; and ( 3 )a decline in the emanation
of the ethylene (Halevy and Mayak 1981, Faragher and Mayak 1984,
Mayak and Halevy 1972).The onset of the second phase, which signals
the terminal stage of senescence (Halevy and Mayak 1981),occurs earlier
in short-lived rose cultivars than in long-lived ones (Mayak et al. 1972). A
rise in ethylene production was evident in rose flowers also during cold
storage. Following cold storage the ethylene production of the stored
roses was higher than that of fresh flowers and their longevity shorter
(Faragher et al. 1986).Ethylene production by rose petals is low, a t most
1-5% of that produced by carnation petals (Mayak 1972, Nichols 1977).
Inhibitors of ethylene synthesis failed to prolong the longevity of rose
flowers despite the reduction in ethylene production (Faragherand Mayak
1984, Wang and Baker 1979).STS, an inhibitor of ethylene action (Veen
1983),applied to rose flowers increased the emanation of ethylene with-
out causing any adverse affects (Faragher and Mayak 1984).STS increased
the longevity of stored rose flowers (Faragher et al. 1986)and prevented
the accelerated abscission of rose petals caused by exogenously applied

ethylene (De Stigter 1980). The role of ethylene in senescence in rose
flowers deserves still further research.
B. Abscisic Acid
Abscisic acid in petals is involved in regulation of senescence (Halevy
and Mayak 1981). The endogenous content of ABA in rose petals fell
during the first 3 days after flower detachment and rose again from the
fourth day onward (Borochov et al. 1976b).The ABA level was higher in
short-lived than in long-lived rose cultivar (Mayak and Halevy 1972).
Exogenous ethylene increased the level of endogenous ABA in rose petals
(Mayak and Halevy 1972).
Kohl and Rundle (1972)reported that ABA in solution reduced water
loss in cut roses, prevented wilting of the peduncle (bent-neck), and
prolonged the vase life. Flower senescence was enchanced, however, when
ABA was applied to rose flowers in the dark, to flowers with defoliated
stems, or to detached petals (Borochov et al. 1976a, Halevy et al. 1974,
Mayak and Halevy 1972). The promotion of rose flower senescence by
ABA was accompanied by suppression of ethylene production ( Mayak
and Halevy 1972).These reports demonstrate the dual effect of ABA: (1)
promotion of flower tissue senescence; and ( 2 ) an aid to longevity by
improving the water balance of the flowers (Halevy et al. 1978).
C. Cytokinin
The level of endogenous cytokinin in rose petals decreased with aging.
I t was higher in long-lived cultivars than in a short-lived one (Mayak and
Halevy 1970).Application of BA improved the water balance of cut roses
(Mayak and Halevy 1974)and increased the life span of a short-lived rose
cultivar (Mayak and Halevy 1970). However, in many rose cultivars,
addition of BA to the vase solution was ineffective (Lester and Durkin
1970).Various studies reported that vase life of roses was promoted by
including BA in preservative solution (Halevy et al. 1978; Zieslin and
Kofranek 1980), but the use of PGR in the preservative solution has
remained limited.
The interactions between ethylene, abscisic acid, and cytokinin plays
an important role in rose flower senescence and vase life. However, these
interactions need further research and clarifications.
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Nutrition of Container-Grown Woody

Nursery Crops
Robert D. Wright and Alexander X . Niemiera
Department of Horticulture,
Virginia Polytechnic Institute and State University,
Blacksburg, Virginia 24061
I. Introduction 75
11. The Container System 77
111. Methods of Nutrient Application 78
I v. Nutrient Requirements for Growth 80
A . Nitrogen 80
B. Phosphorus 82
C . Potassium 84
D . Nitrogen, Phosphorus, and Potassium Ratio 84
E . Calcium and Magnesium 84
F . Micronutrients 85
G. Medium pH 87
V. Soil Testing 88
VI. Tissue Analysis 90
VII. Timing of Nutrient Applications 90
A . Propagation 90
B . During Growth Cycles 91
C . Fall Fertilization 92
D. Mycorrhizae 93
VIII. Conclusions 93
Literature Cited 95
I. INTRODUCTION
During the last 35 years production of woody nursery crops in contain-

ers has gone from a production system of minor importance to the
predominant production method for nursery crops in the United States.
Initially, the growing medium used in container plant production consisted
of mineral soil of varying composition, with fertilization practices similar
to those used in field production of nursery crops. Modern container
media, however, seldom contain mineral soil, but are primarily composed
75
76 ROBERT D. WRIGHT AND ALEXANDER X. NIEMIERA
of organic substances that require frequent irrigation and nutrient appli-

cations. The literature on nutrition of woody nursery crops has been
reviewed by Whalley (1974) with emphasis on field production. Since
1974 a significant amount of literature has been published solely on
container production, but it has not been reviewed. This chapter will
focus on the uniqueness of managing fertility programs for a soilless
container system and the research that has set the stage for current
fertilizer recommendations. A listing of plant material used in this review
is given in n b l e 3.1.
Table 3.1. Scientific Names of Plants Referred to in Text
Genus and species Common name
Acer L. spp. Maple

Acer platanoides L. Norway maple
Berberis thunbergi DC Japanese barberry
Betula L. spp. Birch
Chamaecyparis lawsoniana Parl. Lawson falsecypress
Cotoneaster adpressa praccox Bois + Berthault Cotoneaster
Cotoneaster diuaricata Rehd. + Wils. Spreading cotoneaster
Cotoneaster dammeri Schneid. Bearberry cotoneaster
E u o n y m u s japonicus L. Euonymus
Forsythia Vahl spp. Forsythia
Gleditsia triacanthos L. Honey locust
Ilex crenata Thunb. Japanese holly
Juniperus chinensis L. Chinese juniper
Juniperus conferta Parl. Shore juniper
Ligustrum japonicum Thunb. Privet
Populus L. spp. Poplar
Prunus persica Batsch Peach
Pseudotsuga menziesei Brit. Douglas fir
Pyracantha coccinea R o e m Scarlet firethorn
Pyrus L. spp. Pear
Quercus shumardii Buckl. Shumard oak
Rhododendron obtusum (Lindl.) Planch Azalea
Photenia Xfraseri Dress. Photinia
Syringa uulgaris L. Lilac
Izzxus xmedia Rehd. Anglojap yew
Tilia cordata Mill. Linden
Thuja occidentalis L. Arborvitae
Vaccinium as hei Reade Blueberry (rabbiteye)
Vaccinium corymbosum L. Blueberry
Viburnum burkwoodii Burkwood Burkwood viburnum
Viburnum plicatum Thunb. Doublefile viburnum
Viburnum ruspensum Lindl. Viburnum
Weigela florida A DC Weigela
3. NUTRITION OF CONTAINER-GROWN WOODY NURSERY CROPS 77
11. THE CONTAINER SYSTEM
Fertilizer systems must maintain adequate amounts of nutrients in the

medium solution to promote optimum growth of plants in containers
(Fig. 3.1). Frequent irrigations are necessary to provide sufficient water
for plant growth, but these reduce the nutrient content of the medium due
to its porous nature. Thomas and Perry (1980)found that 85-90% of the
leachable nitrogen was removed from the container when twice the con-
tainer volume of water was applied. Foster et al. (1983) found that the
equivalent of four irrigations of 2.54 cm ( 1in. )would remove nearly 90%of
the leachable NO3 and NHI from columns of pine bark. Thus, within a few
days the quantity of nutrients in a container medium would be below the
level required for optimum plant growth.
Rv-0 approaches have been used by nursery operators to alleviate the
nutrient deficiency problem inherent in container-grown nursery crops.
Controlled-release fertilizers ( CRF ), which provide a gradual release of
nutrients to the medium solution, are preplant incorporated into the
medium or postplant applied to the medium surface. Also water-soluble
fertilizers may be added to irrigation water as a liquid feed (LF),thus
supplying the medium solution with an adequate nutrient supply. &la-
tively high nutrient levels-about 100 ppm nitrogen, 15 ppm phospho-
Fig. 3.1. Interrelationship between plant, medium solution, container medium, and
nutrient supply.
78 ROBERT D. W R I G H T AND ALEXANDER X. NIEMIERA
rus, and 75 ppm potassium-are usually found in the medium solution of

container-grown plants. At these levels nutrient movement to the root
surface is mainly via the convection flow of water (mass flow) created
by plant transpiration, with relatively little via diffusion (Mengel and
Kirkby 1982). However, during periods of heavy rainfall or between
fertilizer applications, the medium solution may be depleted by leaching
and/or plant uptake. In this case the influence of the cation exchange
capacity (CEC) of the medium may play an important role in nutrient
supply. The adsorbed cations in equilibrium with the medium solution
may act as a temporary buffer to the reduction in the nutrient concentra-
tion of the medium solution (Fig. 3.1). Organic media such as pine bark
have the capacity to absorb 1.5 mg NH4-N/g bark (Foster et al. 1983).On
the basis of a 3-liter container, this amount of absorbed nitrogen is
equivalent to twice the amount that an Ilex crenata ‘Helleri’ cutting
would adsorb from the medium during a 4-5 month growing season. Brown
and Pokorny ( 1977)have also demonstrated potassium retention in a bark-
sand medium. There are no studies that have investigated the capacity of
soilless media to supply nutrients to the medium solution. Thus, the
extent to which adsorbed cations play a role in nutrient supply is unknown.
Despite the undesirable leaching of nutrients from the container by
frequent irrigation, one must also recognize that the liquid phase of the
container is the major avenue of nutrient movement to the root surface.
Whether by mass flow or diffusion, maintaining adequate medium mois-
ture levels is necessary for nutrient movement (Nye and Tinker 1977).
Several studies with containerized woody plants have investigated the
relationship between irrigation frequency and plant growth. Jarrell et al.
( 1983)subjected Ligustrum japonicum ‘Texanum’to two irrigation sched-
ules with CRF and found that plants receiving the higher rate of water
grew larger despite a greater leaching of nutrients from this treatment.
Stewart et al. ( 1981)irrigated L. japonicum either daily or every other
day with fertilizer in the irrigation water. Plants irrigated daily were
larger. Ingram and co-workers(1979)have also shown similar results with
Rhododendron obtusum and L. japonicum. While it is difficult to sepa-
rate the influence that container moisture has on nutrient mobilization to
the root surface and plant growth, the influence of moisture on nutrient
availability is obviously important.
111. METHODS OF NUTRIENT APPLICATION
Nutrients are applied to plants grown in containers through the irriga-

tion system, as dry formulations incorporated into the medium preplant,
on the surface of the medium, or a combination of these methods. In any
case, single- and multi-element formulations are available to nursery
operators to meet plant nutrient requirements. In recent years, the most

widely used dry fertilizers have been CRF, which release nutrients over an
extended period of time. Resin- and sulfur-coated materials along with
urea formaldehyde and isobutylidene-diurea comprise the majority of
CRF used by the nursery industry. With CRF, attention must be given
not only to the duration of release as stated by the manufacturer, but
more importantly to the actual period of release, which may be influenced
by temperature, soil moisture, and mode of nutrient release. The properties
and release characteristics of these materials have been reviewed by
Oertli (1980)and Maynard and Lorenz (1979).
Use of CRF has resulted in growth comparable to plants fertilized via
the irrigation system. Yet research has shown that a CRF period of
nutrient release conducive to optimum growth may be less than that
claimed by the manufacturer (Meadows and Fuller 1983; Meadows et al.
1983). Meadows and Fuller ( 1983)compared the release duration of a 8-9
and a 12-14 month CRF and found that nitrogen levels in the medium
solution had dropped below 10 ppm after 4 months. Thus, growth would
likely be limited beyond 4 months for both formulations. Under these
circumstances, supplemental applications of nutrients would be neces-
sary to maximize growth. Consistent with this limitation, growth of I.
crenata fertilized with nitrogen via irrigation was greater than that from
certain CRF (Sharma et al. 1982).This could be attributed to the mainte-
nance of higher nitrogen levels in the container medium late in the
growing season with LF. In support of the contention, van der Boon
(1981) has found enhanced growth of Chamaecyparis, and Pyracantha
when CRF were supplemented with LF late in the growing season. Better
growth of I. crenata, Thuja occidentalis, and Viburnum burkwoodii was
also reported with a LF system than with a number of CRF (Sanderson
and Martin 1974). Other studies have reported no difference in growth of
woody plants between these two nutrient supply methods (Gouin and
Link 1973, Johnson et al. 1981, Hicklenton 1982). Ward and Whitcomb
(1979) in contrast have shown greater growth of I. crenata with CRF
compared to LF. Thus, high-quality plants can be produced from either
LF or CRF regimes provided that adequate nutrient levels are maintained
in the medium solution.
Differences between nutrient use efficiency for CRF and LF depend on
the method of comparison. Efficiency calculations are contingent upon
whether one takes into consideration the unused residual nutrients remaining
in the CRF fertilizer a t the time of calculation. In the latter case the
efficiency of a LF system approaches that of a CRF system (Hershey and
Paul 1982). One factor that may decrease the efficiency of CRF is that
larger quantities of nutrients are released during the beginning of a
growing season when plants are smaller and nutrient requirements lower.
In contrast, LF systems can be managed to supply nutrients a t levels
80 ROBERT D. W R I G H T AND ALEXANDER X. NIEMIERA
commensurate with plant needs. The major disadvantage of the LF

systems is that nutrients are applied to the area between containers and
on aisles and roadways. In addition, this practice leads to nutrient runoff,
which may eventually contaminate ground and surface water. Significant
nutrient leaching from the containers supplied with CRF has also been
demonstrated (Jarrell et al. 1983).These findings demonstrate the need
for further research to develop a CRF in which a release rate parallels
plant nutrient requirements. &search is also needed to refine present LF
practices so that less nutrients are lost by the system.
IV. NUTRIENT REQUIREMENTS FOR GROWTH
There have been many investigations attempting to establish the

nutrient requirements of container-grown woody plants. Direct compari-
sons between these experiments are difficult due to differences in method-
ology, such as growing medium, frequency and method of irrigation,
frequency and method of nutrient applications, plant and container size,
growth rate, and plant material differences. Therefore,no attempt will be
made to establish absolute nutrient levels for optimum growth based on
the available literature, but rather we shall try to give experimental
results that can be used to establish guidelines for nutrient applications.
Regardless of the method of nutrient application-CRF or LF-of
primary concern is the resulting nutrient concentration in the medium
solution. The magnitude of plant growth in response to nutrient applica-
tion is not a direct response to concentration in the medium solution but
to the total nutrient supply or availability. Concentration (ppm) is a
relative representation of the nutrient supply to the plant since the
volume of solution that remains in the container after irrigation and
drainage, multiplied by the concentration, will equal the total amount of
nutrient potentially available to the plant a t that time. Since plant size
and, therefore, absorption of nutrients increase with time, nutrient sup-
ply must be correspondingly increased (Ingestad 1982).Nutrient supply
may be increased either by applying the same concentration more fre-
quently or by increasing the concentration of the nutrient solution applied
to the medium (Ingestad 1982). The same relationship has been demon-
strated in a nutrient flow system (Edwards and Asher 1974) in which
increasing plant nutrient demand may be met by either increasing solu-
tion concentration or the flow rate of the solution through the system.
A. Nitrogen
hsearch results with daily applications of nitrogen have shown that
maximum dry weight of Tilia cordata and Acerplatanoides growing in a
2 turface (calcined clay) : 1 peat moss (v/v) medium occurred a t about
100 ppm nitrogen (Barnett and Ormrod 1985). Maximum dry weight of
Cotoneaster dammeri, Pyracantha coccinea, and Weigela florida grown

in sand culture occurred a t 50 ppm nitrogen (Gilliam et al. 1980a), while
growth of I. crenata in sand culture was greatest at 75 ppm nitrogen
(Niemiera and Wright 1982, Wright and Niemiera 1985).Also two Populus
species grew best in sand culture of 75-100 ppm nitrogen (Einspahr
1971).Plants fertilized on a weekly basis require higher concentrations of
nitrogen. Several studies with Ace5 Ilex, and Pyrus growing in a
predominantly pine bark medium noted greatest dry weight accumula-
tion when 300 ppm nitrogen was applied weekly (Gilliam and Wright
1977a,b, Yeager and Wright 1981b, Gilliam et al. 1980b, 1984).Graca and
Hamilton (1981) grew Cotoneaster diuaricata in a 1 soil: 2 sphagnum
peat : 2 perlite (v/v/v) medium and found significantly more growth at
250 ppm. To illustrate the relationship between frequency of nitrogen
application and concentration further, Yeager et al. (1980) fertilized I.
crenata with 500 ppm nitrogen 2 weeks apart, every 6 weeks. Growth of
these plants was the same as plants fertilized weekly with 300 ppm.
There is considerable experimental evidence of plant growth response
to nitrogen form. Nitrate nitrogen (N03-N) has been shown to produce
greater growth than NH4-N for Viburnumplicatum (Raker and Dirr 1979,
Dirr 1975), and C. dammeri (Gilliam et al. 1982). In contrast, Quercus
shumardii (Ingram and Joiner 1982), Pseudotsuga menziesii (van den
Driessche 1978), and Rhododendron obtusum (Colgrove and Roberts
1956)were found to grow better with NH4-N. Based on a review by Barker
and Mills (1980) covering horticultural crops and three reports with
woody ornamentals (Raker and Dirr 1979, Nelson and Selby 1974, Gilliam
et al. 1980a),growth of plants fertilized with a combination of N03-N and
NH4-N is generally equal to or better than plants fertilized with either
nitrogen source alone.
The growth response to nitrogen source has been shown to be related to
nutritional factors such as iron supply, media pH, plant tissue pH, and
species preference. Colgrove and Roberts ( 1956)noted growth reductions
in azalea with N03-N nutrition to be due to an interference of iron
metabolism. Uptake of N03-N increases the cell pH due to increased
cation absorption (Mengel and Kirkby 1982),causing an inactivation of
tissue iron. Absorption of N03-N by plant roots also increases the medium
solution pH since OH- ions are excreted in exchange for NO3 ions (Mengel
and Kirkby 1982). Since the solubility of iron in the medium is inversely
related to pH, NO3nutrition may reduce iron availability and absorption.
Maintaining acidic conditions in the presence of NO3 nutrition has an
offsetting effect (Colgrove and Roberts 1956, van den Driessche 1978,
Dirr 1975). Additions of an iron chelate to nitrate fertilized plants has
been shown to prevent iron chlorosis (Oertli 1963, Nelson and Selby 1974,
van den Driessche 1978) presumably by offsetting the plant absorption
and tissue inactivation problems.
The form of nitrogen applied and subsequent biotic transformations
(urea hydrolysis and nitrification) influence the chemical form of nitro-

gen in the medium solution. For example, 71% of the urea applied to a
pine bark medium was oxidized to NH4in 24 hr (R. D. Wright, unpublished).
Thus, urea is a desirable nitrogen source since it is rapidly converted to
the readily absorbable NH4form and is generally a less expensive form of
nitrogen. Nitrification of NH4 in the medium solution is also a relatively
rapid process. Niemiera and Wright ( 1986a) found low NH4-N :N03-N
ratios in a medium solution from pine bark that was fertilized with a
NH4-N fertilizer. Niemiera (1985) calculated that a 100 ppm NH4-N
medium solution of a 3.8-liter bark-filled container ( a t 100%gravimetric
moisture) would be nitrified in 50 hr. Nitrification is a pH-dependent
process with maximum nitrification occurring in the range of 7-8 (Focht
and Verstraete 1977).Nitrification in a pine bark medium has been shown
to be stimulated by additions of limestone, which raises the medium pH
and results in a decreased NH4to NO3 ratio (Niemiera and Wright 1986b).
Chrustic and Wright (1983) found lower NH4-N :N03-N ratios in pine
bark a t higher lime rates. In the same study, growth of I. crenata, R .
obtusum, and Juniperus chinensis was inversely related to lime rate
addition. This growth response was partially attributed to a greater
nitrification rate and a low NH4 to NO3 ratio,
Nitrification is an acidifying process and has been shown to result in a
greater availability of calcium, magnesium, manganese, and zinc in the
medium solution (Niemieraand Wright 1986a).These results were attributed
to an increased solubility of salts and a lowered CEC at the lower pH.
Thus researchers should be aware that a growth response to lime may be
a t least in part a response to nitrification, which may change the NH4:NO3
ratio and alter the availability of other cations. The use of a nitrification
inhibitor as a control in nitrogen source experimentation is advisable.
B. Phosphorus
The medium solution phosphorus level required for optimum growth of
woody nursery crops is considerably lower than that for nitrogen. Phos-
phorus applications of about 10 ppm in the irrigation water have resulted
in maximum growth of I. crenata (Yeager and Wright 1982, Wright and
Niemiera 1985), Chamaecyparis lawsoniana (van der Boon 1981), and
Cotoneaster adpressapraecox (Havisand Baker 1985a).There is evidence
that some species such as Rhododendron may grow optimally a t lower
phosphorus concentrations (Havis and Baker 1985a). Relatively high
concentrations of phosphorus have been shown to be toxic to some
species within the Proteaceae (Grundon 1972, Nichols and Beardsell
1981)and for I. crenata (Wright and Niemiera 1985). Greater supplies of
nitrogen and potassium (Grundon 1972) and potassium (Wright and
Niemiera 1985) have been shown to offset the deleterious effect of high
phosphorus concentrations. For example, growth of 1. crenata was
inhibited when phosphorus was supplied a t 40 ppm in the irrigation
water, unless 200 ppm potassium was applied (Wright and Niemiera
1985). The reason for reduced growth a t 40 ppm phosphorus with a low
potassium supply could be explained on the basis of elevated inorganic
phosphorus in the plant (Richards and Flees 1962, Sinha and Singh 1982,
Williams 1948).This toxic condition develops when applied phosphorus
levels are greater than required for optimum growth (Hogue et al. 1970).
Phosphorus toxicity is aggravated under low-potassiumconditions ( Sinha
and Singh 1982,Williams 1948)since potassium promotes the conversion
of inorganic phosphorus to nucleic acids and phosphoproteins. Thus re-
duced growth a t low potassium and high phosphorus supplies could be
attributed to a reduction in the synthesis of organic compounds critical
for growth. Another study has shown that increasing tissue phosphorus
levels are inversely correlated ( r = .93)with the development of Vaccinium
corymbosum leaves (Holmes 1960).The latter study indicates that high
tissue phosphorus levels may interfere with iron mobilization.
Tladitionally, phosphorus has been supplied preplant to container
media as superphosphate followed by regular fertilization with a
phosphorus-containing fertilizer. Yeager and Wright ( 1981a), however,
have shown no advantage to amending a pine bark medium with super-
phosphate if plants were fertilized with either a slow-release or soluble
fertilizer. The level of phosphorus supplied by these complete fertilizers
was shown to be a t least 10 ppm, sufficient for growth of I. crenata.
Leaching studies have also shown superphosphate to be an inefficient
source of phosphorus for soilless container media (Yeager and Wright
1982, Yeager and Barrett 1984, 1985; Marconi and Nelson 1984, Havis
and Baker 1985b). For example, Yeager and Wright (1982)showed that
following superphosphate incorporation the medium solution phospho-
rus concentration ranged from 248 ppm during week 1to less that 10 ppm
in week 6.
The practice of preplant incorporation of superphosphate into currently
used soilless media is a holdover from the time when mineral soil was a
component of container media. Phosphorus fixation to a mineral soil
component of past nursery mixes prevented leaching and supplied phos-
phorus over an extended period of time. Soilless pine bark and peat-based
container mixes have been shown to have a low phosphorus fixation
capacity (Yeagerand Wright 1982, Marconi and Nelson 1984),hence the
rapid leaching of phosphorus from soilless mixes. Despite the apparent
inefficiency of preplant incorporated superphosphate, Yeager and Wright
(1982) found no difference in growth between I. crenata fertilized with
superphosphate and 10 ppm phosphorus in the irrigation water. Tissue
phosphorus levels, however, were lower for the superphosphate-treated

plants. This study was conducted for only 12 weeks; thus growth of
woody nursery plants utilizing superphosphate as a sole source of phos-
phorus for longer periods of time (1-2 years) may be limited compared to
plants with a more continuous phosphorus supply.
Use of superphosphate in conjunction with a complete fertilizer may be
deleterious to plant growth. Growth of photinia (Photinia xfraseri) was
less when grown in a soilless medium amended with superphosphate and
a CRF compared with the CRF alone (Yeager et al. 1985). Reduced plant
growth may be due to the initial rapid release of phosphorus from super-
phosphate, which, as discussed, may be toxic to plants.
C. Potassium
The level of potassium in the medium solution that results in optimum
growth of woody nursery crops has been shown to be intermediate
between levels of nitrogen and phosphorus. Gleditsia triacasthos was
found to grow optimally a t 30 ppm potassium in a sand culture study
(Cannon et al. 1960). Gouin and Link (1966) grew %us media in a
peat-sand medium and found that 45 ppm potassium applied in the
irrigation water every 14 days promoted optimum growth. Spiers (1984)
utilizing solution culture, found maximum growth of Vuccinium ashei to
occur in the range of 6-18 ppm potassium. Maximum dry weight of I.
crenata in sand culture occurred a t 25 ppm potassium (Wright and
Niemiera 1985). Brewer (1967) reported optimum potassium levels for
growth of I. crenata to be 120 ppm. However, a relatively high level of
phosphorus (31 ppm) was used, which as discussed above may require
high potassium levels to overcome phosphorus toxicity with I. crenata.
Thus 25-50 ppm potassium in the medium solution appears to be ade-
quate for growth of woody nursery crops.
D. Nitrogen, Phosphorus, and Potassium Ratio

There is general agreement on the elemental ratio of applied nitrogen,
phosphorus, and potassium for optimum growth of woody nursery crops.
Suggested elemental ratios are 6-1-5 for Betula, Acer, Cotoneaster, and
Berberis (Volden 1979), 5-1-3 for Ilex (Wright and Niemiera 1985)and a
range of 5-1-3 to 8-1-4 for several other woody genera (Ingestad 1979).
Both LF as well as CRF formulations with these ratios are commercially
available to the nursery trade.
E. Calcium and Magnesium

Limited literature is available documenting the calcium and magne-
sium levels required for optimum growth of container-grown woody nur-
sery crops. Despite recommendations for calcium of 200-350 ppm and
magnesium of 75-100 ppm in the medium solution (Davidson and

Mecklenburg 1981), Starr and Wright (1984) have shown 5-10 ppm
calcium and magnesium to be sufficient for growth of I. crenata. This
finding is in general agreement with other studies, where 1ppm calcium
was sufficient for growth of I. crenata (Dunham and n t n a l l 1961) and
0.3 ppm calcium sufficient for growth of Prunus persica (Edwards and
Horton 1981). The study by Starr and Wright (1984)also investigated
the affect of calcium and magnesium ratio on growth. As long as the
applied levels of calcium and magnesium were sufficient for growth, the
ratio of Ca :Mg was of no consequence. Additions of dolomitic limestone
to container media to adjust pH results in elevated calcium and magne-
sium levels in the medium solution and plant tissue. Thus, high tissue
calcium and magnesium levels have perpetuated the concept that nur-
sery crops require additional calcium and magnesium for optimum growth.
In fact, Starr and Wright (1984)found increased tissue levels of calcium
and magnesium even though there was no growth response beyond the
5-10 ppm level.
Preplant incorporation of dolomitic limestone [ CaMg(C03)2]in the
mediumis the predominant sourceof calciumandmagnesiumforcontainer-
grown plants. The amount of calcium in the medium solution cannot be
predicted according to limestone rate, since both Starr and Wright ( 1984)
and Chrustic ( 1982)found that a t 8 kg lime/m3of pine bark there was less
calcium in solution than a t 2 or 4 kg/m3. A reduced dissolution rate of
limestone (Berner and Morse 1974)and an increased CEC a t the higher
pH could account for this effect. There is a closer relationship between the
amount of dolomitic limestone applied and the magnesium concentration
in the medium solution (Starr and Wright 1984, Chrustic 1982). In the
absence of dolomitic limestone Fuller and Meadows (1983)have shown
that calcium and magnesium requirements of eight woody ornamental
species can be supplied with calcium and magnesium sulfates. Similar
results were also found by Dickey (1970) for Viburnum suspensum.
Another study has shown that irrigation water may supply sufficient
calcium and magnesium for optimum plant growth (Whitcomb 1984).
F. Micronutrients
Container-grown woody nursery crops in soilless media have been
shown to respond to micronutrient application (Whitcomb 1984). There
are numerous micronutrient sources available for plants grown in con-
tainers. The most common method of supplying these nutrients to plants
is preplant incorporation into the container medium as sulfates, fritted
oxides, or adsorbed to clay particles. The extractability and presumably
the availability of nutrients from these sources after incorporation into a
soilless medium has been investigated by Broschart and Donselman
(1985). An ammonium acetate (pH 7.0) extraction of the medium indi-

cated a fairly steady supply of iron, copper, zinc, and maganese from all
sources for 18 months. Extractable iron from Esmigran (Mallinckrodt,
St. Louis, Missouri)was low throughout the test, which was attributed to
a high affinity of the Esmigran’s clay base for iron. Extractable manga-
nese and zinc from Micromax Plus (Sierra Chem. Co., Milpitas, Califor-
nia), also a source of superphosphate and dolomitic limestone, was lower
than from other sources. The authors attributed this response to manga-
nese and zinc precipitation with superphosphate. However, lower manga-
nese and zinc levels could have been partially due to the presence of
limestone, which results in a higher medium pH and causes precipitation
and increased adsorption of manganese and zinc to the medium. Other
reports have noted similar reductions in availability of manganese and
zinc in response to increased pH in soilless media (Haynes and Swift
1985, Niemiera and Wright 1986a).
In a study by Broschat and Donselman (1985),iron incorporated as
Na2FeDTPAwas leached from the medium by week 30. However, man-
ganese supplied as NaMn2EDTA remained stable within the medium
throughout the experiment. This was attributed to the substitution of
manganese from the chelate by other cations such as iron, which have a
higher affinity for EDTA. Iron was then leached, since its chelation
prevented adsorption to the medium. Mn2+was then adsorbed to the
medium and therefore was in equilibrium with the medium solution.
Incorporation of fritted and sulfate-based micronutrient formulations
into organic media appears to provide an adequate supply of micronutri-
ents with sufficient longevity.
There is no direct relationship between the ratio of micronutrients
applied in a particular formulation and that which is extractable or in the
medium solution. For example, Broschat and Donselman (1985)showed
that even though five times as much iron as manganese was applied as
sulfates to a peat-based medium, about 200 times more manganese than
iron was extractable with NH40Ac. The same relationship was demon-
strated by Haynes and Swift (1985)using CaC12as an extractant and by
Niemiera and Wright (1986a) using distilled water. The above studies
also showed that the ratio of extracted micronutrients to that applied is
influenced by medium pH. For example, the ratio of extractable manga-
nese to iron was greater a t pH 4 than a t 6. This is due to the fact that
individual micronutrients are differentially complexed with organic com-
pounds in organic substrates (Verloo 1980, Stevenson and Ardakani
1972). Furthermore, the amount of a nutrient extracted by one method
a t a particular pH may differ greatly from another method (Haynes and
Swift 1985).
Documentation of the application rates of micronutrients required for
optimum growth of woody ornamentals is limited. Whitcomb (1984)

varied the ratio of sulfate-based micronutrients to be incorporated into
container medium and determined that the interactions between the
unspecified application rates of the various micronutrients affected the
growth of Pyracantha coccinea and Rhododendron obtusum ‘Hinodegiri’.
The application ratio that resulted in the greatest growth was used in the
formulation of Micromax. Concentrations of micronutrients to be applied
through the irrigation system are not well documented for woody orna-
mentals. However, recommendations for several container-grownforestry
species are as follows: iron, 4.0 ppm; manganese, 0.5 ppm; zinc, 0.05 ppm;
boron, 0.5 ppm; copper, 0.02 ppm; molybdenum, 0.01 ppm (Tinus and
McDonald 1979).Similar concentrationsof these nutrients applied through
the irrigation system have resulted in favorable growth with no detect-
able deficiency or toxicity symptoms for a wide range of woody ornamental
species (personal observation ). Container media amended with munici-
pal compost (Lumis and Johnson 1982)or excessive amounts of micronu-
trients (Gilliam and Watson 1981)may result in micronutrient toxicities.
lbxus media exhibited boron toxicity symptoms when the boron concen-
tration of the irrigation water (applied weekly) was 25 ppm or greater
(Gilliam and Watson 1981).
The results above have indicated that micronutrient availability for
plant absorption may be influenced by the micronutrient under consider-
ation, the form in which it is applied, and medium pH. In addition the
assessment of availability of a particular micronutrient may vary with
the extraction method. In light of these difficulties, research is needed to
establish an extraction method in which levels of extractable micronutri-
ents are closely related with growth.
G . Medium pH
The pH of an organic medium for container-grownwoody ornamentals
has been given considerable attention in recent years. The rationale of
adjusting the pH of container media is likely based on the need to adjust
the pH of mineral soil in the range of 5.5-6.5. The greatest single benefit
of liming acid mineral soils is the prevention of aluminum and manganese
toxicity to plants (Tisdale and Nelson 1975). Most organic container
media are inherently low in aluminum and manganese and therefore
toxicity potential is reduced. Furthermore, maximum nutrient availabil-
ity of soils high in organic matter is 1.0-1.5 pH units lower than for other
mineral soils (Lucas and Davis 1961).Peterson ( 1982)has shown that the
optimum pH range for nutrient availability of organic potting mixes is
4.0-5.2, except for calcium and magnesium, which were less available
below 5.2. Thus there appears to be no advantage of liming organic
container media to adjust pH above 5.0-5.5.
Experimental results often show that more vigorous growth and increased
plant quality can be attained a t pH values of 4-5. Chrustic and Wright
(1983) showed decreases in growth of I. crenata and R. obtusum as pH
was adjusted with dolomitic limestone from 3.4 to 7.2. Juniperus chinensis
growth increased slightly when pH was adjusted to 4.1 from 3.4 but
decreased as pH was increased beyond 4.1. In another study, plant
quality of four azalea cultivars was reduced when 4.7 kg limestone/m3
compared to 2.4 kg/m3 was added to a pine bark :sand medium regard-
less of the fertilizer program (Wade et al. 1983). The best growth of
Berberis thunbergi was found to be between pH 4 and 5 (Volden 1979).
The dry weight of Vaccinium corymbosum grown in a peat medium and
amended with various rates of Ca(OH)z increased as pH was raised from
3.9 to 4.3 but decreased a t higher pH (Haynesand Swift 1985).Growth of
Pseudotsuga menziesii decreased beyond pH 5.5 (van den Driessche
1978).The factors that account for reduced growth a t higher pH are many
and interrelated to the extent that single cause-and-effect relationships
can not be established. Additions of limestone have been shown to
stimulate nitrification (Niemiera and Wright 1986b), which leads to a
decreased NH4-N :N03-N ratio in the medium solution. Subsequent
NO3 absorption by plants results in an increased uptake of cations
(Mengel and Kirkby 1982). High cation tissue content has been posi-
tively correlated with organic acid content, which leads to iron immobili-
zation (Nelson and Selby 1974). Liming also increases medium solution
HC03 levels, resulting in HC03 absorption, which can also lead to iron
immobilization (Rutland 1971, Mengel and Kirkby 1982)-hence the
term lime-induced iron chlorosis. The fact that micronutrients are less
available at high medium pH also adds to the complexity of lime-mediated
growth inhibitions.
The evidence presented above supports the conclusion that there is no
advantage to liming organic container media above 5.5 if all nutrients are
supplied to the medium solution in sufficient, non-toxic quantities. Growth
of eight woody ornamental species when supplied with calcium and
magnesium sulfates was equal to plants supplied with limestone a t
2.97 kg/m3 pine bark-sand medium (Fuller and Meadows 1983). More
research is needed to investigate the interaction between species, media,
and lime additions.
V. SOIL TESTING
Many methods of nutrient extraction for mineral soils based upon

weak acid or organic extractants are available and have been applied to
soilless media. Experimental results have noted relatively high correla-
tions between nutrients extracted by these methods and plant nutrient

uptake (Holcomb and White 1980, Prasad et al. 1981a, b, c). However,
methods based upon water extraction of nutrients may more accurately
represent the actual levels of nutrients in medium solution (Lucas et al.
1972) and thereby provide a better management tool for maintaining
optimum nutrient levels in solution.
An established procedure for extraction of the medium solution is the
solution displacement procedure first described by Parker ( 1921).Subse-
quent studies dealing with the procedure have been reviewed by Adams
( 1974). The displacement method reliably characterizes the solution
surrounding the roots. Solution concentration values from this method
(Pearson 1971) are similar to critical nutrient values for optimum growth
established in hydroponic experiments (Bennett and Adams 1970). A
modification of the solution displacement procedure is the pour-through
( P T )or leachate procedure described by Yeager et al. ( 1983)and Wright
(1984).The procedure entails applying water to the medium surface and
collecting the displaced solution (leachate). Nutrient, soluble salt, and
pH analysis of the leachate can be accomplished immediately or sent to
an analytical lab. This method has been shown to give a reliable represen-
tation of the relative nutrient concentration of the medium solution for
either a pine bark or peat-based medium (Wright 1984).Additionally, the
PT procedure has been shown to be as effective in extracting nutrients
from container media as the saturated soil extract procedure (Yeager et
al. 1983).The advantages of the PT procedure are that the medium is not
handled and there is no danger of rupturing CRF particles, causing
erroneously high nutrient readings. Further, no specialized equipment is
required for extracting the solution from the medium, and the extraction
can be accomplished rapidly in the field. Yet to be established for this
procedure are nutrient sufficiency levels in the medium solution required
for optimum plant growth. Since extractable nutrient levels closely approxi-
mate medium solution levels, nutrient levels for optimum growth should
be similar to those established for plants grown hydroponically.
Another procedure using water as an extractant is the saturated soil
extract method (Warncke 1975).This method involves adding just enough
water to saturate the medium. The water is then removed by vacuum and
analyzed for soluble salts, pH, and other nutrients. Other variations of
this procedure employ different ratios of water to media such as 2 :1,
3 : 1, or 5 :1.Acceptable correlation values between extractable nitrogen,
phosphorus, and potassium levels and tissue nutrient concentrations
have been established for the saturated soil extract procedure (Holcomb
and White 1980, Prasad et al. 1981a,b,c).
In view of the wide range of extraction methods currently being used
for organic substrates, the development or adoption of a standardized
procedure would appear necessary. Experimental results could then be

compared and serve as a basis for the development of nutrient sufficiency
guidelines for various woody ornamental species grown in soilless media.
VI. TISSUE ANALYSIS
Tissue nutrient analysis has limited application as a management tool

for container-grown woody ornamentals. This limitation is brought about
by the wide range of nutrient levels found between species (Smith 1978,
Lumis 1974, 1975) and the lack of experimental results establishing
species sufficiency levels. Tissue nutrient levels may also vary depending
upon nutritional programs (Shear et al. 1948),as the application level of
one or more nutrients may influence the uptake and tissue content of
other elements. Medium pH also affects nutrient availability and uptake
by its influence of nitrification and CEC as discussed in an earlier section.
All of these factors make the interpretation of tissue analysis data a
difficult task.
We propose that tissue analysis be used in conjunction with a soil test
as a diagnostic tool when healthy tissue can be compared with tissue
possessing a nutrient disorder. Maintenance of optimum nutrient levels
in the container medium based upon frequent soil testing and nutrient
additions should be the major approach to nutritional management
rather than tissue analysis.
VII. TIMING OF NUTRIENT APPLICATIONS
A. Propagation
The need to apply nutrients during the root initiation phase of propa-
gation is based on the premise that considerable amounts of nutrients are
leached from cuttings during intermittent mist propagation (Good and
lbkey 1966). However, there is no evidence that nutrients added to the
mist water during propagation will increase or influence root initiation
(Wott and lbkey 1967, Sorrenson and Coorts 1968). Also the incorpora-
tion of CRF in the propagation medium (Ward and Whitcomb 1979,
Carney and Whitcomb 1983, Johnson and Hamilton 1977, Still and Lane
1984) shows, with the exception of Juniperus conferta (Johnson and
Hamilton 1977), that root initiation is not enhanced by fertilizer addi-
tions. In some cases additions of fertilizer prior to rooting have actually
reduced rooting ( Booze-Daniels et al. 1984,Woot and lbkey 1967, Sorenson
and Coorts 1968).
Booze-Danielset al. (1984)studied the effects of timed fertilizer appli-

cations in I. crenata ‘Helleri’cuttings during propagation. Delaying the
initiation of fertilization a t weekly intervals after roots were visible
resulted in lower dry weight per cutting as the delay in fertilization
increased. Nitrogen was found to accumulate in cuttings immediately
after roots were first visible, but not before. Thus, nutrients lost via foliar
leaching during root initiation would rapidly be replaced by fertilizers
applied following rooting without any loss in plant growth. Fertilization
before root appearance was not beneficial. The above studies demon-
strate a definite need for fertilizer applications during propagation, but
only after roots are present.
B. During Growth Cycles

Fertilizer applications may be more efficiently utilized when applied
during specific growth phases. Gilliam and Wright (1978) noted that
fertilization of I. crenata ‘Helleri’plants between shoot growth flushes
resulted in the highest tissue nitrogen accumulation and greater shoot
growth compared to plants receiving fertilizer at other times during the
flush. Mertens and Wright (1978)determined that the optimal time of
fertilization in the above corresponded to a period of root elongation that
occurred after the cessaton of shoot elongation and 1-2 weeks before the
next shoot flush. Hershey and Paul ( 1983) demonstrated that ion absorp-
tion by Euonymus japonica growing in solution culture declined during
periods of shoot elongation but increased when shoot elongation ceased.
The cyclic pattern of uptake is thought to be a result of photosynthates
being preferentially used by shoots during shoot elongation, thus reduc-
ing photosynthate transport to roots for growth and nutrient uptake
(Mertens and Wright 1978). When shoot elongation ceases, roots become
the major sink.
During root elongation the rate of nitrogen applied that maximizes
growth and fertilizer use efficiently was investigated by Yeager et al.
( 1980). Ilex crenata ‘Helleri’fertilized two times 1 week apart a t 500 ppm
nitrogen during root growth were as large as plants fertilized weekly a t
300 ppm. The latter was the control since other experiments had shown
this treatment to be the optimum fertilizer rate. Plants fertilized a t 500
ppm every 6 weeks received 4470 less fertilizer and utilized 18%more of the
fertilizer applied. Factors other than the plant’s increased capacity to
absorb nutrients during root growth may have contributed to the greater
fertilizer efficiency. Plants have a compensatory ability to absorb nutri-
ents a t an accelerated rate following periods of nutrient deprivation
compared to plants that have received nutrients continuously (Drew et
al. 1984). Also, the capacity of pine bark to release adsorbed cations into
solution could have extended the supply of nutrients following the 500
ppm applications. In fact, 3 weeks after the 500 ppm nitrogen applica-
tion, 10 ppm nitrogen remained in the medium solution.
C. Fall Fertilization
Late summer and fall applications of fertilizer to woody ornamental
plants have been given considerable attention to relation to winter hardi-
ness and growth the following spring. A review by Pellet and Carter
(1981) on the relationship between nutrition and cold hardiness con-
cluded that “plants fertilized at levels which promote optimum growth
will cold acclimate a t a similar rate and to the same degree as plants
grown under a lower fertility regime and may even exceed cold hardiness
development of plants grown under severe nutrient deficiencies. Supra-
optimum fertility levels can retard cold acclimation.”
Most studies cited in the above review artificially exposed plants to
various temperatures to determine the temperature at which injury occurred.
Only minor reductions in cold hardiness with increasing nitrogen levels
were found. However, studies evaluating plant survival outdoors under
normal nursery storage conditions have demonstrated a response to
fertilizer application (Kelly 1972,Wright et al. 1978).Kelly ( 1972)found a
significant decline in survival of Pyracantha due to increased nitrogen
application rates, and Wright et al. (1978)found major differences in the
survival of two azalea cultivars grown a t three rates of Osmocote and
overwintered a t two locations with different minimum winter tempera-
tures. Thus methods of evaluating cold acclimation and hardiness of
woody ornamental plants may entail more than assessing the injury of
plants exposed to artificially imposed temperatures.
Late summer and fall applications of fertilizer may influence fall tissue
nutrient levels and spring growth. With %us and Forsythia, spring
growth was correlated with tissue levels of nitrogen, phosphorus, and
potassium, during the preceding dormant season (Meyer and Tukey
19651. Tissue nitrogen and subsequent spring growth of Syringa vulgaris
increased as a result of nitrogen applications made during the previous
year (Meyer and Splittstoesser 1969). Fertilizing three Ilex cultivars a t
different rates in the spring, prior to the first spring growth flush, did not
influence the growth of the first flush but did the second (Gilliam and
Wright 1977b). Spring shoot growth of Pseudostuga menziesii was also
dependent on nutrient reserves accumulated during the previous year
(Kruger 1967).However, if too much fertilizer is applied late in summer,
then growth may be prolonged and freeze damage incurred. This response
is due to a delay in vegetative maturity and subsequent cold acclimation
(Fuchigami and Weiser 1981, Fuchigami et al. 1982).
Nutrient uptake by woody plants occurs if medium temperatures are

above freezing (Good and Tbkey 1969, Meyer and Tbkey 1967, Wright et
al. 1983). Wright et al. (1983) grew I. crenata 'Helleri' a t various day/
night temperatures ranging from 6'/2' to 26'/22'C. Percent tissue
nitrogen increased over time a t all temperatures but a t progressively
slower rates as temperature decreased. Bmperatures of 18'/14' or less
prevented visible shoot elongation, although plant dry weights increased
at all temperatures. Subsequently, Wright and Blazich ( 1983) studied
the uptake of nitrogen a t various temperatures and nitrogen applica-
tion rates. Total plant nitrogen increased over time at progressively high-
er rates as both temperature and nitrogen rate increased. Nitrogen rate
had a greater influence on nitrogen accumulation than temperature. Thus,
fertilizer applications should be reduced in late summer and fall to about
one-half that of summer applications to support root elongation, root and
shoot dry weight increases, and cold acclimation. Fall applications of
fertilizer will also insure sufficient tissue nutrient levels for spring growth.
D. Mycorrhizae
&search investigating the use of mycorrhizae to increase the growth of
container-grown plants has shown both positive and negative results
(Crews et al. 1978, Maronek and Hendrix 1979, Molina 1979, Johnson et
al. 1980, Coxwell and Johnson 1985).Maronek et al. (1981),in a compre-
hensive review of mycorrhizae fungi in horticultural crops, concluded
that the growth response to mycorrhizae infection is dependent upon the
host and mycorrhizae species, nature of the host plant root system, soil
environment, and fertility regime. At present the limited understanding
of these interactive factors prevents the utilization of mycorrhizae to
increase the growth of container-grown plants routinely. However, increased
fertilizer cost and an increased awareness of environmental pollution due
to fertilizer runoff may necessitate the use of mycorrhizae to develop
more efficient fertility programs, but more research is needed before
mycorrhizae can be effectively used in container-grown plant production.
VIII. CONCLUSIONS
The interrelationships between the plant, the medium solution, the

container medium, the method of nutrient application, and the amount
of applied fertilizers affect the nutrition of container-grown woody nur-
sery crops. The limited water and nutrient holding capacity of the
container medium requires frequent water and nutrient additions. The
medium solution is the focal point of nutritional management since it is
the immediate source of nutrients for plant absorption. Thus, adequate
nutrient levels as well as an N : P : K ratio of about 5:1:3 should be

maintained in solution to maximize growth. The medium solution, there-
fore is the primary target for the assessment of nutrient availability to the
plant. The pour-through (leachate) method of medium solution extrac-
tion is recommended by these authors for testing soilless container
media. Its simplicity, quickness, and ease of execution make it a valuable
tool to nursery operators. More research is needed to establish medium
solution nutrient levels for optimum growth for a wide range of genera,
different plant and container sizes, and different growth phases.
The dynamic nature of an organic medium warrants emphasis. Over
time, nutrient levels may increase or decrease depending on plant absorp-
tion, leaching, and nutrient additions. Nitrification in a soilless medium
has a dramatic influence on the NH4 :NO3ratio and decreases pH, which
increases the availability of several cations. The medium solution should
be tested frequently so that fertilization practices may be adjusted
accordingly.
Researchers must monitor medium solution nutrient levels imposed by
treatment to determine nutrient requirements of container-grown woody
nursery crops. Only then will there be a basis for comparing experimental
data in relation to nutrient application rate and methods of nutrient
application. Experiments that investigate the influence of nutritional
treatments that are limited to plant data do not increase the understand-
ing of a growth response to treatment.
Results of several experiments have indicated the need to break away
from traditional fertilization methods, which may have been appropriate
when container media contained mineral soil but are no longer appropri-
ate since present-day container media are soilless. We have questioned
the traditional practice of amending container media with superphos-
phate as a source of phosphorus and limestone to adjust media pH to 6.5.
Superphosphate initially supplies the medium solution with very high
levels of phosphorus and then, within a short period of time, supplies
relatively low phosphorus levels, which over time may limit growth.
Current data indicate that there is no benefit in liming organic media to
adjust pH above 5.5. Additionally, solution levels of calcium and magne-
sium that result in maximum growth of some woody species are relatively
low. In some cases, irrigation water and the medium may supply adequate
calcium and magnesium levels for growth of container-grown woody orna-
mentals. If all nutrients are maintained in the medium solution at suffi-
cient levels and ratios, then pH does not appear to be as important as once
thought. More work is needed to investigate the effect of medium pH on a
broad range of genera as well as increasing the duration of the experiments.
Micronutrient requirements for woody nursery crops growing in organic
media need further investigation. Nursery operators seem to be applying
micronutrients as insurance rather than with assurance that the micronu-

trients are actually helping. There is evidence that sufficient micronutri-
ents are released from an organic medium as it degrades in the container
to meet the demands of the plant. In fact, it may be found that no
micronutrients will need to be added if nitrogen, phosphorus, potassium,
calcium, and magnesium are supplied at the correct rate and proportion
to each other, and the pH is maintained in the 4.5-5.5 range.
Fertilizer delivery systems designed to supply nutrients to the plant to
minimize nutrient leaching from containers and surface runoff are needed.
Further development and use of CRF and the practice of applying
nutrients via trickle irrigation systems should provide a way of supplying
nutrients at rates commensurate with plant demands. More knowledge
on changes in plant nutrient demands during the day as well as changes
in nutrient demands throughout the year are therefore needed.
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WARD, J. D., and C. E. WHITCOMB. 1979. Nutrition of Japanese holly during
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WARNCKE, D. D. 1975. Greenhouse soil testing. Paper presented to 5th Soil-
Plant Analysis Workshop, Bridgeton, MO.
WHALLEY, D. N. 1974. Nutrition of hardy ornamental nursery stock. A D A S
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WHITCOMB, C. E. 1984. “Plant Production in Containers.” Lacebark Publica-
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Elemental Status of
Pine Bark-Based Potting Media
R. J. Ogden, l? A. Pokorny, H. A. Mills,
and M. G. Dunavent
Department of Horticulture, University of Georgia
Athens, Georgia 30602
I. Introduction 103
11. Common Chemical Characteristics of
Soil Organic Matter and Softwood Bark 104
111. Cation Exchange Capacity 105
A . Soil Organic Matter 105
B . Bark and Wood Wastes 106
I v. Nitrogen 108
A . Nitrogen Reactions and 'lkansformations in Pine
Bark-Based Container Media 108
V. Phosphorus 112
VI. Potassium 113
VII. Soil Reaction, Lime, and Calcium 114
A. pH 114
B. Lime 115
C . Calcium 116
VIII. Magnesium 117
IX. Micronutrients 119
A . Boron 119
B. Copper 122
C . Iron 123
D. Manganese 123
E . Zinc 124
I. INTRODUCTION
Legislation by federal, state, and local governments during the 1960s

and 1970s was aimed a t controlling and/or eliminating environmental
pollution. Many industries therefore placed emphasis on conversion of
waste materials into useful and profitable products. Bee bark, a by-product
of the forestry industry, was initially considered useless and was a costly
waste disposal problem. Today, tree bark finds extensive horticultural
use, as a landscape mulch (Aaron 1976; Bollen and Glennie 1961, 1963;
103
104 R. J, OGDEN, F. A. POKORNY, H. A. MILLS, AND M. G. DUNAVENT
Gray 1973; Wright and Fitzgerald 1969), as a potting medium and/or

component (Aaron 1972; Joiner and Conover 1965; Pokorny 1966; Pokorny
and Gugino 1967; Rigby 19631, and as a soil amendment (Bollen and
Glennie 1961, 1963; Giddens and Baxter 1965; Wright and Fitzgerald
1969). Lesser uses for bark are as a packing material for bare-root plants
(Gartner et al. 1971), as a propagation medium (Laiche 1972; Pate1 and
Gammel 1976; Pokorny and Perkins 1967; Pokorny and Thurman 1965;
'lhrner 19731, and as a mulch around bedded container plants (Aaron 19731.
Pine bark is often substituted for moss peat in container production of
woody ornamentals and is sometimes used as a medium component in
greenhouse flower crops because of its low cost (Laiche 1972; Pokorny and
Perkins 1967), widespread availability (Laiche 1972, Pokorny 1979),
pathogen-suppressive benefits (Guginoet al. 1973; Hoitink 1980a,b),and
the slow rate of humification (Allison and Murphy 1963; Maas and
Adamson 1972; Sproull 1969). In contrast to organic soils and many
types of moss peat, pine bark represents relatively undecomposed organic
matter in which the original structure is identifiable (Airhart et al. 1978;
Pokorny and Wetzstein 1984).
Considerable information exists concerning chemical properties and
fertilizer reactions in mineral soils, but comparatively little is known
about these properties and reactions in pine bark-based container media.
Similarities exist between chemical structures of organic fractions found
in natural soils ( Feustel 1939a,b; Lewis and Broadbent 1961; Stevenson
and Ardakani 1972; Tisdale and Nelson 1975) and pine and other soft-
wood barks (Browning and Sell 1957; Fahey and Kurth 1957; Porter 1973;
Sjostrom 1981; Sproull 1969). These similarities suggest ways in which
pine bark-based potting media may interact with added nutrients.
11. COMMON CHEMICAL CHARACTERISTICS OF

SOIL ORGANIC MATTER AND SOFTWOOD BARK
Chemical composition of wood from softwood and hardwood trees is
fairly constant, while that of bark is variable because of chemical changes
that occur after cells are formed (Porter 1973). Bark of Pinus radiata (see
lhble 4.1 for botanical authorities) contains numerous polyphenol poly-
mers, including condensed tannins and phenolic acids, the chemistry of
which is uncertain (Porter 1973). Theoretical structures for lignin and
polyflavenoid compounds in I? radiata bark indicate the presence of
phenolic -OH groups (Porter 1973). Chang and Mitchell (1955) found
significant quantities of these compounds and groups present in bark of
both softwood and hardwood tree species. Slash pine (I? elliottii) bark
contains carboxyl (COOH),carbonyl (CO),and hydroxyl (OH)groups in
various fractions (Browning and Sell 1957),whereas carboxyl and pheno-
lic and alcoholic -OH groups are present in bark extracts of white fir
4. ELEMENTAL STATUS OF PINE BARK-BASED POTTING MEDIA 105
Table 4.1. Botanical Names and Authorities of Plant

Species Referred to in Text
Abies concolor Lindl.

Aglaonema commutatum Schott. ‘Fransher’
Gardenia jasminoides Ellis ‘Mystery’
G . jasminoides Ellis ‘Radicand
Zlex crenata Thunb. ‘Helleri’
I . uomitoria Ait. ‘Nana’
Juniperus chinensis L. ‘San Jose’
J. horizontalis Moench ‘Plumosa’
J. horizontalis Moench ‘Plumosa Compacta’
J . uirginiana L. ‘Sky Rocket’
Ligustrum sinense Lour. ‘Variegata’
Photinia xfraseri Dress
Pinus clausia (Chapm.)Vasey
l? echinata Mill.
l? elliottii Engelm.
I? palustris Mill.
l? pungens Lamb.
l? radiata Don.
l? rigida Mill.
l? serotina Michx.
I! taeda L.
Pittosporum tobira (Thunb.)Ait. ‘Variegata’
Pyracantha koidzumii Xl? coccinea M. J. Roem. ‘Mohave’
Rhododendron obtusum (Lindl.)Planch. ‘Hinodegiri’
R . obtusum (Lindl.)Planch. ‘Rosebud’
R. simsii Planch. ‘Redwing’
Zhxus xmedia Rehd. ‘Anderson’
are similar to those found naturally occurring in soil organic matter and
known to be associated with (1)soil pH (Tisdale and Nelson 1975), ( 2 )
active cation adsorption and exchange attributable to colloids, made
primarily of lignin (Feustel 1939a,b), and ( 3 )chelation (Stevenson and
Ardakani 1972). Chelation in naturally occurring organic materials is
important because it may either increase or decrease availability of
elements, especially plant micronutrients (Miller and Ohlrogge 1958).
However, no direct evidence of chelation by pine bark seems to have been
reported in the literature.
111. CATION EXCHANGE CAPACITY
A. Soil Organic Matter

Organic matter is known to increase soil fertility, and a correlation
between organic matter content and cation exchange capacity (CEC)is
well established (Stevenson and Ardakani 1972).Cation exchange refers
to the relatively weak attraction of cations in soil solution to negatively
106 R. J. OGDEN, F. A. POKORNY, H. A. MILLS, AND M. G. DUNAVENT
to the relatively weak attraction of cations in soil solution to negatively

charged sites on clay micelles or organic matter. The number of negatively
charged sites available for cation adsorption in organic colloids is pH
dependent, with elevated pH inducing greater adsorption because of the
ionization of exposed hydrogen (Buckman and Brady 1969). Optimum
pH for essential element availability is 1.0-1.5 units lower in organic soils
than in mineral soils (Lucasand Davis 1961), with base saturation of 70%
corresponding to a pH of 5.5. Lower dissociation constants for hydrogen
in organic soils (4.5 x 10 for sphagnum peat vs. 4.0 x 10 for clay) have
been proposed as one reason for satisfactory growth a t pH values that
would not adequately support plant growth in most mineral soils. Lewis
and Broadbent (1961) found that CEC of staten peaty muck subsoil is
due to phenolic and carboxyl groups and these groups exhibit a wide
range of pK, values.
B. Bark and Wood Wastes
Evidence of CEC in bark and wood wastes is reported in the literature
(Baxter 1969; Bollen and Glennie 1963; Brown and Pokorny 1975; Davey
1953; Lunt and Clark 1959). A summary of CEC for pine bark, pine
sawdust, and peatmoss is presented in Table 4.2. Tkatment of bark and
wood wastes with nitrogen fertilizer followed by a period of composting
has been shown to increase CEC substantially because the breakdown of
large particles increases the surface area (Bollen and Glennie 1963; Davey
1953). Bollen and Glennie (1963),working with Douglas fir bark, reported
CEC of ground bark to be two to three times that of a silt loam soil.
Phenolic acid polymers, which bark contains, have neutralization equiva-
lents ranging from 100 to 200 meq/100 g.
Table 4.2. Cation Exchange Capacity (CEC)of Pine and Hardwood Barks,
Sawdusts, and Peatmoss Soil Amendments
Soil CEC
amendment ( meq/100 g ) Reference
Pine bark 30-71 Brown and Pokorny (1975),

Neal and Wagner ( 19831,
Self et al. (1967)
Pine sawdust 28 Goh ( 1979)
Hardwood bark 44 Scott and Bearce ( 1972)
Hardwood sawdust 18-77 Baxter ( 1969), Scott and

Bearce (1972)
Peatmoss 30-135 Fushman (1980), Goh (19791,

Scott and Bearce ( 1972)
Table 4.3.Comparison of Cation Exchange Capacity (CEC),Expressed as meq/100 g

and meq/100 cm3, of Several Soils, Artificial Potting Media, and Potting
Medium Components
~ ~~ ~ ~
Soil type, Bulk CEC Sample CEC

potting medium or density (meq/ vol ( meq/
medium component (g/cm’) 100 g ) (cm3) 100 cm3) References
Altavista 1.64 3.97 61 6.5 Perkins

A horizon et al. (1982)

C horizon et al. ( 1982)

Bt2 horizon et al. (1982)
Cecil Btz 1.43 7.02 70 10.0 Perkins

horizon et al. (1982)
Sand 1.56 1.00 64 1.6 Brown and

Pokorny
(1975)
Coarse 0.23 47.0 439 13.0 Brown and

pine bark Pokorny
(1975)
Coarse bark : sand 0.56 23.0 179 12.8 Brown and

( 3 : l v/v) Pokorny
(1975)
Coarse bark :sand 0.97 8.0 103 7.8 Brown and

(1:l v/v) Pokorny
(1975)
Coarse bark :sand 1.29 4.0 78 5.1 Brown and

(1:3 v / v ) Pokorny
(1975)
Peatmoss :sand 1.20 5.8 83 7.0 Joiner and

(1:l v/v) Conover
( 1965)
Peatmoss :perlite 0.10 11.2 1020 1.1 Joiner and

(1:3 v / v ) Conover
(1965)
Cation exchange capacity of various pine bark potting media is found

in the literature (Brown and Pokorny 1975; Joiner and Conover 1965,
1967; Neal and Wagner 1983). Brown and Pokorny (1975)reported that
CEC increases from 1to 57 meq/100 g as the proportion of pine bark in a
bark-sand potting mixture is increased from 0 to 100%. Hoitink and
Poole (1980) proposed that a small quantity of fine bark particles be

retained so as to increase CEC to acceptable levels without adversely
affecting porosity. Addition of lime to bark significantly increases CEC
(Niemiera and Wright 1984).
Several researchers (Brown and Pokorny 1975; Joiner and Conover
1965)have questioned whether the expression of CEC, as it is applied to
native soil, is suitable and meaningful when evaluating artificial potting
media and medium components, e.g., bark, peat, vermiculite, perlite.
Cation exchange capacity of native soils is expressed as milliequivalents
per 100 g dry soil. The advantage of this method of CEC expression for
soils is that different cations are additive when expressed on a weight
basis (Black 1960).Joiner and Conover ( 1965) found large differences in
CEC between 2 bark :1 perlite (v/v mixture) and 1 peat : 1 sand and 1
bark :1sand media when CEC is expressed as meq/100 g. However, when
expressed as meq/100 cm3of potting medium, differences in CEC among
media are minimal. Thus Joiner and Conover ( 1965)concluded that CEC
is valuable only when expressed on a volume basis in container plant
production. Reporting CEC on a weight basis for soilless potting sub-
strates overestimates this parameter because of rather substantial disparities
in bulk density exhibited by artificial substrates and native soils (Thble
4.3). Brown and Pokorny (1975) concurred with Joiner and Conover
(1965) that CEC of artificial potting substrates and components be
expressed on a volume basis (meq/100 cm3).Thus a pine bark medium
with a reported CEC of 57 meq/100 g would have 13 meq/100 cm3(Brown
and Pokorny 1975).Comparable CEC values, expressed as meq/100 cm3,
for a pine bark potting medium are reported by Yeager et al. (1980).
IV. NITROGEN
A. Nitrogen Reactions and Transformations in Pine Bark-

Based Media
Milled pine bark as a container medium has a total nitrogen content of
0.28% (Thble 4.4) with 0.33 ppm water-extractable NH4-N and 0.67 ppm
N03-N (Pokorny 1979). Nitrogen is apt to be the element most limiting
plant growth in container production because large quantities of water,
applied a t frequent intervals, result in extensive leaching. Thus current
greenhouse and nursery practices require frequent periodic applications
of nitrogen and other nutrient elements to sustain plant growth. Dickey
et al. (1978)reported that ‘East Palatka’ holly plants of excellent quality
are produced in containers by applying nitrogen a t 899 kg/ha-yr. Wax-
leaf privet, ‘Formosa’azalea, Japanese pittosporum, and lantana respond
with increased growth to applied nitrogen up to 1685 kg/ha-yr.
Table 4.4. Total and Water-Extractable Elemental Content of Milled Pine Bark
Water-extractable (ppm)
Total (70)
Element Pokorny (1979) Pokorny (1979) Neal and Wagner (1983)
N 0.28 - -
NHI-N - 0.33 -
NOS-N - 0.67 -
P 0.02 9.00 6.9
K 0.10 27.60 6.2
Ca 0.51 7.60 40.6
Mg 0.14 1.60 5.9
Fertilizer programs using dry, liquid, and/or slow-release nitrogen

fertilizers or combinations thereof are found in greenhouse and/or nur-
sery practice. Urea, ammonium, and nitrate nitrogen sources are the only
forms of nitrogen used in container fertilization programs (Dickey et al.
1978; Nelson 1981).
In contrast to mineral soils, little is known about nitrogen reactions
and transformations that occur in pine bark-based potting media.
1. Urea. Urea adsorption in soil is directly related to organic matter
content and is unaffected by CEC, pH, or clay content (Chin and Kroontje
1962).Chin and Kroontje (1962)believed that chemisorption and physi-
cal adsorption are factors in urea retention because urea is known to form
complexes with inorganic salts, fatty acids, waxes, paraffin, long-chain
amines, straight- and branched-chain aliphatic compounds as well as
many nitrogen compounds and resins. Urea adsorption is not affected by
potassium ion concentration; thus Chin and Kroontje (1962) surmised
that adsorption of urea is by a mechanism different than that responsible
for cation adsorption. Physically adsorbed urea is easily desorbed by
water; chemically adsorbed urea, in the form of relatively stable organic
matter-urea complexes, is only partially and slowly dissociated by dilu-
tion (Chin and Kroontje 1962).
Urease activity in soil is independent of either oxygen concentration or
initial urea concentration (Wagenet et al. 1977). Wagenet et al. (1977)
postulated that stimulation of microbial activity may increase the rate of
urease activity over time and that oxygen concentration may then play a
role in conversion of urea to ammonium. Several organic compounds are
known to inhibit urease activity in soils (Bremner and Douglas 1971),
and they and other structurally similar compounds are found in some tree
barks (Chang and Mitchell 1955; Fahey and Kurth 1957; Hergert 1960;
Porter 1973; Sproull 1969).
Urea is easily leached from a 100%pine bark medium unless hydrolyzed
and adsorbed by the potting medium or the plant (Wright 1983a).Hydrol-

ysis of urea to ammonium occurs within 24-40 hr after application
(Wright 1983a).Ogden ( 19821, investigating reactions of applied-nitrogen
sources in a pine bark medium, suggested that some urease activity
occurs in pine bark; however, conversion of urea to ammonium is slower in
pine bark than in soil. Adding lime initially increases urease activity, but
Ogden (1982)found little net difference between limed and unlimed bark
after 10 and 21 days of incubation. In addition, Ogden’s data showed a
rapid decrease in urea solution concentration in contact with bark parti-
cles, suggesting some type of reaction that removes urea from solution.
Ogden postulated that urea reacts directly with bark particles (perhaps
covalent bonding) or is bound by microbial activity without prior conver-
sion to ammonium (Gugino et al. 1973).
2. Ammonium. Ammoniacal fertilizers are used extensively as a nitro-
gen source because of their comparatively low cost in contrast with
N03-N fertilizers; in addition, ammoniacal nitrogen is less subject to
leaching loss and denitrification (Huber et al. 1977). Plants, however,
differ in their response to applied NH4-N fertilizer (Bollen and Glennie
1961; Ingram and Joiner 1982).With many plants, nitrogen uptake in the
ammoniacal form can have deleterious effects on plant growth (Maynard
et al. 1966).Ammoniacal nitrogen will interfere with cation uptake (Kirkby
1968) especially potassium, calcium, and magnesium. Perhaps it is no
coincidence that ammonium toxicity symptoms may resemble deficiencies
of any of these elements (Barker et al. 1967; Maynard et al. 1966; Wilcox
et al. 1973). The suppression of calcium and/or magnesium uptake by
ammoniacal nitrogen (Kirkby 1968)can induce blossom end rot in tomato
fruit, a condition that causes the fruit to be unsalable (Jones 1974;
Wilcox et al. 1973).
No symptoms of ammonium toxicity are reported when tomato plants
are grown in 100% pine bark and fertilized with 100%ammoniacal nitro-
gen (Mills and Pokorny 1978a; Pokorny et al. 1977). However, with
ammoniacal nitrogen, tomato growth is suppressed and plants border on
nitrogen insufficiency (Mills and Pokorny 1978a);this condition is attributed
to ammoniacal-nitrogen binding by pine bark (Mills and Pokorny 1978a).
Thomas and Perry ( 1980)showed that small quantities of ammonium are
bound by pine bark but question the extent to which this binding could
suppress growth. However, Wright and Yeager (1980)reported that 1.5
mg of nitrogen per gram of pine bark is bound when ammonium solutions
are leached through a pine bark medium. Thus, the quantity of bound
nitrogen in a 3-liter container filled with pine bark is greater than the
quantity of ammonium ions used by a rooted holly liner during the course
of a single growing season (Foster et al. 1983; Wright and Yeager 19801.
Extent of ammoniacal-nitrogen adsorption by pine bark is pH depend-
ent. At pH 3.8,20% of applied NH4-Ncan be removed (Foster et al. 1983).

Increased NH4-N adsorption in limed bark is attributed to pH-dependent
charges found on phenolic and carboxyl groups (Browning and Sell 1957;
Chang and Mitchell 1955; Porter 1973).When pH is increased, hydroxyl
and carboxyl groups dissociate, exposing negatively charged sites on
pine bark surfaces (Foster et al. 1983).A competitive effect for exchange
sites between NH4-N and other cations is postulated by Ogden ( 1982).At
pH 3.3 only NH4-N is adsorbed by pine bark particles. With a rise in pH
to 4.4, adsorption of calcium and magnesium increases; continued increase
in adsorption of NH4-Nand potassium occurs a t pH 5.5 (Fosteret al. 1983).
Nitrogen insufficiencies can be anticipated in plants growing in pine
bark media after fertilization with 100% NH4-N unless all available sites
for NH4-N binding are saturated. Foster et al. (1983) reported that 20
days are required for equilibration to occur between a 50 ppm NH4-N
fertilizer solution and adsorption sites on pine bark particles. During this
equilibration period, plant growth is limited by nitrogen insufficiency
unless NH4-N is incorporated preplant in sufficient quantity to saturate
all binding sites (Foster et al. 1983).Leaching of NH4-N and N03-N from
pine bark media is similar once the adsorptive capacity for NH4-N ions
by bark is satisfied (Wright and Yeager 1980).The fate and availability of
NH,-N ions bound to bark particles is still unknown. The creation of amine
or amide bonds to carbon in bark cannot be disregarded (Ogden 1982).
Effects of the nitrification process on pH, NH4-N :N03-N ratio, and
cation availability in a pine bark medium have recently been studied by
Niemiera and Wright ( 1986a,b).Their work shows that liming stimulates
the nitrification process, resulting in a lowering of pH as well as the
NH4-N :N03-N ratio in medium solution. Cation availability, however,
was increased. Niemiera and Wright (1986b) suggest that woody orna-
mental plants, especially those species which achieve maximum growth
with a high NH4-N:N03-N ratio, the nitrification process can be con-
trolled by adding only small quantities of lime to the medium. Thus, the
pine bark remains acid, which slows the nitrification process.
3. Nitrate Nitrogen. Nitrate nitrogen in mineral soils is mobile in soil
solution and is therefore subject to loss by leaching (Huber et al. 1977)
and denitrification (Prasad et al. 1971; Rovira and Davey 1974) unless
immobilized by soil microbes or absorbed by plant roots. In properly
managed cropped soils, loss of N03-N by leaching is not very great (Lyon
and Buckman 1951), while loss by denitrification in many soils is esti-
mated a t 10-3070 of applied nitrogen (Parker 1972). Denitrification in
soils is enhanced by the presence of plant roots (Rovira and Davey 19741,
the form of carbon substrate present (Bremner and Shaw 19581, and
moisture contents above field capacity (Stephenson et al. 1969).
Nitrogen in the N03-N form is readily leached from a pine bark medium
(Foster et al. 1983; Wright and Yeager 1980). Under greenhouse and
nursery conditions, most applied N03-N will be leached from a pine bark
medium after four irrigations in which 2.5 cm ( 1in. ) of water is applied a t
each irrigation (Foster et al. 1983). Thus, plants will be exposed to
nitrogen insufficiency unless N03-N is reapplied or a slow-release nitro-
gen form is initially incorporated into the medium (Foster et al. 1983).
Liming pine bark appears to have little influence on leaching of N03-N
as Foster et al. (1983) recovered 100% of applied N03-N in leachate
regardless of pH. However, Ogden (1982) reported a six- to sevenfold
reduction in N03-N when pine bark is limed. Reduction in water-extractable
nitrogen with liming is attributed to either microbial binding or
denitrification.
Nitrogen loss by denitrification from pine bark media is postulated by
Mills and Pokorny (1978a,b).The porous structure of pine bark (Airhart
et al. 1981; Pokorny and Wetzstein 1984)may provide anaerobic pockets
and favorable conditions for denitrification. Stewart et al. (1981),study-
ing denitrification in several potting media, reported that nitrogen loss is
greatest from pine-sand ( 17.7%)and pine-soil (18.9%)potting media and
least in redwood (9.3%)and peatmoss (8.1%)media.
V. PHOSPHORUS
Southern pine bark contains about 0.02% total elemental phosphorus

in its composition (Pokorny 1979);however, water extracts contain 6.9-9
ppm elemental phosphorus, which is about l/C, the concentration of this
element found in Hoagland’s solution (Jones 1983; Neal and Wagner
1983; Pokorny 1979). Eight to 19 ppm water-extractable phosphorus is
regarded as optimal for tomato plants (Bruce et al. 1980) yielding leaf
tissue levels between 0.5 and 1.0% (Plank 1979). Yeager and Wright
(1982a)reported that 10 ppm phosphorus in soil solution of a pine bark
medium is optimal for growth of Ilex crenata ‘Helleri’. Associated leaf
tissue levels of 0.32% are reported by Yeager and Wright (1982a).
Phosphorus deficiency of greenhouse tomatoes growing in a fresh bark
substrate is reported by Cotter ( 1979).Associated with phosphorus defi-
ciency symptoms are reduced growth and low tissue phosphorus levels.
Plants grown in aged bark made normal growth; leaf tissue phosphorus
content was adequate and plants were free of phosphorus deficiency
symptoms. Cotter ( 1979)attributed suppressed growth in the fresh bark
substrate to microbial competition and demand for available phosphorus.
Superphosphate (0-20-0) is commonly suggested as a preplant phos-
phorus source either used alone or as a component of mixed fertilizers
(Bruce et al. 1980; DeWerth 1971; Spivey et al. 1974). Phosphorus is
usually incorporated during the soil mixing operation a t the rate of

0.7-2.4 kg/m3 (Dickey et al. 1978; Prasad 197913). However, preplant
incorporation of phosphorus in a pine bark medium is questioned (Babcock
et al. 1982; Wright 1983b; Yeager and Wright 1981). Pine bark binds
small quantities of phosphorus (7.5pg/g bark) (Yeagerand Barrett 1984;
Yeager and Wright 1982a). Leaching of phosphorus from pine bark is
similar to that occurring in high organic soils lacking in inorganic col-
loids, where phosphorus is weakly adsorbed and the adsorbed phospho-
rus is highly soluble (Fox and Kampreth 1971). Where slow-release
granular or water-soluble complete fertilizers are an integral part of the
container fertilization program, no growth increase can be expected by
preplant incorporation of phosphorus (Babcock et al. 1982; Yeager and
Wright 1981).
The data of Prasad (1979a)conflict with results of Yeager and Barrett
(1984)and Yeager and Wright ( 1982a, 1982b).Prasad ( 1979a)suggested
that phosphorus retention by bark is considerable, occurring mostly
during the first 6 weeks after phosphorus application. Phosphorus reten-
tion in bark, reported by Prasad (1979a),is correlated with iron content.
The total amounts of iron, copper, and zinc in shoot tissues of I. crenata
‘Helleri’are increased as pine bark substrate phosphorus levels are increased
from 0 to 10 ppm. At phosphorus levels higher than 10 ppm, tissue iron,
copper, and zinc tend to decrease. Manganese concentration of leaf tissue,
in contrast, increases with increasing substrate phosphorus levels up to
30 ppm (Yeager and Wright 1982a). In roots, iron content is erratic,
copper content is unaffected, and zinc decreases with increasing sub-
strate phosphorus (Yeager and Wright 1982a).
VI. POTASSIUM
Potassium is a constituent of tree bark (Koch 1972; Young 1971),but

whether this potassium is available for plant use and to what extent is
uncertain. Matkin et al. (1957)and Rigby (1963)suggested that potas-
sium derived from bark may contribute to the initial nutrition of new
container plantings, but the quantity is considered minimal by Aaron
(1976).Sproull and Pierce (1963)reported traces of potassium in mixed
southern pine bark, while Pokorny (1979)reported total potassium con-
tent of ashed bark of 0.10% (lhble 4.4). Water extracts of unfertilized pine
bark contain 6-28 ppm potassium (Neal and Wagner 1983; Pokorny
1979),which is considered insufficient for growth of greenhouse crops and
tomatoes (Bruce et al. 1980; Warncke and Krauskoff 1983). However, G.
E. Smith (personal communication) states that woody ornamental shrubs
growing in a pine bark potting medium and fertilized with either com-
114 R. J. OGDEN, F. A. POKORNY, H. A. MILLS, AND M. G . DUNAVENT
plete liquid or slow-release fertilizers grow well a t soil test element levels
in the low to acceptable range based on the saturated media extract
method (Warncke and Krauskoff 1983).
Potassium, as a fertilizer element, can be supplied in greenhouse
and/or nursery container fertilization programs by preplant incorpora-
tion of complete slow-release fertilizers (Whitcomb 1984a) or by top
dressing with slow-release fertilizers such as Osmocote (DeWerth 1971;
Whitcomb 1984a). With liquid fertilization programs, potassium chlo-
ride, potassium sulfate, potassium nitrate, or sulfate of potash-magnesia,
all water-soluble potassium sources, may be used (Dickey et al. 1978;
Whitcomb 1984a). Whitcomb (1984a) suggested that 0.5 kg/m3 of K20
applied or released over a growing season is sufficient for plant growth
and to maintain potassium tissue levels in the acceptable range.
An inverse relationship is reported to exist between potassium move-
ment and total porosity and percolation rate in pine bark-sand media
(Brown and Pokorny 1977). Media containing low percentages of sand
exhibit greater total porosity and percolation rates than media containing
high percentages of sand; decreasing sand in the medium is associated
with increasing CEC and residual potassium in the medium profile with
lesser retention a t the 10-20 cm depths (Brown and Pokorny 1977).
Nitrogen source can enhance or suppress potassium retention by pine
bark media. More potassium is retained by pine bark when urea is the
nitrogen source (Ogden 1982) in comparison with NH4-N and N03-N
sources. Potassium retention is similar when plants growing in pine bark
are fertilized with either NH4-N or N03-N (Ogden 1982). Limed media,
fertilized with urea, exhibit reduced potassium retention in contrast to
unlimed bark. This reduced potassium retention is probably the result of
competition for exchange sites between potassium and calcium dissoci-
ated from lime (Ogden 1982). Lime, used in conjunction with NH4-N or
N03-N, enhances potassium retention over unlimed bark (Ogden 1982).
VII. SOIL REACTION, LIME, AND CALCIUM
A. pH
Soil reaction is important because pH is influential in controlling the
availability of essential plant nutrients (Scarseth and Volk 1949).Gener-
ally, as pH rises, micronutrient availability declines; conversely, as pH
declines, micronutrients are more available to the plant (Whitcomb 1984a).
In mineral soils, aluminum and manganese may become toxic and phos-
phorus availability may be reduced a t pH of less than 4.5-4.0 (Buckman
and Brady 1969). Optimal pH for plant production in organic soils is
1.0-1.5 units lower than in mineral soils (Lucas and Davis 1961). Acid-
sensitive crops (sweet clover and alfalfa) can be grown a t soil acidity as
low as 4.1 (Lucas and Davis 1961). Plants can tolerate low pH when
grown on organic soils because of the “acidoid” properties of these soils.

The tolerance of plants to acidity increases with the apparent dissocia-
tion constant of the organic soil (Lucas and Davis 1961).
Pine bark is strongly or very strongly acidic (X-ible 4.5). Brown and
Pokorny (1975)reported a pH of 4.1 for a bark mixture of E! taeda and l?
echinata, while Cotter ( 1974)found a pH of 5.8 with a pine and Douglas
fir bark mixture. Pine bark pH does not change drastically upon aging
(Pokorny 1979).
B. Lime
Preplant incorporation of limestone into the potting substrate is a
common cultural management practice (Dickey et al. 1978; Nelson 1981).
Lime additions may range from 3 to 15 kg/m3 depending upon the
substrate and degree of pH correction required (Nelson 1981; Prasad
1979b),but generally with pine bark media, additions range from 1.8 to 8
kg/m3 dolomite (Goldie 1979; Pokorny 1979; Self and Pounders 1974).
Lime recommendations for peat, organic, and mineral soils are usually
derived from titration curves, from which lime additions are estimated
(Dunn 1943; Goh 1979; Jones and Hoover 1949). Lime additions for pine
bark are often based on recommendations for peat-based media (Riggins
1978). Lime requirement additions for pine bark are given by Natarella
and Pokorny (1977)(Tbble 4.6).
The practice of liming bark media for pH control is questioned by
several investigators (Chrustic and Wright 1983;Whitcomb 1984a; Wright
1983b). Poor plant growth a t low pH is not necessarily due to high
hydrogen ion concentration, but to deficiency of calcium (Albrecht 1941;
Moser 1943). Whitcomb ( 1984a), studying azalea growth in soilless
potting media, controlled pH over a range of 3.0-8.2 with nonessential
compounds to increase or decrease hydrogen ion concentration. Azaleas
grew well over the test pH range provided calcium and magnesium
requirements were met. Chrustic and Wright (1983)report no advantage
to liming 100% pine bark in which I. crenata ‘Helleri’and Rhododendron
Table 4.5. Bark pH of Various Pinus Species
Species PH Reference
l? clausia 3.8 Martin and Gray (1971)

l? echinata 3.7
l? elliottii 3.7
P palustris 3.5
l? pungens 3.8
l? rigida 3.8
l? serotina 3.8
l? taeda 3.8
l? radiata 4.8 Prasad (1979a)
116 R. J. OGDEN, E A. POKORNY, H. A. MILLS, AND M. G. DUNAVENT
Table 4.6. Lime Requirement for a Pine Bark Potting Medium
Amount required to equilibration p H

( kg/m”
Ca/lO g bark Equilibration
( meq ) PH Ca(OHh CaC03 Ca(Mg)COs
0 3.90 0.0 0.0 0.02

2 4.90 1.7 2.3 2.40
4 5.84 3.3 4.5 4.90
6 6.32 4.9 6.8 7.30
obtusum ‘Rosebud’ were grown. Shoot and root growth were, in fact,
suppressed as preplant dolomitic limestone was increased from 0 to 8
kg/m3 (pH 3.4-7.2). Similar findings of either no growth response or
growth suppression were reported for Gardenia jasminoides ‘Radicand
( Laiche 1982),Juniperus horizontalis ‘Plumosa Compacta’ ( Sartain and
Ingram 1984), J. horizontalis ‘Plumosa’ (Yeager and Ingram 1983), R.
obtusum ‘Hinodegiri’ (Yeager and Ingram 1983), R. simsii ‘Redwing’
(Sartain and Ingram 1984), Pittosporum tobira ‘Variegata’ (Cobb and
Zarko 1983),and J. uirginiana ‘Sky Rocket’ (Cobb and Zarko 1983).
Positive growth response in relation to dolomite additions in pine bark
media are reported for G. jasminoides ‘Mystery’ (Fuller and Meadows
1983),G. jasminoides ‘Radicand (Fullerand Meadows 1983),I. uomitoria
‘Nana’(Fuller and Meadows 1983),Ligustrum sinense ‘Variegata’(Laiche
19821,Pyracanthu ‘Mohave’(Laiche 1982),J. chinensis ‘SanJose’(Chrustic
and Wright 1983),and Photinia Xfraseri (Nash et al. 1983).Increases in
plant growth with dolomite additions are not ascribed to changes in pH,
but are attributed to the calcium and/or magnesium supplied by the
dolomite (Fuller and Meadows 1983;Nash et al. 1983)or to a reduction in
the K :Mg ratio to a level where potassium-magnesium antagonism is
not important. Proper plant growth can be achieved with pine bark
media, regardless of pH, by providing sufficient essential elements in an
available form with correct balance.
Niemiera and Wright (1984)reported that changing the hydrogen ion
concentration of a pine bark medium from pH 4.0 to 5.0 results in a 53%
increase in CEC. In addition, these researchers reported that adding
preplant dolomite up to 6 kg/m3 decreases NH4-N, iron, manganese, and
zinc in soil solution. Conversely, N03-N, calcium, and magnesium con-
centration in soil solution is increased (Niemiera and Wright 1984).
C. Calcium
Preplant addition of limestone, dolomitic or calcitic, not only adjusts
hydrogen ion concentration, but also supplies calcium. Generally, increas-
ing the quantity of limestone added to the substrate results in increasing
calcium concentration of the soil solution and elevation of calcium content

in plant tissue (Fuller and Meadows 1983;Starr and Wright 1984).Gypsum
( CaS04)is also used as a calcium fertilizer (Jones 1983)although it may
be too soluble for general use (Starr and Wright 1984;Whitcomb 1984a).
Dissolved salts, especially calcium and magnesium in irrigation water,
contribute significantly to plant nutrition and must be accounted for
when establishing a fertility program. As pointed out by Whitcomb
(1984a), water source should be analyzed so that lime, calcium, and
magnesium additions can be properly adjusted. Failure to account for
calcium and magnesium content of irrigation water may result in sup-
pressed plant growth (lhyrien and Whitcomb 1984). When irrigation
water contains a calcium concentration of about 100 ppm, further calcium
additions are unnecessary (Whitcomb 1984a).
Total calcium content of €? taeda bark from standing trees is 0.214%
(Koch 1972) though calcium content as low as 0.017% and as high as
0.85% has been obtained for milled pine bark potting amendment (F. A.
Pokorny, unpublished data). Of greater interest than total calcium con-
tent is the quantity of calcium available for plant growth. Pokorny ( 1979),
working with a mixture of I? taeda and €? echinata bark, reported a
water-extractable calcium concentration of 7.6 ppm, which is lower than
the 160 ppm in Hoagland’s solution (Jones 1983).However, in a study by
Starr and Wright (1984), I. crenata ‘Helleri’ plants grown in solution
culture thrived when 5-10 ppm calcium was supplied; a t rates higher than
10 ppm shoot length and dry weight did not increase. Unfertilized pine
bark apparently supplies sufficient available calcium for growth of ‘Hel-
leri’ holly (Starr and Wright 1984) since indigenous calcium content in
soil solution is reported to be 21-39 ppm (Starr and Wright 1984).
VIII. MAGNESIUM
Dolomitic limestone, as a nutrient source, contains about 20% calcium

and 10%magnesium (Whitcomb 1984a).Calcium and magnesium imbal-
ances within plant tissue, with subsequent poor plant growth, are reported
to occur when finely ground dolomite is used to supply calcium and
magnesium (Whitcomb 1984b). Differences in solubility constants of
calcium carbonate and magnesium carbonate, which comprise dolomite,
result in different release rates for calcium and magnesium and availabilities
of these elements in soil solution (Whitcomb 1981). Whitcomb (1984a)
found that magnesium concentrations in plant tissue are high during the
first several months after planting in a medium amended with finely
ground dolomite, but rapidly decrease to a low level after 9-12 months.
Thus, soon after planting the Ca :Mg ratio, instead of being 2 :1, approaches
1:50, slowly changing to 2:0.05 or less 9-12 months after planting
(Whitcomb 1984a). Initially, plants are subjected to a magnesium excess

followed by a deficiency. To compensate for the high solubility of magne-
sium carbonate in finely ground (180 mesh plus) dolomitic limestone, a
coarser textured dolomite is recommended in which the magnesium
release rate is slowed (Whitcomb 1984a). Growth of ‘Hetzi’ holly and
‘Blue Pacific’ shore juniper plants is comparable when no dolomite is
added or when 4.5 or 9.0 kg/m3 of very coarse dolomite is incorporated in
the medium ( 3:1: 1by volume pine bark, peat, sand) (Whitcomb 1984a).
Fuller and Meadows (1983), in a study of the effects of powdered vs.
pelletized limestone or CaS04-MgS04in a 4 : 1(by volume) pine bark and
sand medium, found that plants receiving CaS04-MgS04fertilization
(no dolomite added), generally had the highest plant quality ratings in
comparison to no lime additions or when powdered or pelletized dolomite
is incorporated. Poorest plant quality is obtained when no dolomite or
CaS04-MgS04is added (Fuller and Meadows 1983). The increase in
growth of Photiniu xfraseri in response to dolomite additions is attributed
to improved magnesium environment and a favorable K :Mg ratio, so
that potassium-magnesium antagonism is not important ( Nash et al.
1983). If, as several investigators proposed (Chrustic and Wright 1983;
Jones 1983; Whitcomb 1984a), dolomite is unnecessary for pH adjust-
ment and is deleted from the fertilization program, calcium and magne-
sium need to be supplied by other fertilizer sources. Jones ( 1983)suggested
that dolomite be replaced by preplant additions of calcium sulfate (16.8
kg/m3) and magnesium sulfate (10 kg/m3). Whitcomb (198l),evaluating
calcium and magnesium fertilizers for container production, suggested
that combinations of calcium sulfate and magnesium carbonate or cal-
cium carbonate and magnesium oxide, in a 2 :l Ca :Mg ratio, are promis-
ing substitutes for dolomitic lime.
Magnesium content of ashed pine bark is reported as 0.14% (Pokorny
1979), which is similar to that of inner bark of newly harvested I? taeda
trees (White et al. 1970). Water-extractable magnesium of 1.6 ppm is
reported for unamended pine bark (Pokorny 1979), which indicates an
insufficiency of available magnesium for plant growth ( Warncke and
Krauskoff 1983).
For growth of I. crenata ‘Helleri’ 5-10 ppm is adequate (Starr and
Wright 1984). Increasing magnesium content of soil solution above 5-10
ppm does not significantly increase plant growth although magnesium
leaf content is increased (Starr and Wright 1984). Analysis of a pour-
through (Yeager et al. 1983) extract of pine bark receiving no fertilizer
shows 7 ppm magnesium in the extract. Starr and Wright (1984)suggested
bark alone supplies sufficient magnesium for growth of ‘Helleri’holly.
Increasing pH increases magnesium retention by pine bark media
regardless of the source of nitrogen fertilizer applied (urea, NH4-N, or
N03-N).However, the magnitude of this retention in relation to nitrogen

source is urea > NH4-N > NO,-N (Ogden 1982).
Leaching of magnesium from bark fertilized with urea is minimal and
not influenced by pH. In contrast, raising pH reduces leaching of magne-
sium from both NH4-N and N03-N fertilized bark, but pH effect is more
pronounced with N03-N (Ogden 1982).Pine bark has a low attraction for
magnesium in the presence of large quantities of NH4 and calcium
cations because of competition for exchange sites (Ogden 1982).
IX. MICRONUTRIENTS
Micronutrients are essential for plants to make normal growth and are
often provided by using soil and organic matter as a medium component
(Matkin et al. 1957), as contaminants in some irrigation waters (Dickey
et al. 1978; Kamp and Pokorny 1958),as impurities in fertilizer (Gilliam
et al. 198l), and from application of certain fungicides (Pirone 1978;
Smith 1981).
Deletion of soil from the container medium, advancements in under-
standing and control of physical and chemical properties of commonly
used potting substrates, and improved fertilizer practices have created
concerns regarding the micronutrient status of container-grown green-
house and nursery plants. With accelerated growth associated with increased
nitrogen, phosphorus, and potassium fertilization rates, the need for
increased use of micronutrient fertilizer to maintain optimum plant
growth is demonstrated (Whitcomb 1984a). Many micronutrient fertiliz-
ers are commercially available and have been evaluated ( Bonaminio and
Bir 1981; Geer et al. 1978; Laiche 1982; Self 1978; Self and Washington
1978a,b; Whitcomb 1979). While it is suggested that micronutrients be
included in the fertility program for greenhouse and nursery crops grown
in soilless media, little definitive information has been published about
the micronutrient status and interactions in pine bark-based media.
Micronutrient content of milled pine bark is given in Bible 4.7. Pine
bark contains most, if not all, of the micronutrients considered essential
for plant growth, but their availability from pine bark is uncertain (Lunt
and Clark 1959).
A. Boron
The initial water-extractable boron content of milled pine bark ranges
from 0.15 to 1.10 ppm (Bible 4.7), which is approximately '/toi twice the
concentration present in Hoagland's nutrient solution (Jones 1983).Ogden
( 1982), studying interactions between nitrogen source and liming effects
120 R. J. OGDEN. F. A. POKORNY, H. A. MILLS. AND M. G. DUNAVENT
Table 4.7. Micronutrient Analysis ( p p m )of Milled Southern Pine Bark Used
as a Potting Medium or Medium Component
Water-extractable content
Total ( PPm)
elemental Hoagland’s
content Pokorny Neal and Ogden solution
Element (PPm) ( 1979) Wagner (1983) (1982) (PPm)
B 9 0.15 1.10 0.17 0.50

cu 77 0.17 0.33 0.12 0.02
Fe 790 0.61 1.10 5.00
Mn 119 0.01 0.90 1.30 0.50
Zn 119, 0.06 0.36 0.30 0.05
‘Not detectable
on solubility of essential elements in a pine bark potting medium, reported

that addition of lime reduces boron concentration in soil solution regard-
less of nitrogen source applied (urea, ammonium, or nitrate). Thus, it
appears likely that if lime is incorporated in the medium, a plant will be
subject to boron insufficiencyunless boron is supplied by a micronutrient
fertilizer or as an impurity in irrigation water (Kamp and Pokorny 1958).
Similar boron insufficiency is reported by Lucas and Davis (1961)when
organic soils are limed.
Suppression of plant growth by essential element imbalances is well
known. Whitcomb et al. (1980),in an extensive study of iron, manganese,
copper, boron, and zinc fertilizer applications on growth of pyracantha
and azalea ( 2 :1:1by volume pine bark :peat :sand), reported interactive
effects on plant growth among four of the five elements tested. Maximum
growth is achieved with these plants only when iron, copper, and boron
levels in the potting medium are high, zinc is intermediate, and manga-
nese is low (Whitcomb et al. 1980).Changes in the balance of any of these
elements results in diminished growth ( Whitcomb 1984a).Plant quality,
especially of azaleas, is substantially improved by the correct balance of
nutrients in the potting mix. During winter, dormant plants retain dark
green foliage under a balanced micronutrient regime; plants subjected to
a micronutrient imbalance, especially of manganese during the previous
growing season, abscise most of their foliage (Whitcomb 1984a).
Excess boron presents a problem in container plant production -it
often leads to growth suppression and diminished plant quality, a condi-
tion more problematic in soilless media than boron deficiency ( Whitcomb
1984a). Gilliam et al. (1981)suggested the prime cause of boron toxicity
is improper selection and application of micronutrient fertilizers. Selec-
tion of commercial micronutrient fertilizers based on the proper balance
Table 4.8. Mineral Element Composition of Several Micronutrient

Sources ( % by weight of packaged product)ash
Elemental content (%/wt of product)

Commercial
product B Fe Mn Zn Cu Mo
Perk 0.02 3.7 2.2 0.70 0.2 0.0020

Esmigran 0.01 2.0 0.5 1.00 0.3 0.0006
FTE 503 3.10 18.0 7.5 0.05 3.0 0.2000
FTE 504 3.80 14.0 7.0 7.00 7.0 0.0700
Stem 1.45 7.5 8.2 4.50 3.2 0.0460
Lesco-FePlus 0.05 5.0 0.5 1.00 0.5 -
Micromax 0.10 12.0 2.5 1.00 0.5 0.0050
"1%= 10,000 ppm.

'From Gilliam et al. (1981).
of elements in relation to other boron sources is essential to avoid phyto-

toxicity. Mineral composition of seven commercial fertilizers is shown in
Thble 4.8 (Gilliam et al. 1981). Boron concentration in four of these
fertilizers is 1000 ppm or higher and, as Gilliam et al. (1981)pointed out,
were they the sole source of boron, toxicity probably would not occur,
because boron supplied by these fertilizers is released with time (Gilliam
et al. 1981).
Boron is supplied by pine bark (Thble 4.7) as well as by other compo-
nents and fertilizers (Thble 4.9) frequently incorporated preplant into the
Table 4.9. Common Sources and Concentration of Boron in

Fertilizers Used in a Preplant Medium for Container
Production",'
Boron conc
~~
g/z-gal rng/Z-gal
Source kg/m3 container container
Urea formaldehyde 2.6 14.98 0.036

Iron sulfate 0.5 2.70 1.733
Gu 49 (micronutrient) 1.2 6.80 13.260
Dolomitic limestone 2.4 13.60 0.544
'lkiple superphosphate 1.1 6.10 2.001
Gypsum 1.9 10.90 1.526
Total 19.145
"Rate of fertilizer incorporated into the preplant medium.

Samples were digested in 6 N HCl for 4 hrs a t 90°C.
'Adapted from Gilliam et al. (1981).
potting medium. An additional source of boron, frequently overlooked, is

irrigation water (Dickey et al. 1978; Gilliam et al. 1981; Kamp and
Pokorny 1958; Whitcomb 1984a). Kamp and Pokorny (1958) found 0.5
ppm boron in central Illinois irrigation water, resulting in a toxic accumu-
lation of boron in greenhouse rose soils. Dickey et al. (1978) reported
small amounts of boron in irrigation water in Florida and thus did not
recommend adding boron as a micronutrient fertilizer. Additionally, the
problem of boron toxicity may be further accentuated because some
postplant water-soluble and slow-release macronutrient formulations con-
tain traces of boron as impurities (Gilliam et al. 1981).
Gilliam et al. (1981)found boron toxicity symptoms in %us xmedia
‘Anderson’plants when boron content of the medium (4 :1by volume pine
bark :sand) is 25 ppm or above, but no toxicity a t levels of 0.5 or 5 ppm.
According to these investigators, critical foliar boron toxicity levels are
between 85 and 100 ppm.
B. Copper
Total copper in ashed pine bark is 77 ppm (Pokorny 1979). Water-
extractable copper content of pine bark ranges from 0.12 to 0.33 ppm
(Neal and Wagner 1983; Ogden 1982; Pokorny 1979),which is higher than
the concentration in Hoagland’s nutrient solution (Tmble 4.7). Copper
deficiency is rarely encountered in container plant production because
small quantities of copper appear as impurities in fertilizers (Mastalerz
19771, in field soil sometimes used as a potting medium component
(Matkin et al. 19571, and in fungicide sprays (Pirone 1978; Poole and
Conover 1979). With deletion of field soil from the potting medium,
synthetic media utilizing peatmoss, pine bark, sand, perlite, and vermic-
ulite may not supply adequate quantities of copper for plant growth.
Copper deficiency symptoms have been identified and described on sev-
eral container-grownwoody ornamental plants in Florida nurseries ( Dickey
1965), and on Aglaonema commutatum ‘Fransher’ growing in potting
medium of Florida sedge peat, pine bark, and cypress shavings ( 2 :1: 1by
volume) (Poole and Conover 1979). Poole and Conover (1979) reported
that inadequate copper is obtained from the Florida sedge peat, pine
bark, and cypress shavings medium by Aglaonema plants, but all other
micronutrients are adequately supplied by the medium components (no
micronutrient fertilizer applied). Copper deficiency is prevented either by
preplant addition of a commercial micronutrient fertilizer (Dickey 1972;
Whitcomb 1984a)or copper sulfate (70mg/m3) (Dickey et al. 1978).A soil
drench of sequestrine NagCuor foliar sprays of copper fungicides will also
alleviate copper deficiency (Poole and Conover 1979).
C. Iron
Total iron content of milled pine bark is substantial (Tbble 4.7) with
levels as high as 1765 ppm (Ogden 1982).However, the water-extractable
fraction is quite low in relation to the 5 ppm found in Hoagland’s nutrient
solution (Tbble 4.7). The problem of unavailable iron in pine bark, with
accompanying iron deficiency in plants, may be compounded by mixing
pine bark with sand or other chemically inert components, not incorpo-
rating sufficient iron-containing fertilizer into the potting mix, or exces-
sive liming.
Substantial quantities of iron are leached from unlimed and unfertilized
I! rudiutu bark (Prasad 1979a). However, lime additions (3.5 g/liter)
significantly diminish leaching of iron. Prasad ( 1979)recovered about 5%
of applied iron in leachate from limed I! rudiutu bark. Similar results are
reported by Ogden (1982).
Excessive calcium, manganese, and phosphorus also suppress iron
uptake and may cause iron deficiency (Whitcomb 1984a). Interactive
influences between iron, boron, copper, and manganese on growth of pyra-
cantha and azalea plants ( 2 : 1: 1 by volume pine bark :peatmoss :sand
potting medium) are reported by Whitcomb (1984a). Maximum growth
of pyracantha and azalea plants is achieved when iron, boron, and copper
are at high levels and manganese is low (Whitcomb 1984a);an imbalance
of any one of these elements reduces growth. Whitcomb (1984a)suggested
that an Fe :Mn ratio of 5 :1 and an Fe :Cu ratio of 10 : 1 are necessary to
maintain maximum plant growth.
D. Manganese
Total manganese of ashed pine bark is reported a t 119 ppm (Thble 4.7)
by Pokorny (1979)and 194ppm by Ogden (1982).However, water-extractable
manganese ranges from 0.01 to 1.30 ppm, which is about l/5 to twice the
concentration in Hoagland’s nutrient solution (Tbble 4.7).
Manganese is readily leached from both unlimed and limed I! rudiutu
bark (Prasad 1979a). In a study of interactive effects of nitrogen source
and liming practices on micronutrient availability in a pine bark medium,
Ogden (1982) reported that liming (pH 5.5-6.5) reduces manganese in
water extracts when NH4-N and N03-N are fertilizer sources. However,
when urea is applied to pine bark, lime has no effect on manganese
concentration in water extracts. Manganese, in unlimed bark fertilized
with either NH4-N or N03-N and receiving micronutrient fertilizer sup-
plying manganese, may reach toxic levels (Hewitt 1966). Ogden ( 1982)
postulated that some indigenous manganese in pine bark is present in an
exchangeable form.
124 R. J. OGDEN. F. A. POKORNY, H. A. MILLS, AND M. G. DUNAVENT
Dickey et al. (1978) stated that manganese deficiency on container-

grown woody ornamental plants in Florida nurseries is not prevalent, but
can be observed where alkaline soil components are used in formulating
potting media. To prevent manganese deficiency under alkaline condi-
tions, preplant application of finely ground manganese sulfate is recom-
mended (140 g/m3) (Dickey et al. 1978).
E. Zinc
Zinc deficiency is seldom encountered in container culture since small
amounts of zinc occur in potting medium components and as an impurity
in many fertilizers (Cotter and McGregor 1979; Mastalerz 1977; Pokorny
1983; Whitcomb 1984a). Zinc is also supplied when plants are sprayed
with fungicides containing zinc (Pirone 1978). However, Dickey et al.
( 1978) observed zinc deficiency on container-grown loquat and dogwood
plants in Florida nurseries and recommend preplant incorporation of
finely ground zinc sulfate (70 g/m3) into the potting substrate.
Total elemental and water-extractable zinc content of milled pine bark
is given in Thble 4.7. Initial water-extractable zinc, ranging from 0.06 to
0.36 ppm, is reported for pine bark and is adequate to high in relation to
0.05 ppm zinc contained in Hoagland’s nutrient solution (Jones 1983;
Neal and Wagner 1983; Ogden 1982; Pokorny 1979).
Liming unfertilized pine bark does not influence water-extractable
zinc. However, when urea is used as a nitrogen source, liming (pH 5.5-6.5)
significantly increases zinc in the leachate. Using NH4-Nfertilizer increases
zinc content of the leachate in unlimed compared to limed bark (Ogden
1982). Zinc concentration in leachates of unlimed and limed bark is
unaffected when N03-N is used as a fertilizer (Ogden 1982).
Cotter and McGregor (1979)reported that zinc content of bark (mix-
ture of ponderosa pine, Douglas-fir, and white fir) is high but generally
unavailable for plant use. Zinc content of tomato plants cultured in both
fresh and aged bark is high but below phytotoxic levels (Cotter and
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zinc appears to be fixed in a bark medium, rendering plants less respon-
sive to applied soluble zinc. Similar responses to soluble zinc application
are reported by Whitcomb (1984a) for pyracantha and azalea plants
grown in a 2 :1:1 by volume pine bark :peat :sand potting substrate.
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elements in complete trees of eight species in Maine. For. Prod. J. 21:56-59.
5
Iron Deficiency Chlorosis
Ronald R Korcak
Fruit Laboratory, Agricultural Research Service,
U. S. Department of Agriculture, Agricultural Research
Center, Beltsville, Maryland 20705
I. Introduction 133
A. Scope and Previous Reviews 133
B . Overview and Definition 135
11. Soil Iron 137
A. Acidic Soils 137
B . Calcareous Soils 140
111. Iron Uptake 144
A. General 144
B . Phytosiderophores 148
C . Other Considerations 150
I v. Iron 'IYanslocation 153
V. Measures of Plant Iron Status 154
VI. Bicarbonate-Induced Chlorosis 158
VII. Iron Chlorosis: Horticultural Occurrences 161
A. Deciduous 'IYees and Ornamentals 161
B . Grapes 165
C . Citrus 166
D. Other Horticultural Plants 168
VIII. Use of Chelates 169
IX. Conclusion 171
I. INTRODUCTION
Science is a procedure for testing and rejecting hypotheses,

not a compendium of certain knowledge.
Stephen Jay Gould, The Flamingo's Smile
A. Scope and Previous Reviews

Iron deficiency of plants has received voluminous attention and to this
date remains a serious worldwide problem for many crops. As early as
1845 Gris (Gile and Carrero 1920) noted a chlorotic condition of grapes
grown on calcareous soil and associated this to the supply of iron to the
plant. Ferrous sulfate cured the chlorotic condition.
133
134 RONALD F. KORCAK
The historical record of research on iron deficiency chlorosis can be

partitioned into three broad phases:
Mid-1800s to early 19OOs, research centered on observations of

chlorotic plants with the establishment as noted above of the
association with iron supply.
Post-World War I1 to 1970s, a dual approach: the formulation and
use of synthetic chelating agents in correcting iron deficiency
chlorosis, and concentration on species and cultivar differences in
the efficiency of iron uptake.
Both of these research areas were seemingly apparent precursors
for the third and current phase of work on the role of the root, the
role of the rhizosphere (pH modification and exudates) in iron
stress response and the physiology and biochemistry of iron
uptake in plants.
This review will touch upon all three of these general phases of research
and concentrate where possible on the data available on horticultural
and, in particular, fruit tree crops. This, however, does not preclude the
discussion of the important work on agronomic crops.
No single review of iron deficiency chlorosis can cover the enormous
amount of literature available. Many excellent reviews of iron in plant
nutrition and of the factors associated with iron deficiency chlorosis have
appeared. These include factors affecting the capacity of plants to utilize
iron as well as functions, supply, and plant iron requirements (Brown
1956); the use of synthetic chelates in correcting iron supply (Brown
1961); the effects of iron chelates on iron-deficient plant cells (Price 1968);
the occurrence of iron chlorosis in horticultural plants (Wallace and Lunt
1960);and the control of chlorosis in English orchards (Little 1971). The
essential and fundamental roles of iron in plant nutrition have been
reviewed and updated by several authors (Hewitt 1963; DeKock 1971;
Zhiznevskaya 1972; Mengel and Kirkby 1982) as have the uptake and
translocation of iron by Sutcliffe (1971).
Iron nutrition in calcareous soils has been reviewed by Chen and Barak
( 1982) and under English conditions by Schinas and Powell ( 1977).
Wadleigh and Brown ( 1952)and Miller ( 1960)present extended literature
reviews on the role of bicarbonate in iron deficiency. Recently, Chaney
(1984) presented a very useful review on diagnostic practices used in
identifying iron deficiency chlorosis. The proceedings of the first and
second International Symposia on Iron Nutrition (Nelson et al. 1982;
James et al. 1984) provide a wealth of information on all phases of plant
iron nutrition and control. Finally, the mobilization of iron in the rhizo-
sphere of different plant species has recently been reviewed by Romheld
and Marschner (1985).
5. IRON DEFICIENCY CHLOROSIS 135
B. Overview and Definition

Iron deficiency can occur at both extremes of the pH range of agricul-
tural soils: in acid, sandy soils of Florida, citrus trees developed iron
deficiency due to repeated applications of copper fungicides or to an
imbalance in soil levels of zinc, iron, manganese, and copper (Reuther and
Smith 1952, 1953; Smith and Specht 1952). In soils above pH 5.2,
blueberry, an acidophilic plant, often develops chlorosis (Brown and
Draper 1980), and in the calcareous soils of the Ebro and Jalon River
valleys in northern Spain iron chlorosis is a problem in peach cultivation
(Montesinos 1984; Abadia et al. 1985).
Iron chlorosis can occur at most soil pH values. Wallace and Lunt
(1960) and Embleton et al. (1973a) have listed factors that may be
involved either singly or in combination in the development of chlorosis
as follows:
low iron supply

calcium carbonate in soil
bicarbonate in soil or irrigation water
overirrigation or high water conditions
high phosphate levels
high levels of heavy metals
low or high temperatures
high light intensities
high levels of nitrate nitrogen
imbalances in cation ratios
poor soil aeration
certain organic matter additions to soil
viruses
root damage by nematodes and other organisms
Most of these as well as other factors are discussed in detail below.

The symptomology of iron deficiency is usually manifested as an
interveinal chlorosis of young leaves while the veins remain green- hence
the name iron deficiency chlorosis (Fig. 5.1). Color plates of iron defi-
ciency symptoms from a range of crops have been published (Wallace
1951; Sprague 1964; Beyers and Brblanche 1971). The expression of the
symptoms in young leaves is due to the inability to redistribute iron
within the plant. Brown and Holmes (1955)showed that in soybean once
the supply of iron to the plant is interrupted, mobility within the plant
ceases. Iron may, however, become more mobile under stress conditions
(Price 1968), as demonstrated in bean plants (Balba et al. 1980), in to-
bacco (Wallace and DeKock 1965),and in azalea (Rutland 1971).Daven-
port (1983)noted a “shifting” of chlorosis in ‘%hiti’ lime: during a new
flush of vegetative growth, the symptoms of chlorosis shifted to the newly
formed leaves with a corresponding regreening of the previous flush.
Mango trees express atypical lime-induced chlorosis symptoms: initially
136 RONAI,D F. KORCAK
Fig. 5.1. Foliar expression of iron chlorosis: ( A ) grapefruit, ( B ) grape, ( C ) straw-

berry, ( D )blueberry, ( E )apple, ( F )maple.
the entire young leaf blade turns yellowish green, it eventually ceases
growth, and a gradual die-back of the branch occurs (Kadman and Gazit
1984).Horesh and Levy ( 1981)noted that grapefruit tended to develop an
unidentified gummosis on main limbs under iron deficiency stress.
The expression of iron chlorosis may be confounded by the occurrence
of simultaneous micronutrient deficiencies, as with zinc and iron (Dixit
and Yamdagni 1983), and manganese, iron, and zinc in citrus (Bar-
Akiva and Lavon 1968) and apple (Shen and Tseng 1949). McGeorge
(1949) described the symptoms and how to differentiate between iron,
zinc, and manganese deficiencies in citrus. In work on multiple deficiencies
in apple, Zhou et al. (1986) found that when manganese, zinc, or a
combination of the two were deficient and when iron was also deficient,
the expression of iron deficiency predominated. To distinguish chlorosis
due to iron deficiency alone and to provide a workable diagnosis of the
disorder, the definition of iron deficiency chlorosis presented by Chaney
(1984) is most appropriate: any yellowing of leaves that regreens when
treated with FeS04or FeEDDHA (ethylenediaminedi-o-hydroxyphenylacetic
acid), but does not regreen when nitrogen, sulfur, zinc, manganese,
copper, cobalt, or other nutrients are applied alone or in combination.
Scientific names and authorities are listed in n b l e 5.1 for common
plant names used throughout the text.
11. SOIL IRON
Paditionally the forms and plant availability of iron in soils have been
shown to be greatly influenced by soil pH, oxidationheduction status,
and soluble components of organic matter (Oades 1963; Ellis and Knezek
1972; Lindsay 1972; Stevenson and Ardakani 1972). Although iron con-
stitutes about 5% by weight of the earth’s crust, the activity of soluble
iron in soils is very low (Mengel and Kirkby 1982). Chemically the
element iron occurs in two oxidation states: the oxidized form Fe3’ (ferric)
and the reduced form Fez+(ferrous). Ferrous iron is readily oxidized to
ferric, the latter being extremely insoluble in water (O’Connor et al.
1971). These two properties-rapid oxidation (which occurs in “normal”
aerated soils) and insolubility of the oxidized species-are the corner-
stones of the iron deficiency chlorosis problem.
A. Acidic Soils
The solubility of iron in soils is governed by the dissolution and
precipitation of ferric oxides (Lindsay and Schwab 1982):
Fe(OH):<(soil) + 3H’ % Fe3’ + 3H20 5.1)

138 RONALD E KORCAK
Table 5.1. Scientific Names and Authorities of the Common Plant Names
Used in Text
Common name Scientific name and authority
Tkee fruits
Apple Malus domestica Borkh
Plum, European Prunus domestica L.
Plum, Japanese Prunus insititia L.
Peach Prunus persica ( L . )Batsch
Pear Pyrus communis L
Sour cherry Prunus cerasus L.
Sweet cherry Prunus avium ( L . ) L.
Quince Cydonia oblonga Mill.
Apricot Prunus armeniaca L.
Fig Ficus carica L.
Almond Prunus dulcis (Mill.)D. A. Webb
Olive Olea europaea L.
Myrobalan rootstock Prunus cerasifera Ehrh.
Mahaleb rootstock Prunus mahaleb L.
Mazzard rootstock Prunus auium ( L . ) L.
Avocado Persea americana Mill.
Papaya Carica papaya L.
Mango Mangifera indica L.
Citrus
Sweet orange Citrus sinensis ( L . ) Osb.
Sour orange Citrus aurantium L.
Tkifoliate orange Poncirus trifoliata ( L . ) Raf.
Rough lemon Citrus jambhiri
Lemon Citrus limon ( L . ) Burm. f.
Grapefruit Citrus paradisi Macfad.
'Tahiti' lime Citrus latifolia Tan.
Mandarin orange Citrus reticulata Blanco
Tangelo Citrus paradisi X Citrus reticulata
Thngors Citrus reticulata X Citrus sinensis
YuZu rootstock Citrus junos Sieb. ex Tan.
Ornamentals and trees

Azalea Rhododendron spp.
Rose Rosa spp.
Acacia Acacia spp.
Chestnut Castanea spp.
Aspen Populus spp.
White ash Fraxinus americana L.
Austrian pine Pinus nigra Arnold
Lodgepole pine Pinus contorta Dougl. ex Loud
Silver maple Acer saccharinum L.
Pin oak Quercus palustris Muench.
Chrysanthemum Chrysanthemum morifolium Ramat.
Table 5.1. (Continued)
Common name Scientific name and authority
Bush fruits
Strawberry Fragaria spp.
Blueberry Vaccinium spp.
Highbush blueberry Vaccinium corymbosum L.
Raspberry Rubus spp.
Grape Vitis spp.
European grape Vitis vinifera L.
American grape Vitis labrusca L.
Vegetables and garden fruits

Cucumber Cucumis satiuus L.
Watermelon Citrullus lanatus (Thunb.)Matsumot Nakai
Pumpkin Cucurbita spp.
Tomato Lycopersicon esculentum Mill.
Pea Pisum satiuum L.
Peanut Arachis hypogaea L.
Agronomic crops
Soybean Glycine m a x ( L . )Merr.
Bean Phaseolus uulgaris L.
Tobacco Nicotiana tabacum L.
Rice Oryza sativa L.
Sorghum Sorghum bicolor ( L . ) Moench
Sunflower Helianthus annus L.
Maize Zea mays L.
Oat Avena sativa L.
Cotton Gossypium hirsutum L.
Barley Hordeum vulgare L.
Lentil Lens culinaris Medik.
Sesame S e s a m u m indicum L.
Other
Jute Abutilon theophrasti Medik.
Oil radish Raphanus satiuus L.
Since the precipitation of ferric oxides is pH dependent (the activity of

Fe3' decreases 1000-fold for each unit increase of pH), well-aerated acid
soils display higher soluble iron than calcareous soils. At pH levels above
7.5, the solubility of Fe3+falls below lO-'OM (Lindsay and Schwab 1982),
whereas acid soils contain about 10-6M (Chaney 1984). Lindsay and
Schwab (1982)estimated that iron activity must be in the general range
10p7.7M to avoid iron deficiency.
Ferrous iron activity in soils increasesunder reducing conditions (Lindsay

and Schwab 1982):
Fe3' + e % Fe2+ (5.2)
The ferrous form of iron under reducing condition is stable (Uren 1984)
and provides an available source of iron under anaerobic conditions and in
flooded rice fields (Benckiser et al. 1984).
The existence of soluble and stable iron organic complexes in soil
solution and their importance in iron availability to plants has been
hypothesized, but research is needed to identify these forms (Chen and
Barak 1982; Uren 1984; Linehan 1985).
B. Calcareous Soils
An alkaline soil is any soil having a pH (in HzO)greater than 7.0, while
a calcareous soil is defined as any soil containing sufficient calcium
carbonate or calcium-magnesium carbonate to effervesce visibly when
treated with cold 0.1N HC1. Brown (1961)estimated that 25-30% of the
world's land is calcareous a t the surface. The bicarbonate ion, which
forms in calcareous soils (see below), is the most important soil factor
associated with lime-induced iron chlorosis of grapes (Mange1 et al.
1984a);apples, pears, and roses (Boxma 1972);and soybeans (Coulombe
e t a l . 1984).
Bicarbonate activity in high-pH soils will be governed, on a soil by soil
basis, by the following two reactions (Boxma 1972):
OH- + COz % HCOi (5.31
HCOi + OH- % COZ- +2Hz0 (5.4)
According to Eq. (5.3),as soil pH increases,bicarbonate activity increases,

but Eq. (5.4) shows that bicarbonate activity will decrease. These two
reactions will result in a pH vs. HCOi activity curve exhibiting a maxi-
mum HCOS concentration a t a definite pH value. Thus either an increase
or a decrease in pH will result in a lowering of the bicarbonate concentra-
tion. The maxima for any soil would vary, depending upon the solubility
of the carbonate form and the ionic species in Eqs. (5.3) and (5.4).
The importance of MgC03 vs. CaC03 in the potential for chlorosis
development has been studied by Wallace et al. (1976~). The MgC03
content of most calcareous soils is very low; however, the use of dolomite
(Ca-MgC03)might have a greater influence on iron availability than
CaC03 due to its higher equilibrium pH (about 9). Loeppert and Hall-
mark (1985) showed that the soil carbonate composition Mg/Mg + Ca

ratio was better correlated with iron availability to sorghum than total
CaC03. Additionally, the solution phase and exchangeable Mg2+levels
were also correlated with the incidence of iron chlorosis from the 24
calcareous soils tested.
The dynamics of the potential soil bicarbonate concentration change
under conditions of wet calcareous soils. The governing factors under
such conditions are high COZ pressure in the soil, hydrolysis of CaC03,
and soil pH (Boxma 1972).The effects of high COz pressure and hydroly-
sis of CaC03, which are both favored by high soil moisture, can be
expressed as
The importance of Eq. (5.5)lies in the fact that the percentage of CaC03
present in a given soil per se may not be indicative of potential chlorosis
problems, but rather the hydrolysis of CaC03 to form HCOi becomes
predominant, and this is strongly influenced by excessive moisture in the
soil (Thble 5.2). High soil moisture plus CaC03 resulted in a doubling of
the soil bicarbonate content, a 33% reduction in rose leaf iron, and the
development of chlorosis, compared to the same soilhreatment a t low soil
moisture (Boxma 1972).Addition of either 200 or 2000 ppm HC03 from
Mg(HC03)zunder high soil moisture produced relatively small addi-
tional reductions in leaf iron content. Soil pH was actually decreased by
the addition of 2000 ppm HCOi due to enhanced CaC03 formation [see
Eq. (5.5)].
As will be noted later, soil properties and management practices that
control soil moisture can have significant effects on the incidence and
Table 5.2. Effect of Soil 'Reatments on Soil Bicarbonate, Leaf Iron Content, and
Chlorosis of Roses Grown in 'Reated Soil/Peat Potting Mixture"
Soil Leaf
Soil HC03- Fe Plant
water Peatment ( PPm ) ( PPm ) status
Low + CaC03 245 122 No chlorosis
High + CaC03 540 81 Chlorosis
High + CaC03 + 200 ppm HC03- 500 78 Chlorosis
High + CaC03 + 2000 ppm HC03 659 67 Chlorosis
"After Boxma (1972).

142 RONALD E KORCAK
degree of iron deficiency chlorosis. Additionally, the effect of carbon

dioxide and bicarbonate reactions (these buffer p H ) in solution cultures
on chlorosis of bean have been reported by Porter and Thorne ( 1955);pH
was not as important as HCOi activity.
Total soil CaC03 is usually not a good index for the development of
chlorosis (Armson and Williams 1960). Loeppert et al. (1984) investi-
gated “hot spots,” localized variable chlorotic areas within a sorghum
field, which varied from 0.55 to 53.5% CaCOu.They found soil carbonate
and amorphous iron oxide content to be the dominant factors in the
severity of chlorosis. Carter (1981) found that “active” CaC03 was a
better index of potential chlorosis problems for a range of tree species
than total CaC03. The “active” CaC03is an estimate of CaCOZ3 in clay and
fine silt particle sizes.
An inherent problem in testing calcareous soils exists because air-
drying results in the loss of most of the bicarbonate (Yaalon 1957; Clark
et al. 1960). This could have severe implications in soil pot studies if
pre-experimental drying of the soils is performed. Chaney ( 1984)detailed
procedures for handling field-collected soils, either calcareous or non-
calcareous, for use in controlled experiments on iron availability:
maintain soil a t field-moist state, sieved and stored at about 50%

field capacity a t 4OC until needed;
apply sufficient fertilizer to achieve plant biomass expected while
avoiding the NH: form of nitrogen (to prevent excessive rhizo-
sphere acidification) ;
mimic both field water content and bulk density as closely as
possible to the conditions under which iron chlorosis occurs in the
field;
use a mulch (marble chips) on the soil surface to reduce soil
temperature and water distribution errors; and
employ a low plant density to reduce errors from excessive water
removal in pot studies.
Using these criteria Inskeep and Bloom (1984)were able to reproduce

field-expressed chlorosis in pot studies using soybeans representing a
range in iron efficiency. %-ends toward lower plant chlorophyll content
were most consistently correlated with the following: (1) higher soil plant
magnesium; ( 2 )higher plant sodium and chlorine; ( 3 )higher soil Mg :Ca
ratios ( 4 ) higher plant phosphorus; ( 5 ) higher soil moisture; and ( 6 )
higher soil HCOi. The association with higher HCOR was noted in only
one of three soil transects taken from a known chlorosis-inducing field
that contained a complex of three soil types. One of these soils, Harps
silty clay loam, was used in a comparison of chlorosis development in

seedlings of various Malus spp. compared to an iron-efficientand -inefficient
soybean (unpublished data 1. All of the tested apple species proved to be
more sensitive to chlorosis than even the iron-inefficient ‘Corsoy’ soy-
bean. However, there were varying degrees of visual chlorosis among the
apple species Malus micromalus Bailey, M. dornestica Borkh. cvs. York,
Golden Delicious (the latter from tissue culture): all expressed a lower
level of chlorosis compared with M. zumi Mats., M. hupehensis Pampon,
M. honanesis Mill., M. sieboldii Regel, and M. baccata L. (Fig. 5.2).
The effect of high soil salinity, particularly sodium, on the incidence
and severity of iron chlorosis has not received much attention. The most
severe chlorosis in South Africa has been noted on calcareous soils having
either limestone outcrops or high salt concentrations primarily in low-
lying areas within a field (Beyers and Brblanche 1971). The role of
sodium on bicarbonate uptake by plant roots is unknown; however, it has
been demonstrated that bicarbonate uptake by the cyanobacterium
Anabaena variabilis is enhanced in the presence of sodium (Kaplan et al.
1984). Carbon dioxide uptake was not affected. An understanding of the
I
a
a
I
b
a
14 15 14 15 14 15 14 15 14 15 14 15 14 15 14 15 SOIL
BACCA GOLDN HONAN HUPEH MICRO SIEBO YORK ZUMI SPECIES
Fig. 5.2. The degree of chlorosis development of a range of Malus spp. seedlings
grown on two calcareous soils ( 14 and 15) known to induce iron chlorosis in soybeans.
Rating scale: 0, healthy leaves, no chlorosis; 5, severe chlorosis and necrosis. Species
tested: BACCA, Malus baccata; GOLDN, M . dornestica cv. Golden Delicious (from
tissue culture); HONAN, M . honanesis, M. hupehensis; MICRO, M. micrornalus;
SIEBO, M . sieboldii; YORK, M . dornestica cv. York; and ZUMI, M. zurni.
144 RONALD E KORCAK
effect of variable soil sodium levels on the incidence of iron chlorosis may
aid in elucidating the variation in the severity of chlorosis noted between
calcareous soils that appear to have similar pH and “active” CaC03 levels.
111. IRON UPTAKE
A. General
The primary, if not the sole, form in which iron is absorbed by plants
(except gramineous monocots) is Fez+(Chaney et al. 1972). Since the
primary form of iron in most agricultural soils is Fe3+(except under
flooded conditions with rice) plants face a double role in first solubilizing
iron from the Fe3+form and then reducing the solubilized Fe3’ to Fez’to be
absorbed or transported into the root.
The fact that plant species and cultivars within species differ in their
ability to absorb iron has been established. Weiss (1943)demonstrated
that iron uptake differences among soybeans were heritable. Plants can
be differentiated on whether they respond biochemically (iron-efficient)
or not (iron-inefficient)to iron deficiency stress (Brown 1961, 1978).
Differences between mono- and dicotyledonous species in ability to
obtain iron is well known (Zhiznevshaya 1972). Kashirad and Marschner
( 1974)grew sunflower and maize in mono- and mixed cultures. Under iron
stress, sunflower lowered media pH and regreened in either system, while
iron-stressed maize only regreened in mixed or solid phase (sand culture)
systems. Thus an iron-efficient species, sunflower, improved the iron
nutrition of an inefficient species, maize.
Recently, Olsen and Brown ( 1980a,b)demonstrated that iron-inefficient
genotypes of tomato, maize, soybean, and oats all developed chlorosis
when grown in calcareous W p p soil, but iron-efficient genotypes of these
same species remained green. Differences between species (particularly,
dicot vs. monocot) became apparent when both types of genotypes were
grown first in complete nutrient solution and then transferred to a no-iron
solution. The iron-efficient tomato genotype was the only species of these
four to lower solution pH (via H’ efflux from the roots) significantly and
release reductants. The iron-efficient soybeans released reductants but
did not lower solution pH. The monocots maize or oats, either iron-
efficient or inefficient genotypes, neither reduced solution pH nor re-
leased reductants- they all became chlorotic. Further work by Olsen and
Brown ( 1980b)showed that chemical inhibitors of the root-released reduc-
tants were hydroxide, orthophosphate, pyrophosphate, Cu2+,and Ni2+.
Other work has demonstrated that lowering the pH and/or enhanced
reductant release or reduction capacity of iron-deficiency-stressed plants
occurs in pea, jute, peanut and cotton (Kannan 1982; Kafkafi and
Neumann 1985);lentil and sesame (Kannan 1983); sorghum (McKenzie
et al. 1985);and sunflower (VenkatRaju and Marschner 1973).McKenzie

et al. (1985)attempted to utilize measured root reductant levels during
iron stress as an iron-efficiency assay for screening plants but concluded
that a visual rating of the plants was a better index of iron efficiency.
Although research has demonstrated that reduction does take place
and that for certain species the release of reductants is enhanced under
iron deficiency stress, the exact area in, on , or near the root where this
reduction takes place is unknown (Romheld and Marschner 1985).
Chaney et al. (1972)showed it was exocellular. Bienfait et al. (1982,1983)
proposed that the ferric reducing activity in iron-deficient bean plants is
manifested by an enzyme in the plasmalemma of the cortex or epidermis
cells. Romheld and Marschner (1983),working with peanuts, indicated
enzymatic reduction in the plasmalemma of cortical cells of the roots.
Recently, Kannan et al. (1984)isolated dibutyl phthalate from nutrient
media in which iron-efficient sorghum was subjected to iron-deficiency
stress. Although the authors proposed that dibutyl phthalate acts as a
chelator in solubilizing Fe3+,dibutyl phthalate is not a natural product
and is a known contaminant from plastic (R. L. Chaney, personal
communication ).
That reduction in soybeans occurs via an iron reductase in the cell wall
space, with secreted L-malate as the source of electrons was proposed by
Tipton and Thowsen (1985). However, activity of the enzyme was not
related to iron deficiency, while reduction by the roots strongly was.
Venkat Raju et al. (1972) noted that under iron-deficiency stress there
was an accumulation of organic acids in the roots of sunflowers with a
simultaneous decrease in anion uptake in nutrient solution, which
resulted in a lowered media pH.
Plant iron uptake is an energy-requiring process (Moore 1972) and it
has been demonstrated that both oxygen and photosynthetic substrates
are required for iron root reduction in apple (Tong et al. 1985)(Fig. 5.3).
Early split-root solution culture work with maize and rice by Gile and
Camro (1917)showed that the smaller the number of roots exposed to
nutrients, including iron, the smaller the total amount absorbed. Since
that time it has been shown that specific sections of the root may be more
active in iron uptake. The root section between 1and 4 cm from the tip in
barley was shown to absorb and translocate more iron than the rest of the
root (Clarkson and Sanderson 1978). Plant roots under iron stress ab-
sorbed seven to ten times more iron than unstressed roots. Romheld and
Marschner ( 1979),however, demonstrated that most of the proton release
and reductant exudation from sunflower roots occurred in the 0-1.5 cm
region of the root tip. Landsberg (1982a) also working with sunflower
found that the area of enhanced root proton efflux was confined to a
subapical root zone on iron-stressed plants. This area was characterized
as having an outgrowth of numerous epidermal cells into long root hairs
1 a0
80
&
Y 60
4
48
20
-
0
AZIDE CONTROL DIURON DNP ROTENONE
Fig. 5.3. The effect of respiratory and photosynthetic inhibitors on the reducing
capacity of apple seedling roots. Levels applied: azide (0.25 mM); diuron (0.1 mM);
D N P (dinitrophenol)(0.04 mM), and Rotenone (0.025 mM) (After Tong et al. 1985.)
and/or differentiation into transfer cells. Grape rootstock roots under

iron stress conditions displayed enhanced proton release from the root
apical area behind the root cap similar to the distances cited by Clarkson
and Sanderson (1978) (Fig. 5.4). The Mizzardet and SBB rootstocks
reacted differently to the imposed iron stress. Mizzardet produced many
short lateral roots with lowered rhizosphere pH only around the apical
areas of the main roots, while SBB produced lowered pH around both the
main roots and the younger laterals with less lateral proliferation. These
findings indicate a potential difference in the site of iron reduction/
uptake in mono- vs. dicotyledonous species.
Romheld and Marschner (1985, 1986)have proposed that iron uptake
occurs by two species-dependent strategies (Fig. 5.5). Iron mobilization
by dicots and nongramineous monocots in response to iron deficiency
stress is termed strategy I. The roots of these plant types release protons
from an ATPase-driven proton efflux pump (Romheld et al. 1984), often
release phenolics (Romheld and Marshner 1983), and display increased
activity of a NADPH-dependent plasma membrane-bound reductase
(Sijmons et al. 1984a,b).Landsberg ( 1981)found that the ATPase-driven
proton pump is regulated by auxin. The importance of increased proton
release mechanisms is that they not only increase the solubility of Fe3*,
but that there is an increase in the separation and absorption of iron from
Fig. 5.4. Localization of enhanced proton release ( p H lowering) of the rhizosphere

from iron-stressed grape roots grown in low-iron plus calcium carbonate nutrient
solutions. Roots “pressed” into semisolid pH indicator-spiked agar: matrix (black)
p H was 6.3, white halos about p H 4.5. ( A ) SBB rootstock displaying white halos
around main root apical areas and younger laterals; ( B )Mizzardet rootstock displaying
increased lateral formation with white halos only a t main root terminals.
148 RONALD E KORCAK
Strategy I I1
Strategy II
Fig. 5.5. Strategy I and I1 models for solubilization and uptake of iron by higher
plants. Strategy I occurs in most nongraminaceous species (C, chelate; R, an inducible
reductase 1. Strategy I1 occurs in grasses (S,inducible synthesis of phytosiderophores,
from the precursor nicotianamine NA, extrusion of phytosiderophore X?, and a
specific transport system for ferrated phytosiderophores T). (Redrawn from Romheld
and Marschner 1986.)
Fe3'-chelates a t the plasma membrane (Fig. 5.5). Chaney et al. (1972)

demonstrated in soybeans that Fe3'-chelates are reduced to Fez+-chelate;
these readily exchange Fe2', which is absorbed a t the root surface. Dicots
that lack the ability, according to strategy I, to lower the pH (proton
release) and/or enhance reductant levels are classified as iron-inefficient
genotypes. Strategy 11, as proposed by Romheld and Marschner (1985,
1986),entails the release of phytosiderophores by roots under iron stress
conditions. Once released, these chelators bind Fe3+in the rhizosphere
and carry iron into root cells. This response has been found only with
gramineous monocots (Fig. 5.5).
B. Phytosiderophores
&cent work has provided evidence for the existence of a specific iron
uptake mechanism in grasses (Romheld and Marschner 1986). The spe-
cific machanism centers on the occurrence of iron phytosiderophores. It
has been known for some time that many microorganisms assimilate iron
through the release, under iron stress conditions, of siderophores (Emery
1982). Siderophores are powerful, selective, ferric-iron chelators (Nei-
lands 1973).
Evidence on the importance of siderophores is accumulating. Blue-
green algae can suppress other algae by release of a specific iron sidero-
phore (Murphy et al. 1976),iron stressed fungi have been shown to release
iron-chelating siderophores (Emery 1982), and siderophores released
from Pseudomonas ssp. can inhibit Fusarium oxysporum in the rhizo-
sphere of cucumber plants (Elad and Baker 1985). The utilization of
microorganism-produced siderophores, such as ferrated rhodotorulic
acid, by iron-efficient and -inefficient tomatoes was studied by Miller et
al. (1985).They demonstrated that at least some of the iron taken up by
either tomato type was from these microorganism-produced chelators,
under varying degrees of iron stress in solution culture. The exact
amount of the total plant iron and the mechanism of iron uptake involved
still requires additional studies. Reid and Crowley ( 1984)noted that, with
oats, fungal-produced chelators ( siderophores ) may be a more efficient
source of iron than synthetic chelators. However, it has been reported
that the siderophores of soil fungi occur in a range of soils and can be
effective iron chelators over a wide pH range (Powellet al. 1982).They are
present in sufficient concentrations to be a significant source of soluble
iron for plant nutrition.
In support of the potential importance of siderophores in plant iron
nutrition, Clement et al. (1977)studied the CaC03tolerance of Austrian
pine. Seedlings grown without mycorrhizae developed severe chlorosis as
well as disturbed nitrogen metabolism and excessive cation absorption.
Inoculation with mycorrhizae eliminated the chlorosis and resulted in
normal growth. The authors concluded that tolerance to calcareous soils

was not a genetic characteristic, but was due to the associated
mycorrhizae. Although not examined by the authors, the possibility
exists that the mycorrhizae may be releasing siderophores, thus enhanc-
ing the solubility and plant uptake of iron. However, mycorrhizae did not
correct chlorosis in maize grown in flow cultures (Elmesand Mosse 19841.
Phytosiderophores are plant-root-released iron chelators. Grasses re-
spond to iron deficiency stress by releasing enhanced amounts of che-
lating substances (lhkagi et al. 1984), which are nonprotein amino
acids, such as mugineic and avenic acid (Prochazka and Scholz 1984;
Sugiura and Nomoto 1984). Once released from the root, these chelators
bind Fe3' in the rhizosphere and carry it into root cells via a transport
system specific for the Fe3+-mugineicacid complex. Such a system for
solubilizing and transporting Fe3+has not been found in dicots (lhkagi et
al. 1984)and has been labeled strategy I1 (Romheld and Marschner 1986)
(Fig. 5.5). Strategy 11, the release of phytosiderophores by roots under
iron stress, which enhances the solubility of Fe3+,is less inhibited by high
soil pH and is not affected by bicarbonate.
C. Other Considerations
It is apparent that nutrient supply, availability, and uptake differences
that affect the rhizosphere pH can affect the mobilization and uptake of
iron. Aktas and Van Egmond (1979) showed that an iron-inefficient
soybean cultivar displayed more pronounced chlorosis when N03-N was
used as the nitrogen source. A similar effect was noted in peanuts
fertilized with ammonium sulfate (Kafkafi and Neumann 1985). The
explanation offered was the elevation of the rhizosphere pH due to an
increase in either OH- or HCOi efflux from the root (Riley and Barber
1971). This effect of nitrogen source on rhizosphere pH is also seen in
blueberry (Fig. 5.6). Plants subjected to an all NH4-N nutrient solution
produced a lowered pH around the rhizosphere (Fig. 5.6A) compared to
all N03-N fed plants (Fig. 5.6B). Different rates of cation and anion
absorption by the root will influence the direction and amount of pH
change. Different sources of nitrogen alone can produce a pH shift of up
to 2 units (Riley and Barber 1969). Wallace et al. (1976b) summarized
three ways in which differential cation-anion uptake can influence plant
iron status independently of specific ion effects:
(1) Excess cation vs. anion uptake acidifies external medium and
increases iron mobilization (the reverse can also occur).
( 2 ) Excess cations over mineral anions within the plant increase pH
due to higher level of salts of organic acids, resulting in inactiva-
tion of iron (the reverse can also occur).
Fig. 5.6. The effect of nitrogen source on rhizosphere p H of a blueberry hybrid

grown in nutrient solution. Intact roots were placed in a semisolid p H indicator-
spiked agar. The p H of black matrix was 6.8 and t h a t of white areas about 4.5. ( A )
Root system from plant grown in all NH4-N solution displayed lowered p H (white
areas) around entire root. ( B )Plant grown in all N03-N solution indicated no proton
release (lack of white areas).
( 3 ) Unreduced free nitrate in the plant could be accompanied by an

equivalent amount of protons, making iron more available.
Mengel and Steffens (1982)showed that the protons secreted by the roots
into the soil to balance excess cation uptake electrostatically resulted in
alkalinization within the plant cells. This in turn led to the synthesis of
organic anions equal to the amount of protons released. Barak and Chen
( 1984) found that potassium fertilization ameliorated iron chlorosis in
peanuts grown on calcareous soil (63% CaC03). Plants contained two to
three times higher levels of chlorophyll after treatment. The results
obtained were attributed to attainment of a cation-anion balance of iron
uptake and consequent increased rhizosphere acidity.
Similarly, acidophilic plants like blueberry have been shown to develop
chlorosis when supplied with nitrate nitrogen, due to alkalinization of the
rhizosphere or increased tissue pH (Cain 1954; Cain and Holley 1955).
This effect was avoided with azaleas grown with nitrate nitrogen as long
as the pH or base supply was maintained a t a low level (Colgrove and
Roberts 1956). Arnold and Thompson (1982) suggest, however, that
chlorosis of highbush blueberries grown a t their upper pH limit (5.2)is a
consequence of iron inactivation from excessive phosphorus. Iron-
efficient blueberries were capable of lowering solution pH by root proton
release, but iron-inefficient plants did not change the solution pH (Brown
and Draper 1980). Neither blueberry type was found to release reduc-
tants. Iron-efficient blueberry crosses all contained less calcium than the
iron-inefficient crosses.
Chaney and Coulombe (1982) reviewed the effect of phosphate on
regulation of iron stress response. Previous work had indicated that
elevated phosphate levels caused chlorosis. However, by using solution
culture techniques that better mimicked soil phosphate levels, they were
able to show that high solution phosphate levels were inhibiting the
normal iron stress response in genetically iron-inefficient soybeans.
It has been noted that different mechanisms are involved in iron, zinc
and iron, and copper absorption in cucumber, watermelon, and pumpkin
(Swaider 1985). Copper was found to inhibit iron absorption non
competitively. Both copper and manganese have been shown to inhibit
iron uptake competitively in rice (Shim and Vose 1965). Extensive re-
views of the interference of other metal cations on iron uptake are availa-
ble (Zhiznevskaya 1972; Mengel and Kirkby 1982).
The effects of iron stress on root morphology have received limited
attention ( Landsberg 198213). Generally, affected root systems cease
elongation and initiate lateral roots or root hairs (Brown and Ambler
1972; Landsberg 1982b). Coralloid root systems, described as having
numerous short, stubby lateral roots of uniform size, were associated
with lime-induced chlorosis, iron deficiency chlorosis, and chlorosis in-

duced by bicarbonate in a range of plant species (Hutchinson 1967).
Formation of rhizodermal transfer cells occurred as a part of the response
mechanism to iron deficiency stress in tobacco, tomato, and cucumber,
but not all dicots (e.g., soybean) (Landsberg 198213). The formation of
transfer cells may be genetically regulated. Evidence has been presented
that the role of these cells may be the production of reductants-ferric
reduction and proton extrusion-which are both, as noted previously,
mechanisms of iron deficiency stress response in dicots (Landsberg
1984). Furthermore, it has been hypothesized that iron-deficient young
leaves may send a hormonal signal to the roots, which initiates the
formation of transfer cells or at least affects the proton extrusion rate.
This function, as noted, promotes the mobilization and uptake of soil iron.
IV. IRON TRANSLOCATION
Tkanslocation of iron from roots to shoots has been shown to occur as a

ferric-citrate chelate (Tiffin 1972).This is transported to actively growing
shoot regions (Bennett et al. 1982). Under iron stress conditions in pear
(Oserkowsky 1932)and in silver maple (Morris and Swanson 1980)there
was no difference in the pH of the xylary sap compared to that of non-
stressed plants. The latter authors also showed that iron stress did not
change the redox potential of the sap compared to that of nonstressed
trees. Since movement of iron as a citrate complex has been found, the
stability of this complex during transport would be determined, in part,
by both pH and redox potential. Thus under iron stress conditions,
movement as a citrate-chelate should not be affected and Brown et al.
(1967) noted that translocation via citrate in stem exudates of four
soybean genotypes was most nearly related to the available iron supply.
Localization of 59Fealong the veins of chlorotic azalea leaves compared
to a more uniform 59Fe distribution in control plants was noted by
Rutland ( 1971). Tkanslocation of iron from root to leaves was inhibited in
chrysanthemum (Rutland and Bukovac 1971) under conditions of high ( 3
mM)HCOR concentrations. An inhibitory effect of bicarbonate on absorp-
tion and or translocation of iron has been noted by several workers (Brown
et al. 1959; Wallace and Mueller 1966; Coulombe et al. 1984; Kolesch
et al. 1984). Doney et al. (1960)showed that increased HCOR in culture
solution increased the total amount of 59Fetranslocated out of sprayed
maize or bean leaves to other above-ground parts particularly into the
stems and petioles. In leaves, Mengel and Bubl ( 1983)found that HCOR
had a large effect upon the translocation of iron from vascular tissues to
the intercostal cells. These cells in chlorotic grape leaves had extremely
154 RONALD E KORCAK
high calcium and phosphorus contents. The high level of leaf phosphorus
was noted to be an effect and not the cause of the chlorosis. A direct role of
bicarbonate on iron chlorosis development in susceptible soybean culti-
vars was noted by Coulombe et al. (1984)and Fleming et al. (1984).Both
iron stress response and 59Fetranslocation were reduced, with the former
the more important inhibition.
Iron-deficient plants exhibit an increased capacity for iron absorption
(Rutland and Bukovac 1971). This mechanism has been shown to be
specific for iron and cobalt, but not for zinc, manganese, or copper (Young
and Qrry 1984). However, Romheld and Marschner (1985) showed in-
creased zinc and manganese uptake by stressed plants. Thus the particu-
lar response found will vary depending on the test system used. Both iron
and cobalt, once in the leaf, move rapidly across the plasmalemma into
the symplast via nicotianamine, a divalent cation chelator (Young and
W r y 1984). Cobalt is one of the several metals known to induce iron
chlorosis in plants (Hunter and Vergano 1953) and recently has been
shown a t least partially to inhibit the iron stress response in tomato and
soybean (Blaylock et al. 1985).
The yellowing of plant leaves under iron deficiency stress is the obvious
visual symptom. The effects of iron deficiency in the leaves are principal-
ly a decrease in chlorophyll content per chloroplast and abnormal mor-
phological characteristics of the chloroplasts (Vesk et al. 1966; Spiller
and E r r y 1980; Hecht-Buchholtz 1983; Pushnik et al. 1984; Rufner and
Barker 1984; Zhou et al. (1984a). The decrease in chlorophyll content
results in lower photosynthetic productivity by the plant, with resultant
economic losses in yield and production. Zhou et al. (1984a) found
abnormal chloroplasts with dispersed grana and osmosphilic bodies with
reduced size and number of starch granules in iron-deficient apple seed-
lings compared to seedlings containing adequate iron (Fig. 5.7). The
degree of organization of the chloroplasts (Fig. 5.7D) and the leaf net
photosynthetic rate decreased as iron was made less available.
V. MEASURES OF PLANT IRON STATUS
Early investigations of chlorotic apple leaves (Wallace and Mann 1926)

showed that they contained less dry matter, more ash content, ,less
calcium, and more potassium than green leaves. These parameters were
reversed when iron was supplied. Similar trends were noted in apple bark
(Wallace 1928),as well as in plum and pear tree bark. However, iron was
frequently lower in chlorotic leaves, but higher in bark and wood tissues
of chlorotic trees.
Fig. 5.7. Leaf mesophyll cell from apple seedlings grown in nutrient solution
containing either ( A ) adequate iron ( X 17,000) a t p H 5.5, showing well-developed
chloroplasts in cytoplasm along cell wall and large nucleus ( N ) or ( B )no iron ( X 4500 ),
showing large vacuole ( V ) and thin layer of cytoplasm-containing chloroplasts. ( C )
Chloroplast in mesophyll cell ( X 24,000) from same leaf as in 7A showing well-
developed grana, stroma lamallae, and starch granules. ( D ) Chloroplast ( X 16,000)
from iron-deficient mesophyll cell displaying lack of grana lamallae and absence of
starch grains. (After Zhou et al. 1984a.)
Roots of chlorotic ‘Bartlett’ pear trees grown on calcareous soils con-

tained higher iron levels than normal trees with no consistent foliar iron
level trends between chlorotic and nonchlorotic trees (Milad 1924).
Lindner and Harley (1944) examined a range of leaf iron fractions in
chlorotic pears and proposed that lowered Ca :K ratios were a causal factor
in chlorosis development.
A whole range of plant nutrient ratios have been suggested over the
years as better indexes of plant iron status and/or in differentiating
chlorotic from nonchlorotic plants: total phosphorus to iron ( DeKock and
Hall 1955), leaf phosphorus to iron (DeKock et al. 1979), leaf iron to
leaf manganese (%indon and Srivastava 1981), and micronutrient bal-
ances (McGeorge 1948). In all cases, however, ratios or nutrient
inbalances in chlorotic vs. nonchlorotic tissue tend not to hold true for
plants within or among species. One of the reasons for the lack of success
is that under stress conditions, such as with iron deficiency stress, the
availability of and uptake of other nutrients can be altered. Tissue sam-
pling time (McGeorge 19491, in relation to chlorosis development, can
also result in variable tissue nutrient concentrations.
More recently, potassium and K :Ca ratios in the leaf have been used
as potential indices for lime-induced chlorosis (Barak and Chen 1984;
Hamze et al. 1985). The rationale is that potassium builds up in the
leaf due to a reduction in carbohydrate synthesis, which reduces po-
tassium movement from the leaf to the phloem (Hamze et al. 1985).
However, the degree of chlorosis and the time of leaf sampling will have
profound effects on the status of potassium and calcium in the leaf a t
any given time.
There are many reports on the inconsistency of leaf iron levels in
relation to degree of chlorosis or in separating chlorotic vs. nonchlorotic
leaves, e.g., no significant difference in total leaf iron between chlorotic
and nonchlorotic leaves, in apple (Mochecki 1977),in peach ( Abadia et al.
1985), in maize (Rosen et al. 1977), in a range of deciduous fruits (Iljin
1952), and in Eucalyptus sp. (Anderson 1983). Chaney (1984) ascribes
this inconsistency to errors and contamination during sample prepara-
tion. In addition to washing leaves in detergent and/or acid, great care
must be taken to avoid contamination, e.g., dust, soil, and contamination
from such sources as grinder blades and laboratory apparatus. However,
Wallace (1971) showed that chlorotic leaves do contain more iron than
green leaves in leaf samples collected from ornamental trees, acid and
detergent washed and analyzed in triplicate. This agrees with the conclu-
sion of Leeper (1952). It results from metabolism of carbon in dying
chlorotic leaves since inhibition of chloroplast formation is irreversible.
Jolley and Brown (1985) noted that when nutrient solutions were
changed (when the solution reached pH 4) there was an increase in the
concentration of most elements, particularly manganese, in both the tops

and roots of iron-efficient tomato plants. There was no difference in iron
concentration of plants grown in unchanged solutions, but plants in the
changed solution contained less chlorophyll. They concluded that a proper
balance between nutrients is needed in order for iron to function properly.
Sufficiency ranges for leaf iron have been published for some fruit and
nut tree species (Shear and Faust 1980). The reported sufficiency levels
vary with species: (all values in mg/kg on a dry-weight basis) apple
40-500, cherry 20-250, peach 100-200, and pear 100-800. Vose (1982)
presented a generalized breakdown of tissue iron levels: 60-300 normal,
10-30 deficient, and 400-1000 excessive. Recently, Smith ( 1985) deline-
ated sufficiency levels for the ornamental Tolmiea menziesii Pursh of
71-130 mg/kg. The value of these and other reported sufficiency ranges,
particularly a t the lower deficiency levels, in identifying a chlorotic vs.
nonchlorotic plant is questionable and, as noted by Chaney ( 1984),visual
examination probably presents a better index of chlorosis. Pear trees
surveyed in southeastern England (Thble 5.3) for example, could not be
separated into chlorosis classes based solely of leaf iron status not only
within a cultivar but also between cultivars (Little 1971).Thus compila-
tion of leaf analysis values by crop for diagnosing potential iron chlorosis
problems becomes difficult and may have little interpretative value.
Because of the lack of agreement between total iron and chlorophyll in
chlorotic vs. nonchlorotic pear leaves, Oserkowsky ( 1933) studied the
solubility of leaf iron with different extractants. He found the 1 N HC1
Table 5.3. Relationship between Leaf Iron Content

and Degree of Chlorosis from Pear Orchards in
Southeastern England”
Leaf iron
Site Cultivar Chlorosis ( PPm)
1 ‘Conference’ Moderate 60
1 ‘Conference’ Severe 45
3 ‘conference’ Absent 75
3 ‘Conference’ Severe 85
4 ‘conference’ Absent 70
4 ‘conference’ Moderate 50
4 ‘conference’ Severe 45
5 ‘Williams’ Absent 90
5 ‘Williams’ Moderate 90
5 ‘Williams’ Severe 78
“Adapted from Little ( 19711.

extractable iron consistently differentiated green leaves from chlorotic

leaves, and termed this fraction “active iron” since it was thought to
participate in chlorophyll formation. Modification of the original extrac-
tant, such as using 0.1M etherized HC1 (Bolle-Jones 1955), have been
proposed. Although active iron was able to differentiate chlorotic and
nonchlorotic conditions in deciduous fruit trees ( Lindner and Harley
( 1944) and citrus (McGeorge 1949) it is not currently used as a reliable
diagnostic tool (Chaney 1984). Haussling et al. ( 1985)demonstrated that
lower uptake and translocation of iron are responsible for chlorosis in
grapevines rather than an inactivation within the shoots. Determination
of leaf chlorophyll content was considered the most straightforward
means of quantifying iron chlorosis by Chen and Barak ( 1982).This may
not be a valid approach since other elemental deficiencies or toxicities
also cause a lower chlorophyll content. Katyal and Sharma (1980)pro-
posed a ferrous iron leaf assay, which in their work with rice was found to
differentiate between green and chlorotic plants.
Enzyme activities have been proposed as physiological indicators for
diagnosis of copper and iron deficiencies (Brown and Hendricks 1952).
The use of such indicators is of particular importance in citrus, where
multiple micronutrient deficiencies occur simultaneously ( Bar-Akiva
1961).A peroxidase assay was proposed as a reliable means in diagnosing
iron deficiency in chlorotic citrus as differentiated from manganese defi-
ciency ( Bar-Akiva 1965). This assay has yielded significant correlations
with chlorophyll concentrations even before the appearance of visual
deficiency symptoms, thus providing an appraisal of the iron nutritional
status of citrus in the field (Bar-Akiva and Lavon 1968). Garcia et al.
( 1981) studying the role and interaction of metals at the cellular level, used
superoxide dismutase patterns as affected by iron nutrient concentra-
tions. Recently, Sevilla et al. (1984) isolated two iron superoxide
dismutases in lemon leaves, which can be used as markers of functionally
active iron in plant cell metabolism.
VI. BICARBONATE-INDUCEDCHLOROSIS
The common occurrence of iron chlorosis on calcareous soils led to the

term “lime-induced”chlorosis. Juritz ( 1913)reported chlorosis of almond,
apple, apricot, fig, peach, prune, and citrus in South Africa to be
associated with calcareous soil. Other early reports made the same asso-
ciation with melons and cucumbers (Castle l899), with lodgepole pine,
aspen, and a range of other nursery tree species (Korstian et al. 19211,
with pear (Hendrickson 1924),with citrus (Burgess and Pohlman 19281,
and with grape and other woody plants (Wann 1920).
The importance of bicarbonate and its concentration in the soil as

affected by other soil properties have been noted previously. A number of
soil-plant cultural management practices can greatly affect the incidence
and/or severity of lime-induced iron deficiency stress. Among these the
two most predominant factors are practices that affect soil water content
and soil compaction.
The importance of bicarbonate levels in irrigation waters in the devel-
opment of lime-induced chlorosis was initially pointed out by Harley and
Lindner ( 1945),who observed that irrigation waters high in bicarbonate
caused chlorosis of apple and pear grown in north central Washington.
Schinas and Powell ( 1977) warned against using irrigation waters
containing high bicarbonate levels, and Rogers (1978)noted that peach
cultivation in semiarid regions could be affected by irrigation waters
containing high bicarbonate, particularly on high pH soils.
The control of soil moisture content has been observed by many
researchers as a means of reducing or eliminating lime-induced chlorosis
(Juritz 1913; Burgess and Pohlman 1928; Hass 1942; Hilgeman and
Sharples 1957; Boxma 1972). Wet calcareous soils usually have higher
HCO; contents because soil COZincreases. Chapman (1939)grew sweet
orange seedlings in sand culture that were amended with CaC03 in
studies on the use of finely ground magnetite as an iron source. He noted
that frequent flooding of the pots brought on iron chlorosis symptoms.
Similarly, Wallace et al. ( 1976a)found that an excessively wet calcareous
loam soil produced severe chlorosis of iron-inefficient soybeans; the same
soil, but drier, did not induce chlorosis. Frequency and type of irrigation
can also aid in the control of bicarbonate-induced chlorosis. The inci-
dence of lime-induced chlorosis in field-grown grapefruit and orange both
on sour orange rootstock was increased by shortening the interval be-
tween sprinkler irrigations (Levy 1984). Drip irrigation corrected the
condition, and this was verified by increased leaf chlorophyll contents
and peroxidase activity. The proportion of the calcareous soil wetted by
drip irrigation was less than with sprinkler irrigation, which resulted in
less bicarbonate formation. Indigenous poor drainage of some calcareous
western U.S. soils have produced chlorosis in apple (Barney et. al. 1984).
The effect of soil oxygen levels on chlorosis development was studied
by Wallihan et al. (1961). Sweet orange seedlings were grown on an acid
sandy loam soil amended with CaC03 under two aeration levels: 3.5 and
20% oxygen. The calcareous soil under low oxygen produced the most
severe iron chlorosis, and so it was concluded that low oxygen was the
causal factor. However, under such conditions (relatively high CO,) there
may have been a higher concentration of bicarbonate.
The degree of soil compaction has been implicated in the severity of
lime-induced chlorosis (Juritz 1913; Thorne 1941). Mengel and Bubl

(1983) and Mengel et al. (1984b) concluded that the degree of lime-
induced chlorosis in grape was associated with higher soil clay content.
The rationale presented was higher soil clay content + higher compac-
tion of soil thereby producing higher COZ levels, which results in
elevated HCO; concentrations. An associated problem with soil compac-
tion in relation to lime-induced chlorosis is that in many cases plowing,
particularly deep plowing, may increase the problem (Schinas and Powell
1977). Deep plowing may bring more calcareous materials to the surface
and/or it may induce the roots to grow deeper into areas of more highly
calcareous materials, depending upon the soil profile and underlying
materials.
Peach leaf analysis from trees grown on two calcareous soils in Spain
(Abadia et al. 1985) showed increased K:Ca ratios as well as levels of
potassium compared to nonchlorotic trees. Potassium accumulation was
shown to be a result and not a cause of lime-induced chlorosis for peach
seedlings grown in the dark or light under iron stress (Thorne et al. 1950).
Those in the dark contained lower chlorophyll and accumulated leaf
postassium. Contrarily, El Gazzar and Wallace ( 1966) reported lowered
leaf potassium in trifoliate orange and rough lemon seedlings when grown
in solutions containing 20 meq per liter NaHC03, an extremely high bi-
carbonate level.
Some effects of bicarbonate on plant metabolism have been reviewed
by Miller (1960). Altered nitrogen metabolism, and organic acid and
carbohydrate synthesis and translocation were shown to occur from
lime-induced chlorosis (Iljin 1951a,b). Tobacco (Khadr et al. 1966) and
citrus ( McGeorge 1949) exhibited increased leaf citrate to malate levels
after exposure to bicarbonate. However, many types of chlorosis (iron
deficiency, heavy metal induced, aging)have been shown to result in more
citric acid in the leaves relative to malic or oxalic acids (DeKock and
Morrison 1958). Rhoads and Wallace (1960),however, showed that bean
plants had elevated oxalate levels under iron stress. Assimilation of
bicarbonate by roots of different plant species increased the organic acid
content of trifoliate orange roots but less in avocado roots as indicated by
14C-labeledHCOi (Bedri et al. 1960). Iron-efficient soybean and barley
both assimulated less HC03 than either orange or avocado.
Solution-culture-induced bicarbonate chlorosis in sunflower was found
to increase the cytoplasm pH in leaf cells by Kolesch et al. (1984).They
hypothesized that cytoplasm pH plays an important role in establishing
plant resistance or susceptibility to iron chlorosis. The question remains
as to whether this is a cause or effect of iron chlorosis.
VII. IRON CHLOROSIS:

HORTICULTURAL OCCURRENCES
There have been many reports of iron-deficiency stresses in ornamental

and fruit tree species. Most of these reports deal with lime-induced iron
deficiency on calcareous soils but others are rather specific occurrences,
such as nutrient imbalances created in nursery beds. The material that
follows should serve as a compilation of data and observations and a
source of potential new areas for research. n e e species, especially fruit
trees, dominate this section. Work on other horticultural crops, such as
vegetables, has been incorporated in prior sections.
Although most authors have found suitable control measures to ame-
liorate the symptoms of iron deficiency under the soil, climate, and plant
type conditions of their particular concern, all such solutions should be
regarded as temporary. “Control” of iron deficiency chlorosis may last
only a short period (weeks)or up to 2-3 years. Moreover, the source of iron
used and the method of application can have varied affects on foliar injury
to the plant. The work of Joessel et al. ( 1937)on peach and pear illustrates
the range of plant injury that can occur. n u n k injection of peach with
ferric ammonium oxalate, ferric ammonium citrate, or ferric potassium
oxalate resulted in no injury, severe foliar burning, and slower recovery
(compared to ferric ammonium oxalate) with no injury, respectively, for
the three iron sources. Pear tree injection with the same three iron sources
in addition to ferric potassium tartrate and iron pyrophosphate resulted
in some foliar burning and older leaf defoliation independent of iron
source. Ferric ammonium citrate caused a higher degree of burning when
compared to all other sources.
Additionally, the history of iron chlorosis control methodology, as
noted earlier, goes back to Gris in 1845 (Gile and Carrero 1920), whose
work was reproduced and expanded by Von Sachs (1887). He not only
induced and corrected iron deficiency in solution-culture-grown maize
and bean plants with iron chloride or sulfate but also demonstrated that
“painting” iron-deficient bean leaves with a dilute iron solution and
injection of iron sulfate into auger holes in an iron-deficient acacia tree
both were remedies for control of iron deficiency.
A. Deciduous ’Ikees and Ornamentals

Chlorosis of apples in the Bitter Root Valley of Montana was ascribed
to a lack of available soil iron (Burke 1932).Of the control methods tested,
trunk injection of ferrous sulfate gave the best results. McGeorge (1949)
162 RONALD E KORCAK
did a comprehensive study of Arizona orchards in relation to lime-

induced chlorosis. Both apple and peach usually expresssed chlorotic
symptoms during the summer, which agrees with the results of Little
(1971). Deciduous trees grown in similar areas as citrus also displayed a
complex of iron, zinc, and manganese deficiencies. Bunk plugging with
ferric citrate was the best control for deciduous tree chlorosis. lhenty-
three percent of all Utah orchards exhibited some degree of lime-induced
chlorosis (Thorne and Wann 1950). In many of these cases poor soil
drainage increased the severity of chlorosis. A number of possible correc-
tive measures were tested; trunk injection of ferrous sulfate was the
perferred direct control.
Working with ‘Jonathan’ and ‘McIntosh’ apples, Mochecki ( 1977)
recommended trunk injection of ferric citrate, but Barney et al. (1984)
found ferrous sulfate sprays better than injection and Zen-qin et al.
(1979) obtained the best results with 60 liters per tree of 2% ferrous
sulfate applied to the roots. A mixture of ferrous sulfate and horse
manure (or other organic materials) applied in holes around the tree was
an effective control of “greensickness” (iron deficiency) on calcareous
soils in north China (Zhou and Chang-Zhen 1982).The expression of iron
chlorosis symptoms in apples in China is most prominent during the
rainy season, with the symptoms lessening in intensity during the follow-
ing dry season (Shen and Tseng 1949). Foliar application of ferrous
sulfate a t the onset of the rainy season was found to be an adequate
temporary remedy. Chlorotic apples grown on calcareous soils in Pakistan
were regreened by injection of solid ferrous sulfate crystals into the
sap-wood (Ganai 1953).
Recent work by Zhou et al. (1985)confirmed the effect of elevated pH
on chlorosis development in apple seedlings grown under two bicarbonate
and potassium levels. Chlorotic symptoms and reduced growth resulted
from increasing bicarbonate (Zhou et al. 198413). Increasing potassium
tended to ameliorate the effect of bicarbonate both by increasing leaf iron
and by overall growth. Bicarbonate also reduced uptake or translocation
of 45Cato the upper leaves. Tong et al. (1986) found that high chelator
(EDDHA, 10 :1molar ratio) :iron ratios in nutrient solutions reduced the
occurrence of chlorosis in apple seedlings compared to low chelator :iron
ratios ( 1:1 ratio). Iron transport to the tops was apparently facilitated by
the higher chelator :iron ratio. Addition of higher micronutrient (manga-
nese, copper, zinc, molybdenum, and boron) levels to the solutions a t low
chelator :iron ratios resulted in excessive root levels of manganese, zinc,
and copper. The high copper levels may have been a causative factor in
reducing iron uptake per se and/or caused a more direct toxic effect on the
roots. The overall effect of elevated micronutrients on iron chlorosis was
more pronounced than the reduced availability of iron by phosphorus.
Iron chlorosis of pears was identified as early as 1920 in the United

States (Gile and Carrero 1920; Wann 1920) and earlier in France (see
Hendrickson 1924). Hendrickson (1924) studied pear chlorosis in the
Santa Clara Valley of California. Pears grew normally in a noncalcareous
surface soil with a relatively high water table. Over a period of years as
the water table fell, tree roots grew deeper, which initiated a chlorotic
problem, owing to subsurface calcareous soils. His investigation showed
that both rootstock and cultivar displayed varying sensitivities to chloro-
sis. P y r u serotinu Fkhd. and quince were more sensitive than l? communis
L. to chlorosis. The cultivars ‘Hardy’and ‘Clairgean’were relatively more
resistant to chlorosis than ‘Bartlett’,‘Nelis’,‘Cornice’,and ‘Glout Morceau’.
The best corrective measure was surface application of ferrous sulfate,
one application of about 9 kg per 15-year-oldtree lasted 2-3 years. Raese
and Parish (1984) found that trunk injection of 1%ferrous sulfate solu-
tion into ‘Anjou’ and ‘Bartlett’ pears, especially in the spring or fall,
provided adequate iron levels and that injections were more persistent
than foliar iron applications. The quince rootstock has been noted by
others to be more susceptible to lime-induced chlorosis than other root-
stocks for pears (Little 1971; Boxma 1972).Pyrus calleryana Decne. and
l? ussuriensis Maxim. rootstocks were more susceptible to chlorosis than
l? communis seedling rootstock for ‘Anjou’pear in the Rogue River Valley
of southern Oregon (Higdon 1957). European seedling (l? cornmunis)
rootstocks appear to be more tolerant of lime-induced chlorosis than
Oriental pear rootstocks.
Fruit crops and ornamentals seriously affected by chlorosis were listed
by Wallace and Lunt (1960)but apples were not listed. Thorne and Wann
( 1953)list resistant and susceptible ornamentals for planting in Utah and
earlier (1950) ranked deciduous fruit species in order of most to least
severely affected by lime-induced chlorosis in Utah: peaches and
pears > sweet cherries > plums > apricots > apples > sour cherries. This
ranking agrees with that of Barney et al. (1984)but is slightly different
from that of Vose ( 1982),who noted that peaches, plums, and cherries (no
distinction made between sweet and sour) may suffer severely from iron
chlorosis, but apples are relatively tolerant. Kessler (1957) noted the
following order of highest resistance to lime-induced chlorosis:
olive > plum > peach > grape. The plum and peach were both propagated
on almond rootstock and grape was ‘Madeleine Oberlin’ on 41-B root-
stock. However, as noted previously, apple seedlings were found to be
more sensitive to bicarbonate-induced iron chlorosis in controlled soil pot
studies than even iron-inefficient soybean cultivars (unpublished data).
Chlorosis-resistant peach rootstocks have been introduced that are
primarily peach x almond hybrids (such as G.F. 677), although other
Prunus species, such as the Japanese plum, have displayed resistance
and may be useful if graft incompatibility problems can be overcome

(Rom 1983; Syrgiannidis 1985). Field trials over a period of 10 years
showed that the G.F. 677 rootstock exhibited less severe, by about 40%,
symptoms of iron chlorosis compared to peach seedling rootstocks (Fig.
5.8). Although major advances have been achieved in the use of chlorosis-
resistant rootstocks, there has been little if any research on the physiol-
ogy of resistance or tolerance to iron deficiency chlorosis.
The importance of bicarbonate vs. high calcium in calcareous soils on
the incidence of chlorosis of peach was found by Bindra ( 1976)in solution
culture tests. Addition of 1 meq HC03 to solutions induced chlorosis,
while high solution calcium levels had no effect. A nitrogen-induced iron
chlorosis of peaches on a calcareous Colorado soil was noted by Stebbins
et al. (1963).The chlorosis occurred after a 0.4 kg per tree application of
ammonium sulfate and was corrected by an application of FeEDDHA.
Rogers (1978) also corrected chlorosis in peach with FeEDDHA;
postapplication leaf analysis showed higher iron and lower manganese.
When applied in excess, FeEDDHA-induced manganese deficiency in
both peach and apple (Rogers 1978). The potential effect of applied
-
73 74 75 76 77 78 79 80 81 82
YEAR
G.F. 677 **a SEEDLING
Fig. 5.8. Average chlorosis ratings of ‘Vivian’ peach on either peach seedling or
G.F. 677 rootstock grown on a calcareous (1.3-12.2%“active” lime) soil in Greece.
Ratings: 0, no chlorotic symptoms; 10, very strong chlorosis. (Redrawn from Syrgiannidis
1985.)
FeEDDHA on the plant compositional balance between iron and manga-

nese was vividly shown by Moraghan (1985a,b). Manganese toxicity of
flax could be eliminated and an induced manganese deficiency could be
obtained by the application of FeEDDHA to the soil. Thus special
attention should be given to both the iron and manganese nutrition of
plants when FeEDDHA is used as an iron source.
Drip-irrigation-applied FeEDDHA and sulfuric acid resulted in rapid
correction of chlorosis of peach in Chile (Razeto 1982). Quantities of
FeEDDHA necessary for correction are greatly reduced when applied via
drip irrigation. Yoshikawa et al. (1982)corrected chlorosis in plums with
injection of ferrous sulfate; about 400 ml of a 2% solution lasted 2 years.
Ferric ammonium citrate, because of its greater solubility, may be a
better source for tree injection than ferrous sulfate (Wallace et al. 1984).
The larger volumes of ferrous sulfate required necessitate the use of
pressure injection.
The common cherry rootstocks, mahaleb and mazzard, both are sus-
ceptible to lime-induced iron chlorosis (Day 1951). Among stone fruit
rootstocks, almond tolerates calcareous soils best, followed by myroba-
lan, apricot, and peach rootstocks (Day 1953; Kessler 1958). Illarionova
(1980) reported that 43% of 206 cherry cultivars in the U.S.S.R. are
chlorosis resistant.
Various other trees and ornamentals classified as susceptible to lime-
induced chlorosis have been reported in the literature (Korstian et al.
1921; Wallace et al. 1953; Dale et al. 1955; Bennett 1963; Shoulders and
Czabator 1965; Nelson and Selby 1974; Clement 1977; Carter 1980).Gile
and Carrero (1920)noted that chestnut trees were prone to chlorosis in
soils with greater than 3% CaC03; however, chlorosis was absent when
grafted on oak roots. Contrarily, Chadwick (1935)and Messenger (1984)
noted that pin and white oaks were chlorosis susceptible on calcareous
soils, although soil application of sulfuric acid with or without the
addition of ferrous sulfate alleviated the problem.
B. Grapes
I t has long been known that rootstocks from American grapes (Vitis
labrusca) and their hybrids are very susceptible to lime-induced chloro-
sis, while the European ( V ; uinifera) rootstocks are resistant (Gile and
Carrero 1920; Wann 1941; Thorne and Wann 1950). Vitis berlandieri
Planch., a species native to the limestone hills of central and southwest
R x a s (Winkler et al. 1974) and its hybrids have extended the range of
calcareous soils that can be utilized for grape growing ( Spiegel-Roy 1979).
‘Concord’grapes developed the most severe chlorosis during the high-
166 RONALD E KORCAK
light and high-temperature periods of summer; “natural” recovery may

occur later in the growing season (Wann 1941). Later work (Burtch et al.
1948) led to the hypothesis that during periods of high soil water, poor
aeration, and cool temperatures plant iron became inactivated. The small
“shot” berry disorder of ‘MadeleineOberlin’grape was found to be due to
iron deficiency chlorosis ( Samish 1954).Spraying fresh pruning wounds
in early spring with ferrous sulfate alleviated the disorder.
Blanc-Aicard and Drouineau ( 1956) proposed determinations of root
calcium saturation as a quick method to differentiate grape rootstocks a t
an early age. Stocks with a high saturation capacity were very chlorosis
susceptible. It has been demonstrated that grapevine roots have a ferric-
iron-reducing mechanism (Varanini and Maggioni 1982). Saglio ( 1970)
showed that bicarbonate induced chlorosis in a chlorosis-susceptible
grape, while a chlorosis-resistant strain was not affected in solution
culture. The effect of bicarbonate was increased by high levels of solution
phosphate. Mengel et al. (1984a) concluded that bicarbonate was the
primary cause of iron chlorosis in grapes and that high phosphorus levels
usually found in chlorotic leaves are the result and not the cause of iron
chlorosis. Additionally, Booss et al. (1984) considered iron chlorosis in
grapes a complex interaction between bicarbonate and phosphorus. Lists
are available (Galet 1971) for the percentage of soil-“active”lime that can
be tolerated by various grape rootstocks.
Ethylene was the cause of iron chlorosis of bicarbonate-resistant grape
rootstocks in Swiss vineyard soils (Perret and Koblet 1984).The authors
hypothesized that ethylene inhibited both root growth and elongation;
thus iron uptake was inhibited. The incidence of chlorosis was hastened
by incorporation of undecomposed organic matter and high soil water,
which was shown to increase soil ethylene levels. Correction of chlorosis
was found by interplanting oil radish, which reduced the water content of
the root zone.
C. Citrus
An early association was made between the incidence of chlorosis and
irrigation in citrus (McGeorge 1949)as with lime-induced iron chlorosis
of other crops. I t was also noted that the dividing line between chlorotic
and nonchlorotic soils was about 2.5-370 CaC03, with “active” calcium
( 0.2 N ammonium oxalate extractable) significantly higher in chlorosis-
producing soils in Arizona. Sour orange rootstock was less prone to
chlorosis than sweet orange rootstock; orange and grapefruit propagated
on rough lemon root exhibited higher leaf manganese and iron than those
on sour orange root. Sour orange rootstock was suitable under Egyptian
conditions for citrus (El Gazzar et al. 1975). All citrus foliage studied
contained adequate iron, with ‘Washington Navel’ orange having the

highest leaf iron.
n e e s on sour orange rootstock were less susceptible than trees on
trifoliate orange rootstock, although under certain conditions sour orange
was still prone to chlorosis (Wallihan and Garber 1968; Hamze and
Nimah 1982; Levy 1984)as with ‘Washington Navel’ orange as the scion
(Levy and Mendel 1982). nifoliate orange rootstock and its hybrids have
shown tolerance to the tristeza virus but were sensitive to iron deficiency
chlorosis (Horesh and Levy 1981; Levy 1984).Hamze and Nimah ( 1982)
compared trees on sour orange and trifoliate orange rootstocks in con-
trolled tests on three calcareous soils, varying in CaC03 from 0.5 to 8870,
each under three soil moisture regimes. High soil water produced symp-
toms of lime-induced chlorosis on sour orange only on the 88% CaC03
soil. Symptoms appeared on trifoliate orange on soils with 88 or 30%
CaC03 with any of the three soil moisture regimes. These results again
showed that sour orange is not truly resistant to lime-induced chlorosis
under all soil conditions as noted above. Interestingly, foliar iron concen-
trations of chlorotic sour orange seedlings were reduced but still within
the optimum range, about 60 mg/kg. Chlorotic leaves of trifoliate orange
contained more iron than green leaves. Although sour orange seedlings
were more resistant to chlorosis under nursery conditions than ‘Cleopatra’
mandarin seedlings, the former was found to be more susceptible to
salting injury (Cooper and Peynado 1954).The effect of salt level, particu-
larly sodium, has not been fully investigated and may be a confounding
cause of chlorosis development of resistant rootstocks, such as sour
orange, under certain soil conditions. Optimum, high, and excessive
scion leaf sodium levels for citrus have been established (Embleton et al.
1973b). Lemon grown on Yuzu rootstock contained substantially higher
concentrations of iron, manganese, and sodium, and lower calcium than
other lemon rootstocks (Embleton et al. 1962).
The uptake and translocation of iron and zinc of ‘Valencia’orange on
trifoliate orange (susceptible to iron and zinc deficiencies) and rough
lemon rootstock (resistant to iron and zinc deficiencies) was studied by
Khadr and Wallace ( 1964).Under low iron and zinc, rough lemon absorbed
and translocated to the tops more of both elements, while under high
supply the difference between rootstocks disappeared for iron. Khadr and
Wallace concluded that iron and zinc translocation from roots to leaves
may be a more important problem than absorption per se. Both phospho-
rus and bicarbonate inhibited iron translocation in trifoliate orange, while
high calcium supply decreased iron uptake by both rootstocks.
Sour orange and grapefruit rootstocks both were more susceptible than
rough lemon to copper-toxicity-induced iron chlorosis on the acid sandy
soils of Florida (Smith et al. 1950).Redblush grapefruit propagated on 36
citrus rootstocks exhibited varied degrees of chlorosis under field condi-

tions on a calcareous soil (Cooper and Olson 1951).Sour orange rootstock
displayed little if any chlorosis, while sweet orange rootstock was severely
affected. Similar results were obtained with sour orange by Maxwell and
Wutscher (1976),who found tangelo and tangor rootstocks to be intoler-
ant of calcareous soil. Both ‘Kunenbo’ and ‘Cleopatra’ mandarin root-
stocks were found to be more tolerant of calcareous soil than 14 other
rootstocks for grapefruit (Wutscher et al. 1970). An updated listing of
citrus rootstocks and the soils they will tolerate was made by Wutscher
(1979) and reference leaf tissue iron values for diagnosis of potential
chlorosis problems are available (Wallihan 1966, Embleton et al. 1973b).
D. Other Horticultural Plants

The differential susceptibility of two populations of Eucalyptus viminalis
Labill. on acid and calcareous soils has been studied in detail by Ladiges
( 1977). The presence of physiologically distinct populations within a
species has been termed “edaphic ecotypes” ( Snaydon and Bradshaw
1961).Ladiges (1977)showed that the calcareous ecotypes of E. uiminalis
grew better on high pH soil, and when subjected to an iron stress these
plants took longer to develop chlorotic symptoms, developed new roots
faster, and displayed a greater capacity to remove iron from solution, all in
comparison with an acid ecotype. The plants from the acid ecotype
accumulated leaf phosphorus when grown in calcareous soil producing an
elevated leaf P :Fe ratio, symptoms of phosphorus toxicity, and chlorosis.
Similar phosphorus trends were noted by Anderson and Ladiges ( 1982)in
an acid ecotype of Eucalyptus oblique L‘Her. when grown on calcareous
soil. They suggested, for acid ecotype plants grown on calcareous soils,
that the development of severe chlorosis was due either to internal
inactivation of iron by phosphorus or to an inability to lower the external
pH to increase iron availability. Conversely, the calcareous ecotype was
shown to have a greater ability to absorb iron even under high external
phosphorus conditions (Anderson 198213).
The adaptability of the calcareous ecotype may be due to a more
efficient calcium extrusion pump vs. the acid ecotype (Anderson 1983)
and/or the acid ecotype has a lower optimum calcium uptake than the
calcareous ecotype (Anderson 1982a). Interestingly, the acid ecotype
when grown on calcareous soil showed a poor correlation between total
leaf iron and severity of lime-induced chlorosis (Anderson 1983). The
existence of edaphic ecotypes could be used as an important tool in the
determination of physiological and biochemical differences in relation to
lime-induced chlorosis. Such tests have been proposed (Brown 1976;
Voigt et al. 1982)and some information is available on species differences
among ornamentals for sensitivity to high-lime conditions.
Recently, Hershey and Paul ( 1984)classified the evergreen shrub Euon-

y m u s japonica L. as an iron-efficient plant since it responded to iron
deficiency by reducing nutrient solution pH. They noted, however, that
this plant was unusual because the pH reduction occurred during a flush
of shoot elongation when there were no iron deficiency symptoms. This
pH reduction during a time of shoot elongation is fortunate since known
shoot/root relationships in woody plants indicate that during root growth
there is a cessation of shoot growth flushes. Their results are of interest in
relation to the previously cited report by Davenport (1983),which indi-
cated a shifting of chlorosis in lime trees with the onset of a new flush of
vegetative growth. Perhaps, as pure speculation, woody plants have a
mechanism of inducing an iron stress response that is coordinated with
the onset of vegetative growth periods.
Deficiency symptoms and lists of chlorosis-susceptible shrubs, flower-
ing plants, and greenhouse crops are available (Finch et al. 1933; Clapp
1935; Laurie and Wagner 1940; Harbaugh 1986). Nelson and Jolley
( 1984)studied strawberries and raspberries planted on calcareous soils in
Utah. These crops are seldom grown in high-lime areas, but in general
strawberries were more susceptible to chlorosis than raspberries and
there were varietal differences between the strawberry cultivars sampled.
Iron sprays have been used to correct iron chlorosis of strawberries (King
e t a l . 1950).
Rootstock variability in tolerance to calcareous soils has been found in
several evergreen tree species. Mexican and Guatemalan races of avocado
rootstocks generally tended to be less tolerant of calcareous soils than
West Indian races (Halma and Goodall 1952; North and Wallace 1952;
Kadman and Ben-Ya’acov 1982). However, selected rootstock from the
Mexican group exhibited very high tolerance, while some West Indian
rootstock were less tolerant. One of the problems in rootstock populations
is the use of seed as a source of the rootstock, which may inject a great
deal of diversity into the populations. Field correction of avocado chloro-
sis on calcareous soils is best performed by applying 1-2 p g per milliliter
FeEDDHA via irrigation (Kadman and Lahav 1982).Papaya exhibited a
typical iron stress response by reducing solution pH when grown under
negative iron conditions (Kannan 1985).
VIII. USE OF CHELATES
The first successful use of chelated iron ( EDTA, ethylenediaminetetra-

acetic acid) for the correction of iron deficiency in citrus was reported by
Leonard and Stewart (1952). Between that time and the early 1980s a t
least 1300 published papers have appeared on the use of chelating agents
in plant nutrition (Wallace and Wallace 1982). The initial use of EDTA
was to correct iron deficiency in plants growing on acid soils (Leonard

and Stewart 1952; White 1954). The use of chelated iron sources to
correct chlorosis on acid soils not only required substantially less mate-
rial but also resulted in higher foliar iron levels, which were maintained
throughout the growing season (Thble 5.4). It was later noted that
correction was more difficult on calcareous soils, and hence FeEDDHA
was found to be more effective (Leonard and Stewart 1954; Stewart and
Leonard 1957)because of its greater stability a t higher pH levels (Kroll
1957). Fixation of iron from FeEDDHA by calcium has been demon-
strated (Lahav and Zipori 1978), as well as adsorption on clays (Ryan et
al. 1985)and a loss in effectiveness with time (Hamze et al. 1985).Foliar
iron uptake from high-pH (7.2)solution was more rapid with iron chelates
than with ferric chloride (Rutland and Chung 1971).
Many tests have compared the efficiency of various chelating agents
both in soil application (Hill-Cottingham 1957; Kuykendall et al. 1957;
Orphanos and Hadjiloucas 1984) and as foliar sprays (Rutland and
Chung 1971; El-Kassas 1984). The efficiency and inherent problems in
foliar sprays have recently been reviewed by Swietlik and Faust (19841,
the absorption and distribution of iron by Hsu et al. (1982), and the
timing and effect of surfactant on leaf injury by Wallihan et al. (1964).
Current corrective measures for iron deficiency are generally ineffective
and uneconomical (Ponnamperuma 1982). The high cost, potential leaf
injury due to burning, and need for repeated applications of chelating
Table 5.4. Leaf Iron Content of 'Valencia' Orange as Affected by Application of

Chelated and Unchelated Iron Sources to an Acid Soil"
Iron applied Material Flush of Leaf iron

(g h e e ) Condition of tree applied growth (PPm)
0 Deep green, healthy None Spring 62

Deep green, healthy None Summer 124
0 Severely chlorotic None Spring 25

Severely chlorotic None Summer 30
9,000 Most leaves regreen 100 lb Spring 43

Severely chlorotic FeSO4.7H20 Summer 24
30,000 All leaves green 200 lb Spring 82

All leaves regreen FeSO,( anh ). Summer 51
20 All leaves regreen 1/~ lb Fe-EDTA Spring 100

All leaves regreen Summer 95
"From Leonard and Stewart ( 1952).

agents has remained a constant deterrent to their ubiquitous use. Wallace

and Wallace ( 1982)in their review of 30 years experience of using chelating
agents point out six perplexing aspects of their use in plant nutrition:
cost;
poor quality control of some chelating agents, especially FeEDDHA;
foliar sprays give inconsistent results;
plant iron response mechanisms result in differential behavior to
metal chelating agents;
response to chelating agents is inconsistent between soil and
solution culture experiments;
the use of computer-generated chelate distribution in predicting
relative effectiveness of chelating agents may not provide reliable
predictions of their usefulness.
Besides the use of chelates, a number of other corrective measures have

been employed, including sulfuric acid application to the soil, industrial
by-products, pyrite, and organic compounds. Hagstrom ( 1984) recently
reviewed the use and effectiveness of many of these iron sources.
IX. CONCLUSION
This review has covered many aspects of iron in soils and plants
indicating past and current research on deciduous and nondeciduous
plants and the use of chelates for iron deficiency chlorosis. There have
been major advances in our understanding of this unbiquitous plant
disorder, with new information being added almost daily. Most of the
research on uptake mechanisms and translocation within the plant has
been centered on agronomic crops. Horticultural crops, particularly fruit
tree species, have been shown to be as or even more sensitive than
agronomic crops. More research is needed on the uptake mechanism for
iron in fruit trees. &search on perennial tree crops is and has been
confounded in part by the use of seedlings directly or as rootstocks. With
the advent of tissue culture propagation of perennials, researchers can
eliminate this confounding of results.
The role of root/shoot interactions in iron uptake needs to be explored
as well as the effect of an annual crop on iron distribution within the
plant. Certain Prunus species such as peach are known to be more
susceptible to iron deficiency chlorosis than apples or pears. Is this a
reflection of the need for more vegetative growth of these trees for annual
production ( a dilution effect) or is it due to physiological and biochemical
differences between species? Deciduous and nondeciduous trees grown in
172 RONALD E KORCAK
the same locale display different tendencies in developing iron chlorosis

as well as multiple micronutrient deficiencies. This seems to point to-
ward differences in uptake mechanisms and/or reactions of the root to
iron stress.
Since much needs to be known about iron nutrition or tolerance of
bicarbonate, the use of chelating agents as a corrective measure for iron
supply will continue. However, a knowledge of soil- and culture-related
activities could reduce the incidence and severity of chlorosis and will aid
the grower directly.
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6
Ginseng: Industry, Botany, and Culture
J. ?I A . Proctor*
Department of Horticultural Science, University of Guelph
Guelph, Ontario, Canada N1G 2W1
W. G . Bailey?
Department of Geography, Simon Fraser University
Burnaby, British Columbia, Canada V5A 1S6
I. Introduction 188
11. Industry 188
A . Pharmacology 188
B . Production and n a d e 193
111. Botany 195
A. Taxonomy 195
B . Genetics and Cytology 197
C . Morphology, Anatomy, Growth, and Development 198
IV. Culture 206
A . Propagation 206
B . Soils and Nutrition 209
C . Environmental Physiology 211
D. Pest Control 227
E . Weed Control 228
*I am indebted to Dean Louttit for excellent technical assistance. Canadian gin-

seng growers have been very supportive of our efforts and for this I am grateful. The
Ginseng Research Institute, particularly its president, Arthur Rashap, has been
most encouraging to me. Some of the research reported herein was supported by the
Ginseng Growers Association of Canada and by Operating Grant A6697 of the
Natural Sciences and Engineering Research Council of Canada held by me.
+I would like to acknowledge the following: Simon Fraser University, Burnaby,
British Columbia, for providing a President’s Research Grant to assist in the transla-
tion of Korean ginseng manuscripts; Chai-Na-Ta Ginseng Products Ltd., Lytton,
British Columbia, and particularly its President, J. M. M. Latta, for access to ginseng
gardens for research purposes, and for interest and support in the improvement of
ginseng cultivation techniques; R. J. Stathers, Department of Soil Science, Univer-
sity of British Columbia, Vancouver, British Columbia, for assistance in the study of
the micrometeorology of ginseng shade and mulch environments; and A. L. Skretkowicz,
Kwantlen College, Surrey, British Columbia, for assistance in research on ginseng
response to environmental parameters.
187
188 J. T. A. PROCTOR AND W. G. BAILEY
I. INTRODUCTION
Ginseng is an herbaceous perennial that is cultivated for its highly

valued root. It has been used by the Chinese for thousands of years as a
wonder drug, cure-all, and aphrodisiac. Its importance in the New World
is recent (eighteenth century) and can be traced to French Jesuit missionaries
(Carroll 1985; Schorger 1969) who correctly predicted indigenous plant
populations in Canada. Collecting and exporting the dried root eventu-
ally led to cultivation under artificial shade in the late 1800s in the United
States and Canada.
Various aspects of ginseng culture have been discussed in grower
guides (Williams and Duke 1978; Lewis 1980), books (Curran 1983;
Fulder 1980),theses (Carpenter 1980; Polczinski 1982),an indexed bibli-
ography (Rashapet al. 1984),proceedings of six North American ginseng
conferences (Hensley et al. 1979; Jones and Kring 1983; Lapp 1981;
Missouri Department of Conservation 1980; Proctor 1984; Roberts and
English 1982), and proceedings of four international ginseng symposia
(Korean Ginseng Research Institute 1974, 1978, 1980, 1984), but there
never has been a review of North American ginseng production. Our
objective is to review recent western publications and related Oriental
papers (where access and translation permit) on the most important
aspects of industry, botany, and culture of ginseng. In many areas there is
an absence of well-documented research and so we have presented
unpublished research and/or personal observations with the hope that
they will stimulate useful discussion and appropriate research.
11. INDUSTRY
A. Pharmacology
There is tremendous literature on the chemical constituents and associated

pharmacological testing of ginseng. The Ginseng Research Institute’s
indexed bibliography (Rashap et al. 1984)lists some 2320 papers, most of
which deal with plant constituents and their action. A separate review
could easily be written on these topics. The approach taken here will be to
list and discuss briefly the active constituents of ginseng, and then to
discuss how they are thought to work.
1. Medicinal Chemicals. Garriques (1854) in Germany is credited
with isolating the first active principle, a saponin, from ginseng roots.
This was followed by isolation and identification of saponins by Elyakov
and his colleagues at Moscow University (Elyakov and Strigina 1962;
6. GINSENG: INDUSTRY, BOTANY. AND CULTURE 189
Elyakov et al. 1962).At the same time Lin (1961)in lhiwan and Shibata
and his colleagues (Shibata et al. 1962a,b, 1963) a t the University of
Tokyo also reported isolation and identification of saponins. Since that
time, hundreds of papers reporting the chemical details have appeared.
For the interested reader, their titles are listed in Rashap et al. (1984).
The two types of plant saponins or glycosides are the closely related
steroidal and terpenoidal (Bonner and Varner 1965). Familiar steroids in
the animal kingdom are cholesterol, cortisone, estrogen and progesterone
(the female sex hormones), and testosterone (the male sex hormone).
There are many triterpenoid glycosides in the plant kingdom, e.g., in
licorice and thyme.
Advances in the isolation and identification of the ginsenosides of the
ginseng plant have been made possible by improvement in analytical
techniques. mo-dimensional thin-layer chromatography has been used
for separation and identification (Jhang et al. 1974). The ginsenosides
have been quantified using gas chromatography (Brieskorn and Mosandl
1978),gas chromatography-mass spectrometry ( Bombardelli et al. 1978),
high-pressure liquid chromatography ( Sticher and Soldati 1979),droplet
countercurrent chromatography (Otsuka et al. 1977), and spectrodensi-
tometry (Liberti and Der Marderosian 1978).
The Russians (Elyakov et al. 1962) isolated six saponins from r!
ginseng root, which they called panaxosides. They divided these into two
groups, ABC and DEF, based on the structure of the modified aglycones
(nonsugar part of the glycoside). The ABC group has panaxatriol as its
aglycone and the DEF group has panaxadiol. Shibata et al. (1962a,b,
1963) called the saponins ginsenosides rather than panaxosides and
labeled them Ro to Rh from the sequence of Rf values in thin layer
Chromatography. The relationship between the ginsenosides and panaxosides
is shown in n b l e 6.1.
The identity and/or the amount of ginsenosides present in American,
Oriental, and other ginseng plants and products is controversial (Lui and
Staba 1980). Part of this may be due to different environments in which
ginseng is grown, variation among plant species, and the various processes
used to prepare ginseng products.
Lui and Staba (1980)have examined various ginseng plant parts and
plant products for the relative level of ten ginsenosides (lhble 6.2). The
presence of Rgl in American ginseng is contrary to the findings of
Otsuka et al. (1977). I t is interesting that many of the so-called ginseng
commercial products tested contained little or no ginseng. This is one of
the great concerns of the Ginseng Research Institute (Rashap et al.
1984). The Institute would like a set of standards developed relating to
labeling and content of ginseng products, and the placing of a seal on
products meeting the standards.
190 J. T. A. PROCTOR AND W.G. BAILEY
Table 6.1. Some of the Active Constituents of Ginseng“
Ginsenoside‘ Panaxoside“ Aglyconed
Oleanolic acid
F ‘diol
‘diol
E
‘diol
D ‘diol
‘diol
C ‘triol
B ‘triol
‘triol
A ‘triol
‘triol
‘triol
‘triol
“Based on Elyakov et al. (1962)and Shibata et al. (1962a,b, 1963).

‘Name based on the sequence of Rf values in thin-layer chromatography.
‘Name based on aglycone group.
d’diol, panaxadiol; ’triol, panaxatriol.
2. Action of Ginseng. It is now generally agreed that the ginsenosides

are the constituents mainly responsible for the pharmacological action of
ginseng. The primary action is as an adaptogen-a substance that will
correct human physiological imbalances. One of the better texts on
ginseng and its action is “The Root of Being” (Fulder 1980).The Oriental
literature contains many papers (see Rashap et al. 1984)dealing with the
action of ginseng. For example, Saito and Bao (1984) reported that
ginseng helped mice in the forced exercise of rope climbing and protected
the stressed mice against the decrease of sex behavior.
Some of the main claims for ginseng action are summarized by Kim
( 1978)and include “increased performance” during fatigue, “central nerv-
ous system stimulation” during depression, and “increased moistening of
the skin, emollient action on the face,” when the disorder is skin dryness.
3. Toxicology. Ginseng has received some bad publicity. For example,

Siege1 (1979) noted that long-term use of commercial ginseng prepara-
tions resulted in an incidence of hypertension together with nervousness,
sleeplessness, skin eruptions, and morning diarrhea. However, these find-
ings have been severely criticized because of the choice of test materials
and their concentrations, and the patients used in the study (Soldati
1984). Palmer et al. (1978) described a case of mastalgia in a 70-year-
old woman who developed swollen, tender breasts after taking ginseng
Table 6.2. Ginsenosides in Various Ginseng Plants and Productsa
Total
ginsenosides
(W/W%)b Relative amount of ginsenosides'
Material wt Sp Ro Rbl Rbz Rc Rd Re Rf Rg, Rg, Fll

~
Plant root
1. American ginseng (fibered),
I? quinquefolium 28.8 4.7 i ++++ + +++ +++ ++++ - ++ ++ -
2. American ginseng, I? quinquefolium 14.1 5.9 + ++++ + ++ ++++ ++++ - ++ ++ -
3. Wild American ginseng, €? quinquefolium 12.2 8.0 + ++++ + ++ ++ ++++ ~ ++ +++ -
4. Canadian ginseng, P. quinquefolium 17.8 6.1 + ++++ + ++ +++ ++++ - ++ ++ -

5. Korean white ginseng, I? ginseng 11.0 3.7 + +++ ++ +++ ++ +++ ++ ++++ t r -
6. Korean red ginseng, P. ginseng 5.2 2.3 + +++ ++ ++ t +++ + ++++ + +

7. Chinese red ginseng, I? ginseng 7.5 2.7 + +++ ++ ++ + +++ i+ ++++ + +
- - - - - - -
8 . Panax trifolium 3.0 0.8 + + -
9. Sanchi ginseng, I? pseudoginseng 11.3 8.7 tr ++++ tr + + +++ - ++++ - +

10. Panax japonicus 13.2 6.6 ++++ + ++ ++ +++ tr + + - ..
- - - - - - - -
11. Siberian ginseng, Acanthopanax senticosus N N - -
Plant leaves
3.3 - - - - - -
12. Panax trifolium 11.1 ++++ ++++ ++++ ++
13. American ginseng 13.8 6.6 tr +++ - + +++ ++ - + ++ +
continued
Y
Total
ginsenosides
(W/W%)* Relative amount of ginsenosidesc
Material wt Sp Ro Rbl Rbz Rc Rd Re Rf Rg, Rg2 Fll
Commercial products
14. Korean ginseng cigarettes
15. Korean tobacco cigarettes
16. Sliced Korean red ginseng root
17. Chinese Panax ginseng extract
18. Siberian ginseng extract
19. Siberian ginseng root tables
20. Wild American red ginseng root
(Rumex hymenosepalus)
“Adapted from Lui and Staba (1980).

*Total ginsenoside concentration (w/w%) determined by direct weight ( Wt) and spectrophotometrically (Sp). N, None detected.
‘tr, ‘Itace amount. The symbols ++++, +++, ++, + represent the surface area observed after spraying the chromatographed test sample
( 15 pl) and standard ( 10 pl); concentration ( 1 mg ginsenoside/ml) with anisaldehyde reagent. The standard ginsenosides reaction are
estimated as +.
6. GINSENG: INDUSTRY, BOTANY, AND CULTURE 193
powder. Toxicological studies using a standardized ginseng extract on

five kinds of animals showed no toxic effects, no adverse effects on
fertility, no carcinogenic activity, no sex hormone side effects, and no
cardiovascular side effects ( Soldati 1984).
B. Production and Tkade

Ginseng is cultivated in Korea, Japan, China, North America (United
States and Canada), and the Soviet Union. The book by Bae (1978)
includes several relevant chapters such as the history of ginseng produc-
tion (Cho 1978), ginseng cultivation (S.K. Hong 1978),and a history of
ginseng (M. W. Hong 1978). Staba and Kim (1971) have published a
survey of the history of American ginseng. The history of American
ginseng is short relative to that of Oriental ginseng.
Production statistics for ginseng for the above six countries are dif-
ficult to find. World production of ginseng in 1978 was estimated a t
3500 MT (Polczinski 1982). South Korea was the leading producer (2300
MT) and the leading exporter (1600 MT). China was the second largest
producer (600 MT), the United States third (200 MT) and Japan fourth
(125 MT).
Polczinski ( 1982)summarized export data from the United States from
1821 to 1979 and these are reproduced and updated in 'hble 6.3. Ginseng
production in the United States has fluctuated over the last 160 years
('hble 6.3). The drop-off in production around 1900 is thought to have
been due to severe outbreaks of Alternuria blight (Polczinski 1982). A
drop in production and returns in the 1930s is likely a reflection of the
economic depression (Hellyer 1984).This low value lasted through World
War 11. Direct U.S. trade with China ended in 1948 and since 1950
increased trade in ginseng has been through the Hong Kong market.
But and But (1984)have discussed the very vital role of Hong Kong in
ginseng marketing, and particularly that from the United States. The
presence of a very large population of Chinese origin, an experienced and
organized group of herb dealers, and Hong Kong's location and strong
finance and communication system are listed as some of the reasons for
its being a world center for ginseng trade. Changes in the herb industry,
the increasing use of western medications, the return of Hong Kong to
China in 1997, and the cultivation of American ginseng in China may
affect the prominence of Hong Kong. I n addition to Hong Kong, impor-
tant markets for U.S. ginseng in 1981 included 'hiwan and Singapore for
crude ginseng root, and Mexico, Costa Rica, and 'hiwan for processed
ginseng (Patty 1982). Important European markets for processed Korean
ginseng in 1979 and 1980 were West Germany, the United Kingdom, and
Italy (Patty 1982). South Korean ginseng has been promoted by the
Table 6.3. Export, Total Value, and Average Price of American Ginseng Dry Root
from 1821 to 1985"
Total exportedh Total value Average price

Year (MT) (U.S.$ thousands) per kg(U.S.$)
1821 160 171 1.06

1823 175 150 0.86
1845 212 177 0.81
1858 166 193 1.15
1862 286 408 1.85
1868 168 380 2.25
1878 191 497 2.49
1888 140 657 4.69
1890 101 605 5.97
1898 79 836 8.07
1900 73 833 11.46
1905 66 1,069 16.09
1915 47 919 19.64
1916 116 1,597 13.73
1920 73 1,875 25.81
1925 63 1,668 26.61
1929 106 2,766 26.06
1930 92 1,877 20.37
1933 106 844 18.08
1934 105 1,203 11.53
1938 76 1,029 13.56
1964 63 2,731 43.25
1965 53 2,887 54.50
1966 79 4,358 55.40
1967 66 4,507 67.99
1968 61 4,359 7 1.89
1969 66 5,533 83.91
1970 74 5,016 67.97
1971 77 5,827 76.08
1972 103 8,922 86.44
1973 83 8,846 106.48
1976 150 17,910 119.38
1977 173 26,530 153.33
1978 180 24,552 136.38
1979 161 22,216 137.96
1980 192 30,000 133.38
1981 224 35,200 152.93
1982 173 33,752 195.09
1983 185 28,733 155.31
1984 203 35,019 172.51
1985 473 42,086 88.98
"From Polczinski ( 1982) and G. Patty, personal communication.

bAdding 10% to the total exported column provides a n estimate of the total U.S.
production since the United States exports about 90% of its total ginseng production.
Office of Monopoly of the Republic of Korea and is sold worldwide.

Similar efforts by the American ginseng industry might lead to expan-
sion and development of new markets for its ginseng.
111. BOTANY
A. Taxonomy
Ginseng is placed in the genus Panax, Araliaceae, and is a dicotyledon.
According to Lawrence (1960),Araliaceae is placed in the order Umbelli-
floreae, whose main characteristics include flowers in simple or com-
pound determinate umbels (Fig. 6.1),floral parts simple or reduced and
epigenous, and two uniovulate carpels. Araliaceae has about 65 genera
and more than 800 species, most of which are tropical (Lawrence 1960).
The two major centers of distribution are the Indo-Malayan region and
tropical America. Each center usually has genera peculiar to itself. The
three major genera in North America are Araliu, Oplopanax (Echinopanax),
and Panax. The major economically important genera are English ivy
(Hedera helix, indigenous to Europe) and American Ginseng (Panax
Fig. 6.1. ( A ) Drawing of a 9-year-old wild-grown American ginseng plant (From

Lewis and Zenger 1982. Reproduced by permission of W. Lewis and the American
Journal of Botany). ( B )Photograph of a 4-year-old cultivated American ginseng plant.
quinquefolium). Lewis and Zenger (1982) have pleaded for use of the
correct adjective quinquefolium, rather than quinquefolius, which is
often used. Panax trifolium, dwarf ginseng, another ginseng indigenous
to North America may have potential in cultivation because it can
tolerate colder climates, stronger light, and wetter soil than P quinquefolium
(Hu et al. 1980).Inclusion through hybridization and selection of some or
all of these superior characters in the major cultivated species I!
quinquefolium and I! ginseng might overcome some of the major prob-
lems in world cultivation of ginseng (Section 1II.B).
The major Oriental and American ginseng species are listed in Thble
6.4. The number of Oriental ginseng species will vary depending on which
taxonomic treatment is preferred, but for simplicity, treatments by Hu
( 1976)and Lewis (1979)are presented. Other treatments are provided by
Hara (1970), Hoo and Tseng (1978), and Li (1942). Hu's authoritative
treatment includes a key to the Oriental species of Punax used in the
production of medicine for market and local uses. It also contains a listing
of 22 species of fleshy rooted plants, other than ginseng, which are used in
Chinese medicine. These species are from 12 genera in 7 families of
Dicotyledonae, the major family being the Campanulaceae.
The harvesting of indigenous American ginseng has been undertaken
since the eighteenth century. This and its export have raised concern that
the populations in the wild would disappear if indiscriminate harvesting
practices continued and no planned reseeding took place. Although Root
(1905) expressed such concern a t the beginning of this century, it has
only been in recent times that action was taken to protect diminishing
ginseng populations in the wild. The Convention on International n a d e
in Endangered Species (CITES)was ratified in 1973by several countries,
Table 6.4. Panax Species of North America and the Orient'
Botanical name Synonyms Geographical location
l? quinquefolium L. American ginseng, North America

Canadian ginseng,
sang, seng
l? trifolium L. Dwarf ginseng, North America
ground nut
l? ginseng C. A. Meyer Oriental ginseng North China, Korea,
Manchuria, North Japan
l? pseudoginseng Wall. Sanchi ginseng China
l? bipinna tifidum Seem Feather leaf, China
bamboo ginseng
l? japonicum C. A. Meyer Bamboo ginseng China
l? major (Burk.) Ting Big leaf sanchi China
"Based on Hu (1976) and Lewis i1979).

including the United States and Canada. American ginseng is listed in

Appendix I1 on CITES as a species that requires monitoring to ensure
that it does not become endangered. This has resulted in regulation of
trade in wild ginseng in the United States and Canada.
B. Genetics and Cytology

There are no cultivars of ginseng. Establishment and cultivation of
this crop was started by moving wild roots into protected gardens and by
selecting seeds from the plants growing in the wild. Seed from the
domesticated wild roots were used to expand plantings. This practice has
been continued with cultivated plants and still is the only attempt a t
choosing superior strains. Within production areas, e.g., North America,
there are claims that seed from Canadian grown plants are superior to
those grown in Wisconsin; also, the counterclaim is made. With an
essentially free movement of seed, these claims are unfounded. Generally,
the major species (Section 1II.A) have been confined to their centers of
origin. For example, l? quinquefoliumis the major species cultivated in
North America. Panax trifolium, the other species indigenous to North
America, may have potential in a breeding program for inclusion of
desirable characteristics such as its greater cold tolerance, stronger light
tolerance, and ability to grow in wetter soils.
1. Chromosome Number. Chromosomes in American ginseng vary in
length from 1.8 to 4.8 pm and can be arranged in five groups, based on
length, within this range (Blair 1975). n y l o r (1967)reported a chromo-
some number of 2n = 44 for American ginseng growing in Ontario,
Canada, whereas Blair (1975)reported 2n = 48 for Virginia specimens.
Dwarf ginseng, l? trifolium, growing in Massachusetts was found to be a
diploid, 2n = 24, and l? quinquefolium a tetraploid, 2n = 48 (Huet al., 1980).
Chromosome counts for Oriental ginseng, l? ginseng, are 2n = 44
(Kurosawa 1966) and 2n = 48 (Harn and Whang 1963),giving the same
number discrepancy reported for American ginseng. The basic ( x )chro-
mosome number in the Araliaceae varies from 9 to 13, with 12 as the most
frequent (Moore 1971). More detailed cytological studies of ginseng,
particularly the Oriental species ( B b l e 6 . 4 ) ,would be of great interest.
2. Breeding Difficulties. Problems associated with breeding ginseng
stem from the long reproductive and production cycles and difficulties
generally associated with interspecific cross breeding. The inflorescence is
an umbel (Section III.C.B), the seed requires about 20 months before it
will germinate (Section 1V.A.1 ), and the plant is a perennial requiring
about four growing seasons before yield can be assessed. Seed production
is also slow, with 3 to 4 years elapsing between seeding and the collection
of ripe seeds. Also, as Choi et al. ( 1984)have pointed out, “development of

viable seeds in most interspecific crosses of ginseng has been prevented
because of specific inhibition or elimination of key steps in pollination,
pollen tube growth, fertilization,and embryo or endosperm developments.”
Lewis and Zenger (1983)observed major diversity in the floral biology,
breeding, and fecundity among two North Americafi populations of
ginseng growing in the wild in Missouri and New York.
3. Breeding Objectives. When the problems of breeding ginseng have
been reduced or overcome, then it will be appropriate to address specific
objectives in the breeding program. These might include reduced seed
stratification time, increased yield, higher quality roots, and disease
resistance. We know little about how to address any of these objectives
with this crop. The evaluation of yield depends on various factors exter-
nal to the plant including light, air and soil temperature, soil moisture,
and mulch (Section 1V.C).Yield will also depend on a complex of plant
characters including root size, hardiness (Section IV.C.3), and disease
resistance (Section 1V.D).
Root rot is the limiting factor in world ginseng production (Section
1V.D). Alternaria leaf blight is also a very serious problem. We know
nothing about plant resistance to these diseases, or even if it exists.
C. Morphology, Anatomy, Growth, and Development

There is no botanical key for all Punax species although a key is
available for some of the Oriental species (Hu 1976). The following
descriptions are general with specific characteristics noted for some
species.
1. Leaves. The deciduous aerial stem of mature ginseng has whorled
leaves a t its summit. These are known as prongs and each has a petiole
with three to five palmately compound leaves (Figs. 6.1 and 6.2). As
Lewis and Zenger (1982)have pointed out, these prongs are not strictly
equivalent to leaves since they lack axillary buds a t their junctures with
aerial stems. Mature plants usually have five leaflets per leaf, hence
quinquefolium. The two outer smaller leaflets (1and 5 in Fig. 6.2) are oval
to suborbicular in shape with a round base and an acuminate apex
(Hughes and Proctor 1981).The three larger leaves (2,3,and 4 in Fig. 6.2)
are obovate-oblong to obovate with a round to acute base and an acumi-
nate apex. Both leaflet types have serrated margins. Seedlings have three
leaflets. After the first year, these young plants develop four or five
leaflets and one or more prongs. After 4 years most of the plants under
commercial cultivation will have four prongs. Leaflet number and prong
number are correlated with age (Lewisand Zenger 1982))although growth
of native (wild)populations is much slower than growth in cultivation.
Fig. 6.2. A mature ginseng leaf showing the leaflet arrangement. Leaflets are
numbered consecutively clockwise from the petiole to aid discussion in the text.
(From Hughes and Proctor 1981.)
For example, three-pronged plants in native populations averaged 13.5 *

3.3 years before adding a fourth prong (Lewis and Zenger 1982). Maxi-
mum aerial growth of ginseng is reached when the plant has five prongs,
although Hu (1976)has reported six prongs for l? ginseng.
Leaflets of ginseng are green, although we have observed silver leaflet
American ginseng, which may be a mutant. Leaflet shape of ginseng is
variable, but very difficult to compare in situ in the field. The only
attempt we know of to describe differences in ginseng leaf shape was by
Choi and Shin (1982), who categorized leaflets into seven groups and
showed representative drawings of each. The majority fell into the follow-
ing three equal-sized groups:
(1) Three larger leaflets (2, 3, and 4 in Fig. 6.2) were oblong-ovate,
serrate, with acute tip. The length of these leaflets in this group
200 J. T. A. PROCTOR AND W.G. BAILEY
was greater than in other types. The two basal leaflets ( 1and 5 in
Fig. 6.2) were small, serrate, and elliptic.
( 2 ) Three larger leaflets were round-ovate, serrate with a dull tip. The
width of these leaflets in this group was greater than in the other
types. The two basal leaflets were serrate, small, and round-ovate.
( 3 ) Three larger leaflets were dull spear-type-ovate,serrate, with acute
tip and rough margin. The basal two leaflets were small, elliptic,
and serrate.
Stomata1 frequency was greater in leaves of €? quinquefolium (35-43

mm-’) than in leaves of €? ginseng var. atropurpureacaulo (15-26 mm-’)
(Park 1980)in plants grownin Korea. Greenhouse studies of I?quinquefolium
in Canada gave stomata1 frequencies of 40-136 mm-’ (J.T. A. Proctor and
P. Miller, unpublished data). Photographs of stomates of American gin-
seng are shown in Fig. 6.3.
2. Inflorescence. Inflorescence morphology is subject to variation

and a range of inflorescence types occurs within the Araliaceae ( Philipson
1970) where the umbel is characteristic. Choi and Shin (1982) have
described six shapes of €? ginseng inflorescences. The two most com-
mon are a complete hemispherical umbel with all pedicels of the same
length, and a simple umbel in which some pedicels are longer than others
and project outside the hemisphere. Inflorescence morphology was less
variable in cultivated American ginseng with only two shapes: a com-
plete hemispherical umbel with all pedicels of the same length, and a
simple umbel with several branched pedicels below it on the peduncle
(Proctor 1986).
Hu (1980) reported branched inflorescences in €? japonicus and in €?
quinquefolium growing in the wild. However, in the latter case, the result
of branching was not the formation of normal panicles of umbels as is
found in €? japonicus and €? pseudoginseng. Rather a single flower or a
small umbel may diverge from the base of the peduncle, or an umbellet
may arise from the center of the major umbel.
3. Flowers. The nature of the flowers in ginseng appears to vary with

species (Hu 1980).Carpenter and Cottam (1982)reported that American
ginseng grown in the wild had perfect flowers and each flower had mature
anthers or pistils, but not both. In addition, each inflorescence contained
both functionally staminate and pistillate flowers and the flowers were
closely packed on the inflorescence. American ginseng flowers are small,
greenish-white, and pentamerous (Lewis and Zenger 1982). Hu ( 1976)
has provided a more detailed description for 19 ginseng flowers: “small,
Fig. 6.3. Scanning electron micrographs of stomates of American ginseng. ( A )

adaxial leaf surface ( X 820); ( B )stomate enlargement ( X 3240). (J. T. A. Proctor and
P. Miller, unpublished.)
2-3 mm across; sepals 5, green; petals 5, cream yellow, ovate, apex

obtuse; stamens 5, filaments, short; pistil 1, ovary inferior, 2 locular;
styles 2, united a t base; disk cup-shaped.’’
4. Pollination and Fruit Set. Cultivated Oriental ginseng is self-

pollinating with a success rate of about 90% among bagged inflores-
cences (Bae 1978).Pollination occurs between flowers within an inflorescence,
or between plants if pollinators are present (Carpenter and Cottam 19821.
Fruit and seed production does not differ in bagged and unbagged plants
in some populations (Carpenter and Cottam 1982; Lewis and Zenger
1983), but does in others (Lewis and Zenger 1983). Flowers are not
apomictic (Carpenter and Cottam 1982; Lewis and Zenger 1983).
Sweat-bees, particularly Dialictus sp. and Evylaeus sp. (Halictidae),
appear to be the major pollinators of American ginseng (Carpenter and

Cottam 1982; Duke 1980; Lewis and Zenger 1983). These generalist
pollinators probably do not transfer pollen between distant individuals,
and so American ginseng is not an obligate outcrosser (Carpenter and
Cottam 1982).
Little is known about fruit set and development in ginseng, although
seed yield in some years has been worth more than root yield. Information
about fruit set seems to be restricted to plants growing in the wild. For
instance, Lewis and Zenger (1982) reported 80-89% of the flowering
plants set fruit except in a drought year, when only 47% of the flowering
plants produced fruit. Lewis and Zenger ( 1982)attributed lack of fruit set
in flowering plants to abortion following fertilization, incomplete fruit
development due to environmental or other factors, and sterility because
of poor or no pollination. Schlessman (1985)showed that prefertilization
abortion of ovules in one-styled flowers also affected seed production.
Each flower has two locules, and so the usual seed yield is two per fruit
although one and three seeds are often found. Stoltz and Garland (1980)
found that fruit were one-seeded (16.3%),commonly two-seeded (77.0%),
and infrequently three-seeded (6.5%) or four-seeded (0.29’0).A typical
flowering 4-year-old American ginseng plant may carry 30-40 berries in
each inflorescence with an average of two cream white seeds, 5-6 mm long
and 4-5 mm wide, in each berry.
5. Stem. Most ginseng plants have a solitary stem. Multiple stems

from two or more buds formed in the previous year are rare. In a harvest of
5-year-oldcultivated plants in 1983 in Ontario, we found that 85% had a
single stem, 13% had two stems, and 2% had three stems (J. C. Lee and
J. T. A. Proctor, unpublished). In a 1985 harvest of 3-year-oldsin Ontario,
93% had a single stem and 7% had two stems (J.T. A. Proctor, unpublished).
At the base of the stem is a gnarled rhizome (Fig. 6.1), which grows
horizontally or erect. In cultivated plants the rhizome is usually erect
(Fig. 6.1B) and only a few years old. In plants grown in the wild it can be
much older (>50 years; Lewis and Zenger 1982), variable in size, often
horizontal, and sometimes branched (Hu 1976). A main characteristic of
the rhizome is the scars that form as a result of the annual abscission
of the aerial stem. These annual scars on the rhizome allow an estimation
of plant age.
6. Root. The root is a fleshy taproot, often with two to five laterals. It
is light yellowish white in color and narrows apically to the rhizome (Fig.
6.1). The dried root is the economically important part of the plant. Root
dry matter is about 30%, but this varies with age; 1-year-oldswere found
to have 24.9%,2-year-olds26.5%, 3-year-olds28.6%, and 4-year-olds30.9%
(J. T. A. Proctor, unpublished). Kim et al. (1981) reported 25.9% dry

matter for 4-year-old I! ginseng roots harvested in September/October.
Within any one year the proportions of water and dry matter probably
vary with nutrition and environmental factors, particularly water supply,
as has been shown for potato (Burton 1966).In E! ginseng 2-year-oldroot
fresh weight had an inverse relationship with water content (Mork et al.
1981). Roots grown under water-limiting conditions had higher water
contents with an overall range of 55-85%.
The root contains chemical constituents that are claimed to be respon-
sible for the therapeutic effects of ginseng (Section 1I.A 1. These compo-
nents were first isolated from American ginseng in 1854 (Bae 1978) and
identified in the 1960s as saponin glycosides or ginsenosides (Elyakov
and Strigina 1962; Elyakov et al. 1962). Histochemical tests have shown
that these ginsenosides are located in the root outside the cambium (Fig.
6.4) in oil glands.
7. Growth and Development. Data for plants under cultivation are

sparse, and so the description below follows that of Carpenter and Cottam
co
ca
Fig. 6.4. If-ansverse section of a root of American ginseng ( X 43) showing the
epidermis (el, cortex (co),resin canal ( r ) ,phloem ( p ) ,cambium (ca),and xylem ( x ) ,
primary ( p r )and secondary (se).(Prepared by A . Tsai.)
( 1982)for plants grown in the wild with inclusion of comparable available

data for cultivated plants.
Plants emerge in late April-early May and grow to full height and leaf
size in about one month. Leaf area per plant was about 1100 mm2 in
seedlings and increased to about 35,700 mm2 in the fourth-year plants
(Hughesand Proctor 1981). Leaf area index in a commercial planting was
0.17 in year 1and 1.30 in year 2, but increased dramatically to 4.80 in year
3 and 6.78 in year 4 (Hughes and Proctor 1981).
Leaf growth did not occur a t temperatures below 5OC (Lee et al. 1985).
Above this temperature leaflet length and width of 3-year-old plants
grown in a growth chamber a t a continuous loo, 15O, 20°, and 23OC was
not influenced by temperature (Lee et al. 1986). In contrast, in green-
house root-zone warming experiments using soil temperatures of 13O,
1B0, and 23OC, seedling leaflet length was greatest a t 1B0C, but leaflet
width was not affected by temperature (Lee et al. 1986).
Root weight increased with plant age and was influenced by plant
spacing (Konsler and Shelton 1984) and, in Oriental production, by row
location on the bed (Fig. 6.5). Roots spaced very closely in the row (25
mm) increased only slightly in the second to fifth years and did not
increase between the fifth and sixth years (Fig. 6.5A, curve a). Roots at
the widest spacing in the row (229mm) increased in weight a t a constant
rate between the second and fifth years, but between the fifth and sixth
years this rate was somewhat reduced (Fig. 6.5A, curve d). In Oriental
production, root weight increase was greatest in rows 1 and 2 (Fig. 6.5B,
curves 1 and 2), which is a reflection of the greater light interception by
these plants (Sections 1V.C and IV.B.4). The data in Fig. 6.5 indicate
greater individual root weights in Oriental production than in North
American production in spite of greater yields per hectare in North
America (Patty 1982).
In reproductive plants (usually with three prongs) the flower buds are
small and clustered tightly on a short peduncle (Fig. 6.1) as leaves
expand. Toward the end of leaf expansion and into early June the pedun-
cle elongates and flower buds enlarge. Flower opening in the terminal
hemispherical inflorescence, the umbel, usually starts in the outermost
part in early June. The flowering period is usually about 6 weeks, although
Lewis and Zenger ( 1983)have reported 8 weeks for plants in Missouri and
3 weeks for plants in New York. Similarly, the number of flowers open per
plant can vary from one to three or as many as eight, and seems to be
related to the length of the flowering period. Shortly after the flower buds
open, the stamens and petals dehisce. Fruit are set between late June and
late July and grow and develop until mid-August. The new terminal bud
on the rhizome is initiated in July and enlarges during the remainder of
the growing season.
40
-- 30
t
0
e
-+
0
I
p 20
3
>
E
n
5
2
10
n I I I I I
2 3 4 5 6
YEAR
40
--
30 -
c
e
-+
0
I
p 20 -
3
zn
5
2
10 -
0 &
I I I I I
Fig. 6.5. Individual root dry weight. ( A )R quinquefolium. Rows were 152 mm apart
and plants spaced a t 25 ( a ) ,76 ( b ) , 152 (c), and 229 ( d )mm apart. (From Konsler and
Shelton 1984.) ( B ) R ginseng. The numbers denote the row number in the bed,
numbered from the front row. (From Kim 1964a.)
Fruit ripening, indicated by the exocarps turning red, first in the

outermost fruit of the umbel and progressing to the center, usually starts
about mid-August. Fruit are picked when all exocarps in the umbel are
red and prior to abscission, probably in early September. Yellow and
orange-yellowberried mutants of Korean ginseng have been found and are
being studied by the Korean Ginseng Research Institute (Bae 1978; Park
1980).Leaf color also starts to change from green to yellow to rust-colored
in August with a general senescence of the aboveground parts completed
in October, leaving the rhizome to perennate.
IV. CULTURE
There are essentially two different ways of growing ginseng: woods

grown and artificial shade grown. Woods grown (Lewis 1980)refers to the
use of the natural forest canopy for shade. The associated cultural tech-
niques can vary from simulating forest conditions to practices similar to
those used in cultivating ginseng under artificial shade (Curran 1983;
Currie 1980; Williams and Duke 1978).Artificial shade growing is the most
common worldwide (Fig.6.6). S. K. Hong ( 1978)has reviewed cultivation
methods in Korea, Japan, China, North America, and the Soviet Union.
The common features in both cultural methods are shade (natural or
artificial), a mulch (natural leaf fall, applied straw, or other suitable
material), and the growing of the plants on raised beds. The plants are
obtained either by direct seeding or by seeding into a nursery bed and
transplanting roots after one year of growth to a growing bed. Once the
crop is established there is need each year for weed control and spraying
for diseases and insects. The crop is dug by hand or mechanically with a
modified potato digger usually after three or four growing seasons.
In Ontario and Wisconsin, the roots are harvested after growth has
ceased (lateAugust/early September) and the tops removed or died down
(Curran 1983; Currie 1980). The ginseng extract quality was higher for
the summer harvests in Korea, China, and Japan than for the later
autumn harvests. For example, Kim et al. (1981) found that summer
harvested roots had >20% saponin concentration, whereas late autumn
harvested roots had <lo%.
A. Propagation
1. Seed. Seeding is the principal method of propagating ginseng.
Unfortunately, most of the papers on this topic are in Japanese and
Russian. Baronov ( 1966)has cited some of these papers and reported that
B
Fig. 6.6. Comparison of shade materials used in the Korean

culture of ginseng ( A )with t h a t used in North America (B,C).
( A ) Impermeable thatch shade of straw or reeds used in
Korea. ( B ) 'Ikaditional wooden lath screens used in North
America since about 1890. ( C ) Woven black polypropylene
shade t h a t has come into more extensive use in the last decade
in North America.
N
s C
208 1. T. A. PROCTOR AND W. G. BAILEY
some seeds may germinate as early as 8 months following dissemination

and others may germinate only after prolonged periods in the soil. Lewis
and Zenger ( 1982) recognized the seed afterripening and germination
period as a highly vulnerable stage in the life cycle of ginseng growing in
the wild because only 0.6% of the seeds germinated after 20 months.
Generally, afterripening of 18-22 months is required for Panax seeds
before germination.
In North America seeds are afterripened in stratification beds out-
doors after they are collected in August/September (Polczinski 1982).
They will not germinate and grow until the second spring following
harvest (the 18-22 months referred to above). Splitting of the endocarp
has usually occurred after 12 months of stratification. This is the time
when direct seeding takes place. Peatment of the endocarp with sodium
hydroxide increases splitting and stimulates embryo growth (Lee et al.
1983,1984).Growers have observed that in some cases, particularly when
high concentrations of captan were used as a seed dressing, seed germina-
tion was delayed or inhibited (Curran 1983). Possibly the captan has an
adverse effect on the microflora, which are necessary for the weakening of
the endocarp prior to splitting (J. C. Lee, personal communication).
When ginseng seeds are harvested they have an immature embryo that
grows during the subsequent afterripening period (Lee et al. 1983; Stoltz
and Garland 1980; Stoltz and Snyder 1985).A fluctuating temperature
regime of cool-warm-cool temperatures over the 18-22 months stratifica-
tion were reported necessary for embryo growth and maturation and
germination of seeds (Lee et al. 1983; Stoltz and Snyder 1985).Stoltz and
Snyder (1985)suggested that the second cool period is needed to release
the embryo from an endogenous dormancy.
2. Tissue Culture. Tissue culture of P ginseng was attempted by

Russian scientists in 1967 (Slepyan et al. 1967).Since then ginseng callus
and organized structure have been successfully grown from various tis-
sues including leaves (Butenko et al. 1968; Choi et al. 1982a;Jhang et al.
1974),cotyledons (Choi et al. 198l),stem (Butenko et al. 1968; Jhang et
al. 1974),and roots (Butenkoet al. 1968; Chang and Hsing 1980a; Choi et
al. 1981, 1982a; Jhang et al. 1974). Plant regeneration through somatic
embryogenesis has produced embryoids from root-derived callus (Chang
and Hsing 1980a,b; Choi et al. 1982a).However, it took most of a year to
get embryoids and shoots and they were unable to induce plantlets
similar to ginseng seedlings. Attempts to obtain ginseng plant regenera-
tion from protoplasts have been initiated and viable protoplasts isolated
from root tissue and callus (Choi et al. 1984).
Cultures of ginseng produced the same kinds of saponins as intact
plants, but only about 0.4% of their weight in extractable saponins,
which is about ten times less than the dried roots (Jhang et al. 1974;
Soldati and Tanaka 1984). The various media and their components
including nutrients and growth regulators are given in the above-listed
references. Inclusion of ginseng powder in the culture medium for
Peperomia viridis consistently increased the plantlet regeneration per-
centage, the number of shoots formed per leaf explants, and increased
plantlet quality (Hui and Zee 1981).
In summary, although tissue culture of ginseng has been studied for
about 20 years little progress has been made in propagation and in vitro
ginsenoside production. We would challenge the optimistic projection by
Fulder ( 1980)that “it will not be too long before the production of ginseng
saponin becomes a purely industrial matter without any need for dirtying
hands by growing the root.”
B. Soils and Nutrition

1. Soils. “By proper treatment, almost any fairly good soil can be
conditioned for ginseng growing” (Williams and Duke 1978).This quota-
tion suggests considerable flexibility in selecting a suitable soil for grow-
ing ginseng but appropriate research to support or refute this statement
does not exist. There have been a number of surveys of soil types used by
ginseng growers in Kentucky (Roberts and Richardson 1980),in Wiscon-
sin (Polczinski 1982), and in Arkansas (Fountain 1982). Roberts and
Richardson (1980)found that the soils in which ginseng was growing had
a wide pH range-from 4.55 to 7.44. The soil test readings showed that in
61 of the 64 plots the phosphorus was low, and the potassium was
medium or high.
Polczinski (1982) showed that the major soils in which ginseng was
growing were Marathon silt loam, Moberg gravelly loam, Fenwood silt
loam, Rozellville silt loam, and Chetek sandy loam. In general, these
soils are rich in humus, have textures ranging from silt loam to sandy
loam, are underlain by porous subsoils, and are well drained. Further, the
gentle sloping gardens had slopes of 2-15% with the majority in the range
of 2-690 slope. Polczinski’s findings (1982)are in agreement with the site
selection criteria set out by Root (1905), Harding (1972), and Lewis
(1980).
Fountain’s (1982) survey found that the typical soil was a moderate
clay soil with moderate to high fertility. The pH range was wide-from
4.6 to 6.8. The typical locations of the natural populations were on slopes
of gentle to moderate steepness and a t well-drained midslope positions.
2. Nutrition. Konsler and Shelton ( 1984)found that oak bark/sawdust
mulch was the most active of six tested in affecting soil status with
relatively higher levels than most mulches in soil pH, organic matter,
phosphorus, potassium, calcium, and magnesium and a tendency toward

lower levels of manganese, sulphur, and zinc. In earlier work, Konsler
( 1979, 1980) reported that lime and phosphorus applications increased
root weight, and soil pH of 5.5 was superior to pH 4.4or 6.5.
Stoltz (1982)studied the effects of several nutrient deficiencies on the
expression of leaf symptoms and on fresh weight of roots using sand
culture. Foliar deficiency symptoms of calcium, including collapse of
5-10 mm of the tissue a t the tip of leaflets, occurred first, followed by
those for magnesium and phosphorus. Nitrogen deficiency symptoms
occurred very gradually. Root fresh weight gain was most restricted by
omission of calcium, phosphorus, or magnesium from the nutrient solution.
An accurate assessment of the ginseng plant nutrient status is essen-
tial for an effective, efficient fertilization program. Information about soil
types and soil testing for ginseng is very limited. Similarly, plant analysis
data are lacking. Lee et al. (1980b)investigated the distribution pattern
of mineral elements in different parts of 5-year-old i? ginseng plants
grown under four different nutritional regimes. In the roots nitrogen and
phosphorus showed a positive correlation ( r = 0.77) with each other and
with magnesium and copper, while all other elements (potassium, calcium,
magnesium, iron, manganese, zinc, copper, and boron) were highly corre-
lated with each other. The number of nutrient pairs having significant
correlations was much less in shoots than in the root.
In Konsler and Shelton’s( 1984)mulch experiments with P quinquefoliurn,
hardwood leaf mulch gave very high magnesium and pine needles low
magnesium in leaves of 1-year-old ginseng. Straw induced low manga-
nese, poplar bark/sawdust low zinc, and pine needles low iron in the
foliage. The mineral content of 1-year-oldroots was more stable than in
the leaves, and the levels of calcium, magnesium, manganese, and zinc
were much lower than in the leaves. Nutrient element content of 6-year-old
roots was of the same order of concentration as in 1-year-old roots,
although nitrogen, potassium, and iron were appreciably lower and cal-
cium higher than in the older roots. There was little correlation between
content of mineral elements in soil and in the roots except for zinc. Root
size at 6 years was positively correlated with soil pH, potassium, calcium,
and magnesium. Phosphorus, calcium, and magnesium in tissue were
positively correlated, and manganese and zinc negatively correlated with
root size.
Foliar analysis of ginseng and its application to fertilizer recommenda-
tions is just now beginning. Khwaja et al. (1984) reported the leaf
concentration of 11 elements and suggested their relationship to visual
deficiencies and plant growth (‘hble 6.5). These data are based on Sam-
ples from commercial growers and not from designed experiments.
Table 6.5. Ranges of Nutrient Concentrations in Mature Leaves of 2- to 7-year-old

American Ginseng Plants (Based on 35 Analyses)"
Foliar element ranges
Element Deficiency* Low' Sufficiencyd High' Excessive'
<1.50 1.51-3.0 3.01-5.0 5.1-7.5 >7.5

<0.20 0.20-0.30 0.31-0.95 0.96-1.5 >1.5
<1.5 1.5-3.0 3.1-5.0 5.1-7.5 >7.5
<0.2 0.21-0.50 0.51-1.0 >1.0
<0.10 0.11-0.25 0.26-2.5 >2.5
<0.11 0.12-0.2 1 0.22-0.55 >0.55
<15 16-25 26-75 76-150 >150
<10 11-25 26-35 36-55 >55
<25 26-45 46-175 176-350 >350
- <40 41-375 476-500 >500
<5.0 5.0-8.0 8.1-35 36-50 >50
"From Khwaja et al. (1984).

bPlants with visible symptoms of deficiency.
cPlants with normal appearance but likely to respond to fertilization with this
element.
dPlants contain adequate amount of this element for maximum yield.
eOptimum yield can be expected; however, concentration of this element is higher
than normal.
'Nutrient level too high; plants may show nutritional disorder or stunted growth.
3. Soils, Plant Nutrition, and Diseases. Ohh (1981)has reviewed this

topic and reported that the outbreak of sclerotinia was most severe at pH
4.7, of root rot at pH 6.5-7.5, and red rot at pH 6.0-6.5. Both root rot and
red rot were increased with high applications of nitrogen.
Controlled studies using a combination of soil testing and foliar anal-
yses for assessing ginseng nutritional status and its relationship to
root yield and quality are needed. Such testing and analysis must be
done using controlled, uniform procedures with all possible sources of
error known.
C. Environmental Physiology
Both I? ginseng and I? quinquefolium were originally harvested from
the wild and the scarcenessof the wild root stimulated cultivation. Detailed
research on the growing environments of ginseng has only recently
received consideration. Daditionally, decisions about the growth and
management of the crop have been based primarily on practical experi-
ence with little organized research applied to ginseng culture ( Hartman
1979). The art of ginseng growing was empirical, with growing secrets
passed on from generation to generation. This has led to confusing and
sometimes conflicting information about the environmental resources
required for optimal ginseng production.
Ginseng requires very specific enviromental conditions to survive and
grow well. The native environment of this perennial herbaceous root crop
is in the understory of deciduous and mixed forests in eastern Asia and
eastern North America. It normally grows where diffuse solar radiation
and adequate soil moisture reserves occur during the summer months.
The natural tree cover shades the plants from excessive solar radiation.
The lack of shade will result in leaf necrosis in a few days and, over
extended periods, plant death. The leaf mulch under the forest canopy
plays an important role in the survival of ginseng as it aids in conserva-
tion of soil moisture during dry periods in the summer and provides
protection for roots from excessively low soil temperatures during the
winter season.
Commercial growers of ginseng attempt to emulate the environmental
conditions of the forest in their ginseng gardens. In Korea, Japan and
China, there has been a long history in the development of growing
practices (M. W. Hong 1978). The cultivation technique employed in
these countries involves the use of both nursery and mature plant beds.
The beds are covered by an elevated canopy of a thatched material such as
woven rice straw. Wherever possible the gardens face the northeast to
minimize solar radiation (M. W. Hong 1978). Blinds can be employed at
the front (north side) to minimize solar radiation, as well as to control air
flow and the temperature and humidity regime. In addition, side blinds
can be employed a t the east, south (back or rear), and west sides.
Recent Korean literature (Choi et al. 1982b; Lee et al. 1980a; Park 1980)
has recognized the limitations of this traditional growing environment.
As a consequence of the nature of the canopy structure, solar irradiance
levels in the back of the beds are too low for optimal photosynthesis. Lee
et al. (1980a) reported that the relative light itensity in a traditional
five-row planting was 8-970in the front rows, 2-3% in the middle rows, and
1-2% in the back row. Use of different shade material (pine board, styro-
foam board, and polytex fabric)and an improved structure design resulted
in higher solar radiation levels, particularly in the rear rows of the beds,
and increased root yields.
In North America, both woods-grown (Jenkins 1980) and artificial-
shade (Currie 1980) cultivation techniques are employed. Much of the
literature available on the culture of I? quinquefolium (Currie 1980;
Hartman 1979; Jenkins 1980; Williams and Duke 1978),although some-
what explicit on the cultivation methodology, is quite limited in defining
the environmental conditions for the successful growth and development
of the plant and in the assessment of the potential of the crop for new
production areas. Only recently has research been directed to overcoming
this dearth of information.
1. Climate Resources. a. Radiation, energy, and water balances. A
thorough understanding of the modified growing environment for gin-
seng will aid in the optimization of its production. The development of a
theoretical treatment of the radiation and energy balances of this envi-
ronment has been derived by Stathers and Bailey (1986).The solar and
longwave radiation balances above and below a suspended polypropylene
shade fabric are illustrated in Figs. 6.7 and 6.8. An analogous treatment
can be developed for wooden lath shade and has been presented for the
traditional Korean shade environment by Kim (196413).From this work it
is recognized that a shade canopy significantly influences energy receipt
a t the ground surface by absorbing and reflecting a large portion of the
incoming radiation. Only the radiation that is transmitted through the
canopy or is emitted from the canopy influences the radiation regime
beneath the canopy.
Proctor (1980) presented a summary of radiation data collected for
polypropylene shade and wooden lath environments (Fig.6.6).The results
Fig. 6.7. Schematic of solar radiation balance for a ginseng garden. (From Stathers
and Bailey 1986. )
4
I
1tbl ;
I
1
'; 14.1 t <.ltb'
I
I
I
SHADE CANOPY
I
I
I LEGEND: 1' = net longwave rodtotion
lll I ll = incoming longwove radiotion
I
I "b l t = outgoing longwove rodiation
ltb
I I = tronrrnirsivilyof shadecanopy
l o longwove rodiotaon
I
I SUBSCRIPTS: D refers to above canopy
I c referr lo conopy
I b refers lo below canopy
I
I r l t +11
l*b= -Itb
I
I
GARDEN SURFACE
I
Fig. 6.8. Schematic of longwave radiation balance for a ginseng garden. (From
Stathers and Bailey 1986.)
are summarized in Thble 6.6. The wooden lath shade, a traditional type of
shade in North America, had a surface area of 70% and permitted
radiation penetration of only 18%.This discrepancy (12%) is accounted
for by the interception of solar radiation by the width of the wooden lath
material (thickness 9.5 mm). The polypropylene had an estimated sur-
face area of 72% and the irradiance data confirm this. I t is noted that the
net radiation for the wooden lath is less than that for the polypropylene
above the canopy. This is likely accounted for by the higher albedo of the
wood surface and the greater longwave radiation losses from the wood
'bble 6.6.
Solar and Net Radiation Flux Density Dataa for Seven Days in July
and August 1977 for Black Polypropylene and Wooden Lath Shadeb
Solar radiation
Location Total Diffuse Net radiation

~
Above shade 100 33.6 100.0(polypropylene)

77.2 (wood)
Under shade
wooden lath 16.4 6.6 21.9
black polypropylene 26.6 11.4 23.2
aData are expressed as a percentage of the radiation above the shades.

'From Proctor (1980).
due to its slightly higher surface temperature. As a consequence of its

high heat capacity and its low mass, the temperature of the polypropyl-
ene is quite similar to the air temperature.
Figure 6.9 presents the solar and net radiation both above and beneath
a black polypropylene shade canopy at Lytt.cn, British Columbia ( Stathers
and Bailey 1986)that had an estimated surface area of 78%. The differences
r
between the trends of solar and net radiation are accounted for by the net
longwave exchanges. As is evident from this figure, the net longwave
radiation beneath the canopy is small. This results from the temperatures
of the shade canopy, air, and garden surface being quite similar.
-
-
K 1 above canopy
-
'O0O K below canopy
Q above canopy
900 Q ' below canopy
0 2 4 6 8 10 12 14 16 18 20 22 2
TIME (h, PST)
Fig. 6.9. The diurnal course of solar ( K J ) and net ( & * ) radiation above and below a
black polypropylene shade canopy on August 1, 1984, at Lytton, British Columbia.
(From Stathers and Bailey 1986.)
The consequences of this microclimate modification must affect both

the thermal and moisture regimes of the ginseng garden. Figure 6.10
illustrates the temperature regime of the Lytton ginseng garden during
the extremes of day and night. The soil temperature regime is effectively
separated from the atmosphere even with air temperatures exceeding
3OoC. Bble 6.7 illustrates a comparison between the temperature regimes
of wooden lath and black polypropylene shade canopy environments. The
differences are minimal during the day for this study in southern Ontario
(Proctor 1980). The temperature beneath the canopy in Ontario was
cooler than that above the canopy environment a t night. For the Lytton
data, the opposite was quite marked. These differences merit further
study, particularly as it affects potential spring frost events. The differ-
ences found likely reflect differences in the general thermal environments
and ventilation in the two locales.
The significant microclimate modifications in ginseng gardens were
reflected in the seasonal trend of soil temperature and moisture (Figs.
6.11 and 6.12). The soil temperature was significantly lower in the gin-
3.0- 11
SHADE CANOPY
2.0.
Temperature profile Temperature profile

for OSOOh PST for 12OOh PST
1.0-
-
v
E PLANT CANOPY
.
I
I- 7 - -
-
I
(30
Lu
A
I STRAW MULCH
SOIL
-1.0-
I I I I I I I I I I I I
10 12 14 16 18 20 22 24 26 28 30 32
Fig. 6.10. Temperature profiles above and below a black polypropylene shade can-
opy during day and night on August 2, 1984, a t Lytton, British Columbia. (From
Stathers and Bailey 1984.)
6. GINSENG: INDUSTRY. BOTANY, AND CULTURE 217
Table 6.7. Mean Air Temperatures above and below a Wooden Lath and Black
Polypropylene Shade for Days (0800 to 1900) and Nights (1900 to 0800)
for the Period 1630 on July 27, 1977, to 1530 on July 29, 1977"
Mean temperatures (OC)
D aY Night
Shade Location Air Leaf Air Leaf
Woven black polypropylene Above 23.1 - 14.0 -
Below 23.3 24.2 12.8 12.6

Wooden lath Above 23.7 - 13.4 -
Below 24.1 23.7 12.6 13.4
"From Proctor (1980).
seng garden when compared with the unaltered environment (native

pasture) (Figure 6.11) and the soil moisture regime in the ginseng garden
was kept near field capacity throughout the growing season (Fig. 6.12).
The shade canopy and mulch effectively separated the soil from the
30
-
u
h
- NATIVE
v
w
E - -
E 20-
2 --
w
c
=!
g -
GINSENG
10
GARDEN
0 I I I I I
APR. ' I l l ) )
MAY ' 1 1 1 1 1
JUNE
DATE
' , I ) ) I
JULY ' 1 1 1 1 1
ALJG. ' I ( I I I
SEPT.
Fig. 6.11. The trend of soil temperature a t a depth of 0.15 m during the 1984 growing
season a t Lytton, British Columbia in a native pasture and in a second year ginseng
garden. (W. G . Bailey, unpublished data.)
Fig. 6.12. The trend of volumetric soil moisture for depth 0-0.25 m during the 1984
growing season at Lytton, British Columbia in a second year ginseng garden and in a
native pasture. (W. G. Bailey, unpublished data.)
atmospheric environment. This is even more dramatic when one consid-

ers that air temperatures routinely exceed 4OoC during the summer, and
over the period April 1 to October 1, only 90 mm of precipitation were
received (no irrigation was applied). Clearly the microclimate modifica-
tion conserves soil water and moderates soil temperature. This results in
an environment where soil water is readily available to the growing
plants. However, stressful conditions also arise in Lytton, as leaf temper-
atures can exceed 4OoC whereas the root is usually at less than 15OC. In
addition, the large temperature gradients from the top of the mulch into
the soil when coupled with the abundant moisture conditions, create host
environments for plant diseases. The predominant root diseases in gin-
seng likely reflect the impacts of these conditions.
b. Soil moisture. The experiments on the role of soil moisture on
ginseng growth are limited to the work of Mork et al. (1981). Using
2-year-old roots of P ginseng in a pot experiment, four levels of soil

moisture were maintained (30,45,60,and 80% of field capacity). The 60%
soil moisture level produced optimal aerial and root growth. A deficit or
overabundance of soil water resulted in decreased aerial and root growth,
including an increase in the number of missing plants. Statistical analy-
sis indicated that the relative growth rate was maximal a t 65.5%of field
capacity and was zero a t 31.5 and 100%of field capacity. Unfortunately,
field trials were not undertaken and the nature of the soil was not
specified to allow equivalent soil water potentials to be derived. As
expected, evapotranspiration rates were higher for the higher soil mois-
ture levels.
c. Temperature. The role of temperature on r! quinquefolium embryo
development was addressed by Stoltz and Garland ( 1980).Under constant-
temperature conditions in a controlled environment, the most rapid and
uniform development occurred at 2OOC. At O O C , no development occurred;
at 5' and 10°C, little development occurred; and, a t 3OoC the seed rotted.
In an outside plot that was subject to daily variations in temperature, a
wide range of results were obtained.
This research was extended by Lee et al. (1983)for I! ginseng and I!
quinquefolium. As opposed to using constant temperatures like Stoltz
and Garland (1980),optimal temperatures for embryo growth and devel-
opment were found to vary with different stages of embryo growth (Fig.
6.13).Embryo growth before seed coat splitting was maximum a t E°C,
30 -
-
oa
25
Y
?l 20
-
a
c
2
9)
15-
Q
c
E 10-
5-
I I I I 1 I
-
Initial Middle Late
--
Seed Plant
Germination Growth
Embryo Growth
Fig. 6.13. Predicted optimum temperatures for embryo growth, seed germination,
and plant growth. (From Lee et al. 1983.)
while temperatures of 10°C and over 25OC restricted embryo growth.

After splitting, temperatures higher than 2OoC restricted embryo growth
and low temperatures were more favored a t the full-grown embryo stage.
It is suggested that low temperatures are needed to break dormancy.
Germination percentages increased gradually with temperatures below
2OoC, while temperatures higher than 25OC caused rapid deterioration of
seed. A t temperatures of 5OC, shoot growth was restricted and a t temper-
atures higher than 3OoC, all seed deteriorated.
Dormancy and growth of r! quinquefolium roots is influenced by
temperature. Lee et al. ( 1985)examined the role of four temperatures (5,
10,15, and 2OoC)on roots stored a t 5 * 2OC. Maximum growth patterns
were found a t 15OC and after 100 days. At this temperature, stem growth,
leaf growth, stem dry weight, and plant growth rate increased with time
after 50 days storage a t 5OC.
Panax quinquefolium seedling growth was studied in chamber envi-
ronments a t 10,15,20, and 23OC (Lee et al. 1986). In addition, seedlings
and 3-year-old roots were grown a t constant soil temperatures of 13, 18,
and 23OC and the aerial parts of the plants were subjected to air tempera-
tures of 21OC during the day and 13OC a t night. In the chamber, maxi-
mum growth occurred a t 15OC and it was slow below 10°C. In the
controlled-root-temperatureexperiment, maximum leaf area, shoot, and
dry weight occurred a t 18OC. Maximum root growth occurred a t 15-18OC.
2. Mulch Effects. In the North American culture of ginseng, surface

coverings of mulches are employed as illustrated in Fig. 6.14. The mulch
emulates the leaf litter found in the native habitat of ginseng. It helps to
conserve soil water in the summer and protect the roots against damag-
ing cold in the winter season. In addition, mulches alter the characteris-
tics of the soils and impact on the leaf and root tissue constituents (Zito
et al. 1984).
Research conducted in North Carolina by Konsler (1979, 1980) and
Konsler and Shelton (1984) has documented the changes to the soil
characteristics attributable to mulch degradation. In dryland environ-
ments, these changes are less as mulch degradation is minimal (W. G.
Bailey, unpublished).
Konsler and Shelton ( 1984)presented summary tables (Thbles 6.8 and
6.9) on the role of six different mulches on ginseng root growth over ages
1-6 years and on seed yield over 4-6 years. For root growth, oak bark/
sawdust and poplar bark/sawdust mulches had the largest roots and the
greatest yields. Seed production was greatest with oak bark/sawdust
mulch. The low seed yield in year 6 was associated with the drought
period during the summer of 1983.
Zito et al. (1984) examined mulch effects on ginsenosides (Section
Fig. 6.14. A photograph of Canadian dormant season ginseng production showing

one method of wooden lath shade storage, and the mulch of straw to prevent low soil
temperature injury to the overwintering roots.
Table 6.8.Ginseng Root Fresh Weight Response to Six Bed Mulches from
Plant Ages 1-6 Years in North Carolina"
Fresh weight per root ( g )a t year:
Mulch 1 2 3 4 5 6
Straw 0.9 3.3 12.9 18.2 27.4 32.9

Pine needles 1.0 4.1 13.5 18.4 30.5 26.1
Poplar bark/sawdust 1.1 4.4 17.6 25.2 31.3 43.9
Oak bark/sawdust 1.1 4.2 15.2 26.9 37.0 51.7
Pine bark/sawdust 1.1 2.9 11.2 23.2 24.5 29.7
Hardwood leaves 1.1 3.5 11.9 19.6 26.3 26.1
LSD (0.05) 0.1 0.3 2.1 4.8 6.6 6.9
"From Konsler and Shelton 1984.
1I.A) in l? quinquefolium. Four-year-old plants grown in North Carolina

were used for the study and six mulch types employed. These were the
same as used by Konsler and Shelton (1984). For leaf material, total
ginsenosides were higher for straw, oak and hardwood leaf mulches.
Table 6.9. Ginseng Seed Yield Response to Six Bed Mulches at Plant Ages 4, 5,
and 6 Years in North Carolina"
Seed yield ( g/m2) at year:
Mulch 4 5 6
Straw 95 120 99
Pine needles 79 125 61
Poplar bark/sawdust 159 175 126
Oak bark/sawdust 162 232 193
Pine bark/sawdust 78 161 153
Hardwood leaves 79 137 115
"Adapted from Konsler and Shelton 1984.
Individual analysis for ginsenosides A1, Rgl, Rd, Rc,and Rb2 supported
this except for oak which showed a decrease in Al, Rgl, and Rb2. Root
tissue showed no significant differences in either total or individual
ginsenosides due to mulch treatments.
3. Hardiness. American ginseng is native to the southern part of

eastern and central Canada, the northern limit of its growth in North
America. I t survives low winter temperatures under natural mulches in
deciduous woodlands and under applied mulches in cultivation ( Currie
1980; Section IV.C.2).
A number of herbaceous perennials successfully overwinter in cold
climates. For example, fern rhizomes growing in deciduous forest floors in
Hokkaido, Japan, resisted temperatures ranging from -5.0 to -17.5'C
(Sato 1982). Strawberry crowns overwinter, but may be injured by low
temperatures of about -23'C, or by alternating warm and cool periods
where the lower temperatures may range from about -4 to -9'C (Daubeny
et al. 1970).We have been unable to find similar information for ginseng.
Lewis and Zenger (1982) reported low and variable seed survival in
indigenous populations and low-temperature injury a t this time is possi-
ble. Hu et al. (1980) have reported very high death rates in natural
populations; low-temperature injury could be inferred.
In Ontario, we have observed instances where cultivated ginseng has
not grown in the spring, implicating low-temperature injury to the rhi-
zome in the previous winter, particularly in the absence of a snow cover.
Also, we have seen examples of low-temperature injury to ginseng rhi-
zomes after they were accidently exposed to temperatures of -2OOC.
In controlled freezing tests, Proctor and Lee ( 1983)showed that Ameri-
can ginseng roots were not damaged by temperatures above -1OOC. At
-ll°C the small fine roots were damaged and at -12OC 1-year-oldrhizomes
were damaged. The main root resisted damage until -17OC. Alternating
temperatures were more damaging than continuous low temperatures.
For example, if roots were kept a t -3OC for 12 hr, then a t room tempera-
tures (2OOC)for 12 hr, and then returned to -3OC for 12 hr, they all died.
On the other hand, roots kept continuously for the 36 hr at -5OC were
not damaged. The practical implication of these studies is that care
with mulches must be exercised, particularly during the spring thaw
when air temperatures, and therefore soil temperatures, are liable to fluc-
tuate widely.
In other controlled-freezingtests, Proctor and Lee (1983)also reported
that germinated seeds were damaged a t temperatures below -9OC;
nongerminated seeds were more resistant to low temperatures and were
not injured until temperatures were below -13OC. The lower the water
content of the seeds, the lower the temperature a t which freezing injury
occurred. Ginseng seeds survived low temperatures of -15OC provided
they had intact endocarps. Seeds without endocarps, and particularly
those with high water content, were likely to be damaged by temperatures
between -3' and -1OOC.
4. Influence of Radiation on Photosynthesis and Stornatal Conductance.

a. Photosynthesis. Both I! ginseng and I! quinquefolium are shade
species and in the wild are components of the forest floor environment.
Hence, photosynthesis response to solar irradiance involves the adapta-
tion to the shade environment. The research on ginseng photosynthesis
was pioneered by Kim (1964a,c),who conducted experiments on severed
leaves from 6-year-old ginseng plants. Photosynthesis light saturation
curves were evaluated and a decrease in photosynthesis with air tempera-
tures exceeding 25OC was noted. This research served as a major contri-
bution to the review of photosynthesis response to irradiance that was
prepared by Park ( 1980).A wide range in photosynthesis light saturation
was found in the literature and it is evident that a high degree of
uncertainty exists in our understanding of the photosynthesis process.
Figure 6.15 presents a summary of photosynthesis response to photo-
synthetically active radiation (wavelengths 0.4-0.7 pm) compiled from
Proctor (1980)and Lee et al. ( 1980a).For comparison purposes, the light
and radiation measurements were converted to a standardized base. The
Proctor ( 1980)results present the influenceof increasing photosynthetically
active radiation on the net photosynthesis of attached leaves on 3-year-old
I! quinquefolium in a laboratory experiment using potted plants. The
*
temperature regime was maintained a t 25 2OC. Maximum photosyn-
thesis rate was 0.175 COa m2/h and the saturation irradiance was 40
W/m2 ( 10%of full sunlight). No inhibition of photosynthesis was obtained
224 J. T A. PROCTOR AND W. G. BAILEY
0 20 40 60 80 100 120 140 160 180 200
PHOTOSYNTHETICALLY ACTIVE RADIATION (Wrn-')
Fig. 6.15. The photosynthesis response of ginseng leaves to photosynthetically

active radiation. Curve A , I! quinquefolium, temperature 25' *
2'C. (From Proctor
1980.) Curve B, temperature 20'C; Curve C, temperature 30'C. (From Lee et al.
1980a.)
up to 100 W/m2. Similar data have been presented by Lee et al. (1980a)
for €? ginseng. The photosynthesis results for the 2OoC temperature
regime are comparable to those of Proctor ( 1980)in form. With a tempera-
ture increase to 3OoC, inhibition is seen to occur. This suggests that the
effects of temperature during the summer months can result in photosyn-
thesis reductions even during periods of optimal irradiance. Similar
findings have been presented for both I? ginseng and I? quinquefolium by
Park (1980).However, a large difference between the photosynthesis rate
between the two ginseng species is evident in Fig. 6.15. The high rates
evident in the Korean research suggest limitations in the experimental
techniques. In the only direct comparison of I? ginseng and I? quinquefolium
responses (Park 1980),both species were found to have high photosynthe-
sis rates. At low irradiances, I? quinquefolium exceeded €? ginseng.
However, a t higher irradiances, the reverse was found.
It has been recently recognized by Korean researchers (Lee et al.
1980a; Park 1980; Choi et al. 1982b)that the traditional shade environ-
ment created by a rice straw canopy limits photosynthesis in the back
rows of the garden bed. This has led to the examination of new shade
materials (wooden lath, Styrofoam boards, woven fabric) to enhance the
radiative regime in the growing environment.
Proctor (1980)found that I! quinquefolium had a low total chlorophyll

content per leaf area. This was comparable to other shade species and less
than that for sun species. However, expressed on a weight basis, it was
found to be more comparable to sun species than to shade species.
Whereas shade species usually have a higher chlorophyll b than chloro-
phyll a content, I! quinquefolium has a chlorophyll a to b ratio of 3 to 1,
comparable to sun species. In studies of I? ginseng, Korean researchers
(Choi et al. 1982b; Lee et al. 1980a; Park 1980) found similar results to
those of Proctor ( 1980),but the ratio of chlorophyll a to chlorophyll b was
2 to 1. Both Park (1980)and Choi et al. (1982b)have illustrated that total
chlorophyll as well as chlorophyll a and b decreased with increases in the
illumination levels of the crop. Further, Lee et al. (1980a) have shown,
using a limited data set, that there is a linear relationship between net
photosynthesis and total chlorophyll content.
b. Stornatal conductance. Photosynthesis is highly regulated by sto-

matal activity, which is crucial to both the efflux of HzO and the influx of
COz of the leaf. Proctor (1980) measured stomatal conductance on the
abaxial surface of exposed leaves of I? quinquefolium in the field using a
diffusive porometer and related stomatal conductance (the reciprocal of
stomatal resistance) to photosynthetically active radiation (Fig. 6.16 1.
..
I .
"d-;1
0 ON-.
a .
,"
-I: - 0
0 100 200 300 400 500

PHOTOSYNTHETICALLYACTIVE RADIATION
(Wm-2)
Fig. 6.16. The relationship between stomatal conductance and photosynthetically

active radiation for 3-year-old €? quinquefolium growing under wooden lath shade ( 0 )
and under woven black polypropylene (0). (From Proctor 1980.)
Stomata1 conductance reached a maximum a t approximately 225 W/m2,

but a wide scatter in the data indicated the role of other parameters. Chief
among these are internal plant water status and the influence of soil
moisture availability. Neither of these was measured, but the soil mois-
ture was not believed to be severely limiting. In a further experiment in a
laboratory environment, ginseng showed relatively slow closure when
light intensity decreased, but a rapid response when exposed to light.
Proctor ( 1980)found the cuticular conductance on the adaxial leaf surface
to be 5-1070 of stornatal conductance, indicating it plays a minor role in
water efflux.
Park ( 1980)has illustrated that stornatal conductance increased with
increasing light. As a direct consequence, transpiration also increased. In
a comparison of l? ginseng and l? quinquefolium, the latter was found to
have more stomata per unit leaf area. In transpiration experiments, this
was reflected by higher water efflux from l? quinquefolium than from
l? ginseng.
5. Endogenous and Exogenous Growth Regulators. Little published

information is available on the manipulation of ginseng growth and
development using plant growth regulators nor have endogenous growth
regulators been extracted and identified. The root dormancy period of
American ginseng has been shortened by exogenous application of gib-
berellic acid to the rhizome (J. T. A. Proctor, unpublished data) and Park
et al. (1979)have reported a similar effect for Oriental ginseng. Recently,
various fungicides have been reported to have growth-regulating effects.
For instance, Church et al. (1984) reported that triadimefon increased
apple fruit set, paclobutrazol decreased it, and both chemicals reduced
eventual leaf size. Root rot is the limiting factor in world ginseng pro-
duction (Hildebrand 1935; Ohh 1981; Section 1II.E). A growth regu-
lator that would protect ginseng against devastating fungi and a t the
same time increase root yield would be a great asset to the world gin-
seng industry.
6. Air Pollutants. Proctor and Ormrod (1981)have shown that expo-

sure of ginseng plants to 20 pphm (parts per hundred million) ozone for 6
hr daily under controlled conditions for 4 days induced upper-surface
leaflet stippling along the veins and interveinally. Similar time exposure
to sulfur dioxide a t 50 pphm induced mild chlorosis to irregular necrotic
areas. This report established that ginseng may be classified as moder-
ately sensitive to ozone and sensitive to sulfur dioxide. However, the
impact of these findings on ginseng production in the field is unknown
and very difficult to assess.
D. Pest Control
1. Diseases. “The average yield of ginseng per acre in the United
States is not more than one-sixth to one-third of what might reasonably
be expected, the shortage being caused almost entirely by the numerous
diseases which attack the crop” (Whetzel et al. 1930). This statement
summarizes a situation that prevailed over 50 years ago and that is
similar today not only in the United States, but in Canada and other
producing areas around the world. Ohh (1981) reviewed the ginseng
disease problem, particularly for the Orient and his data are summarized
in Bble 6.10. Description of the various ginseng diseases can be found in
Whetzel and Rosenbaum (1912),Whetzel et al. (1930),and Curran (1985).
The latter guide is written by a layman, but it does contain a lot of correct
and useful information gleaned from various sources. An authoritative
guide or bulletin on ginseng diseases written by plant pathologists is
much needed.
Disease control remains the central problem in world ginseng produc-
tion. In addition to the diseases in Bble 6.10, there are others that we
have not recognized and identified. The root rot complex of diseases is
particularly difficult to control because of the many factors involved and
the perennial nature of the crop. Much more needs to be known about the
root rot complex and the other major diseases so that effective control
strategies can be formulated. A brief concerted study on one disease has
been made. Hildebmd (1935)worked on root rot, particularly “disappearing-
rot,” aptly named because in a relatively short time the roots either
Table 6.10. Some Diseases of Ginseng and Associated Damage and Loss“
Damage or
Disease Pathogen(s)* losses (70O)c
Anthracnose Colletotrichum panacicola 20-47

Damping-off Phytophthora cactorum, Pythium sp. 5-50
Rhizoctonia solani
Root rot Cylindrocarpon des tructans, 1-60
Fusarium solani, Pseudomonas sp.,
Ramularia sp., Sclerotinia sp.
Phytophthora Phytophthora cactorum 2-30
Alternaria blight Alternaria panax 10-20
Missing rated 10-50
“From Ohh (1981).

some cases the disease may be caused by several pathogens.
‘Several estimates for the same disease are listed in Ohh (1981).
dRefersto percentage of plants remaining a t harvest taking the number a t planting
as 100%. Disease is one of the causal agents in this missing rate.
completely disappear in the soil or leave as evidence of their presence only

a peridermal shell enclosing fragments of vascular tissue. Many more
such studies are needed on a continuing basis. They must be integrated
with studies on cultural practices and the screening of suitable pesticides
for disease control in ginseng. In North America one initiative has been
taken by the Wisconsin Ginseng Growers Association and the University
of Wisconsin at Madison. Field test sites have been set up and laboratory
back-up put in place (refer to examples of fungicide testing results below).
2. Pesticides. There are no pesticides registered for use on ginseng in

Canada, but there is limited use of some registered chemicals in the
United States (Heise 1984). Ginseng could not be cultivated economi-
cally without pesticides. There are a number of recent pesticide screening
tests with ginseng (Curran 1985; Putnam 1984; Putnam and Mitchell
1984; Rahimian and Mitchell 1984).Putnam and Mitchell (1984)found a
reduction in Phytophthora root rot and an increase in yield from a single
treatment of metalaxyl. Rahimian and Mitchell (1984)obtained a reduc-
tion in percentage of infected plants and leaf area infected with leaf blight
caused by Alternaria when plants were sprayed with maneb, mancozeb,
or iprodione.
3. Insects and Other Pests. Relative to diseases, insects and other

pests (aphids, leafhoppers, lygus bugs, cutworms, scale, slugs and snails,
and mice) are minor problems but should not be ignored. Curran (1985)
has reviewed the various control measures currently available for the
above-mentioned pests.
E. Weed Control
Tkaditionally weed control in ginseng has been by hand, a labor-
intensive and expensive operation ( Love 1982). Elimination of perennial
weeds and soil fumigation prior to seeding keep weeds in new plantings a t
a low level. Subsequently, weeds in ginseng usually occur where straw
mulch containing weed seeds is used, or where weed seeds are blown in
from adjacent weeded areas. As ginseng production has now expanded
into new growing areas, like British Columbia, new weed problems have
arisen. Dodder, Cuscuta campestris Yunker, a parasitic vine weed usually
associated with alfalfa, will kill P quinquefolium if left unattended (W. G.
Bailey, unpublished data). Present control is by hand weeding. Weed
control is less of a problem in 3- and 4-year-old plantings, where the leaf
area index is higher (Hughes and Proctor 1981) and weeds have little
opportunity to become established and compete with the ginseng plants.
There are no herbicides registered for use in ginseng. Some North
American ginseng growers have used paraquat or glyphosate in the

spring when weed seeds have germinated and grown. This is done before
the ginseng has grown through the mulch.
Herbicide research trials in each of two years with fluazifopbutyl or
sethoxydim as early crop postemergence treatments at the 3-5 leaf stage
of grasses killed annual grasses and inhibited growth of perennial
quackgrass, with no crop phototoxicity (Souza Machado and Ali 1984,
19851. Complementary trials aimed a t controlling broadleaf weeds, in
addition to annual grasses, have shown very good weed control with
trifluralin and linuron pre-emergent to the crop and weeds (Zilkey and
Capell 1984, 1985). Adding extra water (simulated rain) to the trifluralin
gave good control of germinating broadleaf weeds. Chlorothal-dimethyl,
chloramben, and diethatyl ethyl applied postemergent to the crop, but
pre-emergent to the weeds also gave very good weed control. Other
herbicides that have been tested and found phytotoxic, and therefore
unsuitable, included diphenamid, oxyfluorofen, ioxynil, and bromoxynil.
These trials in Canada indicate that expensive hand weeding can be
replaced with suitable herbicides. The precise weed management pro-
gram may vary in different production areas and will depend on factors
such as herbicide availability, soil types, rainfall, and weed species.
V. CONCLUDING REMARKS
Ginseng research over the last century has been concerned primarily
with the constituents of the plant and their uses rather than on the
growth, development, and cultivation of the crop. Nevertheless, pioneering
research in the latter area has taken place. In the first quarter of this
century, H. H. Whetzel contributed much to our understanding of gin-
seng diseases. His work was extended in the 1930s by A. A. Hildebrand.
Studies on photosynthesis and growth relationships of ginseng by J. H.
Kim in Korea occurred in the 1960s. In the last decade or so, we have seen
a more concerted effort in North America and the Orient. In North
America we have seen the ecological studies of S. Y. Hu and W. H. Lewis,
the cultivation studies by T. R. Konsler and L. P. Stoltz, and the disease
work by J. E. Mitchell and others a t Wisconsin. A Ginseng Research
Institute has been established by A. W. Rashap. Contributions to exten-
sion have been made by people such as G. F. Hartman, L. Martin, and C.
R. Roberts. In the Orient the Koreans at the Korean Ginseng Research
Institute have undertaken many different research projects. A few exam-
ples are in tissue culture and breeding (K. T. Choi), in environmental
studies (H. Park and J. C. Lee), and in diseases (S. H. Ohh).
The ginseng plant and its cultivation are very complex. Much research
remains to be conducted to understand fully the mechanisms control-

ling the overall growth and development of the ginseng plant. The in-
creased interest and research in the last decade, particularly in the western
world, augurs an increase in research reports in the remaining years of
this century.
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The Honey Bee Pollination Component

of Horticultural Crop Production Systems
G. DeGrandi-Hoffman*
Carl Hayden Bee Research Center
U.S. Department of Agriculture,
Agricultural Research Service, lhcson, Arizona 85719
I. Pollination in Agroecosystems 237

A . Weather and Honey Bee Foraging 238
B . Floral Attractiveness and Foraging Behavior 239
C . Nectar, Pollen, and Foraging Activity 241
11. Pollination and the Production of Horticultural Crops 243
A . Deciduous Fruit Crops 244
B . Citrus 247
C . Melons 247
D. Berries 248
E . Nuts and Oilseeds 250
F . Vegetables 251
G . TkopicalCrops 254
111. Mechanisms for Cross-Pollination 256
I v. Concluding Remarks 259
I. POLLINATION IN AGROECOSYSTEMS
When bees (and other organisms) visit flowers to gather nectar and
pollen, they often transfer pollen between reproductive structures, and
initiate the seed set process. Both plants and pollinators clearly benefit
from this mutual relationship. The coevolution of plants and their polli-
nators has insured reproductive success for the former and a food supply
for the latter. In agroecosystems though, plant/pollinator relationships
are often viewed as a major constraint in the production of crops requiring
insect-mediated pollination for seed/fruit set.
Insect pollination is a requirement for the production of many crops,
*I thank Stephen Buchmann, Clarence Collison, Eric Erickson, Marshall Levin,
Gerald Loper, and Gordon Waller for their review of this manuscript and for their very
valuable suggestions. I especially thank Cristina Bramley for great help in the
preparation of this manuscript.
237
238 G. DEGRANDI-HOFFMAN
but in agroecosystems native pollinators are often too scarce to insure

adequate pollination. Insecticides, herbicides, and cultivation practices
have reduced feral bee populations in many areas to the point where they
are insufficient for pollination of commercial plantings. Growers of horti-
cultural crops rely almost entirely upon honey bee (Apis melliferu L.)
colonies to meet their pollination needs (McGregor 1976).For this reason
this review is limited to honey bee pollination of horticultural crops.
The practice of renting colonies for pollination began about 1900, after
orchardists reported increased fruit yields when colonies were present
(Todd and McGregor 1960).Honey bees have attributes that make them
especially valuable for crop pollination. Populations can be housed in
manageable colonies, and so literally thousands of potential pollinators
can be introduced and removed. In addition to pollination services, colo-
nies can be managed to produce surplus honey, beeswax, propolis, and
royal jelly, which are all salable products. Currently, about one-third of
U.S. agriculture is dependent upon or benefits from honey bee pollina-
tion (McGregor 1980).The contribution of honey bees to agriculture has
been estimated to be about $20 billion per year (Levin 1983).
The number of foraging bees in a colony is directly proportional to.the
colony’s overall population size (Farrar 1931; Gooderham 1950).Regard-
less of the colony’s size, weather factors -particularly temperature, wind
velocity, solar radiation, and relative humidity -determine the percent-
age of a colony’sforaging population that will leave the hive to forage a t
any given time.
When colonies are introduced for pollination, a variable percentage of
the population will forage the “target crop.” Once foragers leave the
colony, there is competition among plant species for pollinators. Plants
with distinctive odors and colors that attract bees improve their chances
of being visited. The quantity and quality of the nectar and pollen then
determine whether the food source is suitable for additional foraging. If
nectar and/or pollen are readily available and nutritious, foragers will
return to the hive and recruit additional bees. Thus, pollination can be
considered a dynamic process, whose rate is dependent on weather,
available foragers, and floral “attractiveness” relative to other blooming
plants in the honey bee colonies’flight range.
A. Weather and Honey Bee Foraging

The major factors influencing honey bee flight initiation are tempera-
ture and solar radiation (Brittain 1933; Butler and F’inney 1942; Burrill
and Dietz 1981).Honey bees will not fly if temperatures are below 9OC.
Flight and temperature are positively and linearly related, a t least for the
temperature ranges encountered in these studies (14O-22OC)(Szabo 1980;
7. HONEY BEE POLLINATION AND HORTICULTURAL CROP PRODUCTION 239
Burrill and Dietz 1981). Even when suitable temperatures exist, flight
will not occur without sufficient light. Some bees fly on cloudy days, but
they tend to stay close to the hive (Phillips 1930a,b).In the morning and
afternoon, flight activity is positively related to solar radiation. During
the solar noon period (i.e., when the sun is at its zenith) though, flight
activity appears to be negatively related to solar radiation (Lundie 1925;
von F’risch 1965; Gary 1967; Burrill and Dietz 1981).Difficulty in locat-
ing and communicating food sources when the sun (honey bees’ principal
means of orientation) is directly overhead may explain the decline in
foraging activity.
Flight activity is negatively related to humidity, as would be expected
since temperature and humidity are inversely related (Brittain 1933;
Szabo 1980). Wind velocity is also negatively related to flight. Honey
bees fly a t about 6.3 m/sec (14 mph), and so it is reasonable to assume
that wind speeds greater than or equal to that average negatively affect
flight (Williams and Sims 1977).Lundie ( 1925)reported that the thresh-
old wind speed that causes a reduction in flight varies between 4 and 9
m/sec (9-20 mph). Observations of honey bees foraging apple blossoms
indicated that foraging activity declines at wind velocities greater than
3.1 m/sec ( 7 mph) (Brittain 1933).
The influence of weather conditions on foraging activity is dependent
on colony size. During unfavorable weather, a lower percentage of the
foraging population of larger colonies leaves the hive compared to smaller
colonies (lhranov 1952; Free and Preece 1969).Hence, a colony’s foraging
population must be considered a variable that is dependent upon both
climatic and colony conditions.
B. Floral Attractiveness and Foraging Behavior

The three major biochemical components of flower recognition are odor,
color, and the nutritional value of the nectar and pollen (Harborne 1982).
When honey bees (and other pollinators) approach flowering plants, one
stimulus they receive to search for rewards is odor. Pollinators can respond
to odors from a considerable distance. Honey bees detect odors with their
antennae, and will not respond to scents without them (Brett and Sullivan
1972). Newly recruited foragers seeking a given food source for the first
time rely on odor (Wells and Wenner 1971, 1973; Wenner 1974) and are
unable to locate any food source that is completely unscented ( Wenner
et al. 1969; Wells and Wenner 1971; Friesen 1973). Once a pattern
of foraging a t a source is established, memory of location and visual
cues such as landmarks become more important (Levin 1966; Wells and
Wells 1985).
Plants release scents at times and temperatures when their pollinators
are most active. In alfalfa florets, the initial emanation of volatiles is

photoperiodically induced, and there is a cyclic release of volatiles during
the day (Loper and Lapioli 1971). Odors are composed of mono- or
sesquiterpenes, simple aliphatic alcohols, ketones, or esters and can
originate from petals, leaves, or flowers. Normally, a mixture of these
compounds provides a flower’s scent (Harborne 1982; Williams 19831.
As pollinators get closer to flowers, the next cue directing them to the
nectar and pollen may be the contrasting color of the petals against the
usually green leafy background. Flower color results from the reflection
and refraction of light from cell surfaces. The most important group of
floral pigments are the flavonoids, which make up the cyanic colors
(orange, red, and blue), yellows, and white (Harborne 1967, 1976). The
other major group of floral pigments is the carotenoids, which provide
primarily yellows, and some oranges and reds. Minor classes of pigments
are chlorophylls (greens),quinones (occasional reds and yellows), and
betalain alkaloids (reds, yellows, and some purples) (Harborne 19821.
Much is known about color preferences of bees largely due to the work
of von Frisch (1950)and others. Bees are attracted to flowers that appear
blue or yellow to the human eye. Bees also discern differences in absorp-
tion in the ultraviolet (UV)region of the spectrum, and are sensitive to
the intensely UV-absorbing flavones and flavonols (Kevan and Baker
1983). These pigments, present in practically all white flowers, occur as
co-pigments in others. Although bees are insensitive to red hues, they
still will visit some red flowers if the petals have UV-absorbing flavones
(Harborne 1982).For a discussion of the influence of color on bee behavior
see Kevan (1983).
Experiments using artificial patches of blue and yellow “flowers” show
that some honey bees are constant to color and others to odor (Wells et al.
1981, 1983; Wells and Wells 1983). When visual cues are limited or
uniform, bees may rely more on odor and become odor-constant foragers
(Wells and Wells 1985).
In addition to odor and color, the outline of a flower’s petals is an
important sensory cue to pollinators. Flowers with highly broken outlines
are more stimulating than those with smooth outlines (Kevan and Baker
1983). Once pollinators alight on a flower, they may receive additional
directions to the nectar by visual and olfactory nectar guides on the
petals (Manning 1956; Free 1970; Jones and Buchmann 1974). Nectar
guides are created by a differential distribution of pigments within the
flower tissue.
Additional information concerning species identification and reward
location may be provided to pollinators by microsculptural features on
the petals (Kevan and Lane 1983, 1985). These tactile cues (previously
used solely for plant taxonomy) are species specific, and may provide a
means for pollinators to discriminate between plants of similar floral

morphologies. Microstructural patterns differ from one end of the petal
to the other, and may also function as nectar guides to pollinators.
Attempts have been made to increase a plant’s attractiveness to polli-
nators by accentuating the biochemical cues, particularly odor. Applica-
tions of sugar water solutions containing forager attractants have been
applied to crops, but to date no compound has increased plant attractiveness
to honey bees or other pollinators (Free 1965; Waller 1970; Burgett and
Fisher 1979; Mayer and Johansen 1982; Margalith et al. 1984).
C. Nectar, Pollen, and Foraging Activity

The primary reason honey bees (and many other pollinators) visit
flowers is to collect nectar and pollen for the colony’s nutritional needs.
Nectar may have no other role in angiosperms than to attract pollinators.
The three sugars most commonly found in nectar are glucose, fructose,
and sucrose, although other sugars are often present in small amounts
(Baker and Baker 1983).The occurrence of these sugars has been defined
for over 800 species by Percival(l961). There are three groups of nectars:
sucrose dominant, those containing primarily glucose and fructose, and
nectars with approximately equal amounts of glucose, fructose, and
sucrose. Honey bees prefer nectars containing sucrose to those with
either glucose or fructose. Fructose is preferred over glucose. Sugar solu-
tions containing 30-5090 sucrose are more attractive than solutions
containing higher or lower sugar concentrations ( Waller 1972). Wykes
(1952a,b) reported that flowers that are attractive to honey bees are
characterized by approximately equal amounts of glucose, fructose, and
sucrose. Sugar solutions containing equal parts of sucrose, glucose, and
fructose, however, are less attractive to honey bees than solutions containing
only sucrose,or solutions where sucrose is the predominant sugar (Bachman
and Waller 1977). In addition to sugars, amino acids and proteins are
commonly present in nectar in trace amounts (Baker and Baker 1973a,b 1.
The threshold temperature for nectar secretion and the amount of
nectar produced are species dependent. Nectar secretion may vary between
cultivars of the same species (Beutler 1953). Using an artificial “flower”
patch, Wells et al. (1981)reported that the number of flowers visited per
trip is negatively related to the mean nectar yield per flower, and that the
relationship is logarithmic. The relationship was found whether all flow-
ers provided reward or if some were empty.
The amount of nectar in a flower and its sugar concentration are strongly
influenced by environmental factors. In opened dish-shaped flowers, nec-
tar concentration can vary greatly with relative humidity (Vansell 1934,
1942; Corbet et al. 1979a,b). Nectar can also be diluted by dew or rain.
Conversely, nectar can be concentrated by dry winds, and increase (or in

some cases decrease) floral attractiveness (Kevan and Baker 1983).More
nectar is secreted on sunny than cloudy days, indicating that nectar sugars
are a direct product of photosynthesis, which is influenced by sunlight
(Shuel1955a).Soil moisture and composition, atmospheric pressure, size
of the nectary, flower age, available carbohydrates, and position of the
flower on the plant also influence the amount of nectar secreted (Butler
1945; Percivall946; Wykes 1952c; Beutler 1953; Shuel1955b,1957; Free
1970;Loper et al. 1976).Thus, it can be concluded that nectar secretion is
an extremely dynamic plant process, and the amount of nectar occurring
in a flower a t any point in time is a response to the climatic, edaphic, and
biotic conditions a t that time. For a discussion of techniques and consid-
erations used to study nectar secretion, see Beutler (1953)or Cruden and
Hermann (1983).
Nectar provides a carbohydrate source for honey bees (and other
Apoidea), while pollen supplies the remainder of their nutritional needs
(Haydak 1970). Pollen is primarily protein, but also contains starch,
sugars, fatty acids, and trace amounts of inorganic salts (Todd and
Bretherick 1942; Harborne 1982). For a discussion of pollen chemistry
see Barbier (1970)or Stanley and Linskens (1974).
Anther dehiscence and subsequent pollen shedding occurs with a
rhythm that is species specific, but influenced by weather conditions
(Free 1970).For example, apple blossoms will not open and anthers will
not dehisce if temperatures are below 10°C (Percival 1955).Pollen collec-
tion by honey bees is also influenced by weather. Following days of un-
favorable weather, pollen collection is more intense than when conditions
are uniformly favorable. This may be partly because bees respond more
readily to improvement of conditions, and partly because the pollen needs
of the colony are greater following a period of confinement (Free 1970).
Foragers’ propensity to collect pollen is largely dependent on colony
needs. The presence of brood (i.e., eggs and larvae) and a queen in
coloniesis a strong stimulus for foragers to collect pollen (Free 1967;Todd
and Reed 1970; Al-Tikrity et al. 1972; Moeller 1977). Overwintered
colonies generally have larger populations and more frames of brood than
do package colonies in early spring (Farrar 1931; Filmer 1932;Gooderham
1950). Consequently, overwintered colonies will contain more foragers
(Szabo 1980) than package colonies. Increased pollen foragers make
overwintered colonies more valuable for pollinating crops that bloom
early in the spring (e.g., apricot or cherry), especially those where pollen
collectors have a greater chance of contacting stigmata than nectar
collectors (e.g., apple or almond) (Roberts 1945; Free 1966b; Robinson
1979; Thorp 1979). The percentage of pollen collectors in the foraging
population can also be increased by installing pollen traps that remove
pollen loads from the legs of foragers as they enter the colony (Webster et
al. 1985).Hence, pollen traps might increase the pollinating efficiency of
colonies, but continuous pollen trapping can drastically reduce brood
rearing (Moeller 1977).
Honey bees generally forage within 5 km of their hive, but are capable
of greater distances if adequate resources are not available (and weather
conditions are suitable) closer to the colony. Although individual honey
bees are usually species specific (for a discussion of foraging constancy,
see Grant 1950;F’ree 1963,1970;Waddington and Holden 1979;Waddington
and Heinrich 1981; Waddington 1983), colonies often distribute their
foragers among several plant species, thereby satisfying the colony’s
nutritional requirements (Gary 1979).
Honey bee foraging behavior is a balance between maximizing individ-
ual efficiency in food collection and maximizing the use of available food
resources by the colony as a whole (Nunez 1982). Between‘ 5 and 35%
(depending on forage availability) of a colony’s foragers are scout bees
( Seeley 1983),whose role is to find previously unexploited food sources.
Scouts communicate information on the location of the food source,
nectar abundance, and concentration to nestmates through the dance
language (von Frisch 1967, 1974; Lindauer 1971; Gould 1975, 1976;
Gould et al. 1970). While nectar concentration determines the species
that will be foraged, nectar abundance influences the size of the foraging
population on the species (Butler 1945).On any given day, the majority of
a colony’s foragers are distributed over a relatively small number of food
sources (Visscher and Seeley 1982). As nectar quantity from a given
species declines, the number of foragers that dance when they return to
the hive is reduced (Frisch 1967). Foragers that find species with higher
sugar concentrations dance with more vigor, and more successfully recruit
new foragers than those finding nectar with lesser sugar concentrations
( Lindauer 1948). Apparently, honey bee colonies constantly readjust
their allocation of foragers to various food sources, so that on any given
day foragers are focused on the most rewarding food sources the colony
has found (Visscher and Seeley 1982).
11. POLLINATION AND THE PRODUCTION OF

HORTICULTURAL CROPS
This section will present a brief discussion on the pollination require-

ments of horticultural crops (Thble 7.1). An overview of the subject is
provided by Free (1970),McGregor (1976),and Crane and Walker (1984).
In general, honey bee pollination not only affects crop yields, but also
quality. Many crops require most if not all ovules to be fertilized for
Table 7.1. Horticultural Crops Included in This Review
Family Species Common name
Actinidiaceae Actinidia chinensis Planch. Kiwifruit (Chinese

gooseberry)
Amaryllidaceae Allium cepa L. Onion
Anacardiaceae Anacardium occidentale L. Cashew
Mangifera indica L. Mango
Caricaceae Carica papaya L. Papaya
Cruciferae Brassica oleracea L. Brussel sprouts
(gemmifera group)
Cucurbitaceae Citrullus lanatus (Thunb.) Manf. Watermelon
Cucumis melo L. Muskmelon or cantaloupe
Cucumis satiuus L. Cucumber
Cucurbita spp. Pumpkin and squash
Ericaceae Vaccinium ashei Reade Rabbiteye blueberry
Vaccinium angustifolium Ait Lowbush blueberry
Vaccinium corymbosum L. Highbush blueberry
Vaccinium myrtilloides Michx. Sour-top bilberry
Vaccinium macrocarpon Ait Cranberry
Leguminosae Arachis hypogaea L. Peanut
Phaseolus lunatus L. Lima bean
Vicia faba L. Broad bean
Myrtaceae Psidium guajaua L. Guava
Palmaceae Cocus nucifera L. Coconut
Pedaliaceae S e s a m u m indicum L. Sesame
Proteaceae Macadamia integrifolia Maiden and Macadamia
Betche
Macadamia tet raphyl la L .A .D . Macadamia
Johnson
optimum fruit size and shape (e.g., apple, berries, watermelon). Honey
bees can assure maximum crop size and quality if enough colonies are
introduced, sufficient pollen is available (if the cultivar is self-incompatible),
and good honey bee flight weather occurs during bloom.
A. Deciduous Fruit Crops

Many deciduous fruit trees (with the exception of those that are
anemophilous or produce fruit parthenocarpically) require insect-mediated
pollination for seed formation and fruit development. Most apple and
plum cultivars and practically all commercial sweet cherry are genetically
self-incompatible and require cross-pollination. Apple cultivars such as
‘Golden Delicious’ or ‘Northern Spy’, tart cherry, and most peach culti-
vars, are a t least partially self-compatible (i.e., pollen from blossoms of
the same clone can fertilize ovules and initiate fruit formation), but still
Family Species Common name
Rosaceae Frageria Xananassa Duch. Strawberry

Malus aldenhamensis Crab apple (dark
crimson petals)
Malus domestica Borkh Apple
Malus floribunda Crab apple (pink
petals)
Prunus auium L. Sweet cherry
Prunus cerasifera Ehrh. Plum (myrobolan or
cherry)
Prunus cerasus L. Tart cherry
Prunus domestica L. European plum
Pyrus communis L. Pear
Prunus dulcis (Mill.) D.A. Webb Almond
syn Prunus amygdalus Batsch
Rubus spp. Raspberries
Rubiaceae Coffea arabica L. Coffee
Coffea robusta Linden Robusta Coffee
[ C. canephora Pierre ex Froehner]
Coffea Ziberica Bull. ex. Hiern. Liberian coffee
Rutaceae Citrus limon ( L . )Burm. f. Lemon
Citrus sinensis ( L . ) Osbeck Sweet orange
Citrus paradisi Macf. Grapefruit
Citrus reticulata Blanco lhngerine
Sapindaceae Litchi chinensis Sonn. Litchi (lychee)
Umbellifera Daucus carota L. Carrot
PetroseZinum crispum (Mill.) Nym. Parsley
require bees to transfer pollen from anthers to stigmas. Conversely,

self-incompatible pear cultivars such as ‘Bartlett’ (Griggs and Iwakiri
1954) and ‘Packhams Tliumph’ (Langridge et al. 1976) can set fruit
parthenocarpically in some years and sites. For a complete discussion of
fruit trees that require pollination, see McGregor (1976),Free (19701,and
Crane and Walker (1984).
Fruit trees that require cross-pollination must have a cultivar planted
nearby to serve as a compatible pollen source ( pollinizers). Pollinizers
must bloom annually and produce pollen that is genetically compatible
with the main cultivar (Free 1960). To be consistently effective, polliniz-
ers should begin to bloom 1-2 days before the main cultivar, although this
characteristic may differ from one location to another (Dennis 1979).
Bloom phenology is influenced by temperature: cool weather prolongs
bloom, while warmer temperatures accelerate it (Morris 1921). This rela-
tionship causes the period of overlap between pollinizer and main variety
blooms to differ from year to year, depending on temperatures.
Recently there has been increased interest in the use of ornamental

crab apple (Malus spp. ) as pollinizers for commercial apple cultivars
(Williams 1972). There are several advantages to using crab apple as
pollinizers. Some cultivars produce more flowers and consequently more
pollen per unit length of branch than commercial apple cultivars (Church
et al. 1974). Hence, fewer crab apple trees may be needed per hectare to
optimize compatible pollen availability. The small size of crab apple fruit
makes it readily distinguishable from main cultivar fruit, though crab
apples are rarely harvested. To be an effective pollen source, the crab
apple cultivar must be attractive to bees, produce compatible pollen, and
consistently bloom in synchrony with the commercial cultivar. Three
ornamental species that differed in blossom petal color (dark crimson
Malus aldenhamensis, pink Malus fZoribunda Hillierii, and white-flowered
Malus cv. Golden Hornet) were equally attractive to bees (Kendall and
Smith 1975).The relationship between the bloom patterns of several crab
apple and commercial cultivars was reported by Crassweller et al. ( 1980),
along with the crab apple’s growth habits and susceptibilities to disease.
The behavior of honey bees on fruit trees has been the subject of
considerable study (see Free 1960,1970; McGregor 1976).Bee visitation
rates are positively related to blossom density. Honey bees can in some
cases discriminate between cultivars, and tend to show cultivar-specific
fidelity (Free and Spencer-Booth 1964a; Free 1966a; Robinson 1981). In
studies using ‘Cox’s Orange Pippin’ and ‘Golden Delicious’, most bees
visited flowers with dehiscing anthers. Apetalous flowers were often
visited for nectar, while newly opened flowers (without dehisced anthers)
were not visited by honey bees (Williams and Brain 1985).
Honey bees that establish foraging areas on a cultivar tend to continue
visiting it on consecutivetrips. Cultivar fidelity occurs less frequently and
foraging areas become larger in the early and later stages of bloom, when
nectar and pollen rewards on a single tree (or cultivar) are more limited
(Free 1966a).
Although honey bees readily visit fruit trees, in some instances nectar
collectors fail to contact the stigma and hence do not pollinate blossoms.
Honey bees may stand on petals and reach between the stamina1 fila-
ments into the nectaries (sideworking). Honey bee sideworking behavior
has been used to explain poor fruit set in apple cultivars such as ‘Deli-
cious’ (Roberts 1945; Robinson 1979),but this is probably not the case.
If the cross-pollination rate of ‘Delicious’blossoms is plotted against the
percentage of honey bees employing sideworking behaviors over the
bloom period, it becomes apparent that by the time a majority of foragers
become sideworkers,most blossoms are already cross-pollinated(DeGrandi-
Hoffman et al. 1985).
B. Citrus
For many years almost all commercial citrus (Citrus spp.) cultivars
were either self-pollinating or produced parthenocarpic fruit. Bees were
unnecessary. The selection of self-incompatible chance seedlings and
interspecific hybrids necessitates insect-mediated cross-pollination in
some cultivars (Krezdorn 1970). For example, ‘Fairchild’tangerine (Cit-
rus reticulata) is self-incompatibleand requires cross-pollination for fruit
set (Moffett and Rodney 1971; Moffett et al. 1981).
Citrus flowers are highly attractive to bees because they produce
copious amounts of nectar (Vansell et al. 1942). Honey bees collect
pollen and nectar from citrus, and beekeepers often move colonies near
citrus groves because this nectar source produces a high-quality honey.
There are conflicting reports on the need for bees in some citrus
cultivars. Richter (1916)and Webber (1930)reported that bee pollination
is of negligible importance in lemon (Citrus limon), while Zavrashvili
(1967) and Moffett and Rodney (1975) increased lemon production on
trees that were open-pollinated or caged with bees, over those where bees
were excluded. F’rancke et al. (1969) stated that bees have no effect on
‘Valencia’ orange (Citrus sinensis) yields, but Cameron et al. (1960)
reported that fruit size and seed number are positively related, and that
cross-pollination with ‘Pearl’tangelo pollen improved fruit size and possi-
bly number of fruit. Cross-pollination is not needed in the production of
grapefruit (Citrusparadisi)(Coit 1915;Webber 1930;Soost 1963;Krezdorn
1970, 1972).
Because of inconsistent reports, it is difficult to make generalizations
about the need for bee pollination in citrus orchards. What can be
concluded is that depending on the cultivar and conditions a t the site,
insect pollination may increase fruit set, fruit size, and seed number.
C. Melons
Insect pollination is essential for the production of watermelon (Citrullus
lanatus) and muskmelon (Cucumis melo) (Bohn and Davis 1964; Mann
1953; McGregor et al. 1965). Watermelon plants are generally monoe-
cious, but sometimes bear hermaphroditic instead of pistillate flowers
(Rosa 1925; Goff 1937). Honey bees are often rented for commercial
watermelon production, because pollen must be transferred from stami-
nate to pistillate flowers (or anthers to stigma on the same flower). At
least eight bee visits are needed to optimize pollen loading on the stigma
( Adlerz 1966). If sufficient amounts of pollen are not deposited on every
stigma lobe, misshapen melons develop (Mann 1943).Adequate amounts
of pollen are also needed to maximize seed number, which is highly cor-
related to melon weight (Brewer 1974).
Most American muskmelon cultivars are andromonoecious, with sta-
minate and hermaphroditic flowers on the same plant. The ratio of
staminate to hermaphroditic flowers is fruit set dependent. The absence
of fruit stimulates hermaphroditic flowering ( McGregor 1951). Herma-
phroditic flowers are incapable of self-pollination, and fruit set does not
occur in the absence of pollinators (Mann 1953; Bohn and Davis 1964;
McGregor et al. 1965). Multiple bee visits greatly enhance melon size
and shape. McGregor et al. (1965)reported that to obtain high-quality
marketable melons, flowers ought to receive a t least 12 bee visits. Infield
placement of colonies a t a rate of 7.5 colonies/ha resulted in the highest
quantity and quality of melons, when compared with lower colony/hectare
rates and placement of colonies around the periphery of the field (Atkins
and Kellum 1979).
D. Berries
Honey bee pollination is required for the production of blueberries
( Vaccinium spp. ), strawberries (Frageria xananassa), raspberries (Rubus
spp. ), and cranberries (Vaccinium macrocarpon). Insect (primarily honey
bee) pollination is also needed for optimum highbush (V corymbosum)
and lowbush ( V angustifolium and V myrtilloides) blueberry produc-
tion. The highest quality fruit results from cross-pollination (Shaw et al.
1939; Aalders and Hall 1961; Eck and Childers 1966; Marucci 1966;
Martin 1966; Wood 1968). Blueberry growers have been advised to plant
a t least two cultivars to ensure that compatible pollen is available (Boller
1956; Darrow and Moore 1962). There is a positive relationship between
berry size and seed number (Brewer and Dobson 1969a). Flowers that are
cross-pollinated produce berries that are larger and earlier maturing than
those from self-pollinations (Morrow 1943; Martin 1966).Because 50430%
of the flowers must set berries for a good commercial crop (McGregor
1976),inadequate pollination can be considered a major constraint in the
production of some blueberry cultivars.
The rabbiteye (V ashei) blueberry cultivar ‘Tifblue’ can set fruit
parthenocarpically and from self-pollination ( Ambrose and Mainland
1979). Whatley and Lackett (1979)reported similar findings and added
that ‘Menditoo’( a rabbiteye cultivar ) can also set fruit parthenocarpically,
although a t a lower rate than ‘Tifblue’.
Honey bees discriminate between blueberry cultivars, and show strong
preferences for some over others (Rajotte and Roberts 1979). For exam-
ple, honey bees prefer ‘Rubel’ over the major commercial cultivar ‘Jer-
sey’ (Brewer et al. 1969). Studies were conducted to determine if nectar
7. HONEY BEE POLLINATION OF HORTICULTURAL CROP PRODUCTION 249
secretion and composition, time and rate of pollen production, or flower

morphology differed significantly between the cultivars. Interestingly,
nectar sugar concentration was higher in the less attractive ‘Jersey’than
‘Rubel’, although nectar volume was not significantly different. ‘Rubel’
blossoms produced more pollen. Differences in blossom morphology did
not seem to influence bee visitation rates (Brewer and Dobson 196913).A
similar study was conducted by Rajotte and Roberts (1979)comparing
‘Coville’(the preferred cultivar) to ‘Jersey’. The sugar content of ‘Coville’
nectar was significantly greater than ‘Jersey’particularly early and late
in the bloom period. ‘Jersey’showed a possible resorption of nectar sugars
late in bloom, which might be related to the setting of low-quality
parthenocarpic fruit.
The importance of bee pollination in the production of cranberries has
been reported by Gates (1911), Franklin (1911),Farrar and Bain (1946),
Marucci and Moulter (1977),and Kevan et al. (1983).Pollen is shed prior
to stigma receptivity, which occurs 24-36 hr after anther dehiscence.
When stigmata become receptive, stamens have usually stopped shed-
ding pollen (Rigby and Dana 1971).Hence, pollen must be moved between
flowers of different ages for berry set to occur. For an excellent discussion
on the energetics of honey bee and bumble bee foraging on cranberry, see
Roberts (1979).
Strawberry originally was a dioecious species, but today all commer-
cial cultivars produce hermaphroditic flowers ( Darrow 1966; Wilhelm
1974). The first flowers to open on a stalk (primary berry) have the
greatest number of achenes and produce the largest and highest quality
fruit, especially if flowers are cross-pollinated (Jaycox 1970). Some polli-
nation occurs by wind movement or gravity when anthers dehisce, but
the arrangement of stamens is such that pollen is not usually deposited
on all stigmata (Conner 1970). For maximum berry size, all stigmata
must be pollinated, and this requires insect pollination (Free 1968a).
When all stigmata are not pollinated, small irregularly shaped berries
develop (Free 196813; Nye and Anderson 1974). Berries produced from
cross-pollinations are usually larger than those from self-pollinations
(Jaycox 1970).
Many insects visit strawberry flowers, but only bees effectively transfer
pollen. Growers often introduce honey bee colonies into strawberry plantings,
because feral bee populations usually are not sufficient to supply flowers
with the 16-25 visits needed to maximize berry size (Skrebtsova 19571.
The benefits of honey bee pollination to strawberry flowers are well
documented (Skrebtsova 1957; Mommers 1961; Hughes 1961; Petkov
1965; Couston 1966; Rejput and Singh 1967; Free 1968a,b; Moore 1969;
Conner and Martin 1973; Lackett and Burkhardt 1979). Bee pollination
greatly improves berry set, weight, achene development, and berry shape.
Bee pollination is most important in cultivars with stamens that are

shorter than the receptacles, and least important in those with taller
stamens whose pollen can drop on to stigmata (Connorand Martin 1973).
Because strawberry flowers are not highly attractive, fields must be
saturated with bees (Skrebtsova 1958),and competitive plants kept to a
minimum to force foragers on to strawberry flowers. A list of references on
strawberry pollination work conducted prior to 1969 can be found in
Anderson (1969).
Raspberries were originally thought to be produced from cleistoga-
mous pollinations (Wellington 1913; Hardy 1931), but Johnson (1929)
showed significant increases in berry production between plants exposed
to pollinators and those where pollinators were excluded. Couston (1963,
1966) demonstrated that not only was berry number increased when
raspberry flowers were pollinated by bees, but berry size and weight were
also significantly increased, especially when flowers received multiple bee
visits (Eaton et aZ. 1968).Shanks (1969)concluded that honey bees were
by far the most abundant pollinating insect found on raspberries, and
probably the most important. Raspberry flowers are highly attractive to
bees, because they produce abundant amounts of nectar (about 13 mg
per flower) ( Haragsimova-Neprasova 1960; Petkov 1963) and pollen.
E. Nuts and Oilseeds

Most tree nut crops are anemophilous (e.g., pistachio, walnut, pecan).
Almond (Prunus dulcis)is not wind pollinated, but is completely depend-
ent upon insect movement of pollen. Almost all almond cultivars are
self-incompatible and require cross-pollination for nut set ( Kester 1969).
The honey bee is the only pollinating insect commercially used for
almond (McGregor 1976; Thorp 1979).Other bee species such as Osmiu
Zignaria can pollinate almond blossoms (Torchio1979),but their manage-
ment has not yet been developed for commercial orchard pollination.
Almond trees bloom very early in the spring, and in most areas are the
first commercial crop to produce the large amounts of pollen and nectar
needed for colony population increase. Almond trees are extremely attrac-
tive to honey bees (McGregor 1976; Thorp 1979).
Honey bees use different behaviors to collect pollen and nectar from
almond blossoms. As in some apple cultivars, pollen collectors climb on
to the anthers and almost always touch the stigma during a visit.
Foragers most often collect nectar by clinging to a petal and probing
between the bases of the stamens to the nectaries. In California orchards,
nectar collectors rarely contact stigma or stamens during a visit, and
usually do not pollinate blossoms (Thorp 1979).
While foraging almond, honey bees tend to confine their visits to small
areas during most of the foraging trip, until pollen and nectar rewards
become limited. When honey bees fly between trees, it is often between
trees of the same cultivar (Thorp 1979). Similar behaviors have been
observed in other crops (Free 1970).Honey bees are more likely to move
between trees of different cultivars when bee density is high, and food
resources are limited. Hence, cross-pollinations from tree to tree move-
ment occur in the later foraging hours each day, and early and late in the
bloom season (Thorp 1979).
Macadamia nut (Macadamia integrifolia and M. tetraphylla) also
require insect (primarily honey bee) pollination for set, because most
macadamia flowers are a t least partly self-sterile (Urata 1954; Schroeder
1959; Sedgley 1983).The anthers of macadamia flowers dehisce 1-2 days
prior to anthesis, but stigmata do not support pollen tube growth until
1-2 days postanthesis (Sedgleyet al. 1985).Honey bees pollinate madacamia
blossoms primarily while foraging blossoms for pollen (Urata 1954;Gary
et al. 1972).
Peanut (Arachis hypogaea) is largely self-fertilized (Stokes and Hull
1930), but bee visitation and subsequent pollination increases yields
(Girardeau and Leuck 1967). Peanut plots excluding insects had lower
yields, smaller seed size, and fewer seeds per pod than those that were
open pollinated by bees (Rashad et al. 1979a). Bee visitations occur
frequently on peanut flowers and a high level of flower “tripping” occurs.
Repeated tripping causes a compaction of pollen in the flower keel tip,
and a thrusting of the stigma through the mass causes ejection of pollen
through the tip opening. Thus, it appears that even though peanut
flowers are self-fertilizing, the tripping action of pollinators (such as
honey bees) assists and possibly increases self-pollination (Leuck and
Hammons 1965; Girardeau and Leuck 1967).
F. Vegetables
In crops such as cucumbers (Cucumis sativus) and squash (Cucurbita
sp.), insect pollination is essential for optimum fruit set and quality. For
other vegetables, pollination is needed for the production to seed. The
development of male-sterile lines and the exploitation of hybrid vigor has
made cross-pollination essential for production of seed for onion (Alium
cepa),carrot (Daucus carota), brussel sprouts (Bmssica oleracea gemmifera),
and other Cole crops (Brassica spp. ).Parsley (Petroselinum crispum) and
beans (Phaseolus spp., Vicia spp.) can also benefit from bee pollination.
Honey bee pollination is needed for cucumber (particularly pickling
cucumbers) production, because plants are usually monoecious (though
some cultivars are gynoecious), and pollen must be transferred between
male and female flowers for set to occur. Flowers are receptive to pollina-
tion for only one day (Connor et al. 1975). Bees that forage cucumber
flowers exhibit preferences that change during the day. Staminate flow-
ers are relatively more attractive than pistillate for most of the day,
probably because of their high nectar sugar concentration (Collison and
Martin 1979). Bees do not collect cucumber pollen in great amounts
(Olsen et al. 1979). After a visit, flowers will replace their nectar some-
times within 5 minutes. Flowers should receive a t least 10 visits to insure
good cucumber fruit set and shape (Connor et al. 1975).
During the 1960s the technology of cucumber production changed
from sequential hand pickings to once-overdestructive machine harvests
(Connor and Martin 1970; Connor et al. 1975). Hence, the timing for
introduction of colonies to maximize set over a limited period of time
became critical. Ideally, colonies should be introduced to cucumber fields
5-11 days after the first flowers appear (Connor and Martin 1971).Cucum-
bers are not highly attractive to bees, and so this delay should insure that
enough flowers exist to attract foragers. Seed blends of gynoecious and
monoecious cultivars to produce a pistillate to staminate flower ratio of
2 :1 will result in the highest potential fruit set and dollar yields (Connor
and Martin 1971).
Squash plants are usually monoecious, but hermaphroditic flowers
sometimes occur. Pollen transfer between flowers (orwithin hermaphroditic
flowers) is facilitated by many species of bees, including squash bees
(Peponapis spp., and Xenoglossa spp.), Bombus spp., Mellissodes spp.,
and most often honey bees (Jaycox et al. 1975).Honey bees forage squash
flowers most intensively in the morning particularly, between 8:OO and
9:00 (Sanduleac 1959).Fruit set rates and size increase with multiple bee
visits (Jaycox et al. 1975).
Onion is largely self-compatible, but relies on insects for pollination
(Free 1970). When insects are excluded, onion seed yields are reduced
substantially (Jones and Emsweller 1933;Jones and Davis 1934; Benedek
and Gaal1972). Many species of bees and flies are capable of pollinating
onion flowers. Honey bees forage onion almost entirely for nectar, but
become heavily dusted with pollen in the process (Benedak and Gaal
19721. Stigmata of onion flowers become receptive to pollen only after the
anthers have dehisced (Rodrigo et al. 1936; Parker and Hatley 1979).
The discovery of male sterility in onion and the subsequent production
of hybrid onion seed has increased the need for insect-mediated pollina-
tion of this crop. Higher pollinator populations are required to transfer
pollen effectively from male-fertile to -sterile lines. Individual honey bees
tend to restrict their visits to either male-sterile or -fertile lines, thus
reducing pollen movement to male-sterile flowers (Waller 1975).Nye et al.
(1971) reported that seed yields of male-sterile lines decrease as the
distance from the male-fertile lines increases.
Although onions produce sufficient amounts of nectar with sugar con-
centrations that are attractive to bees, foraging honey bees a t times

appear to avoid onion flowers. Apparently the cause for this avoidance is
that sometimes levels of potassium in onion nectar can be high enough to
reduce flower attractiveness. The presence of high potassium levels in
onion nectar occurs over a wide range of geographic, climatic, cultivar,
and cultural conditions, suggesting that high potassium may be charac-
teristic of the species (Waller et al. 1972).
Carrot flowers are highly attractive to various insects, and so pollina-
tion is not a problem in commercial seed fields of open-pollinated cultivars
(Jones and Davis 1944).Carrot is self-fertile,but autogamy is rare because
carrots are protandrous. Pollen dehiscence is usually complete before the
first stigma is receptive (Erickson et al. 1982).As in many other commer-
cial seed crops, honey bees are the principal pollinator of carrot (Bohart
and Nye 1960).
Although adequate pollination is easily achieved in open-pollinated
cultivars of carrot, production of seed from cytoplasmic male-sterile
carrots has been hampered by pollination problems ( Erickson and Peterson
1979). The ineffectiveness of honey bee pollination for hybrid carrot seed
production is largely due to inherent floral diversity between parents. The
loss of pollen from male-sterile lines results in loss of aroma and nectar
production for 5-10 days after anthesis. Hence, male-sterile lines produce
aroma and nectar later than male-fertile lines (Erickson et al. 1979;
Erickson and Peterson 1979), causing as much as a sixfold increase in
honey bee visitation to male-fertile lines (Erickson et al. 1982). There is
also a wide range of variability in the onset and duration of floral events
between male-sterile and -fertile lines, which leads to an asynchrony of
pollen dehiscence and stigma receptivity. Nectar quantity can also differ
significantly between the two lines, causing honey bees to confine their
foraging to the most rewarding lines (Erickson and Peterson 1979).
Because of differential attractiveness, pollen transfer from male-fertile
to -sterile lines usually ranges between 3.3 and 10% (but has reached
23.6%) (Erickson et al. 1982).
Honey bees are also important in the production of seed for hybrid
brussel sprout cultivars. Pollen can be transferred between flowers on the
same plant, within the same cultivar, and between cultivars. Plants
produced from self-pollination lack vigor and give reduced yields of
marketable sprouts ( Faulkner 19761. Thus, honey bee cross-pollination
and resulting hybrid seed production are imperative for maintenance of
high-quality brussel sprout production. Cross-pollination may be limited
and hybrid seed production reduced because honey bees discriminate
between brussel sprout cultivars based on plant height (Faulkner 1976)
and flower color (Faulkner 1976; Free and Williams 1983). When plant
height between cultivars differs but flower color is the same, the rate of
honey bee movement in self- vs. cross-pollinating directions is 23 : 1.
When plant height is the same but flower color between cultivars differs,
the self- to cross-pollination ratio is 33 :1 (Faulkner 1976).These results
suggest that to optimize honey bee cross-pollination between inbred
lines, plants should be of similar height and flower color.
Parsley has protandrous flowers (Knuth 1908),with anther dehiscence
occurring 1-2 days prior to stigma receptivity (Burgett 1980).Pollinator
exclusion demonstrates the need for insect pollination in seed produc-
tion. Honey bees and syrphid flies are the most common pollinators of
parsley, with pollen-collecting honey bees the more efficient pollinators
(Burgett 1980).
Honey bee pollination can increase yields of lima beans (Phaseolus
lunatus), and broad beans (Vicia faba). Lima beans (unlike many other
beans) secrete large amounts of nectar, making this plant attractive to
bees. Honey bees visit lima bean blossoms throughout the day (Vansell
and Reinhardt 1948) and can significantly increase yields over those
where pollinators are excluded (Amos 1943).Likewise, broad beans caged
with bees produce more seeds per pod and more seeds per plant than
those without bees (Free 1966~).
G . Tropical Crops
Honey bees play an important role in the production of several tropical
horticultural crops. Plants such as coconut (Cocus nucifera), litchi (Litchi
chinensis), kiwifruit (Actinidia chinensis), and papaya (Carica papaya)
require bees to transfer pollen between their staminate and pistillate
flowers. Others such as cashew (Anacardium occidentale),mango (Mangifem
indica), and guava (Psidium guajava) have hermaphroditic flowers, but
need bees to transfer pollen from the anthers to stigma. Crops such as
sesame (Sesamum indicurn) and certain coffee species (coffea spp.) are
either wind or self-pollinating, but set larger crops when flowers are
pollinated by foraging bees. Apis mellifera is not present in some tropical
areas, but crops are pollinated by other honey bee species: A. florea, A.
dorsata, and A. cerana.
Coconut is monecious, but can set fruit if pollen is transferred among
flowers on the same tree (Sholdt and Mitchell 1967).Wind, insects, and
birds have been mentioned as pollinating agents (Davis 1954; Kidavu
and Nambiyar 1925). Free et al. (1975)reported that honey bees collect
both pollen and nectar from coconut flowers,pollinating them in the process.
Litchi plants can have staminate, pistillate, and sometimeshermaphroditic
flowers (Butcher 1957).Staminate flowers appear first, and in some years
and cultivars are the only flower type produced. Reasons for this are not
known. In Florida, flies and honey bees are the most common pollinators
of litchi flowers, but in the Indian Himalayas A. florea is the most
abundant pollinator. Apis cerana, A . dorsata, and Vespa spp. are also
litchi pollinators in the Himalayas (Dhaliwal et al. 1977).
Kiwifruit (Chinese goosberry ) is produced on dioecious shrubs, with
vines that are usually trained upon a trellis in commercial plantings
(Bailey and Topping 1950). Wind and insects are the most important
means of pollen exchange between plants. Honey bees are generally relied
upon for pollination of commercial plantings. In tests where honey bees
were excluded, only 20% of the fruit was of marketable size, compared to
90% on limbs exposed to honey bees (Palmer-Jones and Clinch 1975).
Similar results were obtained by Marletto (1980), who reported larger
fruit with more seeds and lower fruit abscission rates on branches exposed
to honey bees and other pollinators. Kiwifruit flowers are not highly
attractive to honey bees, and so competitive plants should be kept to a
minimum in and around orchards during bloom (Clinch 1984).
Papaya trees can produce staminate and pistillate flowers on the same
tree or in some cases on separate trees. Sometimes trees change from
producing one type of flower to another, or from male to hermaphroditic
flowers (Free 1970).There is also variation among hermaphroditic flowers
(Storey 1937,1941,1958).Although there are some parthenocarpic culti-
vars (McGregor 1976), and hermaphroditic flowers can set fruit even if
pollinators are excluded (Harkness 1967),generally pollen must be trans-
ferred between flowers for fruit set to occur. Papaya flowers can be polli-
nated by wind and insects such as bees and moths (Story 1941; n a u b et
al. 1942;Marin-Acosta 1963).Allen (1963)reported that honey bees were
the primary pollinating agents of papayas in South Africa.
Flowers on cashew trees are very attractive to bees because they
produce abundant nectar (Morton 1961) and have an extremely strong
floral scent (Free 1970).Inflorescences contain male and hermaphroditic
flowers. Hermaphroditic flowers are self-compatiblebut not self-pollinating
(Northwood 1966). Flies and ants are the primary pollinators of cashew,
but Polistes wasps (Polistes spp.) and honey bees also forage flowers.
Cashew production can be increased by introducing honey bee colonies
(Free and Williams 1976).
Mango is similar to cashew in that trees produce staminate and
hermaphroditic flowers that have a strong fragrance and produce abun-
dant nectar (Mukherjee 1953).Mango flowers are not particularly attrac-
tive to honey bees. Anthesis occurs in the early morning, and stigmata
are receptive when flowers open (McGregor 1976). Insects, particularly
bees, are needed to transfer pollen. Yields are significantly lowered if
pollinators are excluded (Free and Williams 1976).Some mango cultivars
are self-compatible (Young 1942), while others set fruit only if cross-
pollinated (Singh et al. 1962).
Guava is a shallow-rooted, many-branched shrub or small tree that
produces hermaphroditic flowers. The stigma is positioned above the

anthers, and so insects are needed to transfer pollen (Hamilton and
Seagrave-Smith 1954). Bees visit flowers for nector and pollen, and
although flowers are self-compatible, cross-pollination gives higher fruit
yields (Malo and Campbell 1968; Hamilton and Seagrave-Smith 1954).
In some areas honey bees are the primary pollinating agents (Soubihe
and Gurgel 1962).
About 90%of the coffee produced worldwide is Coffee arabica ( McGregor
1976).This species produces self-compatible hermaphroditic flowers that
are capable of self-pollination (Wellman 1961). Yields are increased, how-
ever, when honey bees are present (Raw and Free 1977).W o other species
of coffee are grown commercially but to a lesser extent: C. liberica and C.
robusta. Both are self-sterile and pollinated by wind and insects (Carvalho
and Krug 1950).
Sesame is a self-pollinatedcrop, but seed yields and quality are increased
by insect pollination (Rashad et al. 1979b). Syrphid flies, Vespid wasps,
and Halictid and Anthophorid bees can pollinate flowers, but if sufficient
populations of these insects are not present, honey bee colonies should be
introduced. In many areas, honey bees are the most common foragers on
sesame flowers (Langham 1944; Rashad et al. 1979b).
111. MECHANISMS FOR CROSS-POLLINATION
Crops such as apple, almond, cucumber, and sweet cherry present

special pollination problems, because pollen must be transferred between
relatively large plants for fruit set to occur. Honey bees are conservative in
their foraging movements, preferring to establish foraging areas of lim-
ited size when flowers are rewarding (Phillips 1930a; Butler et al. 1943;
Ribbands 1949; Singh 1950; Moffett et al. 1974). Honey bees expand
their foraging areas when little nectar or pollen is available (Ribbands
1949; Weaver 1957; Free 1960). In apple orchards with dwarf trees, bees
visit flowers along about 3 m of a continuous row during a foraging trip.
Relatively few foragers move from one row to another (Free and Spencer-
Booth 1964a). In orchards of standard-sized apple trees, most honey bees
establish foraging areas on a single tree, and a t best visit two adjacent
trees (usually in the same row) on a foraging trip ( h e 1966a). Honey
bees also can discriminate between some apple cultivars, and if the
cultivar retains or increases in attractiveness, most of its foragers will
demonstrate fidelity to it (Free 1966a). These behaviors would lead to
more self- than cross-pollinations.
Although most honey bees limit their foraging areas, and demonstrate
species fidelity, competition among foragers or inconsistent rewards due
to waning bloom cause some honey bees to move from plant to plant to
obtain a full nectar or pollen load. If movement is between different plants
(or in self-incompatible plants, different cultivars) of the same species,
fruit set can occur. Evidence for cross-pollinations resulting from tree to
tree movement in fruit orchards has been reported by Williams ( 1959),
Free (1962),and Free and Spencer-Booth (196413).In these reports, trees
adjacent to pollinizers set significantly more fruit, in some instances,
than those two or more rows away. The higher fruit set was attributed to
honey bees having a greater likelihood of moving to an adjacent row (i.e.,
from a pollinizer to an adjacent main cultivar row) to find a foraging area,
then wandering several rows away.
Although tree-to-tree movement can explain greater fruit set on trees
adjacent to pollinizer rows, it does not account for how fruit set occurs in
trees surrounded by incompatible pollen sources. In addition, fruit set
does not always decrease with distance from pollinizer trees (Zibles 7.2
and 7.3) (Free and Spencer-Booth 1964b; DeGrandi-Hoffman et al. 1984).
Table 7.2 Mean Percentage of Flowers That Set

Fruit (Initial S e t ) on Dwarf Pear ’Ikees a t Different
Distances from P e e s with Pollinizer Graftsa
Flower set ( % )
Distance from
nearest tree ( m ) Cornice Conference
1.2 9.4 69.5

2.4 9.4 75.5
3.7 7.5 60.9
4.9 8.9 65.5
6.0 9.8 69.2
“From Free and Spencer-Booth ( 1964b).
Table 7.3. Influence of Distance from the Pollinizer Row on ‘Delicious’ Fruit Set
in Two Michigan Orchards“
Number of rows away Blossoms setting fruit

Site from pollinizer Number of limbs (70)(mean 2 S D ) b
Orchard A Adjacent 15 19.8 f 3.0

1 10 16.5 k 4.0
Orchard B Adjacent 16 16.4 i 3.0
1 16 15.0 -+ 3.0
2 16 17.0 * 3.0
3 16 *
14.8 3.0
4 8 *
16.7 4.0
“From DeGrandi-Hoffman et a[. (19841.

*No significant differences between means within sites by F test.
Honey bees carry many types of pollen on their bodies (Free and Williams
1972; Kendall and Solomon 1973; DeGrandi-Hoffmanet al. 1984), although
most collect pollen from only one species on a foraging trip (Betts 1920,
1935; Brittain and Newton 1933,1934; Free 1963). These incongruencies
suggest that honey bees acquire compatible pollen (and consequently
cross-pollinate flowers) by a means other than plant-to-plant movement.
The suggestion that honey bees may acquire pollen in the hive from
nestmate contacts has been made by several authors (Betts 1920; Karmo
and vickery 1954; Lukoshus 1957; Free and Williams 1972; DeGrandi-
Hoffman et al. 1984, 1986). Evidence for in-hive pollen transfer has been
supplied by Free and Williams (1972),who placed newly emerged pollen-
free bees in colonies and later examined them for pollen. Results of these
experiments are summarized in lhble 7.4. Similar experiments were
conducted by DeGrandi-Hoffman et al. (1986), who confirmed Free and
Williams' (1972)results and demonstrated that honey bees could acquire
enough viable compatible pollen from nestmate contact to set fruit on
apple trees. Thus, it appears that cross-pollinations may result from
in-hive pollen transfer as well as compatible pollen acquisition from tree-
to-tree movement.
Cross-pollination from tree-to-treemovement is apparently influenced
by competition for nectar and/or pollen, and a plant's distance from its
nearest compatible pollen source. Constraints on cross-pollinations from
in-hive pollen transfer are the availability of compatible pollen, and the
percentage of a colony's foraging population collecting pollen from that
source. The more compatible pollen entering the hive, the greater the
chances of it being transferred in amounts sufficient for foragers to
cross-pollinate the first few flowers visited on a foraging trip.
liable 7.4.
Amount of Pollen That Accumulated on the Bodies of Dead Bees
Exposed in the Hive"
~
Exp. 1 (19-20 June), Exp. 2 (17 July),

bees exposed in: bees exposed in:
Number of pollen Storage Brood Storage Brood

graindbee chamber chamber chamber chamber
0 37 14 5 3
c 1,000 27 8 11 7
1,500-5,000 5 2 5 5
5,500-10,000 2 3 0 0
11,000-50,000 6 7 0 3
51,000-100,000 1 3 0 0
110,000-150,000 1 0 0 0
2 150,000 0 1 0 0
Mean 5437 8205 857 4111
"From Free and Williams (1972).

IV. CONCLUDING REMARKS
Current agricultural practices have caused an acute need for honey bee
pollination. Large-scale monocultures, reliance on pesticides for crop
protection, and cultivation practices that reduce foraging areas and
nesting sites for feral pollinator populations in many areas have made it
imperative that honey bee colonies be introduced. Agriculture is experi-
encing a steady decline in colonies for pollination. Pesticide contamina-
tion (see Erickson et al. 1983; Erickson and Erickson 1983a,b, 1984;
Crane and Walker 1983; Johansen 1977),the entry of the tracheal mites
(Acarapis woodi Rennie) into the United States (see DeJong et al. 1982),
and the inevitable migration of the Africanized bee (A. mellifera scutelhta)
from South America (see Michener 1975; Taylor 1985) along with its
parasitic mite ( Varroa jacobsoni Oudemans) will reduce the number of
colonies available for pollination. The profit margin on the wholesale
price of honey also influences the availability of bees for pollination
(Taylor 1985). The American beekeeping industry is supported largely
from honey sales, but imports of less expensive foreign honey will make it
difficult for this relationship to continue. Yields of crops discussed here
will undoubtedly reflect shortages of colonies for pollination, particularly
in areas where native bees are scarce.
Future efforts in pollination research need to be directed a t increasing
the pollinating efficiency of colonies. This will require an interdisciplinary
effort among apiculturists and floral biologists, ecologists, horticulturalists,
plant breeders, and pest management specialists. If bee-pollinated crops
are to be managed for optimum production, the relationship between
insect pollination and plant reproduction needs to be considered. This
review has concentrated on pollination, but plant-based factors such as
pollen viability, and stigma and ovule receptivity determine whether
seed will set. The time during a plant’s bloom period when pollination will
most likely lead to ovule fertilization and seed set needs to be determined
for bee-pollinated crops, so that colonies can be strategically introduced.
Crops that rely on cross-pollination need to be attractive and rewarding
to pollinators. Compatible pollen sources must be equally attractive, and
bloom in synchrony with the main cultivar. Frequently plant-breeding
procedures leading to the development of hybrid seed parents have altered
nectar secretion, flower aroma, synchrony of floral events, i n d visual cues
used by pollinators to locate rewards (Erickson et al. 1982). Factors
affecting cross-pollination rates must be considered during plant selec-
tion and cultivar release, or the best traits of plant quality, resistance
(to insects or diseases), or vigor will be lost due to inadequate fruit or
seed set.
Pollinators must be recognized as integral components of crop produc-
tion systems, since their presence strongly affects crop size and potential
260 G. DEGRANDEHOFFMAN
value. This review has concentrated on honey bee pollination of crops, but
there are many other species of pollinators. At present, the contributions
of native bees to pollination has been suppressed because crop protection
measures have not been sensitive to the survival of feral bee populations.
If honey bee colony availability becomes limited, pest management prac-
tices that are compatible with the survival of native bees will be essential.
An understanding of pest management in relation to pollinator preserva-
tion will require communication between pollination ecologists and pest
management specialists.
While problems facing the beekeeping industry in the United States
may limit colony availability, we hope they will spur the beginning of
a new era of compatibility between pollinator population maintenance
and crop protection. Cultural practices that conserve pollinators will
help ease the stress shortages in colony availability will have on crop size
and quality, and insure the productivity that is vital for sustaining
agroecosystems.
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8
Tissue Culture of Temperate Fruit
and Nut Trees*
James R Hutchinson
Department of Agriculture and Rural Affairs, Horticultural
Research Institute,
Knoxfield, Ferntree Gully, 3156 Victoria Australia
Richard H. Zimmerman
Fruit Laboratory, United States Department of Agriculture,
Agricultural Research Service, Agricultural Research Center,
Beltsville, Maryland 20705
I.Introduction 273
11.Micropropagation 274
A . General Methodology 275
B . n e e Fruits 281
C . n e e Nuts 310
D. Problem Areas 315
111. Virus Elimination 318
IV. Genetic Improvement 318
A . Regeneration through Adventitious Pathways 319
B . Protoplast Culture 322
C . Haploids 323
D. Embryo Culture 324
V. Germplasm Conservation 324
VI. Commercial Application 325
VII. Conclusions 327
I. INTRODUCTION
Tissue culture has a number of potential uses for temperate fruit and
nut tree species. These include the micropropagation of rootstock or scion
cultivars in short supply, resulting from breeding or virus elimination
*We would like to thank t h e word processing section of the Department of Agricul-
ture and Rural Affairs and Pauline Nichols for typing the first draft and Julie Stubbs
for final typing. Sylvia Wilson assisted with the literature research.
Mention of a trademark, proprietary product, or vendor does not constitute a
guarantee or warranty by the Department of Agriculture and Rural Affairs, Victoria,
or the United States Department of Agriculture, and does not imply their approval to
t h e exclusion of other products or vendors t h a t may also be suitable.
273
274 JAMES E HUTCHINSON AND RICHARD H. ZIMMERMAN
programs; virus elimination; genetic improvement through in uitro mutant

selection, and the regeneration of plants from protoplasts or callus via
somatic embryogenesis or androgenesis ( Ramming 1983; Zimmerman
1983a);and finally to provide material for physiological and biochemical
studies and insight into the processes examined in such studies.
Micropropagation represents the greatest use of tissue culture, espe-
cially for rootstocks. For many temperate fruit and nut species, orchards
are planted with trees propagated by budding or grafting the desired
scion cultivar onto a rootstock. Rootstocks are used because (1)many
scions are difficult to propagate by cuttings, ( 2 )they can confer dwarfing
or vigor characteristics to the scion, or (3) they have resistance to soil
pathogens, insects, or stress. With experimental systems tending toward
ultra-high-density plantings, such as meadow orchards ( Luckwill and
Child 1973; Erez 1982) and 'Ihtura trellis (Chalmers et al. 1978),budded
or grafted trees may be too expensive and the use of these planting
systems may come to depend on alternative means of propagation.
General reviews on temperate fruit tm tissue culture have been published
(Skirvin 1981;Jones 1983),on the propagation of deciduous fruit and nut
tree species (Boxus and Druart 1985; Lane 1982; Hammerschlag 198613;
Zimmerman 1986),and on the following specific crops or groups of crops:
apple (Zimmerman 1984a; Skirvin et al. 1986),pear (Singha 1986),cherry
(Ivanicka and Pretova 1986),peach (Hammerschlag 1986a),plum (Druart
and Gruselle 1986),stone fruits (Skirvin 1984),and olive (Rugini 1986).
There are also two relevant conference proceedings (Zimmerman 1980;
Boxus 1981).
11. MICROPROPAGATION
A range of tissue culture techniques have been developed that will

effectively achieve rapid propagation. The various procedures to produce
plants through in uitro techniques are illustrated in Fig. 8.1 and have
been reviewed a number of times (de Fossard 1981; Hussey l978,1983a,b;
Lawrence 1981; Bonga and Durzan 1982; Krikorian 1982; Bhojwani and
Razdan 1983; Hu and Wang 1983; Styer and Chin 1983; George and
Sherrington 1984).The method of choice to maintain the clonal integrity
of a species is axillary shoot proliferation, in which shoots develop from
preexisting vegetative meristems. Murashige ( 1974, 1977, 1978) has
described micropropagation as involving three stages: stage I, the estab-
lishment of aseptic cultures; stage 11, shoot proliferation; and stage 111,
shoots ready for establishment in potting medium. Debergh and Maene
(1981) have proposed a four-stage process: stage 0, the preparation of
stock plants under hygienic conditions; stage I, the establishment of
8. TISSUE CULTURE OF TEMPERATE FRUIT AND NUT TREES 275
EXPLANT
PROTOPLASTS
REGENERATED
Fig. 8.1. Morphogenetic pathways to produce plants in tissue culture.
aseptic cultures; stage 11,the induction of meristematic centers and their

development into rapidly proliferating buds; stage I1Ia, elongation of
buds to form shoots and the preparation of uniform shoots; stage IIIb,
the rooting and initial growth of the in vitro produced shoots, usually in a
soilless potting mixture. The majority of fruit and nut species have been
rooted in vitro.
A. General Methodology*
1. Disinfestation and Explantation. The shoot-tip or lateral bud is the
explant of choice for clonal propagation in vitro. Suitable growth can be
achieved, provided the chilling requirement, if necessary, has been satis-
fied. Washing explants under running tap water for 30-90 min can assist
in obtaining sterile cultures by physically removing some of the contami-
nants, especially with field-grownmaterial (de Fossard 1976;Jones et al.
1979).Surface sterilants that have been most commonly used are calcium
or sodium hypochlorite (Jones 1983).Calcium hypochlorite is used a t con-
centrations from 4 to 8% (w/v) and sodium hypochlorite a t about 0.5%.
Other surface sterilants such as mercuric chloride have been less com-
monly used ( Borkowska 1983). After appropriate surface sterilization
and rinsing in sterile water, the explant is isolated. Explant size varies
from the meristem-tip with one or two leaf primordia (Walkey 1972) to
apical shoot-tips 2-3 cm in length (Snir and Erez 1980).Contamination is
less of a problem with smaller explants, but successful establishment of
cultures is more difficult compared to larger explants where the reverse
*Some authors use mass and others molar units for hormone concentrations. Molar
units have the advantage that they allow direct comparison of hormone types. If
authors have used mass units they have been converted to molar units and rounded-
off and placed in parentheses. See Thble 8.1 for abbreviations used.
Table 8.1. Abbreviations Used in This Chapter
Media Auxins
AS Zhao ( 1982) IAA Indole-3-acetic acid
B-5 Gamborg (Gamborg et al. 1968) IBA Indole-3-butyric acid
D Druart ( 1980) NAA a-Naphthalene acetic acid
DKW Driver and Kuniyuki ( 1984) NOA /j-Naphthoxy acetic acid
LS Linsmaier and Skoog ( 1965) 2,4-D 2,4-Dichlorophenoxy
MS Murashige and Skoog (1962) acetic acid
PM Fiorino and Leva ( 1983)
Other media components
WPM Woody plant medium
PG Phloroglucinol
(Lloyd and McCown 1980)
GAS Gibberellic acid
Cytokinins
BA Benzyladenine
2iP iso-Pentenyladenine
4-PU N-(2-Chloro-4-pyridy1)-
N-phenylurea
applies. Details for isolating shoot-tips and meristem-tips are described

in Broome and Zimmerman (1984).
2. Shoot Proliferation. The proliferation of shoots for micropropagation
is derived from the research of Wickson and Thimann ( 1958) and Sachs
and Thimann ( 1964),who found that exogenous application of cytokinins
could release lateral buds from apical dominance, and from Morel’s ( 1964)
discovery that Cymbidium orchid buds could be dissected from pseudobulbs
and induced to form continuously multiplying protocorms that could be
maintained more or less indefinitely. Since the discovery that BA could
stimulate shoot elongation (Jones 1967)and shoot proliferation (Pieniazek
1968),this cytokinin has been routinely used for temperate fruit and nut
tree species. Zeatin is the preferred cytokinin for Olea europaea (Rugini
and Fontanazza 1981) and Diospyros KaKi (Cooper 1983; Cooper and
Cohen 1984).Occasionally combinations of cytokinins have been benefi-
cial, e.g., with Corylus avellana (Anderson 1983) and Pyrus communis
(Shen and Mullins 1984). Other factors influencing shoot proliferation
include light level (Hutchinson 1984a),explant type (Hutchinson 1984a;
Shen and Mullins 1984, agar concentration (Singha 1982, 1984), inver-
sion of explant (Lane 1979; Vieitez and Vieitez 1983),and the use of liquid
medium (Snir and Erez 1980).
3. Rooting. Adventitious rooting involves two stages: the initiation
phase, usually in response to an auxin, and an elongation phase (Haissig
1974; Mohammed and Eriksen 1974; Smith and Thorpe 1975;Mitsuhashi-
Kato et al. 1978). I n uitro rooting of fruit trees has been most widely
studied with Malus cultivars. Initially, only an auxin-enriched medium
was used (Jonesand Hatfield 1976; Jones et al. 1977),although problems
8. TISSUE CULTURE O F TEMPERATE FRUIT AND NUT TREES 277
associated with excessive callus formation occurred (Jamesand Thurbon

1979). Another approach was to use two rooting media, one with auxins
for initiation and another, usually hormone free, for root elongation
(Jones et al. 1979; James 1983a). A medium with a composition that
stimulates root initiation, does not inhibit root elongation, and does not
enhance callus formation has been used successfully ( Zimmerman and
Broome 1980; Hutchinson 1984a). Most recently rooting and acclimati-
zation as one process in potting medium is being employed (Simmonds
1983; Zimmerman and Fordham 1985).
Many factors have been studied that can influence rooting in vitro for a
range of species, e.g., salt concentration of the medium (Welander 1983;
Simmonds 1983),auxin type and concentration (Lane 1979; Sriskandarajah
and Mullins 1981; Barghchi and Alderson 1983a), temperature (Lane
1978; Rosati et al. 1980; Zimmerman 1984b), sucrose concentration
( Sriskandarajah and Mullins 19811, wounding the base of the stem (Snir
and Erez 1980; Snir 1984),and the physical support on which explants are
grown (Zimmerman and Broome 1980; Hutchinson 1984a). It is known
that some phenolic compounds, especially when applied together with an
auxin, can stimulate rooting (Goodwin 1978).Jones ( 1976)and Jones and
Hatfield (1976)first showed with apple shoots in vitro that the addition
of 1mM PG or phloretic acid with IBA more than doubled the percentage
of rooting in the apple rootstock M.7. Since then it has been studied in a
range of species with variable results (Jones 1983).While a large range of
variables have been investigated that affect rooting in vitro, not a great
deal is known about the process. Working with the difficult to root apple
cultivars ‘Jonathan’ and ‘Delicious’,Sriskandarajah et al. ( 1982)noted an
increase in rooting as the number of shoot proliferation cycles increased,
but only when cultures were grown a t 26 * 2OC with continuous illumina-
tion. This increase in rooting response was associated with a subsequent
decrease in levels of gibberellins in shoots (%keno et al. 1982/83).
Gaspar ( 1981) developed a model system using the variation of peroxidase
activity as a marker of the rooting process. During the inductive phase
total peroxidase activity of the whole cutting increases, with root primordia
initiation occurring following the peak of activity. The technique com-
prises evaluating chemical (e.g., auxin) and/or physical (e.g., environ-
ment) factors that favor an increase of peroxidase activity during the
inductive and a decrease during the initiation phases, and has been of
value in improving the rooting efficiency of Prunus persica (Mosella Ch.
and Macheix 1979), Malus ‘Jonagold’ (Druart et al. 1982), and Cynara
scolymus (Moncousin and Gaspar 1983).With ‘Jonagold’shoots, the use
of continuous darkness during both phases, instead of the commonly
used 16 :8 hour (light :dark) photoperiod, favored alteration in peroxidase
activity and induced a higher percentage of rooted plantlets.
278 JAMES F. HUTCHINSON AND RICHARD H. ZIMMERMAN
4. Acclimatization. Micropropagation depends upon the ability to

transfer plantlets to potting medium and to acclimatize them success-
fully to free living conditions. Many researchers have not gone beyond the
demonstration that a few plants produced from tissue culture would
survive ex vitro, and descriptions of the techniques used for acclimatiza-
tion are often minimal. Although acclimatization has not been the object
of careful study, the potential problem areas have been pinpointed and are
primarily associated with poor water relations (Grout and Aston 1977a;
Wardle et al. 1979,1983b;Conner and Thomas 1981; Conner and Conner
1984). Epicuticular wax is reduced or absent on the leaves of in vitro
cultured plants compared to glasshouse- or field-grown plants (Grout
1975; Sutter and Langhans 1979, 1982; Conner and Conner 1984). Dur-
ing acclimatization, the density of waxes increases as the humidity is
lowered (Wardle et al. 1983b).The wax component determines the rate of
water loss through the cuticle (Denna 1970; Martin and Juniper 1970)
and the susceptibility of tissue-cultured plants to desiccation has been
attributed to a reduction or absence of wax. In a comparison of the
chemical composition of epicuticular wax of micropropagated and green-
house plants of Brassica oleracea (capitata group) ‘Market Prize’, Sutter
(1984)found a greater percentage of polar compounds (fatty acids, pri-
mary alcohols, aldehydes, and esters) and fewer alkanes and secondary
alcohols in wax from micropropagated plants. The composition of the
wax may also affect water loss from plants as the wax components most
permeable to water flow are those containing polar functional groups:
fatty acids, alcohols, and aldehydes (Bakerand Bukovac 1971;Grncarevic
and Radler 1967).
Stornatal functioning has considerable bearing on successful acclima-
tization. Brainerd et aL ( 1981)showed that excised leaves of micropropagated
Prunus insititia ‘Pixy’ lost more than 50% of total leaf water content
within 30 min compared to those of greenhouse plants, which lost 50% of
their water content in 90 min. Furthermore, the water loss from leaves of
cultured plants was entirely abaxial (Fuchigami et al. 1981). Additional
studies with cultures of ‘Mac 9’ apple rootstock showed that water loss
was linearly related to stornatal closure (Brainerd and Fuchigami 1981);
plantlets were grown a t 30-40% relative humidity for 6 days: for the first 3
days stornatal closure was low and water loss high, but after 4 days the
stornatal closure mechanism developed. Later, Brainerd and Fuchigami
(1982) showed that treatments known to induce rapid stornatal closure,
such as darkness, mannitol-induced water stress, ABA, and increased
COaconcentration, were not successful with leaves from cultured plants.
Delayed stornatal closure has also been found in newly transferred B.
oleracea Botrytis group (cauliflower)(Wardle et al. 1979) and Chrysan-
themum morifolium (Wardle and Short 1983).The exact reasons for lack
8. TISSUE CULTURE OF TEMPERATE FRUIT AND N U T TREES 279
of stomatal closure in vitro are not known; however, it has been found
with cultured cauliflower that the concentration of potassium relative to
other elements was reduced in guard cells (Wardle et al. 1979, 1981)and
since potassium is important for stornatal function (Outlaw 1983),it may
be involved. In addition, there may be carryover effects of cytokinins,
since they promote stomatal opening (Jewerand Incolll980; Zeiger 1983).
Other differences in development of micropropagated plants have been
noted. Grout and Aston (1977a)found excessively high water loss from
B. oleracea, which they attributed to reduced wax and incomplete vascu-
lar development between roots and the shoot. Leaf anatomical studies of
micropropagated plants have revealed altered palisade and spongy meso-
phyll cell layers with large air spaces compared to established micro-
propagated and greenhouse or field-grown plants for a range of species
including I? insititia ‘Pixy’(Brainerd et al. 1981),Liquidambarstyraciflua
(Wetzstein and Sommer 1982),and Rubus idaeus (Donnelly and Vidaver
1984a),which can result in excess water loss.
Another problem associated with micropropagated plants is that they
are heterotrophic. During culture of cauliflower it has been shown that
there is no net COz uptake and very low levels of photosynthesis, with
growth totally dependent on an exogenous carbon source in the medium.
I t was not until 14 days after transfer that they became autotrophic
(Grout and Aston 1977b; Grout and Crisp 1977). Fkduced levels of COZ
uptake have also been found with R. idaeus and one month after trans-
plantation leaves retained from culture contributed less than 10%of the
COz uptake (Donnelly and Vidaver 1984b). Wardle et al. (1979)postu-
lated that leaves formed during culture are only a transitionary form and
act as storage organs until leaves with normal characteristics have been
produced. To test this, Wardle et al. ( 1983a)prepared shoot cultures of C.
morifolium ‘Snowdon’containing rubidium as a tracer for potassium and
found that each successive new leaf formed after transplantation con-
tained proportionately less rubidium.
Since both the anatomy and morphology of leaves and the physiology
of plants in culture are different from those of normally cultured plants, it
is important to consider these differences when acclimatizing plantlets.
The most commonly adopted procedure is to remove the plantlets from
the culture vessel, gently wash off the agar, if present, transfer to potting
medium, and maintain them under high relative humidity, usually achieved
by frequent misting, until growth has commenced (Dunstan 1981; Dunstan
and mrner 1984).
Antitranspirants may be of use during acclimatization. Wardle et al.
( 1979)evaluated a polyvinyl resin (S600)applied immediately after trans-
planting cauliflower. They found that treated plantlets had higher
cuticular resistance, but that there was no effect on stomatal resistance.
These reasons may explain why Hutchinson (1984b) found applications

of Folicote ( a hydrocarbon wax) to be useful for acclimatization of Malus
‘Northern Spy’. Folicote was applied immediately after transferring plantlets
to either coarse sand :peat moss ( 3: 1, v/v) or vermiculite :perlite ( 1 : 1,
v/v) and placing them in a greenhouse. Untreated plantlets were kept
under intermittent mist in a greenhouse. High survival rates were obtained
both with potting media with Folicote and with vermiculite :perlite under
mist, indicating that potting medium is important. Evaluating a number
of antitranspirants applied to C. morifolium and Dianthus caryophyllus,
Sutter and Hutzell (1984) determined that most were ineffective due
principally to phytotoxicity a t recommended concentrations and lack of
effect at lower concentrations. Most antitranspirants reduced plant growth
and photosynthesis as well as transpiration (Davies and Kozlowski 1974;
Davenport et al. 1974). Although antitranspirants have been ineffective
when used in conjunction with high-humidity tents (Sutter and Hutzell
1984),they may be effective if applied to plantlets transferred directly to
the greenhouse (Hutchinson 198413). I t is important to note their poten-
tial phytotoxicity and species differences.
It has been observed that I! insititia ‘Pixy’ growth was poor after
acclimatization. The problem could be overcome either by chilling rooted
plantlets in uitro at O°C for 2 months or by spraying with 200 mg/liter
(0.6 mM) GA3 after transfer (Howard and Oehl 1981).
Mycorrhizal fungi are beneficial symbiotic microorganisms that increase
growth and yield of most crop plants through increased nutrient uptake
(Harley 1970; Menge 1983). Their use in the establishment of micro-
propagated plants has not been extensively studied. When Wooi et al.
( 1982) established Phoenix dactylifera plantlets in sterilized soil, they
found that the plants grew poorly. Examination of roots from plantlets
established in unsterilized soil revealed fungal hyphae within the root
cortex that were absent in plantlets in sterile soil. They concluded that
poor growth may be attributed to failure to form an endotrophic mycorrhizal
association. Inoculation of sterile soil with mycorrhizal fungi was reported
by Morandi et al. (19791with R. idaeus. Inoculated plantlets were more
vigorous and uniform in size than noninoculated controls. Similar results
were obtained by Granger et al. (1983) using the Malus rootstocks MM
111and M.7. With M.7, plantlets inoculated with Glomus epigaeum were
taller after 15 weeks; however, there was no difference in MM 111. The
fungus caused significantly increased amounts of foliar copper of both
rootstocks and an increased amount of phosphorus in M.7, but it had no
effect on calcium, magnesium, and zinc, and actually depressed the
amount of potassium in M.7. Recently, Pons et al. (1983) produced
plantlets of the Prunus auium rootstock F12/1 that were infected with
Gigaspora margartia in uitro. Strullu et al. (1986) have successfully
inoculated in uitro cloned seedlings of chestnut (Castanea satiua) with

the fungus Paxillis inuolutus. Inoculated plants reportedly grew better
than noninoculated controls, although no data were presented.
The status of micropropagation research with temperate zone fruit and
nut crops is summarized in n b l e 8.2. Details on procedures for each of the
crops and cultivars listed can be obtained from the references cited.
B. n e e Fruits
Malus, Pyrus, and Prunus spp. are the most widely grown fruit tree
species. Climatic requirements restrict their cultivation to temperate
zones, usually 30'-50' of latitude north and south of the equator. Pro-
duction may extend to lower latitudes a t high elevations and to higher
latitudes where there is a moderating influence of the climate, as in
western Europe by the Atlantic Ocean (Westwood 1978).
1. M a h s (Apple). The cultivated apple has resulted from hybridiza-
tion of a number of Malus spp. over many years; consequently there is
some dispute concerning the correct botanical name. In a historical
analysis of apple nomenclature, Korban and Skirvin (1984) concluded
that the legitimate epithet for the cultivated apple is Malus xdomestica
Borkh.
a. Surface sterilization and establishment. The first report on aseptic
culture of apple buds was by Jones (1967),who reported that they were
difficult to surface sterilize since they were damaged by commonly used
sterilants. However, they became more resistant to damage after a short
period on a holding medium. A two-step procedure was developed in
which 2- to 3-cm-long shoot-tips from actively growing glasshouse mate-
rial were collected into sterile water and surface sterilized in sodium
hypochlorite (0.14%available chlorine for 1 min). After rinsing in sterile
water, they were incubated overnight on a simple basal medium and then
surface sterilized again for a longer time in sodium hypochlorite before
isolating tips about 1cm long for experiments. This method consistently
produced 90-100% undamaged sterile explants. A two-step surface steril-
ization procedure is time consuming and can be avoided by keeping
shoots in running water for about 1 hr prior to a single hypochlorite
treatment (Joneset al. 1979).The most commonly used surface sterilants
are chlorine-containing solutions in the form of calcium or sodium hy-
pochlorite and the choice is arbitrary. Calcium hypochlorite has been
used by Liu et al. (1978),Nemeth (19Sl),and Zimmerman and Broome
(1980, 1981), and sodium hypochlorite by Werner and Boe (1980) and
Sriskandarajah et al. ( 1982). Occasionally only dips in ethanol (Lane
1978; Welander and Huntrieser 1981) or no surface sterilization (Walkey
Table 8.2. Summary of Micropropagation h s e a r c h with Temperate-Zone Tree

Fruits and Nuts"
Shoot Root E x uitro

Estab- prolif- initi- acclimati-
Clone lishment eration ation zation References
Malus, rootstocks
A2 + + + + Welander and
Huntrieser ( 1981)
Antonovka 313 + + + 'Ravers et al. ( 1985)
Antonovka KA 313 + + + + Cheng (1978, 1979)
KSC-3 + + + + Embree and Hicks (1985)
Marubakiado + + ? ? Ishihara and Katano
(1982)
M.4 + + + ? Liu et al. ( 1978)
+ + + + Dunstan (1981),
Dunstan et al. (1985)
+ + + - Ochatt and Caso (1983a)
M.7 (EMLA-7) + + + - Jones and Hatfield (1976)
+ + + + Cheng (1978, 1979)
+ + + + Werner and Boe ( 1980)
+ + + + Dunstan (1981)
- + + - Meng and Zhou ( 1982)
M.9 (EMLA-9) + + + + Cheng (1978, 1979)
+ + + + James and Thurbon
(1979, 1981a,b)
+ + + - Strahlheim and Cailloux
(1981)
+ + James and Wakerell
( 1982)
+ + + - Lane and McDougald
( 1982)
f +
- * -c Loreti and Morini ( 1982)
M.25 (EMLA-25) + + + + Cheema and Sharma
(1983)
M.26 + + + + Jones et al. (1977)
+ + + + Dunstan (1981)
+ + + - Nemeth (1981)
+ + + ? Ishihara and Katano
(1982)
+ + + - Lane and McDougald
(1982)
f % f f Loreti and Morini ( 1982)
'Notations used in table:

+ successful ? difficult to determine from
+ ( 1 ) successful but limited response published data
f successful but details not S.C. sterile cultures
reported n.s. not successful
~ not reported S.S. sterile seedling
Shoot Root Ex uitro

+ + + Welander (1982, 1983)

- + + Simmonds ( 1983)
M.27 + + - Cheng (1978)
+ + + Loreti et al. ( 1981)
+ + +(1) Nemeth (1981)
+ + + Lane and McDougald
(1982)
f i- i- Loreti and Morini ( 1982)
MAC-9 (Mark) + + + Cheng (1978, 1979)
MM 104 + + + Snir and Erez ( 1980)
+ + + Nemeth (1981)
MM 106 + + + Snir and Erez ( 1980)
f f * Loreti and Morini ( 1982)
MM 109 + + + Snir and Erez (1980)
MM 111 + + + Lane and McDougald
(1982)
i- i- i Loreti and Morini ( 1982)
Ottawa 3 + + + Strahlheim and
Cailloux (1981)
+ + + Pua et al. ( 1983)
+ + + Chong and Pua ( 1985)
P. 16 + + + Liu et al. (1978)
Robusta No. 5 + + + Pua and Chong ( 1984)
M+s, scions
Akero + + + Welander ( 1985)
Cox’s Orange + + + Abbott and Whiteley
Pippin (1976)
+ + + Jones et al. ( 1979)
Delicious + + + Sriskandarajah et al.
(1982)
+ + +(1) Zimmerman ( 198313)
+ + + Zimmerman (198413)
+ + + Zimmerman and
Fordham ( 1985)
Oregon Spur I1 + + + R. H. Zimmerman,
unpublished
Redchief + + +( 1 ) Zimmerman and
Fordham (1985)
Redspur + + + Zimmerman (1984b)
+ + + Zimmerman and
Fordham (1985)
Starkspur Red + + +(1) Nemeth (1981)
(continued)
Shoot Root Ex vitro

Starkspur + + + + R. H. Zimmerman,
Supreme unpublished
Supreme Red t + + ~
Anderson (1981)
'Ikiple Red + + + + Zimmerman ( 1984b)
or Royal Red + + + t Zimmerman and
Fordham (1985)
Vermont Spur + + + + Zimmerman (198413)
Wellspur i- + + ~
Anderson (1981)
+ + + - Jacoboni and Standardi
(1982)
Fuji + + + + Ishihara and Katano
(1982)
Gala + + + + Zimmerman (1981,
198313)
+ + + + Zimmerman and
Fordham ( 1985)
Golden Delicious + + + + Jones et al. (1979)
+ + ~ ~
Lundergan and Janick

(1980)
+ i- t % Loreti and Morini ( 1982)
+ + + + Fiorino and Leva ( 1983)
+ + + + Zimmerman (1981,
198413)
+ + + + Zimmerman and
Fordham (1985)
Golden Delicious t t i t Loreti and Morini (1982)
B
Goldspur + + + + G. Suttle, unpublished;
R. H. Zimmerman,
unpublished
Perleberg * t i c Loreti and Morini (1982)
Granny Smith + ~ - - Elliot ( 1972)
+ + + + Sriskandarajah and
Mullins (1981)
Greensleeves + + + + Webster et al. (1985)
James Grieve + + + + Jones et al. (1979)
Jonagold +
- + + - Druart et al. (1982)
Jonathan + + + + Huth(1978)
+ + + + Zimmerman (1981,
198313)
+ + t ? Sriskandarajah et al.
( 1982)
Malling Kent + + + + Jones et al. (1979)
Malling Suntan + + + + Jones et al. (1979)
McIntosh i L L ~
Walkey (1972)
(Seedlings)
Shoot Root E x vitro

McIntosh + + + Lane (1978)

Macspur + + - Lane et al. (1982)
+ + + Pua and Chong ( 1985)
Imperial + + + Zimmerman and
Fordham (1985)
Wijcik + + - Lane et al. ( 1982 1
Paladin0 Spur + + + Zimmerman and
Fordham ( 1985)
Summerland Red + + ~
Lane et al. (1982)

Mutsu ( Crispin) + + + Zimmerman (1981)
+ + + Zimmerman and
Fordham ( 1985)
Norcue * ? ~
McIntyre and Dinkel

(1983)
Norda +
- ? - McIntyre and Dinkel
(1983)
Noret -c ? ~
McIntyre and Dinkel

(1983)
Northern Spy + + + Zimmerman (1981,
1983b)
+ + + Zimmerman and Broome
(1981)
+ + + Hutchinson (1982,
1984a,b)
Nugget + + + Zimmerman (1981,
198313)
(1981)
Ozark Gold + + + Zimmerman ( 1981,
198313)
( 1981)
Prairiefire k + +- Joung et al. ( 1983)
Rome Beauty + + + Zimmerman (1981,
1983b)
Spartan + + + Zimmerman (1981,
1983b)
(1981)
+ + + Zimmerman and
Fordham ( 1985)
Spuree Rome + + + Zimmerman (1981,
1983b)
(1981)
Stark Jumbo + + + Cheng (1978)
-
continued

Starkspur Golden + + Fiorino and Leva ( 1983)

Stayman t + Zimmerman (1981,
198313)
+ + Zimmerman and Broome
(1981)
'Railman t ? McIntyre and Dinkel
(1983)
Yellow spur + + Fiorino and Leva ( 1983)
York Imperial + t Zimmerman (1981,
198313)
+ + Zimmerman and
Fordham ( 1985)
Pyrus, rootstocks
Old Home X
Farmingdale 51 + + Cheng (1978, 1979)
I? calleryana (D-6) + + J . F. Hutchinson,
unpublished
Pyrus, scions
I? communis
Abate Fete1 f t Loreti and Morini (1982)
Bartlett + + Lane (1979)
f t Loreti and Morini ( 1982)
+ + Shen and Mullins ( 1984)
Beurre Bosc + + Shen and Mullins (1984)
Conference i + Loreti and Morini ( 1982)
Decana del t f Loreti and Morini (1982)
Com izio
Jinfeng + + Zhao ( 1982)
Kaiser i f Loreti and Morini (1982)
Packhams + + Shen and Mullins (1984)
Tkiumph
Seckel + + Singha (1980, 1982, 1984)
Zaosu + + Zhao ( 1982)
I? pyrifolia (syn.
I? serotina)
Hosui + + Bhojwani et al. (1984)
Kosui + + Bhojwani et al. (1984)
Nijusseiki + + Bhojwani et al. (1984)
Shinseiki + + Bhojwani et al. (1984)
Shinsui + + Bhojwani et al. (1984)
Prunus (cherry),rootstocks
Cer W 10
(I? cerasus) + + Paul and Feucht ( 1985 1
8. TISSUE CULTURE O F TEMPERATE FRUIT AND N U T TREES 287

_ _ _ ~ ~ _ _ _ _ ~
~

Colt ( P auiurn x + + Cheng (1978, 1979)

P pseudocerasus) * zk Loreti and Morini ( 1982)
S.C. + Wilkins and Dodds
( 1982/83)
F12/1 ( P . auiurn) + + Jones and Hopgood
(1979)
+ + Dunstan (1981)
+ + Nemeth (1979, 1982)
+ + Ancora et al. ( 1982)
+ + Paul and Feucht ( 1985)
Mahaleb X + + Cheng (1978, 1979)
Mazzard 14
Prunus auiurn (sweet cherry), scions
Bing + + Snir (1982a,b)
Black 'Ihrtarian + + Snir (1982a,b)
Burlat n.s. - Snir (1982a)
Early Rivers + + Skirvin et al. (1979)
Early Ruby ? ? Snir (1982b)
Hedelfinger + + Paul and Feucht ( 1985)
Renier n.s. - Snir (1982a)
Royal Ann + + Snir (1982a)
Sam + + Snir ( 1982a,b)
+ + Paul and Feucht ( 1985)
Seedling selections + + Sauer (1983)
Prunus cerasus (sour cherry), scions
Ben Zion + + Snir (1983)
Boscha + + Paul and Feucht ( 1985)
Chios + + Snir ( 1983)
F1 + + Ponchia and Roselli
( 1980)
Marasca di Braza + + Cossio (1981)
Marasca Luxardo + + Cossio ( 1981)
Montmorency + +(1) Skirvin et al. (1979, 1982)
Schattenmorelle + + Cossio ( 1981)
+ + Borkowska (1983),
Borkowska and
Zajac ( 1985)
+ + Paul and Feucht ( 19851
Schwaebische + + Paul and Feucht (1985)
Weinwechsel
Skubinka + + Popov et al. (1976)
Stevnsbaer + + Cossio ( 1981)
Vladimir + + Ponchia and Roselli
(1980)
+ + Cossio et al. ( 1981)
continued
Shoot Root Ex vitro

Prunus (cherry),scions
Durone I t t t -r
- Loreti and Morini ( 1982)
Durone I1 + + & i- Loreti and Morini ( 1982)
Germella + % t i- Loreti and Morini ( 1982)
Vittoria t i t f Loreti and Morini ( 1982)
Prunus (peach),rootstocks
G F 43 i; i 37 i- Loreti and Morini ( 1982)
G F 655/2 + + +
- i- Loreti and Morini ( 1982)
G F 1869 f f % f Loreti and Morini ( 1982)
Love11 + + + + Almehdi and Parfitt
(1986)
Red Leaf + + +(Z) - Ochatt and Caso (198315)
l? persica X
l? amygdalus
G F 557 + + + - Nemeth (1979)
G F 677 f t~ -i + Zuccherelli (1979)
i- t t +
+ + + t Kyriakidou and Pontikis
( 1983)
l? persica X
I? davidiana
Nemaguard + + + + Miller et al. (1982)
+ + +(I) (1) Reeves et al. ( 1983)
+ + + + Almehdi and Parfitt
(1986)
+ + + + Hammerschlag et al.
(1986)
Hybrid 41-6-22 + + ? ? Vertesy (1980)
Hybrid 41-5-3- + + ? ? Vertesy ( 1980)
Hybrid VII. 20/3 + + ? ? Vertesy (1980)
Hybrid 65-2 + + ? ? Vertesy ( 1980)
Hybrid 41-4-19 t + ? ? Vertesy ( 1980)
Hybrid 41-5- + + ? ? Vertesy (1980)
Hybrid 41-6-2 + + ? ? Vertesy ( 1980)
l? amygdalus X
$? persica
Hansen 536 + + + + Martinelli ( 1985)
Hansen 2168 + + + + Martinelli ( 1985)
l? amygdalus
‘Nonpareil’ X
P: persica
0607 + + ? - Tabachnik and Kester
(1977)
l? amygdalus
0609 + t(1) ? ~
Tabachnik and Kester

( 1977)
Shoot Root En uitro

Prunus (peach),scions
Armking + Loreti and Morini (1982)
Candor t Hammerschlag ( 1982b)
Compact Redhaven + Hammerschlag ( 198213)
+ Hammerschlag et al.
( 1987)
Dixired + Hammerschlag ( 198213)
Early Red Free + Vertesy ( 1980)
Fayette t Hammerschlag ( 1982b)
Firebright ? Loreti and Morini ( 1982)
Flavorcrest f Loreti and Morini ( 1982)
Harbelle + Skirvin et al. (1979)
Harbrite + Skirvin et al. (1982)
Jerseyqueen + Hammerschlag ( 198213)
( 1987)
Madeleine P + Vertesy ( 1980)
Madison + Skirvin et al. ( 1979)
Maria Bianca 2 Loreti and Morini (1982)
Maycrest * Loreti and Morini ( 1982)
Raritan Rose + Vertesy ( 1980)
Redhaven + Skirvin and Chu (1977,
1978), Skirvin et al.
(1979)
+ Hammerschlag ( 1982b)
( 1987)
Redskin + Hammerschlag (198213)
( 1987)
Rio-Oso-Gem + Hammerschlag (1982b)
( 1987)
Springold + Hammerschlag ( 198213)
Suncrest 2 Loreti and Morini ( 1982)
( 1987)
Sunhaven + Vertesy ( 1980)
Sunhigh + Hammerschlag ( 198213)
( 1987)
Sunred * Loreti and Morini ( 1982)
Vesuvio + Vertesy ( 1980)
Vivid + Hammerschlag ( 1982b)
I? persica (from s.s Feliciano and de Assis
embryo cultured ( 1983)
seedlings)
continued
Shoot Root Ex uitro

Prunus (plum),rootstocks
I? cerasifera + + + + Hammerschlag ( 1982a)
G F 31 + + + - Nemeth (1979)
Mr.S. 2 15 + + + + Loreti et al. ( 1982)
I? insititia
Pixy + + + + Cheng (1978, 1979)
+ + + + Jones and Hopgood
(1979)
i + t f Loreti and Morini (1982)
I? domestica
St. Julien X + + + + Cheng (1978)
St. Julien
hybrid No. 2 + + + ? Nemeth ( 1982)
Prunus (plum),scions
I? dornestica
d’Agen + + + +(1) Baleriola-Lucas and
Mullins (1984)
Early Golden i f i f Loreti and Morini ( 1982)
d’Ente 707 +
- f f f Loreti and Morini ( 1982)
+ + + + Baleriola-Lucas and
Mullins (1984)
Laroda f f f + Loreti and Morini ( 1982)
President f + f f Loreti and Morini ( 1982)
Santa Rosa + f +
- f Loreti and Morini ( 1982)
Shiro f f f +
Sorriso di f f f f Loreti and Morini (1982)
Primavera
Stanley f f f +
2 f f f Skirvin et al. (1980, 1982)
+ + + + Pietropaolo and Reisch
(1984)
I? salicina
Calita + + + + Rosati et al. (1980)
Prunus armeniaca (apricot), scions
Caldesi I k f f f Loreti and Morini ( 1982)
Canino + + + + Snir (1984)
Defarges f f f f Loreti and Morini ( 1982)
Reale d’Imola f f f f Loreti and Morini ( 1982)
S. Castrese +
- f f f Loreti and Morini ( 1982)
Stella + +([I +(1) ~
Skirvin et al. (1979)
Diospyros kaki (Japanese persimmon)

Gailey + + + ? Cooper and Cohen ( 1984)
Fuyu + + ? ? Cooper and Cohen ( 1984)
Maekawa Jiro + + ? ? Cooper and Cohen ( 1984)

~~~~
Shoot Root Ex uitro

Hiratanenashi + +(O Cooper and Cohen (1984)

Izu + +(1) Cooper and Cohen ( 1984)
Olea europaea (olive)
Dolce Agogia + + Rugini and Fontanazza
(1981), Rugini (1984)
Frantoio + + Rugini (1984)
Moraiolo + + Rugini ( 1984)
Morus indica (mulberry)
Unnamed cultivar + +(1) Pate1 et al. (1983)
Unnamed selections + + Mhatre et al. (1985)
Seedling + + Kim et al. (1985)
Ficus carica (fig)
Calimyrna + ? Muriithi et al. (1982)
Caprifig + ? Muriithi et al. (1982)
Excel + ? Muriithi et al. (1982)
Kalamon + + Pontikis and Melas
(1986)
Osborne Pacific + ? Muriithi et al. (1982)
Pasquale + ? Muriithi et al. (1982)
Tena + ? Muriithi et al. ( 1982)
Zidi + ? Muriithi et al. (1982)
Carya illinoensis (pecan)
Seedlings + + Wood ( 1982)
L + Lazarte ( 19831, Hansen
and Lazarte (1984)
Castanea (chestnut)
C. mollissima + + Yang et al. (1986)
(seedling)
C. satiua (seedling) f + Vieitez and Vieitez
(1980a)
S.S. + Rodriguez ( 1982c,d)
f + Vieitez and Vieitez ( 1983)
+ + Strullu et al. (1986)
Marigoule
HV
+
+
+ . Chevre et al. (1983)
+(1 ) Vieitez et al. ( 1983)
T-13 + + Vieitez et al. (1983)
43 1 + + Vietiez et al. (1983)
Cardaccio + n.s. Biondi et al. (1981)
Mozza + + Biondi et al. ( 1981)
Poli tora - n.s. Biondi et al. (1981)
Corylus auellana (filbert, hazelnut)
Daviana +(I) +
- Anderson (1983)
Ferlile de Coutard * +(1 ) Kai et al. (1984)
continued
Shoot Root E x uitro

Kai et al. (1984)

Kai et al. ( 1984 1
Seedling + + i + Anderson ( 1983)
+ + + ? Perez e t al. ( 1985)
duylans (walnut)
J . hindsii X J . regia + + + t Driver and Kuniyuki
Paradox (1984)
J . regia (seedling) Rodriguez ( 198213)
+ + +
~~
I
‘hlecke and McGranahan
(1985)
Pistacia (pistachio)
I! terebinthus i + I t Pontikis (1984)
‘Tsikoudia’
I! uera (seedling) s.s + + t Barghchi and Alderson
(1983a,b, 1985)
P r u n u s amygdalus (almond)
Ferraudel +- ~ + + ~ + Loreti and Morini ( 1982)
Ferragnes + t I i Rugini and Verma ( 1982,
1983)
Nonpareil + + ‘? ~
Tabachnik and Kester

( 1977)
S. Caterina * * * * Loreti and Morini ( 1982)
nono + * i * Loreti and Morini ( 1982)
Seedling s.s i i ? Hisajima (1982)
1972) have been used successfully, but only if meristem-tip explants are
isolated. Agitation has been shown to be beneficial during the calcium
hypochlorite stage (Zimmerman and Broome 1980),presumably by mak-
ing better contact between the sterilant and the contaminants.
After surface sterilization and during establishment, browning of ex-
plants, due presumably to oxidation of phenols, can be a problem that
may be reduced by a number of approaches: Keeping explants in running
water will leach out water-soluble phenols and has been used with suc-
cess (Jones et al. 1979; Hutchinson 1984a); Walkey ( 1972) incorporated
-
polyvinylpyrolidone ( PVP, mol. weight 40,000) and found it essential
for satisfactory growth, although it could be eliminated after 4 to 8 weeks.
Reduction of MS salt concentration to half strength considerably reduced
the browning problem with M.7 (Werner and Boe 1980).
Explants can be successfully established from both glasshouse- and
field-grown material a t virtually any time of the year using explants of
various types. Cultures established from field-grown material of ‘North-

ern Spy’ during midspring or summer grew more rapidly, and contamina-
tion and explant browning were less of a problem, compared to cultures
established in midautumn or winter (Hutchinson 1984a).The majority of
workers have used shoot-tips, 1-2 cm long, from actively growing plants
(Jones and Hatfield 1976; James and Thurbon 1979; Lundergan and
Janick 1980),whereas others have used apical meristem-tips or meristem-
tips from axillary buds with up to four leaf primordia (Walkey 1972;
Abbott and Whiteley 1976; Welander and Huntrieser 1981).
Although most workers have used agar-gelled medium for establish-
ment, Walkey (1972)and Jones and Hatfield (1976)used liquid medium
with explants supported on filter paper bridges. Zimmerman and Broome
(1980) found liquid rotated a t 1 rpm for 2-4 days to be superior to
agar-gelled medium with better growth and fewer losses of explants. Pua
and Chong (1984) found a requirement for sorbitol for the successful
establishment of M. robusta No. 5 rootstock. Good survival was obtained
after 4 weeks with glucose, sucrose, sorbitol, or fructose as the carbon
source, but after 8 weeks only cultures on sorbitol medium survived.
b. Shoot proliferation. (i) Basal medium. The macro- and micronu-

trients described by Murashige and Skoog (1962)have been used by most
workers. Jones and Hatfield (1976)used Knop’s solution with MS micro-
nutrients and 20 mg/liter NaFeEDTA. While there are no dose-response
curves for macronutrients, Dutcher and Powell (1972) reported that if
they were omitted little or no growth resulted. With regard to micronu-
trients, they state that growth was best if manganese and zinc were
present, but they were not able to demonstrate conclusively that growth
inhibition, albeit slight, was due to any particular element. The organic
addenda of MS, or their modification by Linsmaier and Skoog ( 19651,
have been commonly used. Fiorino and Leva (1983)compared three basal
media and concluded that PM medium was superior to either MS or Cheng’s
(1978). No reference or details of PM medium were given.
(ii) Cytokinins. Jones (1967) reported that 1 mg/liter (4.4 p M ) BA
would promote stem elongation and leaf formation in the rootstock M-26,
but a t 10 and 50 mg/liter ( -44 and 222 p M ) extension was inhibited and
profuse callus growth resulted. Pieniazek (1968) also showed that BA
could induce growth and, a t 2 mg/liter (8.9p M ) ,it could induce growth of
lateral buds, but the leaves were narrow and small. The most commonly
used cytokinin to induce and maintain shoot proliferation is BA. Lundergan
and Janick (1980) found that BA was the most effective cytokinin for
stimulating proliferation of ‘Golden Delicious’ shoot-tips, with 2iP the
least effective and kinetin intermediate between BA and 2iP. They noted
that growth (elongation) was inversely related to proliferation, with 2iP
the most and BA the least stimulative. A single tall shoot was produced
by 2iP, with BA producing a dwarfed rosette type of growth. In experi-
ments combining BA and 2iP, shoots responded similarly to shoots
growing on BA alone. Lundergan and Janick concluded that 2iP had
little or no effect on the response of the shoots to BA at the concentra-
tions tested. Extending this work using ‘Northern Spy’, Hutchinson
(1984a) reported that zeatin was more effective than kinetin, but less
so than BA, and that the response was dependent on cytokinin concen-
tration and light level. Similar results were presented by Fiorino and
Leva (1983).
Lane (1978)published a dose-response curve for BA using ‘McIntosh’:
5 pit4 was optimal, 0.5 pit4 was slightly less effective, and 50 pit4 killed the
explants. Although BA is a suitable cytokinin for shoot proliferation, it
can cause problems. Dunstan et al. (1985) tested a range of cytokinin
concentrations and found that 1.125mg/liter (5.OpM)BA combined with
0.15 or 0.20 mg/liter (0.7-1 pit4) IBA produced the maximum number of
usable shoots of M.4 rootstock. After about 4 months culture, Werner
and Boe (1980) observed that several cultures showed BA toxicity and
periodic substitution of 2iP for BA eliminated the problem. Occasionally
kinetin has been used as the sole cytokinin source; Walkey ( 1972) used
1.25 mg/liter (5.8 pit4) but presented no quantitative details. Abbott and
Whiteley ( 1976)also used kinetin and noted that concentrations less than
0.1 or greater than 5 mg/liter (-0.5 and 23 pit4) completely inhibited
shoot proliferation and that concentrations between 0.5 and 1 mg/liter
(2.3 and 4.6 pit4) could induce satisfactory proliferation.
Thidiazuron (N-phenyl-N’-1,2,3-thiadiazol-5-ylurea) has been shown to
have potent cytokininlike activity for stimulating shoot proliferation in
cultures of apple cultivars in vitro (van Nieuwkerk et al. 1986; R. H.
Zimmerman and I. Fordham, unpublished data). Concentrations as low
as 0.1 p M stimulated as many new shoots per explant as 4.4 pit4 BA,
although the shoots proliferated with thidiazuron had about half the
length of those on 13A (van Nieuwkerk et al. 1986). Shoots rooted readily
once they had been grown without thidiazuron for 4-8 weeks.
(iii)Auxins and gibberellins. While some authors have used only a
cytokinin to induce and maintain proliferation ( Abbott and Whiteley
1976; Werner and Boe 1980; Sriskandarajah and Mullins 1981; Simmonds
1983) others have incorporated IBA, NAA, or GA,. Most have not
presented quantitative details on the influence of these hormones (Jones
et al. 1977,1979; James and Thurbon 1979; Snir and Erez 1980). Using a
standard concentration of 5 pit4 BA Lane (1978) showed that adding 5
pit4 GA3caused a slight, although probably not significant, reduction of
shoot proliferation compared to the addition of 0.5 pit4 NAA, which
reduced proliferation. If both NAA and GA3 were added, proliferation
was further reduced, indicating that shoots probably produce sufficient

endogenous auxin for growth. This was supported by Lundergan and
Janick (1980), who found that incorporating 1 mg/liter (4.9 p M ) IBA
significantly reduced shoot proliferation of ‘Golden Delicious’, but GA,
was without effect. James and Thurbon (1981a) showed with M.9 that
adding various concentrations of IBA had no significant influence on
proliferation and BA alone was the most important factor.
(iv)Phloroglucinol. Using the rootstocks M.7 and M.26 Jones (1976)
showed that incorporating 1mM PG into a medium with BA, IBA, and
GA, increased shoot proliferation dramatically. The effect was described
further for M.26 by Jones et al. (1977),who found that single shoot-tips
produced between two and five shoots over 4 weeks on 10 ml of medium;
however, after transfer to larger containers with 125 ml of medium for a
further 8 weeks, cultures had produced between 20 and 42 shoots. The
effect of PG on proliferation is not universal. Snir and Erez (1980)found
that PG gave no advantage in preliminary experiments, Whiteley and
Abbott (1977) reported that it inhibited growth of their cultures, and
Zimmerman and Broome (1980)observed that it had no consistent effect.
James and Thurbon ( 1981a)found no effect of PG on multiplication rates
per se with M.9, but considered it important, as shoots grown in its
presence subsequently had a higher percentage root initiation and a
greater number of roots per rooted plantlet. Adding PG to the medium
during establishment and first five subcultures increased shoot prolifera-
tion for the first four subcultures, after which it was without effect
(Hutchinson 1984a).
( v ) Agar-gelled vs. liquid medium. During proliferation all authors
have used agar-gelled medium, with the exception of Snir and Erez
( 1980),who found that shoots grew faster in liquid medium on an orbital
shaker. Although no quantitative details were provided, this more rapid
growth may have been due to increased absorption of nutrients and
hormones through a greater surface area. Agar is usually used at a
concentration of 0.6-0.9%, but Singha (1982, 1984)found that a concen-
tration of 0.3% resulted in maximum shoot proliferation and growth with
the ornamental Malus ‘Almey’. These results may be explained by the
presence of inhibitors in agar (Kohlenback and Wernicke 1978),the effect
of concentration on the diffusion of molecules through the medium
(Romberger and n b o r 1971), or reduction in the availability of water a t
higher agar concentrations (Stoltz 1971). Agar brand and type can also
influence proliferation. Singha (1984)found that a t 0.3% there was very
little difference between brands but as concentration increased Bacto-
agar (Difco Laboratories) was more inhibitory than TC agar ( KC Biologi-
cals, Inc.)
(vi) Carbon source. As with explant establishment, sucrose is the
sugar of choice for virtually all shoot proliferation studies. However, Pua
and Chong ( 1984,1985)have reported that sorbitol produced the greatest
number of shoots of M. robusta No. 5, ‘MacSpur’, and seedlings of
‘Macspur’. With M. robusta No. 5, 30 g/liter sorbitol produced nearly
four times as many shoots as 30 g/liter sucrose, although fresh weight
was only twice as great and dry weight just 30% greater (Pua and Chong
1984). When concentrations were increased to 50 g/liter, differences in
shoot number disappeared. Although similar results were obtained with
‘Macspur’, a combination of 25% sorbitol and 75% sucrose was best for
shoot production (Pua and Chong 1985).
(vii) Explant type. Once a culture line is established it is usually
maintained by subculturing shoot-tips onto fresh medium a t regular
intervals and often discarding the remainder. Hutchinson ( 1984a)divided
an established culture into shoot-tips, nodes, and the basal mass (mate-
rial remaining after removal of other explants). He evaluated each for
shoot proliferation and showed that the commonly used shoot-tips were
not the most productive source of additional shoots. The main advan-
tages of using alternatives to shoot-tips for routine proliferation is that all
shoot-tips can be used for rooting studies and that the time to produce a
large number of shoots can be considerably reduced. However, care must
be taken since adventitious shoots can form from the basal mass (Nasir
and Miles 1981; Welander 1985).
c . Root initiation. ( i )Auxins. Both auxin type and concentration in-

fluence rooting in Malus cultures. The preferred auxin for root initiation
is IBA. Jones and Hatfield (1976)demonstrated that IBA is superior to
IAA and NAA, and Lane (1978)found that NAA decreased root initiation.
The use of NAA led to more rapid root initiation in ‘Ozark Gold’, but after
4 weeks the difference in percentage root initiation between IBA and
NAA had disappeared (Zimmerman and Broome 1980).In a comparison
of IBA, IAA, and NAA, Zimmerman and Fordham (1985)round IBA to
be the most effective and IAA the least effective for a range of scion cul-
tivars. Using liquid medium, Sriskandarajah and Mullins (1981) found
IBA and NOA equally good, with both better than 2,4-D a t 10p M . 2,4-D
was inhibitory and caused discoloration of the medium. Both IBA and
NOA up to 10 p M promoted root initiation, but a t higher concentrations
shoots produced few roots and soft white callus. They preferred IBA since
the roots were thicker and more numerous and formed a t the base of the
cutting as well as along the stem. Nemeth (1981) reported that substi-
tuted 2-chloro-3-phenylpropionitriles were more effective than 5 p M IBA
in the rootstocks M.26, M.27, MM 104, and the scion cultivar ‘Stark
Spur’ (sic).Of those tested, 5 p M 2-chloro-342,3-dichloro-phenyl)propionitrile
significantly increased percentage root initiation and root length, but not
root number compared to IBA.
The concentration of auxin used varies with cultivar. Using M.7 root-
stock Werner and Boe (1980)found no difference in percentage rooting
between 18 and 35 days with 1mg/liter (4.9 p M ) IBA, but if the concen-
tration was increased to 3 mg/liter (15 p M ) , percentage rooting was
initially low but increased with time. Welander and Huntrieser (1981)
reported that 15 p M IBA resulted in lower percentage rooting than 5
and 10 p M in studies involving the rootstock A2; however, there were
significant interactions between the growth phase of the plant and the
use of PG.
(ii) Inorganic salt formulation and concentration. The inorganic salt
formulation of Murashige and Skoog (1962)or variations of it have been
used by most workers, with some using it at full strength (James and
Thurbon 1979, 1981a; Jones et al. 1977; Loreti et al. 1981)and others a t
reduced concentration (Werner and Boe 1980; Sriskandarajah and Mullins
1981; Welander and Huntrieser 1981; Lane and McDougald 1982; Joung
et al. 1983).In a comparison of MS macronutrient concentration, Simmonds
(1983)found that lowering concentration to half or quarter strength had
little effect on percentage rooting, but rooting intensity was enhanced on
a medium with quarter-strength macronutrients and 1% sucrose. Com-
paring full- and half-strength salt concentration, Hutchinson ( 1984a)
found a significant increase in percentage rooting with half-strength
salts, but only when 1p M IBA was used. The effectiveness of a reduction
of salt concentrations is not fully understood, but the salt concentration
in a medium can affect its osmotic potential, which can influence the
uptake of nutrients by plant tissues or the release of some substances into
the medium (Thorpe 1978; Barghchi and Alderson 1983a). In a detailed
study of rooting of Rosa ‘Improved Blaze’, Hyndman et al. (1982)con-
cluded that improved rooting at reduced salt concentration was probably
attributable to a more favorable nitrogen concentration. Comparing half-
strength MS macronutrients and Lepoivre medium (Quoirin et al. 1977),
Welander ( 1983)found no significant difference between the two media in
terms of percentage root initiation with M.26, but a slight increase in root
number per rooted shoot and significantly better survival of plantlets
with Lepoivre medium, which was attributed to reduced callus formation.
(iii)Phloroglucinol. Jones and Hatfield ( 1976)first showed with apple
shoots in uitro that the addition of 1mM PG or phloretic acid more than
doubled percentage root initiation in M.7 when cultured with 5 p M IBA.
Phloroglucinol was also found effective with M.26 (Jones et al. 1977)and
with five scion cultivars (Jones et al. 1979). The general method was
refined and improved subsequently by James and Thurbon (1979) and
James (1983a)with M.9 shoots in which they found that an exposure of

only 4 days to IBA and PG was sufficient to induce roots. Subsequent
root development was allowed to take place in hormone-free medium.
James and Thurbon (1981a) optimized the concentration for M.9 a t 3
mg/liter (15 p M ) IBA and 10 mM PG.
The response of apple cultivars to PG is not consistent. Cheema and
Sharma (1983) found it deleterious. Working with nine scion cultivars,
Zimmerman and Broome (1980, 1981) found a positive effect only with
‘Spartan’. With the rootstock A2, both the PG concentration and the
growth phase of the plant influenced the response (Welanderand Huntrieser
1981). For shoots from trees in the adult phase, 0.1 mM PG promoted
rooting, whereas 1mM markedly inhibited it. However, with shoots in the
juvenile growth phase, both 0.1 and 1 mM PG stimulated rooting. In
addition, 1mM PG reduced callus formation with all IBA concentrations
used. With ‘Delicious’and a number of its strains, Zimmerman (198413)
found varying results. When proliferating shoots were incubated for 30
days in light or dark, PG tended to reduce subsequent rooting of ‘Deli-
cious’, but the effect on ‘Redchief Delicious’ was inconsistent. When
shoots of ‘Redchief Delicious’were incubated in darkness during rooting
for varying lengths of time in the presence or absence of PG, the presence
of PG increased rooting, with maximum response occurring with 1week
of darkness. Increasing temperature from 25’ to 3OoC increased percent-
age rooting of ‘Royal Red Delicious’ in the presence of PG, but only with
4.9 p M IBA; if IBA was used a t 1.5 p M there was no effect. Comparing
rooting between M.9 and M.26, James and Thurbon (1981b) found the
presence of 1 mM PG in both the shoot proliferation and rooting media
increased both rooting percentage and number of roots per rooted shoot
with M.9, but was generally without effect on M.26. Similarly, variation
has been seen in the rooting response of some dwarf clones to the presence
or absence of PG in both shoot proliferation and rooting medium (Jones
etal. 1985).
The exact mode of action of PG is unknown. Using ( 2-14C)-IAAin the
presence or absence of PG, James (1983b) found no difference in the
uptake of IAA and very little IAA movement from the basal end of the
shoot with M.9 and M.26, which indicates that PG effects are not
mediated through enhanced IAA uptake and that differences in rooting
performance were likely to be due to difference in metabolism of exoge-
nous IAA. Alternatively, PG may be an auxin protector by acting as an
alternative substrate for IAA oxidase and/or peroxidase (James and
Thurbon 1981b).
(iv)Agar concentration. Agar a t a concentration of 0.6-0.870 is the
most commonly used support for explants. Werner and Boe ( 1980)found
that by reducing agar to 0.2% (and MS salts to one-third),they were able
to obtain 80-100% root initiation in M.7 with 5-15 p M IBA. This com-
pares favorably to the maximum of 60-7070 of Jones and Hatfield (1976)
working with the same rootstock, but using Knop’s solution with liquid
medium and filter paper bridge supports. Agar concentrations between
0.45 and 0.8070 had no effect on the cultivars tested by Zimmerman and
Broome (1980).
( v )Non-agar-basedsupports. Agar-gelledmedia have long been favored
by most workers, presumably because they are simple to prepare and
have given satisfactory results. However, the choice between agar and
liquid formulations should not be made arbitrarily (Murashige 1977).
When testing a range of physical supports, Sriskandarajah and Mullins
( 1981)found an improvement in rooting with ‘Granny Smith’ when shoots
were grown in liquid medium with low levels of continuous light. Filter
paper bridges have been used successfully for the scab-immune crabapple
‘Prairiefire’by Joung et al. ( 19831,who found them better than agar, and
by Sriskandarajah et al. ( 1982)for ‘Jonathan’and ‘Delicious’.Vermiculite,
perlite, and sand were tested successfully with a number of scion culti-
vars by Zimmerman and Broome ( 1980)and perlite was used routinely by
Machnik and Orlikowska (1981). In testing nine different physical sup-
ports with ‘Northern Spy’, Hutchinson ( 1984a)found good root initiation
with agar, coarse sand, perlite, and liquid-rotated medium, but poor root
number and elongation with agar. He attributed the improvement in root
number and elongation to better aeration of nonagar medium.
(vi)Wounding of stem. Wounding the base of the stem has had no
significant effect on the overall percentage root initiation, but has
significantly increased the number of roots per rooted shoot in ‘Granny
Smith’ (Sriskandarajah and Mullins 1981).Snir and Erez ( 1980)were able
to achieve 100% rooting using the stem-wounding technique as a routine
procedure with MM 104, MM 106, and MM 109. They exposed their
shoots to IBA-enrichedmedium for 6-8 days and then transferred them to
hormone-free medium with 0.25% activated charcoal and were able to
prevent callus formation and produce an average of 10 roots per shoot.
(vii) Carbon source. Sucrose is the most commonly used carbon
source, usually at a concentration of 2-370, but there are only a few
reports on the effect of concentration. Sriskandarajah and Mullins (1981)
found that ‘Granny Smith’shoots died within 1week without sucrose and
that the optimal concentration was 170,in which up to 80%root initiation
occurred. Similar results were obtained by Simmonds (1983) for M.26.
Fkducing sucrose concentration below 2% resulted in greener and health-
ier shoots, but root initiation decreased in proportion to the lowering of
sucrose concentration and rooting decreased with concentrations above
5.2% (Lane 1978). The difference in response may have been due to the
genotype or to the fact that Sriskandarajah and Mullins (1981) were
using liquid medium and so getting better absorption. Snir and Erez
( 1980)found no roots formed in the absence of sucrose and routinely used
290, but noted that root initiation could be partially restored by transfer
of shoots to sucrose media, even after 2 weeks. Varying sucrose concentra-
tion between 1.5 and 690 had no effect on a number of scion cultivars
(Zimmerman and Broome 1980). Rooting in M. robusta no. 5 was satis-
factory with either sucrose or sorbitol (Pua and Chong 1984).
(viii)Temperature. Most workers incubate cultures a t or about 25OC.
Lane ( 1978) found that the optimal day :night temperature for ‘McIn-
tosh’ was 28:22OC; lowering it to 23:17OC or 18:12OC reduced both
percentage rooting and the number of roots formed. Elevating tempera-
ture has also been found effective with a number of other apple cultivars.
In testing ‘Delicious’,some of its strains, and ‘Golden Delicious’,Zimmerman
(1984b)found that by imposing a dark treatment and increasing temper-
ature to 3OoC, percentage rooting was increased in ‘Delicious’,‘Vermont
Spur Delicious’, and ‘Royal Red Delicious’,but not in ‘Golden Delicious’.
Increasing temperature to 35OC further improved rooting in ‘Royal Fted
Delicious’. Temperatures between 22 and 29OC had no effect on rooting of
M.9 (James 1983a).
(ix)Time on multiplication medium. An interesting observation was
made by James and Thurbon (1980, 1981b),who found with M.9 that as
the time that shoots were left on shoot proliferation medium within a
subculturing cycle increased, percentage root initiation and root number
increased. They argued that, since exogenously supplied BA can accumu-
late in lateral buds before they grow into shoots (Woolley and Wareing
1972),then with increasing time in culture there was a fall in BA concen-
tration in the shoot bases that subsequently provide the sites for root
formation upon removal of the cytokinin and the addition of an auxin.
( x )Light and darkness. Cultures are normally incubated with a 16 :8
hour (light :dark) photoperiod. Recent research has shown that a contin-
uous dark period or continuous light can be beneficial. With ‘Supreme
Red Delicious’ and ‘Wellspur Delicious’, Anderson (1981) found that
incubation of shoots during proliferation for 2 weeks in darkness followed
by a regreening stage improved subsequent rooting. However, there were
problems of low vigor and poor survival of the plantlets upon acclimati-
zation. Also working with ‘Delicious’types, Zimmerman ( 1984b)observed
that shoots of ‘Delicious’proliferated in the dark rooted better, but the
reverse was true of ‘Redchief Delicious’. In addition, shoots proliferated in
the dark were weak and difficult to acclimatize. Darkness during the
rooting phase has been more effective, but the response in ‘Delicious’
types is dependent on cultivar, temperature, and the presence or absence
of PG (Zimmerman 1984b)and the duration of the dark period (Zimmerman
and Fordham 1985). Exposure to darkness with auxin-enriched medium
followed by transfer to light with hormone-free medium has resulted in

increased root initiation with M.9 (James 1983a) and M.26 (Welander
1983). In contrast, Sriskandarajah and Mullins ( 1981) have found good
rooting with ‘Granny Smith’, but only with continuous light and con-
tinuously agitated liquid medium.
The reasons why darkness may be beneficial are unknown, but Druart
et al. (1982) have found with ‘Jonagold’ that with darkness applied to
shoots during elongation, due to the addition of GA3, there is an increase
in peroxidase activity and a reduction in phenol content.
The development of roots on apple shoots in uitro is variable with
considerable genotypic difference and is not well understood physiologically.
I t is difficult to generalize about the response of a cultivar to any particu-
lar root-inducing treatment.
2. Pyrus (Pear). The most commonly cultivated pear is Pyrus commu-
nis, although I! pyrifolia (syn. l? serotina), the Asian pear or Nashi, is
widely grown in Japan and is gaining importance in Australia, New
Zealand, and the United States. Research to date has concentrated on
scion cultivars.
a. Surface sterilization and establishment. Cultures have been suc-
cessfully established from explants collected a t different times of the
year. Lane (1979) isolated meristem-tips from terminal buds of field-
grown trees of ‘Bartlett’ in spring; however, Bhojwani et al. (1984) ex-
perienced high contamination rates from both terminal and nodal buds
of similar material of Asian pear. Others have used glasshouse-grown
material. Singha (1980)used dormant terminal buds of ‘Seckel’collected
just after active growth ceased, and Bhojwani et al. (1984) and Shen
and Mullins (1984) used nodal buds from actively growing shoots. So-
dium hypochlorite has been used by all workers. Casein hydrolyzate was
incorporated during culture establishment (Lane 1979), presumably to
detect contamination rapidly since it was omitted for subsequent shoot
proliferation.
b. Shoot proliferation. A basal medium of MS has been used by all
workers except Zhao (1982), who found better growth with AS medium
(no details provided) compared to MS. The cytokinin of choice has been
BA. Lane (1979) and Singha (1980) established dose-response curves.
For ‘Bartlett’ a concentration of 5 or 10 puM BA resulted in maximum
proliferation, but at 10 p M shoots failed to elongate and were often
fasciated (Lane 1979).With ‘Seckel’maximum proliferation occurred a t 2
mg/liter (8.9 p M ) BA, but there was an inverse relationship between BA
concentration and shoot elongation ( Singha 1980). Others have noted
cultivar response to BA (Shen and Mullins 1984). Adding auxins and
gibberellins has resulted in varying responses. Incorporating NAA and

GA, individually with BA resulted in a slight reduction in proliferation
compared to BA alone, but adding both restored proliferation (Lane
1979). Both Singha (1980)and Bhojwani et al. (1984) found maximum
proliferation in BA medium, but incorporated NAA (Bhojwani et al. 1984)
and NAA and GA, ( Singha 1980) since shoot elongation was better. Poor
survival of shoots on BA medium has been found (Shen and Mullins 1984).
Low shoot proliferation and poor extension growth appear to be com-
monly encountered problems and ways to overcome them have varied. Tip
removal and inversion or horizontal placement of explants have been used
(Lane 1979). In addition, adding zeatin and 2iP to BA has resulted in
shoot elongation (Shen and Mullins 1984).Proliferation from basal explants
has been better than shoot-tips (Shen and Mullins 1984) and with the
rootstock l? calleryana D-6 (J. F. Hutchinson, unpublished). Agar brand
and concentration can also influence proliferation. With TC agar prolifer-
ation increased with increasing concentration up to 12 g/liter, whereas
with Bacto-agar proliferation increased up to 6 g/liter and then decreased
with higher concentrations ( Singha 1984).
c . Root initiation and growth. Both NAA and IBA have been used
successfully with Pyrus spp. Lane ( 1979)found that 10,uMNAA resulted
in 70% root initiation, but IBA a t the same concentration tended to be
toxic. Root morphology varied with auxin type; roots induced by NAA
tended to be thicker and more numerous than those induced by IBA,
which were longer and more fibrous. Although root elongation was poor
with NAA, Lane (1979) considered it an advantage since less damage
resulted upon transfer to potting medium. Callus formation around the
base of the stem has been observed with both NAA and IBA (Lane 1979),
but it was not considered a problem since roots arose from the stem and
not from callus tissue. Bhojwani et al. (1984)also noted callus formation
on an adult seedling clone, but the addition of 1mM PG reduced the callus
and increased both the percentage rooting and subsequent survival of
plantlets upon acclimatization. Singha ( 1980) reported poor root initia-
tion and callus formation with IBA and better rooting with 2 mg/liter ( 11
p M ) NAA, but the roots developed calluslike tissue and had poor lateral
development. However, a t lower NAA concentrations, root development
improved and there was little or no callusing. In contrast to Lane (1979),
Shen and Mullins (1984),using either 10,uMNAA or IBA, obtained good
root initiation with roots arising from the basal callus; plantlets were able
to be successfully acclimatized. Rooting response in Asian pear cultivars
was poor (Bhojwani et al. 1984), although a range of treatments was
tested, including auxin type and concentration, the presence of PG, and
attempts to root shoots in uiuo. Humidity can also influence successful
rooting (Lane 1979), since single shoots in unsealed tubes resulted in

shoot-tip death and necrosis of basal leaves with only 50% root initiation;
adding three shoots per tube raised humidity sufficiently to reduce stress
and resulted in 70% rooting.
3. Prunus (Stone Fruits). The genus Prunus contains a number of
species of horticultural importance, all of which have been micropropagated
with varying degrees of success.
a. Prunus armeniaca (apricot). Only a limited amount of information
is available for apricot. Cultures have been established from actively
growing shoots using sodium hypochlorite as a surface sterilant. Basal
media have varied. Skirvin et al. (1979)used MS salts and Staba (1969)
vitamins, whereas Snir (1984) evaluated a number of media and found
continual growth only on WPM (Lloyd and McCown 1980). Lower light
levels (-40 pmol/m2/sec) were better than higher (-200 pmol/m2/sec)
for establishment and maintenance of cultures (Snir 1984). For shoot
proliferation Skirvin et al. ( 1979)used BA plus NAA and Snir ( 1984)used
2iP. The cytokinin 2iP has been used successfully only for l? cerasifera
‘Newport’ (Garland and Stoltz 1981). Both groups reported problems,
either slow proliferation (Skirvin et al. 1979)or almost none (Snir 1984);
however, the latter overcame it by using nodal buds for culture mainte-
nance. Only limited rooting was obtained by Skirvin et al. (1979),but
Snir ( 1984) used a stem-wounding technique and 0.5 mg/liter (2.7 p M )
NAA to obtain a good rooting.
b. Prunus avium (Sweet Cherry). Scion cultivars of l? avium are
usually budded onto a clonally propagated rootstock, such as Mazzard
F12/1, which is difficult to propagate from cuttings (Feucht and Dausend
1976), or the semidwarfing Colt (l? avium xl? pseudocerasus).
Cultures of F12/1 have been established from actively growing shoots
from glasshouse-grown material (Jones and Hopgood 1979) and from
scion material that had been given its chilling requirement (Snir 1982a,b1.
Surface sterilization procedures have differed, with Jones and Hopgood
( 1979)using running tap water followed by sodium hypochlorite and Snir
( 1982a) using isopropanol. For establishment Snir (1982a) evaluated a
number of media and found Knop’s solution as used by Tabachnik and
Kester (1979)superior to that of Jones et al. (1977)or Boxus and Quoirin
(1974),both of which contain the macronutrients of MS. However, it was
not suitable for all cultivars, since ‘Burlat’ and ‘Renier’ degenerated.
After successful establishment, MS with 1 mg/liter (4.4 p M ) BA and 1
mg/liter (5.4 p M ) NAA has been used for shoot proliferation. Cultures of
F12/1, ‘Hedelfinger’, and ‘Sam’ were more difficult to establish than
those of various l? cerasus cultivars, although cultures grew satisfacto-
rily on the LS medium plus 0.5 g/liter casein hydrolyzate that was used
(Pauland F e x h t 1985).Removing the tip of the shoot improved prolifera-
tion, but F12/1 proliferated more readily than either cultivar.
In order to improve shoot proliferation Snir (1982a,b)used decapitated
shoots and was able to induce two to three times as many shoots to form
than with nondecapitated controls. In testing a range of concentrations
of various growth regulators with Colt, Wilkins and Dodds (1982/83)
obtained proliferation with BA alone, but adding IBA nullified the
response; NAA was a better supplement with BA because it increased
leaf production.
Root initiation has been achieved in F12/1 with M S and 3 mg/liter (15
p M ) IBA (Jones and Hopgood 1979),and with scions using half MS and
1mg/liter (4.9 p M ) IBA or 0.5 mg/liter ( -2.7 p M ) NAA with or without
wounding (Snir 1982a).Root formation was faster with NAA and wounding.
Interestingly BA has been used to induce roots in F12/1 (Nemeth 1979)
and Colt (Wilkins and Dodds 1982/83).
c. Prunus cerasus (Sour Cherry). As with I? avium, several approaches

to micropropagation have been used with r! cerasus. Cultures have been
established from expanding buds or actively growing shoots ( Borkowska
1983) and from dormant buds (Snir 1983). A problem associated with
dormant buds was the formation of a rosette of leaves with very little
shoot elongation that was overcome (Snir 1983)by transferring for shoot
proliferation to LS medium with 1mg/liter (4.4p M ) BA, 1 mg/liter (4.9
p M ) IBA, and 0.1 mg/liter (0.3p M ) GA,. A similar medium was used by
Borkowska (1983). Cultures of cultivars and a rootstock were readily
established from either actively growing shoots or from dormant buds,
but the multiplication rate during the first 8 months was higher when
cultures were established from the shoots (Paul and Feucht 1985).Culti-
vars proliferated as readily in uitro as the rootstock Cer W 10.
For rooting, Borkowska (1983)evaluated a number of basal media and
found half-strength MS or D medium (Druart 1980) the most suitable,
with the addition of PG having little effect with half-strength M S but
improving rooting with D medium. The use of the ammonium salt of IBA
improved shoot quality compared to free IBA (Borkowska 1983). In
contrast (Snir 1983) used an auxin-enriched medium for root initiation
and a hormone-free medium for root elongation. The presence of PG in
root initiation medium delayed root formation upon transfer to hormone-
free medium with ‘Chios’, but both treatments eventually resulted in
100% successful rooting, whereas with ‘Ben Zion’ rooting was delayed by
the presence of activated charcoal in root elongation medium. The use of
an auxin dip [50 mg/liter (246 p M ) IBA for 18 hr] and then transfer to
hormone-free medium was used successfully by Popov et al. ( 1976).
d. Prunus persica (Peach). Scion cultivars are usually budded or

grafted onto clonal or seedling rootstock although progress has been
made on propagating scions from cuttings (Issell 1976; Issell and Bolch
1976; Issell and Chalmers 1979; Couvillon and Erez 1980).
( i )Rootstocks. There is increasing use of clonal rootstocks for peaches.
Popular clonal rootstocks include the G F series ( Ppersica X P amygdalus )
from the Institut National de la Recherche Agronomique (INRA) in
France, which are compatible with a range of scions and adapt well to dry
and calcareous soils, and Nemaguard ( P persica X P dauidiana), which
has resistance to some nematodes.
Cultures have been established from actively p w n shoot-tips (Zuccherelli
1979; Miller et al. 1982; Kyriakidou and Pontikis 1983),nodes of actively
growing shoots (Reeves et al. 1983), and shoot-tips from dormant buds
(Ochatt and Caso 198313). Both calcium and sodium hypochlorite have
been used as surface sterilants, although an antibiotic and/or fungicide
mix has been used prior to sterilization (Vertesy 1980; Reeves et al. 1983).
Basal media have varied, the only consistency being that MS macro-
and micronutrients are used, although Kyriakidou and Pontikis ( 1983)
found Anderson’s ( 1978)macro- and micronutrients superior, resulting in
faster growing and greener cultures compared to those on MS, which were
slow growing and showed signs of degeneration. Anderson’s medium
(1978)has only one quarter the total nitrogen and potassium and twice
the phosphorus and iron content of MS. Various organic addenda have
been utilized, e.g., LS (Kyriakidou and Pontikis 1983),MS plus ascorbic
acid (Zuccherelli 1979), and Staba (1969) vitamins (Miller et al. 1982;
Reeves et al. 1983). Similarly, hormone types and concentrations vary.
Zuccherelli (1979)used 0.6 mg/liter (2.7 p M ) BA, 0.01 mg/liter (0.05 p M )
NAA, and 0.1 mg/liter (0.3 p M ) GA3 for shoot proliferation, but reduced
the BA concentration to one-sixth and increased GA3 fivefold for shoot
elongation with GF-677. Kyriakidou and Pontikis ( 1983)used 2.5 mg/liter
( l l p M ) B A ,0.1 mg/liter(0.5pM)IBA,andO.l mg/liter(0.3pM)GA3for
the same rootstock. Ochatt and Caso ( 1983b)used a constant concentra-
tion of 1 mg/liter (4.4p M ) BA and various concentrations of IBA and
GA3 with Red-Leaf. For Nemaguard, Miller et al. (1982)used 2 mg/liter
(8.9 p M ) BA and 0.1 mg/liter (0.5 p M ) NAA, while Reeves et al. (1983)
used 1 mg/liter (4.4p M ) BA and 0.01 mg/liter (0.05 p M ) IBA. Mainte-
nance of cultures appears to be a problem and there are few detailed
reports on long-term shoot proliferation. Reeves et al. (1983)tested the
effects of physical support and pH of the medium on survival, growth,
and condition of Nemaguard cultures and reported that agar medium a t
pH 5.8 was best, although mortality of cultures increased steadily with
increasing time in uitro.
Rooting is a problem, with the only success being reported by Zuccherelli
(1979),who used 0.1 mg/liter (0.5p M ) NAA. Nemaguard rooted poorly

with 2 mg/liter (9.8p M ) IBA (Reeves et al. 1983),although a considerable
improvement was found with 0.1 mg/liter (0.5p M ) NAA by Miller et al.
(1982), who observed that the vitamin content of the medium had a
dramatic effect. No rooting was found if all the Staba (1969) vitamins
were used; an assessment of individual vitamins attributed rooting
inhibition to the presence of riboflavin. The use of riboflavin in tissue
culture media is uncommon (de Fossard 1976),since plants in culture can
synthesize it in uitro (White 1951; Gautheret 1955). Riboflavin has been
reported to sensitize the in uitro photooxidation of a number of auxins
including IAA and IBA (Galston 1949), NAA (Gortner and Kent 1953),
and 2,4-D (Bell 1956). Recently Gorst et al. (1983) have shown with
Eucalyptus ficifolia that there is a degradation of exogenous IBA by
exogenous riboflavin in the presence of light. Thus, the effect of riboflavin
may have been due to light-induced degradation of NAA.
(ii)Scions. Early preliminary reports on scion cultivars ( Skirvin and
Chu 1977, 1978; Skirvin et al. 1979)involved the use of actively growing
shoots from field-grown trees, but these reports could not be reproduced
(Hammerschlag 1980)due to contamination and unsuitability of media.
Hammerschlag ( 198213)used a surface sterilization procedure consisting
of ethanol, hypochlorite, and antibiotics with sterile water rinses and
reduced contamination from 100 to 3% in ‘Sunhigh’. For this she used
actively growing shoot-tips from both field-grown trees and forced buds,
although shoot-tips from the field collected early in the season were
sensitive to antibiotics. Currently, Hammerschlag (private communica-
tion) uses shoot-tips from pathogen-indexed stock plants grown in the
greenhouse. These shoot-tips are sterilized in 0.5% sodium hypochlorite
plus 0.01% ’heen 20, followed by sterile water rinses. Shoots are culture
indexed twice (Knauss 1976)before going into the multiplication phase.
Vertesy (1980)obtained low contamination rates with many cultivars by
using a two-step surface sterilization procedure involving the initial use
of a low concentration of calcium hypochlorite, holding the cultures in a
refrigerator for 24 hr, followed by another treatment with a higher calcium
hypochlorite concentration. This procedure was not suitable for all culti-
vars tested. Skirvin et al. ( 1979)also used a two-step procedure, consisting
of hypochlorite with sterile water rinses followed by ethanol and further
sterile water rinses.
Macro- and micronutrients of MS have been used by all workers;
however, organic addenda have included those of MS plus ascorbic acid
(Skirvin and Chu 1977), Staba ( 1969) vitamins (Skirvin and Chu 1978),
Jacquiot (Vertesy 1980),and MS with elevated concentrations of thiamine-
HC1 and p-aminobenzoic acid (Hammerschlag 1982b). Hormonal supple-
ments have ranged from 2 mg/liter (8.9p M )BA and 0.1 mg/liter (0.5p M )
(Skirvin and Chu 1977, 1978) to 1 mg/liter ( 5 p M ) each of BA and IBA

and 0.1 mg/liter (0.3 p M ) GAS (Vertesy 1980). Hammerschlag (1982b)
tested various BA and IBA combinations and obtained best growth with
0.2 mg/liter (0.9 p M ) BA and 0.01 mg/liter (0.05 p M ) IBA; higher IBA
concentrations resulted in callus formation along the stem and higher BA
concentrations resulted in stem necrosis. Physical environment is also
important for successful establishment (Hammerschlag 1982b). Liquid
media, either static with filter paper bridges for support or agitated
(rotated),were better than agar, and 21' or 24OC were better than 26' or
28OC. Successful establishment of cultures is difficult with I? persica
scion cultivars and there is considerable genotypic variation (Vertesy
1980; Hammerschlag 198213)indicating that additional work is required
to improve the media or the preculture environment.
Preliminary results indicate that a reduced concentration of MS macro-
and micronutrients in solid medium favors rooting, although the addition
of 0.1-2 mg/liter (0.5-11 p M )NAA may be beneficial (Skirvin et al. 1980).
Later these workers (Skirvin et al. 1982)used a medium (MS-H)developed
for rooting of 'Harbrite' composed of MS macro- and micronutrients, with
a complex organic supplement including two auxins, two cytokinins, and
GA3. The use of MS-H resulted in multiple roots in about 50% of cultures
with extensive shoot growth, although it was not suitable for all Prunus
spp. Hammerschlag et al. (1987)recently reported that 65-100% rooting
could be achieved for several scion cultivars. Shoots were cultured on
half-strength MS medium for 6 weeks at 4OC in the dark followed by
culturing on half-strength MS containing 28.5 p M IAA plus 100 p M PG
for 2 weeks a t 26OC in the light. They also reported that length of time in
uitro increased percentage of rooting for most cultivars. Cytogenetic
analyses of micropropagated plants indicated that all plants were diploid,
2n = 2x = 16.
e. Prunus spp. (Plum). Several Prunus spp. make up the group com-
monly known as plums. Most scion cultivars, which include P domestica
and I? salicina, are budded or grafted onto rootstocks that include I?
insititia (e.g., Pixy) and I? cerasifera (Myrobalan)seedlings or selections
(e.g., Mr. S. 2/5).
( i )Rootstocks. Cultures have been established from actively growing
glasshouse material (Jones and Hopgood 1979)or actively growing forced
shoots (Hammerschlag 1982a; Loreti et al. 1982). Sensitivity to sodium
hypochlorite concentration has been observed with Mr. S. 2/5 (Loreti et
al. 1982). Basal media have consisted of MS macro- and micronutrients
with organic addenda varying, although most are based on the MS
formulation. Additional components have included ascorbic acid ( Loreti
et al. 1982) and p-aminobenzoic acid (Hammerschlag 1982a). Different
hormones have been used, e.g., 1mg/liter (4.4 p M ) BA, 0.1 mg/liter (0.5
p M ) IBA, and 0.1 mg/liter (0.3p M ) GA, with Pixy (Jones and Hopgood
1979). For Myrobalan two approaches and hormonal supplements have
been tried. With shoot-tips from seedlings, Hammerschlag ( 1982a)used
1 mg/liter (4.4 p M ) BA and 0.01 mg/liter (0.05 p M ) IBA. For Mr. S. 2/5
Loreti et al. (1982)used 0.6 mg/liter (2.7pM)BA, 0.01 mg/liter (0.05p M )
NAA, and 0.1 mg/liter (0.3p M ) GA3 for shoot proliferation; they reduced
the BA concentration to one-sixth and increased GA3 five-fold for shoot
elongation, as done earlier by Zuccherelli (1979)for the peach rootstock
G F 677. Agar-gelled media have been used (Jones and Hopgood 1979;
Loreti et al. 1982),but liquid medium for 4-7 days followed by transfer to
agar medium was necessary for maximum shoot proliferation (Hammer-
schlag 1982a). Incorporation of 1 mM PG enhanced shoot proliferation
and elongation of Pixy (Jones and Hopgood 1979).
Good rooting has been obtained with Pixy (Jones and Hopgood 1979)
using 3 mg/liter ( 15 p M ) IBA with the inclusion of 1mM PG improving
number and length of both shoots and roots. For Mr. S. 2/5 shoots, 1
mg/liter (4.9 p M ) IBA resulted in good, but slow, rooting; the type of
medium that shoots had been on previously influenced the process (Loreti
et al. 1982). Hammerschlag (1982a) evaluated the effects of a range of
factors on rooting of explants derived from 1-year-old Myrobalan seed-
lings. She found that a 2-week dark period prior to transfer to light was
essential for maximum response, and that temperature determined the
rate of root formation, with 26’ being better than 21’C. She obtained a
better response with IAA than with IBA, but IAA concentration was
important. The presence of GA3or alteration of macro- and micronutrient
concentration was not important for rooting.
(ii) Scions. Cultures have been established from actively growing
glasshouse material using shoot-tips (Rosati et al. 1980)ornodes (Baleriola-
Lucas and Mullins 1984)and from field-grown material (Pietropaolo and
Reisch 1984). The time of the year is important. Material collected in
autumn is more difficult to establish than that collected in summer
(Pietropaolo and Reisch 1984). Basal media have been based on MS,
although the form of chelated iron (Rosati et al. 1980) and the concentra-
tions of organic addenda (Pietropaolo and Reisch 1984)have differed. For
shoot proliferation the hormones have varied not only in terms of concen-
tration, but also with the species and cultivar being tested. Rosati et al.
(1980)reported that growth of P salicina ‘Calita’was satisfactory with 1
mg/liter (4.4 p M ) BA, 0.1 mg/liter (0.5 p M ) IBA (as ammonium salt),
and 0.1 mg/liter (0.3puM)GA3 ( a sammonium salt).With P domestica 1.1
mg/liter (4.9 p M ) BA only was used for ‘Stanley’(Pietropaolo and Reisch
1984),but 1 mg/liter (4.4 p M ) BA and 0.1 mg/liter ( -0.5 p M ) IBA were
used with ‘d’Agen’and ‘d’Ente 707’ (Baleriola-Lucas and Mullins 1984).
During each culture period 10-20 shoots of ‘Calita’ formed. Increasing

IBA concentration tenfold shortened internode length, making subcultur-
ing difficult, but increasing GA, concentration five-fold had no effect on
shoot elongation (Rosati et al. 1980).Shoot proliferation was satisfactory
on BA media for ‘Stanley’ (Pietropaolo and Reisch 1984), but with
‘d’Agen’and ‘d’Ente 707’ the presence of 1 mM PG was essential for
growth and survival of cultures (Baleriola-Lucas and Mullins 1984).Omit-
ting PG resulted in a rosette of leaves and subsequent death of cultures.
Short photoperiods appear to be more favorable for shoot proliferation
with plums. Either a 10 :8 or a 16 :8 hour (light :dark)cycle was suitable
for ‘Stanley’ (Pietropaolo and Reisch 1984). Photoperiods in excess of
16 hr resulted in weak growth and chlorotic cultures for ‘d’Agen’and
‘d’Ente 707’, with the optimum cycle being 12 : 12 ( Baleriola-Lucas and
Mullins 1984).
Several different approaches have been used to initiate roots on plum
scions. For ‘Calita’ the effects of media modification and temperature
have been assessed (Rosati et al. 1980).Temperature mainly affected the
time required for root initiation and the length of the roots produced. At
15OC both the time needed for root initiation and the length of the roots
were less than a t 21’ and 26OC; the addition of activated charcoal
reduced root initiation a t all temperatures. Adding GA3 overcame the
delay of rooting a t 26OC, but had no effect a t 15’ or 21OC. With I!
dornestica cultivars a reduction in concentration of macro- and micronu-
trients has been used for rooting. Pietropaolo and Reisch (1984)reported
that after 3 weeks exposure to 2-6 mg/liter (9.8-29 p M ) IBA, the percent-
age rooting, root number, mean root length, and subsequent plant sur-
vival were better than if lower auxin concentrations were used. In contrast,
the incorporation of 1 mM PG and the use of agar-gelled medium were
essential for root initiation with ‘d’Agen’ and ‘d’Ente 707’ (Baleriola-
Lucas and Mullins 1984). The number of subcultures through which
shoots had passed during stage I1 was also important, with rooting
occurring only after five or more subcultures; shoots were able to form
roots in the presence of BA ( Baleriola-Lucasand Mullins 1984).Condition-
ing of shoots by growth in a 12 :12 (light :dark) photoperiod and the
importance of subculture number may explain why Nemeth (1979)was
able to obtain rooting in the presence of BA with a number of species.
4. Minor Fruits. a. Diospyros kaki (JapanesePersimmon). Early re-

ports emphasized callus studies; Yokoyama and Thkeuchi ( 1976)induced
callus on hypocotyls from embryos of immature fruit and were able to
regenerate roots and shoots. Later, Yokoyama and Thkeuchi (1981) ob-
tained callus from cambial tissue of adult trees but were only able to
induce roots and some abnormal structures.
Micropropagation using shoot-tips and dissected buds from stock

plants of several cultivars was achieved on MS basal medium supplemented
with 1 mg/liter ( 5 p M ) zeatin and 40 mg/liter (0.1 mM) adenine sulfate
(Cooper 1983; Cooper and Cohen 1984). Other cytokinins were unsatis-
factory. BA resulted in explant death and 2iP caused excessive callus
formation. Proliferation from nodal explants was superior to shoot-tips.
Root initiation in vitro was difficult and best rooting was obtained by
dipping the shoot bases in an aqueous solution of 1000 mg/liter (4.9 p M )
IBA and transferring the treated shoots to a potting medium of fine
pumice in a high-humidity chamber. The use of other potting media was
less successful.
b. Olea europaea (Olive). Preliminary work on 0. europaea (Rugini and
Fontanazza 1981)using sucker explants and MS basal medium resulted
in unsatisfactory shoot growth. A new shoot proliferation medium was
developed, based on chemical analyses of shoots and mature embryos, to
which 4 mg/liter (20p M ) zeatin or 2iP was added (Rugini 1984).Satisfac-
tory rooting was obtained with either half-strength MS or Bourgin and
Nitsch (1967)medium using 1 mg/liter ( 5 p M ) NAA.
c. Morus alba, Morus indica (Mulberry). Shoots have been induced
from hypocotyls cultured on MS containing the cytokinin-active urea
4-PU (Ohyama and Oka 1982)and from nodal buds of mature trees with
MS plus 1 mg/liter (4.6 p M ) kinetin and 0.5 mg/liter (2.7 p M ) NAA
(Patelet al. 1983),which induced both a shoot and roots to form. Adventi-
tious shoots developed on leaves from established shoot cultures that had
been soaked for 48 hr in MS liquid medium containing 2.2-8.9 p M BA
before being cultured on solid medium of the same composition (Mhatre
et al. 1985). These shoots were then rooted and the resulting plants
established in soil.
d. Ficus carica (Fig). Shoots have been induced from apical meri-
stems of a number of cultivars (Muriithi et al. 1982) using M S with 0.1
mg/liter (0.4 p M ) BA, 0.18 mg/liter (0.1 p M ) NAA, and 0.03 mg/liter
(0.1 p M ) GA3. While shoot proliferation was apparently not achieved, it
was noted that medium browning was a problem that could be alleviated
by weekly transfer to fresh medium. Rooting was obtained with MS
supplemented with 0.5 mg/liter (2.5 p M ) each of IBA and NAA, where
incubation in darkness was more rapid and superior to continuous light.
Plants produced in this manner were virus symptomless for fig mosaic.
C. n e e Nuts
This is a botanically diverse group of plants growing mainly in temper-
ate regions. They are generally characterized by their edible seeds and
high oil content (Janick 1986).
1. Castanea sativa (Chestnut). Chestnuts are difficult to root from

cuttings (Vieitez 1974); initial research looked upon tissue culture as a
means of propagation. Callus has been induced from cambial tissue
(Jacquiot 1950) and cotyledons (Vieitez et al. 1978). A basal medium
supplemented with 1or 10 mg/liter (4.5or 45 p M ) 2,4-D plus 0.5 mg/liter
(2.3p M )kinetin or 0.5 mg/liter (2.2 p M ) BA was best for callus initiation.
Root initiation occurred with 10 mg/liter (49 p M ) IBA plus 0.5 mg/liter
(2.3 p M ) kinetin or 0.5 mg/liter (2.2 p M ) BA, and with 10 mg/liter (54
p M ) NAA plus 0.5 mg/liter (2.3 p M ) kinetin.
Several reports have been published concerning initiation and mainte-
nance of shoot proliferation, mostly from seedling material. Using
3-month-old seedlings and nodal explants, Vieitez and Vieitez (1980b)
induced shoot proliferation on M S medium with 1mg/liter (4.4 p M ) BA
in which 50% of explants produced 10-20 shoots 3-5 cm tall. Zeatin
induced a higher percentage of buds to proliferate and the resulting
shoots were more vigorous, but the proliferation rate was reduced. This
was confirmed by Vieitez and Vieitez (1980a),who reported that shoots
were stunted with 5 mg/liter (22p M )BA, but elongated when the BA was
reduced to 0.1 mg/liter (0.4 p M ) . Rooting could be achieved by exposing
shoots to 1 mg/liter (4.9 p M ) IBA for 8 days followed by transfer to
hormone-free medium, in which roots emerged in 2-3 weeks, with about
50% rooting occurring after 8 weeks. An interesting observation was that
rooting was best with initial cultures and poorer with shoots that had
been subcultured. These general trends were confirmed by Rodriguez
(1982c,d)and Vieitez and Vieitez (1983).
In a more detailed study, Chevre et al. (1983)found that a modified M S
medium, with calcium and magnesium concentrations doubled, 1mg/liter
(4.4 p M ) BA, 3.5% sucrose, and a pH of 4, was suitable for shoot
proliferation of both juvenile and adult growth phases. Shoot elongation
was poor with adult material and it was necessary to add 1mg/liter (5.7
p M ) IAA, 1 mg/liter (5.3 p M ) adenine, and 0.5% activated charcoal.
Using adult material of various clones of C. sativa XC. crenulata hybrids
with resistance to Phytophthora cambivora and I? cinnamomi, Vieitez et
al. (1983)evaluated a range of treatments affecting culture establishment
and shoot proliferation. Experimental trees were pruned to ground level
and shoots were allowed to grow; establishment was best with shoot-tip
explants from shoots collected in winter, stored for about 3 months a t 4OC
and then sprouted in the glasshouse. Storage of shoots for longer periods
led to explant necrosis. Cultures were established on MS with half-
strength nitrate and with 0.1-1 mg/liter (0.4-4.4 p M ) BA. Continued
maintenance on M S with half-strength nitrate resulted in succulent
shoots with flaccid tips and elongated dark green leaves, and proliferation
capacity was rapidly lost. By screening a range of macro- and micronutri-
ent basal media, it was found that Blayde’s (1966) medium, Heller’s
( 1953) medium + 1 mM NH,NO, and Lepoivre’s (Quoirin and Lepoivre
1977) medium with 0.1-0.5 mg/liter (0.4-2.2 p M ) BA were suitable.
Although limited success has been achieved with mature material,
there are still a number of problem areas. Explant browning during
establishment has been observed (Jacquiot 1950; Vieitez and Vieitez
1980b; Chevre et al. 1983; Vieitez et al. 1983). This has been overcome
by soaking explants in sterile water after surface sterilization and prior
to transfer to media, a method also successfully employed by Creswell
and Nitsch (1975)for Eucalyptus grandis. Rooting was not achieved in
adult material (Chevre et al. 1983) and to only a limited extent (Vieitez
et al. 1983) by using auxin dips followed by transfer to hormone-free
medium. In addition there are considerable differences among cultivars
in response to shoot proliferation and rooting (Chevre et al. 1983; Vieitez
et al. 1983). Micropropagated plants from chestnut seedlings have been
used to study the formation of mycorrhizal syntheses with the fungus
Paxillus involutus in vitro, but no attempt has been made to evaluate
the effect of this association on survival and growth of acclimatized
plants (Strullu et ul. 1986).
2. Juglans spp. (Walnut). Early studies involving Juglans spp. were

concerned with root initiation from callus as a means of solving problems
associated with the rooting of cuttings. Jacquiot (1951) obtained no
callus on J. regiu (English walnut) explants using an auxin-free medium.
Later, callus was induced and maintained for a short time (Cummins and
Ashby 1969) from stem tissue of J. nigra (black walnut) using White’s
( 1963)medium with NAA and kinetin, although no differentiationoccurred.
Callus and root initiation have been achieved from cotyledon explants of
J. regiu (Rodriguez 1982a) using 2,4-D plus kinetin for callus initiation
and NAA plus kinetin or BA for root initiation.
Shoot proliferation has been induced and maintained from sterile
seedlings of J. regiu (Rodriguez 1982b)using BA and IBA. Research with
the hybrid rootstock ‘Paradox’ (J. hindsii XJ. regiu) has been more
successful. In testing a range of basal media, Driver and Kuniyuki ( 1984)
found that performance was better on WPM or B-5 than on MS or Cheng
(1978) media with 4.5 p M BA and 5 p M IBA. However, continuous
maintenance of cultures could not be achieved and after ten subcultures
the shoots formed were reduced in size and could no longer proliferate.
%sting individual components of the medium a t several levels resulted in
a medium (DKW)on which growth was faster and could be sustained.
Rooting could be achieved on WPM with 30 p M IBA or by dipping
shoots in 5 mM IBA followed by direct rooting and acclimatization.
3. Prunus amygdalus (Almond). Cultures have been established from

dormant shoot-tips (Kester et al. 1977; Tabachnik and Kester 1977) and
actively growing shoots (Rugini and Verma 1982, 1983).A basal medium
based on Knop’s macronutrients and MS micronutrients and organics
has been used with 0.7-1 mg/liter (3.1-4.4 p M ) BA for culture establish-
ment. Incorporating low concentrations (0.01 mg/liter, -0.05 puM) of
auxins tended to produce callus from ‘Nonpareil’ shoot-tips (Tabachnik
and Kester 1977),whereas 0.1 mg/liter (0.5p M ) NAA was used routinely
with ‘Ferragnes’(Rugini and Verma 1983).For shoot proliferation, Knop’s
macronutrients with MS micronutrients and organics and l.mg/liter (4.4
p M ) BA have been used; half-strength MS medium produced chlorotic
shoots (Tabachnik and Kester 1977). In contrast, Rugini and Verma
(1983) used MS medium for both shoot proliferation and elongation,
achieving elongation by emitting NAA and reducing BA from 0.7 mg/liter
to 0.2 mg/liter (3.1 to 0.9 p M ) .
Rooting remains a problem. Limited rooting occurred using IBA plus
dark incubation, or NAA plus light, or sterilized vermiculite in tubes
with no auxin (Tabachnik and Kester 1977). Better rooting was achieved
by Rugini and Verma (1982, 1983) using the basal medium of Bourgin
and Nitsch (1967)with 1mg/liter (5.4p M ) NAA and dark incubation for
14 days. Afterwards, shoots were transferred to hormone-free liquid
medium with vermiculite as a physical support. Alternatively, a root
induction treatment with NAA for 4 days was followed by transfer to a
medium containing IAA for root elongation.
4. Corylus avellana (Hazelnut, Filbert). Research on micropropagation

has concentrated on developing procedures with seedling material and
adapting these techniques to cultivars. Anderson ( 1983)tested a range of
macro- and micronutrient formulations with LS organics plus adenine
sulfate and found Anderson (1978)medium superior to M S or WPM. On
the other hand, Kai et al. ( 1984)used MS macro- and micronutrients with
Zuccherelli (1979) vitamins. A combination of 2 mg/liter (8.9 p M ) BA
and 1 mg/liter (4.9 p M ) 2iP was used by Anderson (1983), whereas 5
mg/liter (22 p M ) BA, 0.01 mg/liter (0.05p M ) NAA, and 0.1 mg/liter (0.3
p M ) GA, was used by Kai et al. (1984). Anderson (1983) observed the
development of pale yellow leaves on plants grown on MS medium, which
he felt was similar to salt toxicity. Kai et al. ( 1984)also observed chlorotic
shoots on MS medium, but alleviated the problem by changing the iron
source from FeS04. 7H20 and Na2EDTA. 2 H z 0 to Sequestrene 138 Fe
( Ciba-Geigy ).Shoot proliferation was better using basal explants instead
of shoot tips (Anderson 1983). More recently, Perez et al. (1985)found
that optimum shoot initiation from seedling shoot and cotyledonary
node segments was attained by culturing them on Cheng’s medium plus

25 p M BA for 15 days, then transferring them to the same medium with
0.5 or 2.5 p M BA.
Preliminary experiments on rooting (Anderson 1983) showed that
half-strength Anderson ( 1978)medium with 0.5 mg/liter (2.5pM)IBA or
0.5 mg/liter (2.9 p M ) IAA can be used. Kai et al. (1984) found 0.1
mg/liter (0.5p M ) IBA superior to IAA or NAA, with higher concentra-
tions of auxins tending to produce callus. In contrast, rooting was
achieved in 80% of the shoots obtained from seedlings when they were
incubated in Cheng’s medium with 50 p M IBA for 5 days, and then
transferred to the same medium without IBA (Perez et al. 1985).
Bchniques developed for seedlings have been adapted to ‘Daviana’,
although contamination rates were high (Anderson 1983). A number of
French cultivars have also been micropropagated, but the procedures
require further refinement before this technique is commercially viable
(Kai et al. 1984).
5. Carya illinoensis (Pecan). Pecans are native to southern United

States and Mexico, where production is largely from wild and seedling
trees. Selected cultivars are propagated by grafting (Janick 1986).
Initial attempts to culture pecans resulted in failure due to internal
contamination of buds with Alternuria, but cultures established from the
spring flush of growth were free of contaminants (Knox and Smith 1980).
Endophytic contaminants could be partially controlled by incorporating
200 mg/liter streptomycin and 40 mg/liter filter-sterilized pimarcin in the
culture medium (Wood 1982).
Cultures have been established from seedlings, the pretreatment of
which was apparently important (Hansen and Lazarte 1984). No shoot
proliferation occurred with explants from etiolated stock plants, but
proliferation was satisfactory on explants from stock plants grown on
16-hr photoperiods in a greenhouse. Furthermore, successful culture
establishment required daily transfer to fresh liquid medium for 4 days
followed by maintaining cultures on filter paper bridges in darkness for 2
weeks prior to transferring to low-intensity light ( 11pmol/m2/sec) for 1
week before growing them on 16-hr photoperiods a t 43 pmol/m2/sec.
Shoot proliferation has been induced from nodal explants using WPM
(Wood 1982; Hansen and Lazarte 1984). Hormonal supplements have
varied. Wood ( 1982)determined that a combination of 4 mg/liter ( 18p M )
BA and 0.001-1 mg/liter (0.005-4.9pM) IBA was best for shoot prolifera-
tion and elongation, whereas Hansen and Lazarte ( 1984) found best
shoot proliferation and elongation with 3 mg/liter (13 p M ) BA.
Rooting has been achieved (Hansen and Lazarte 1984) either in uitro
using a liquid medium enriched with IBA for 6-10 days followed by
transfer to hormone-free liquid medium or by in vitro exposure to 10

mg/liter (49 p M ) IBA for 8 days followed by transfer to potting medium
for acclimatization.
6. Pistacia Vera (Pistachio). Cultures have been established from ster-
ile seedlings (Alderson and Barghchi 1982; Barghchi and Alderson 1983a,
b, 1985)and from actively growing shoot-tips of the rootstock I? terebinthus
‘Tsikoudia’(Pontikis 1984).Explant browning has been a problem (Pontikis
1984),but could be controlled by weekly transfers to fresh medium during
the first 6 weeks. The assertion that techniques were developed for in
vitro clonal propagation of two commercial cultivars ( Barghchi and
Alderson 1985)is incorrect, since the 1-to 2-year-oldplants cultured were
derived from seed of the commercial cultivars.
Different culture media for shoot proliferation have been used. Barghchi
and Alderson (1983a)tested MS with various hormonal supplements and
-
found maximum response using between 2 and 8 mg/liter (8.9 and 36pM)
BA with low levels of NAA (0.25 mg/liter, 1.3p M ) ,but preferred to use
4 mg/liter (18 p M ) BA only as subsequent rooting was better. Pontikis
( 1984)used Anderson ( 1978)macro- and micronutrients with LS organics
and 2.5 mg/liter (11.1p M ) BA, 0.1 mg/liter (0.5 p M ) IBA, and 0.1
mg/liter (0.3p M ) GAS. Both groups reported tip necrosis after 4-6 weeks
on proliferation medium on some cultures, the cause of which is un-
known. More frequent subculturing of shoots may be needed to overcome
this necrosis.
A range of factors affecting rooting has been tested (Barghchi and
Alderson 1983a,b)in which half-strength MS with 2.5 mg/liter ( 12.3p M )
IBA with darkness for 7 days followed by transfer to hormone-free
medium has proved satisfactory. Pontikis (1984)used LS medium with 1
mg/liter (4.9 p M ) IBA and 89 mg/liter (0.55 mM) PG.
D. Problem Areas
1. Vitrification. The presence of vitrified, glassy, or translucent shoots
has been reported in a number of fruit trees, including Malus domestica
(Hegedusand Phan 1983;Vieth et al. 1983;Hutchinson 1984a; Zimmerman,
1984a),Prunus auium (Phan and Letouze 1983), P persica (Zuccherelli
1979), and other Prunus spp. (Quoirin and Lepoivre 1977; Druart et al.
1981; Kevers et al. 1984).In addition, the problem has been found in other
species, including Cynara scolymus (Debergh et al. 1981; Debergh 1983),
Dianthus caryophyllus (Hakkaart and Versluijs 1983; Leshem 1983a,b;
Ziv et al. 1983; Kevers and Gaspar 1985), Picea abies (Bornman and
Vogelmann 1984; Von Arnold and Eriksson 1984),Pinus radiata (Aitken-
Christie and Jones 1985),and Salk babylonic (Letouze and Daguin 1983).
Vitrification is a physiological disorder in which leaves are broad,
316 lAMES E HUTCHINSON AND RICHARD H. ZIMMERMAN
thick, translucent, and often wrinkled and fragile (Kevers et al. 1984).
Microscopic examination of vitrified leaves from Cynara scolymus has
revealed that the palisade layer is absent and only spongy mesophyll is
present (Debergh et al. 1981).Biochemical analysis of Prunus avium has
shown reduced levels of chlorophyll a and b, lower protein content, and
lower hydroxycinnamate-CoA ligase activity ( Phan and Letouze 1983).
In Dianthus caryophyllus lignin content was reduced (Kevers and Gaspar
1985).In other species there is a higher water content, generally increased
total peroxidase activity, reduced phenylalanine ammonia lyase activity
with reduced ethylene levels in established cultures, and an increase in
ethylene production when explants were transferred to vitrifying condi-
tions (Kevers et al. 1984).An hypothesis has been proposed (Kevers et al.
1984) that vitrification is due to a burst of ethylene as a result of some
stress, causing a decrease in phenylalanine ammonia lyase activity, thus
hindering the lignification process that would allow more water uptake
due to reduced cell wall pressure.
Various ways to reduce the problem have been adopted, such as a
reduction of ammonium (Letouze and Daguin 1983) or chloride ions
(Quoirin and Lepoivre 1977),change from liquid to solid medium (Sutter
and Langhans 1977;J. F. Hutchinson, unpublished results), and increased
agar concentration (Debergh et al. 1981; Debergh 1983; Ziv et al. 1983).
llansferring cultures from BA to 2iP-containing medium ( Zimmerman
1984a)has also been shown to reduce vitrification; the BA toxicity that
Werner and Boe (1980)alluded to might have been vitrification.
Studying vitrification has been hindered because the condition is
difficult to reproduce consistently, but by growing apple cultures on
medium solidified with Gelrite@,an agar substitute, the condition can be
induced a t will (Pasqualetto et al. 1986).Combining a reduced amount of
Gelrite@with a smaller than usual quantity of agar produced a mixture
with which vitrification could be eliminated a t no loss in rate of prolifera-
tion. At low concentrations of gelling agent, 4.4 p M BA generally caused
more vitrification than 2.2 pM, but as gelling agent concentration increased,
the difference between the two concentrations of BA disappeared.
2. Acclimatization. Many fruit trees grown in in vitro have been diffi-

cult to acclimatize compared to herbaceous plants, such as strawberries
or grasses, in part due to reasons discussed in Section II.A.4 and possi-
bly since the shoot-tip is exposed and not enclosed in a sheath of foliage.
Although many authors have successfully acclimatized tissue-cultured
plants of fruit trees, few reports exist with detailed information. However,
the use of high-humidity chambers (Jones et al. 1977; Machnik and
Orlikowska 1981)and misting (Zimmerman and Broome 1980; Hutchinson
1984a) is common. Recent research (Dunstan and n r n e r 1984) has
8. TISSUE CULTURE OF TEMPERATE FRUIT AND N U T TREES 31 7
shown that the behavior of tissue cultures during shoot proliferation can
influence subsequent rooting and successful acclimatization. During
acclimatization, important factors include the growing environment, the
potting medium, fertilization, and control of disease. From a commercial
standpoint the initiation of roots in culture is expensive, due to the
additional labor costs and the occupation of culture room space. Simulta-
neous rooting and acclimatizationof plantlets (Simmonds 1983;Zimmerman
and Fordham 1985) has considerable economic implications.
3. ’Iiueness-to-Qpe. For micropropagation to be adopted commercially
it is imperative that regenerated plantlets, whether they be rootstocks or
scions, be phenotypically identical to the original cultivar and as geneti-
cally stable as plants produced by conventional means. The problem is
additionally complex with fruit and nut species, since long-term field
testing is required to determine flowering and fruiting characteristics of
either self-rooted scion cultivars or micropropagated rootstocks that are
budded or grafted. Field experiments have been established using a
number of cultivars of micropropagated fruit trees ( Boxus and Quoirin
1977; Martin et al. 1983; Zimmerman 1981) and no abnormalities or
mutations have been reported. In a more detailed study Webster et al.
( 1985)evaluated the orchard establishment and performance of four scion
cultivars and found they were more difficult to establish, which was at-
tributed to lack of experience with micropropagated plants. Performance
varied with cultivar and there was no sign of mutation or difference in
fruit characteristics. Other studies now indicate that the treatment of
micropropagated trees prior to planting and their size and growth status
a t planting can have significant effects on the subsequent growth, flower-
ing, and fruiting in the orchard (Rosati and Gaggioli 1987; Zimmerman
1986; Zimmerman and Miller 1985). I t would be desirable to have meth-
ods to assess clones in uitro and analysis using isoenzymes (Aoki et al.
1974; Kuhns and Fretz 1978; Vinterhalter and James 1983) or immunol-
ogy (Raff et al. 1979) may be required.
4. Bacterial Contamination. It has generally been considered that
plants in uitro are sterile but it is becoming common to see the sudden
appearance of bacterial contaminants from freshly cut surfaces some
months after the establishment of cultures. Acinetobacter calcoaceticus,
which is present in soil and water and is often isolated from animals and
humans, has been isolated from Malus in which it is not considered to be
pathogenic; chlortetracycline has been used to control the infection, but
does not eliminate it (Zimmerman 1984a). Rinsing of affected shoots in
dilute sodium hypochlorite was more effective to control the organism.
An unusual strain of Xanthomonas campestris has been isolated from
in uitro apple shoots grown from meristem-tip explants of ‘Ftedchief
Delicious’apple ( Maas et al. 1985).Apparently this organism exists in or

on apple trees grown outdoors, but damage to plant tissue occurs only in
vitro. In addition Bacillus pumilis has been tentatively identified in
Malw cultures (Constantine et al. 1980)and was eliminated with vancomycin
in some cases. Young et al. ( 1984)eliminated contaminants from Malus
and other woody plants using a mixture of cefotaxime, tetracycline,
rifampicin, and polymyxin a t 25, 25, 6, and 6 mg/liter, respectively.
Bacterial contamination may be more widespread than commonly thought,
since early detection can be difficult with agar-gelled medium due to its
opaqueness. The use of Gelrite,@an agar substitute that is virtually
transparent, may allow earlier detection of contaminants.
111. VIRUS ELIMINATION
It is well known that many fruit trees are infected with a range of
viruses, resulting in a reduction of vigor, quality, and yield (Aitkinson
1971; Childers 1976; Westwood 1978). Virus distribution and/or concen-
tration is not uniform in a plant (Limasset and Cornuet 1949)and virus is
often absent or not detectable in the apical meristem, which led Morel
and Martin (1952) to postulate that it might be possible to isolate the
apical meristem to obtain virus-free plants. This is now common practice
and often used in combination with heat-treatment (Nyland and Goheen
1969). Several recent reviews on the use of meristem-tip culture for virus
elimination have been published (Quak 1977; Walkey 1978; Wang and
Hu 1980).
Meristem-tip culture has been used with fruit trees (Walkey 1972; Lane
1978)but is rarely practiced for virus elimination, thermotherapy being
the favored method (Fridlund 1980), although latent viruses have been
eliminated (Campbell 1962). An alternative method has been in vitro
grafting of the meristem-tip onto seedlings (Alskieff and Villemur 1978;
Huang and Millikan 1980; Jonard et al. 1983).
IV. GENETIC IMPROVEMENT

A range of in vitro techniques has been suggested as having potential
for plant improvement, such as regeneration from callus (Skirvin 1978;
Larkin and Scowcroft 1981, 1983; Reisch 1983) or protoplasts to induce
somaclonal variation (Vasil and Vasil 1980; Cocking 1983; Evans and
Bravo 1983a,b; Larkin et al. 1983; Karp and Bright 1985; Scowcroft,
1985),embryo culture (Yeung et al. 1981; Raghavan and Srivastava 1982;
Williams et al. 1987), and the production of haploids via androgenesis
(Nitzsche and Wenzell977; Vasill980; Bajaj 1983)or gynogenesis (San
Noeum and Ahmadi 1982; Yang and Zhou 1982; Evans et al. 1984). In
addition, protoplasts can be used for fusion (Schieder and Vasil 1980;
Evans 1983; Evans et al. 1983)or transformation studies (Manzara and
Lurquin 1983). Generally, little work has been done on using novel
breeding techniques for improvement of fruit and nut species compared
to other groups of plants, due in part to technical difficulties, their long
breeding cycle, and the relatively few research groups working on them.
A. Regeneration through Adventitious Pathways

A range of adventitious regeneration systems can be achieved in vitro.
Shoots or embryos can be formed directly on the explant (direct adventi-
tious development)or via a callus intermediary (Fig. 8.1).The morphogenetic
pathway used has implications regarding the “trueness-to-type’’ of
regenerants; if adventitious shoots or embryos are produced directly on
the explant, they are more likely to retain clonal fidelity (Broertjes and
Keen 1980)compared to those organs induced to form from callus (Larkin
and Scowcroft 1981).
Callus of pome fruits was first produced from immature fruit of Malus
(Letham 1958), Pyrus and Cydonia (Letham 1960) and since then has
been used to study a range of physiological and biochemical problems.
Examples include using Malus callus to study vigor in rootstocks (Messer
and Lavee 1969; Schneider et al. 1978), sorbitol metabolism (Chong and
B p e r 1972,1974),hormone metabolism (Epstein and Lavee 1975; Epstein
et al. 1975), and growth kinetics (Pareilleux and Chaubet 1980). Prunus
callus has been used to study response to cyanide (Heuser 1972)and the
effects of hormones (Feucht et al. 1974). More recently research has
evaluated the potential of adventitious regeneration systems.
1. Malus. Limited regeneration of shoots from callus induced from

various explants of open-pollinated ‘Golden Delicious’seedlings has been
achieved (Mehra and Sachdeva 1979; Liu et al. 1983a). In a series of
experiments, Kouider and co-workers were able to induce shoots from
cotyledon callus of ‘Delicious’. Using seed collected from mature fruit
from which the two seed coats were removed, they found that when the
embryonic axis or intact embryo was present with all or part of the
cotyledons, a single shoot and root were produced and adventitious
shoots were formed from callus at the cotyledon-callus interface. Shoot
formation was related to the size of the explant and occurred on cut
surfaces proximal to the embryonic axis (Kouider et al. 1984a). In addi-
tion, they found that shoot formation was delayed if explants were
incubated in 16 hr of light, as opposed to an initial 4-day dark treatment,
and that the response was dependent on the absence of the seed coats and
the presence of BA, coconut water, and malt extract in the medium
(Kouider et al. 1984b).Later it was shown that explant age influenced the
potential to form shoots (Kouider et al. 1985), with immature embryos
(up to 4 weeks postanthesis) only producing callus, older embryos ( u p to
about 10 weeks postanthesis) producing multiple shoots, and mature
embryos producing a single shoot and root. For cotyledon explants,
adventitious shoot number increased with explant age until seed taken
from fruit that had been stored for 6 months formed only callus. These
results indicate that the potential for regeneration is dependent upon the
presence of the embryonic axis and is absent in cotyledons from very
young embryos and from mature stratified seed. Korban and Skirvin
( 1985)reported that the plumule and hypocotyl sections of the shoot axis
of mature embryos from trees of ‘Delicious’and ‘Winesap’apple developed
shoots, but only those from ‘Winesap’ developed shoots from the root
section of the axis or from intact axes. niploid plants have been regener-
ated from endosperm callus (Mu et al. 1977).
Shoots a t low frequencies have been regenerated from root-induced
callus of micropropagated plants of M.25 (Jones et al. 1984) and inter-
node callus of M.9 (Wei-lunet al. 1979). In attempting to repeat the work
of Wei-lun et al. (1979),James et al. (198413)were not able to regenerate
M.9, but could regenerate internode callus of M.25 and M.27 by inducing
callus in the dark in the presence of NAA and regenerating in the light in
the absence of NAA. In addition, leaf disk callus of M.27 could be
regenerated by incubating in the dark, but not under low levels of light
(James et al. 1984b). Callus induced to form on internode stem sections of
in uitro-cultured ‘Akero’ apple grew best on full strength MS medium
with 1 mg/liter (4.9 puM) IBA and no BA (Evaldsson 1985). A few
cultures (about 4% of the total) formed shoots. Embryoidlike structures
have been produced on callus from seedlings of ‘Golden Delicious’(Mehra
and Sachdeva 1980)and from cotyledon callus of ‘Golden Delicious’ and
‘Phiriki’ (Rubos and Pryke 1984)’ although development to plants has
not been achieved.
In addition to regeneration via callus, direct adventitious shoot and
embryoid formation has also been reported. A range of explants (leaf,
cotyledon, and hypocotyl) from sterile seedlings of ‘Golden Delicious’
produced more shoots when incubated in darkness for 3 weeks and
transferred to light, with the exception of hypocotyl explants, where
initial darkness had no effect (Liu et al. 1983a). Adventitious shoot
formation is to some extent under phytochrome control, since red light
suppressed shoot formation on leaf explants, but this was negated by
subsequent exposure to far-red light (Liu et al. 1983a). Shoots were
produced from cotyledon explants of mature fruits of ‘Golden Delicious’
and ‘Phiriki’ (Rubos and Pryke 1984).Embryoids have been formed from
the micropylar half of ‘Golden Delicious’ nucellus tissue, 50 days

postanthesis, cultured in darkness (Eichholtz et al. 1979; Eichholtz and
Robitaille 1980). Extending this work James et al. (1984a) were able to
induce embryoids on nucellar tissue of four scion and four rootstock
cultivars on both basal and hormone-supplemented medium. When pol-
len from a red-leaved Malus cultivar was used for pollination, purple
pigmentation was noted in zygotic embryos and in shoots multiplied
from embryos (James et al. 1986). If the pollen were irradiated (30 kRad)
first, embryos and cultures without pigmentation were obtained, indicat-
ing origin from the gametophytic tissue. The only report on the direct
production of embryos on nonreproductive tissue is that of Liu et al.
( 1983b).They stated that globular to heart-shaped embryoids and shoots
were produced on leaf tissue of sterile seedlings, but no histological
evidence on the origin and development of the adventitious structures
was presented.
To date most regeneration has been from tissues of seedling origin and
this is of limited value to existing scions or rootstocks. However, the
important groundwork has been laid for future development and some
trends are starting to develop. The use of an initial dark incubation
followed by light has been important for many systems (Liu et al. 1983b;
James et al. 1984b; Kouider et al. 1984a,b, 1985),as has callus induction
on hormone-supplemented medium followed by transfer to hormone-free
medium for regeneration (Wei-lunet al. 1979; Matsuta et al. 1983; James
et al. 198413).There are important cultivar differences in the pattern of
response that may be under genetic control (James et al. 1984b).Regener-
ation from material of known genotype is important for plant improve-
ment and has been possible (James et al. 198413; Jones et al. 1984).
Regeneration from nucellar tissue is equally important since it is obtained
from immature seed, but is maternal in origin (James et al. 1984a).There
are, however, some anomalies. Kouider et al. ( 1984b)found the incorpora-
tion of malt extract (as well as BA and coconut water) essential for shoot
regeneration, whereas James et al. (1984a) found it inhibitory for
embryogenesis in all tested cultivars. Mu et al. (1977)found that growth
regulators were necessary for growth of endosperm callus, whereas James
et al. (1984a)were able to obtain hormone-autotrophic endosperm callus
in eight cultivars.
2. Pyrus. Callus has been induced from immature fruit (Lethan 1960)
and various parts of seedlings, and in the later case was reported to have
formed shoots (Mehra and Jaidka 1979) and embryoids (Mehra and
Jaidka 1980).Janick ( 1982)reported the formation of an adventive embryo
from an explant of endosperm plus nucellar tissue of ‘Old Home’pear, but
the embryo did not completely develop.
3. Prunus. Limited regeneration of shoots has been observed on callus

from leaves, cotyledons, and embryos of I! amygdalus seedlings (Mehra
and Mehra 1974), from callus on leaves, petioles, and internodes of I!
incisa X I! serrula (GM9)and I! cerasus ‘Griotte de Schaerbeek’ (Druart
1985a),from root callus of micropropagated I! dawyckensis, I! incisa X I!
canescens (Druart 1980, 1985a),and I! auium X I! pseudocerasus ‘Colt’
(Joneset al. 1984),and from internode callus of ‘Colt’(Jameset al. 1984b).
With I! mahaleb, rhizogenesis was found on leaf callus, but no shoot
formation occurred (Hedtrich 1977), whereas with I! lannesiana ( a root-
stock used for cherries in Japan),Matsuta et al. ( 1983)were able to induce
shoot formation from leaf callus on transfer to hormone-free medium.
Repeatable plant regeneration has been achieved from callus derived
from immature embryos of Prunus persica (Hammerschlag et al. 1985).
Chronological age of the explant as well as period of time in uitro was
shown to influence organ formation. A specific type of callus (smooth,
white, nodular) was identified as being highly regenerative and this
callus could be maintained for several months without losing morphogenetic
potential. Hammerschlag ( 1986c) recently reported that bacterial leaf
spot-resistant plants could be regenerated from this highly regenerative
callus following in uitro selection of cells for insensitivity to a toxic
metabolite produced by the leaf spot pathogen (Xanthomonascampestris
pv. pruni). Embryoid production from immature embryos of ‘Nonpareil’
almond occurs in response to 2,4-D and coconut water (J.Janick, unpub-
lished data).
4. Others. Callus initiation and rhizogenesis from cotyledon explants
(Vieitez et al. 1978) and adventitious buds and rhizogenesis from hypo-
cotyl and epicotyl (San Jose et al. 1984) of Castanea satiua has been
obtained. Embryogenic callus was produced on ungerminated cotyledons
of a C. satiua x C. crenata hybrid and this formed embryoids that
developed as far as the torpedo stage, but no complete plantlets were
recovered (Gonzalez et al. 1985). Embryogenesis has been achieved from
callus initiated from immature zygotic embryos of Corylus auellana
(Radojevic et al. 1975).
Somatic embryogenesis has been induced from cotyledons of imma-
ture embryos of five cultivars of J. regia and one of J. hindsii ( Tblecke and
McGranahan 1985). Adventive somatic embryos would then form on
roots, hypocotyls, and cotyledons of older somatic embryos. Embryos
were grown to maturity and the resulting plants were established in soil.
B. Protoplast Culture
Early attempts to isolate protoplasts from leaves of Pyrus serotina
were unsuccessful (Wakasa 1973),but they have been isolated from in uitro
and field leaves of f? communis (Ochatt and Caso 1986). Cell colonies
recovered from the protoplasts were grown to obtain callus from which
shoots could be regenerated. Rooting of these shoots was very limited. In
addition, protoplasts have been isolated from Malus ‘Golden Delicious’to
study ethylene production (Anderson et al. 1979), from young root-tips
of Prunus spp. to obtain mitotic chromosomes (Salesses and Mouras
1977), and from callus and suspension cultures of a number of Malus
cultivars (Hurwitz and Agrios 1984). For developmental studies proto-
plasts have been isolated from haploid callus and callus subsequently
formed from Malus ‘Orei’(Niizeki et al. 1983),where it was found that the
use of Pectolyase Y-23 with Onozuka cellulase R-10 was essential for
successful isolation. Kouider et al. ( 1 9 8 4 ~tested
) a range of protoplast
sources, including callus, suspension cultures, leaves, petioles, and stems,
and were able to obtain protoplasts from leaves, callus, and suspension
cultures and callus from callus and suspension cultures with leaf proto-
plasts failing to divide. Embryolike structures were obtained that formed
roo$. The yield of protoplasts obtained from young tissue-cultured leaves
of ‘Akero’and M.26 apple was increased two- to six-fold when 0.5 mit4
methionine was included in the medium (Wallin and Welander 1985).
Protoplasts isolated from apple endosperm or stem internode callus or
cell suspensions divided and formed microcalli of 30-40 cells, but did not
develop further (James et al. 1986). Protoplasts could also be isolated
from mesophyll cells obtained from leaves of several cultivars taken from
axenic shoot cultures and plantlets rooted in uitro. Cultures or plantlets
grown under a light intensity of 63W/m2 produced twice the yield of
protoplasts compared to cultures or plantlets grown under 18 W/m2. At
the lower light intensity, protoplast yield from leaves of rooted plants was
30 times that from leaves of shoots in axenic culture. Protoplasts from cell
suspensions were fused with mesophyll protoplasts, but no wall forma-
tion or division was observed in the surviving heterokaryons.
Protoplast technology is still in its infancy for fruit trees. Much has yet
to be learned about the isolation, culture, and development of protoplasts
before they can be manipulated to the extent of some herbaceous genera.
C. Haploids
Haploid plants are of considerable value to breeders, since homozygous
diploids can be obtained in a single generation and this is of particular
value to fruit and nut tree breeders because of the long life cycles.
Pollen grains from Malus ‘Jonathan’have been reported to form torpedo-
stage embryoids (Kubicki et al. 1975; Milewska-Pawliczuk and Kubicki
1977)and haploid callus in Malus ‘Orei’( Hidano 1982).Moderate success
has been claimed in China, where haploid plants of a crabapple (Wu
198l),‘Delicious’(Fei and Xue 1981),and several scions including ‘Ralls’,

‘Rainier’,and ‘Golden Delicious’ (Xue and Niu 1984)have been obtained.
Limited success has been obtained with the culture of anthers of
Prunus spp. Haploid callus has been obtained from I! arrneniaca ( Harn
and Kim 1972; Michellon et al. 1974), I! auiurn (Jordan 1974, 1975;
Jordan and Feucht 1977; Legeida et al. 1976),and r! persica (Michellonet
al. 1974; Hammerschlag 1983). Diploid plants have been regenerated
from anthers of I! auiurn (Seirlis et al. 1979).
A haploid apple, originated by in uitro gynogenesis with the chromosomes
doubled by colchicine treatment, has been propagated using standard
micropropagation techniques and plants have been established in soil
and grown to flowering (Duron and Lespinasse 1985).Similar results were
obtained with second-generation inbred plants of ‘Golden Delicious’.
D. Embryo Culture
Embryos were the first plant organs to be successfully cultured in uitro
on artificial medium (Hannig 1904)and one of the first fruit tree organs to
be cultured (Tukey 1933, 1937). In a series of papers Tukey (1934, 1938,
1944) described embryo culture techniques in relation to fruit develop-
ment for a number of genera including Prunus, Pyrus, and Malus.
Hybrid embryos from crosses of early-maturing I! persica cultivars were
frequently found to abort before maturity and it was suggested by
Davidson (1933,1934)that these could be cultured in uitro to extend the
range of materials available for breeding. The technique and media were
improved by Smith et al. (1969),Abou-Zeid (1973),and Ramming (1985).
Problems of poor root growth and acclimatization difficulties have been
overcome by establishing proliferating shoot cultures and using adventi-
tious root initiation (Ivanicka and Pretova 1980), thereby producing a
number of plants from a single cross. The following I! persica cultivars
have been released as a result of embryo culture: ‘Collins’, ‘Fillette’,
‘Goldcrest’,‘Mayfire’,and ‘Summerglo’in the United States and ‘Culemborg’,
‘Van Piebeeck’, and ‘Swellengrebel’in South Africa (Ramming 1983; D.
Ramming and R. Scorza, personal communication). In addition, I! persica
(nectarine) cultivars have been released in Argentina from embryo cul-
ture programs (Torroba and Frangi 1979; Torroba et al. 1980).
V. GERMPLASM CONSERVATION
The preservation of germplasm either for or from breeding programs is

important and can be done by storing seed or vegetative material. With
fruit and nut tree species, this is done with vegetative material as
plantations or in insect-proof greenhouses. This technique is expensive

and time consuming, but essential to prevent the spread of insects or
pollen-transmitted viruses. I n uitro techniques have considerable advan-
tages since labor and space requirements are considerably reduced. ?tyo
in uitro methods are available: ( 1) the use of minimal growth conditions
where the temperature is reduced, the culture medium modified (Kartha
1985), or the atmosphere modified (Bridgen and Staby 1983), which
offers short- to medium-term storage, and ( 2 ) cryopreservation (Withers
1985), where metabolic activity is suspended by immersion and storage
in liquid nitrogen, which offers long-term storage.
To date only a limited amount of work has been published on in uitro
germplasm storage of fruit trees and little is known of the response of nut
trees. Storage of proliferating shoot cultures of Malus ‘Golden Delicious’
(Lundergan and Janick 1979) and ‘Northern Spy’ (J. F. Hutchinson,
unpublished results) is possible for 12 months by incubating cultures in
darkness a t 1’ or 4OC. Cultures of three Prunus rootstocks in the prolifer-
ation stage were stored successfully for up to 10 months a t constant
temperatures ranging from -3’ to 8OC (Marino et al. 1985). Growing
freshly transferred cultures for 1-2 weeks before storing enhanced sur-
vival only at -3’ in the dark. Druart (1985b) reported that defoliated
shoots (terminal bud removed) of several Prunus and Malus species and
cultivars could be stored in uitro in the dark a t 2OC for up to 4 years; the
shoots would begin growing again after transfer to fresh medium and
placement in the light under normal cultural conditions.
There are no reports on successful cryopreservation of cultured mate-
rial although dormant buds of some Malus cultivars have been preserved
in liquid nitrogen for about 2 years and cultures subsequently established
(Sakai and Nishiyama 1978; Katano et al. 1983; Sakai 1985). Shoot-tip
explants of ‘Jonathan’ apple taken from in uitro proliferating cultures
survived freezing to -196OC, but regrowth following freezing was only
as callus, indicating tissue damage by the freezing and/or rewarming
treatment (Kuo and Lineberger 1985).
VI. COMMERCIAL APPLICATION
Research work on the micropropagation of fruit and nut trees has

lagged behind much of that on herbaceous species and consequently the
adoption of such techniques for micropropagation by commercial labora-
tories has only occurred in recent years. A number of laboratories have
been established specializing in fruit and other woody tree species in the
United States (Frecon 1980),Canada (Dunstan 19811, and Italy (Zuccherelli
et al. 1978; Loreti and Morini 1982). The major factors limiting the use of
326 TAMES F. HUTCHINSON AND RICHARD H. ZIMMERMAN
micropropagation for fruit and nut tree species are technical and eco-
nomic (Navatel 1980). Rchnical problems are solved by the appropriate
developmental research, but economic problems require more imagina-
tive thinking. Labor costs constitute 60-8070 of the total costs of produc-
ing in uitro plants, whether herbaceous (Anderson et al. 1977; Strain
1980) or woody (Brown and Sommer 1982). These costs are primarily
associated with media preparation, subculturing, and attention during
and after acclimatization. Media preparation time has been reduced in
commercial laboratories by the use of a cold dispensing technique, where
agar is added to the liquid medium and stirred so that the agar forms a
suspension during dispensing to the culture containers. Agar varies in
chemical composition (Bridson 1978)and quality, and is the most expen-
sive single ingredient in a medium. Recently an agar substitute, Gelrite,B
has become available that is similar in price to agar but has three
advantages: ( 1) it can be used a t about one-quarter the concentration, ( 2 )
it forms a clear gel allowing for easier detection of contaminated cultures,
and (3)it can be cold dispensed. Although research with a number of
Malus cultivars and ornamental species has shown that it is equal to or
better than agar for both shoot proliferation and root initiation (J. F.
Hutchinson, unpublished results 1, it can cause vitrification in cultures of
some species, e.g., Malus, which negates the advantages (R. H. Zimmerman,
unpublished data). Mixtures of agar and Gelrite@have been found to
retain most of the advantages of both gelling agents ( Pasqualetto et al.
1986). Subculturing is a manual task and will remain so for some time;
however, the time needed can be indirectly reduced by the use of
Bacticinerabrs@or hot bead sterilizers, where instruments do not have to
be held in a bunsen burner flame but are instead placed in the sterilizer
while other instruments are being used. The future may see the use of
nutrient replenishment (Maene and Debergh 1985; J. Aitken-Christie,
personal communication), robotics, or chemostats. The development of
root initiation and acclimatization as one process considerably reduces
time because a subculturing step is eliminated and culture room space
may not be required. Such techniques have been used with some Malus
cultivars (Simmonds 1983; Zimmerman and Fordham 1985). Some sys-
tems, e.g., preformed peat plugs, have been developed for seedling pro-
duction, but have been adapted for use in micropropagation. Other
systems have been and are being developed especially for use in tissue
culture. High humidity during acclimatization is usually achieved with
misting, although systems such as fogging have certain advantages since
the droplet size is smaller and insecticides, fungicides, and foliar fertiliz-
ers can be applied through the system (Press 1983). The use of a fogging
systems for root initiation and acclimatization for micropropagated shoots
is described in Read and Fellman (1985).
VII. CONCLUSIONS
The use of tissue culture for fruit and nut tree species has increased
substantially since the early 1970s. Virtually all of the major temperate
fruit tree species have been micropropagated with various degrees of
success and the important research areas pinpointed. Micropropagation
of most nut tree species is still a t the developmental stage, although
many have been micropropagated from juvenile material. Micropropaga-
tion, particularly of scion cultivars to produce self-rooted plants, will
open up new areas of research and allow for changes in traditional fruit
tree horticulture. These have been summarized by Faust and Fogle ( 1980)
and include the more economic use of high- and ultrahigh-density orchards,
since plants should be less expensive, the elimination of the graft union
may increase translocation of minerals such as calcium, and the more
rapid turnover of trees in an orchard will take advantage of new cultivars.
The elimination of rootstocks and the use of virus-tested clones will
require better orchard management, because trees may be more vigorous
and researchers will have to develop methods to control tree size with the
use of growth regulators, deficit irrigation management (Chalmers et al.
198l), or root restriction (Richards 1986). The primary application of
novel breeding techniques to fruit trees with success has been embryo
culture. The recent report by Hammerschlag ( 1 9 8 6 ~on ) in uitro selection
of disease-resistant cells followed by regeneration of peach plants from
the resistant cells illustrates the potential for some of these modern tech-
niques for plant improvement.
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9
Summer Pruning of Apple and
Peach Trees
Richard I? Marini and John A . Burden
Department of Horticulture, Virginia Polytechnic Institute
and State University,
Blacksburg, Virginia 24061
I. Introduction 351
A . History 351
B . Terminology 352
11. Peach 353
A . General 353
B . Vegetative Growth 354
C . n e e Physiology 355
D. Light Penetration 356
E . Fruit Production 356
F . Cold Hardiness and n e e Longevity 357
G . Summer Pruning in Intensive Orchards 358
H. Commercial Applications of Summer Pruning 359
111. Apple 360
A . Vegetative Growth 360
B . n e e Physiology 361
C . Light Penetration 363
D. Yield 363
E . Fruitsize 364
F . FruitQuality 365
IV. Summary 367
INTRODUCTION
A. History
Fruit trees are pruned to restrict tree size, ensure adequate penetration
of sunlight into the tree canopy, remove dead, diseased, or broken branches,
adjust crop load, facilitate orchard operations such as spraying and
harvesting, and maintain a balance between vegetative and reproductive
growth. Waditionally, commercial fruit orchards in North America have
been pruned during the dormant period. Dormant pruning invigorates
351
352 RICHARD P. MARINI AND JOHN A. BARDEN
shoot growth in the vicinity of the pruning cut (Gardner et al. 1952;
Elfving and Forshey 1976),and may aggravate tree crowding and reduce
fruit quality in orchards where tree spacing is not adequate. In contrast,
European gardeners have practiced summer pruning for more than 250
years (Blake 1917). Summer pruning has traditionally been thought to
induce fruiting, while dormant pruning induces vegetative growth (Saunders
1863; %key 1964).
During the early 19OOs, American pomologists were very interested in
summer pruning, primarily as a method of inducing flowering in young
apple trees. Evaluation and comparison of those experiments are difficult
because they usually lacked statistical designs and analyses and detailed
description of methods. Results of these early studies have been reviewed
(Blake 1917; Gardner et al. 1952; Gourley and Howlett 1941; Auchter
and Knapp 1946) and indicate that the response to summer pruning
depends on time and type of pruning, cultivar, rootstock, tree vigor, and
environmental factors. Summer pruning sometimes suppressed tree growth
more than comparable dormant pruning, and yield usually was not
adversely affected. Because of the variable responses, however, the gen-
eral conclusion was that summer pruning was not suitable for commercial
use (Gardner et al. 1952).
Most of the early researchers used trees on vigorous rootstocks in
widely spaced plantings. ll-ees in recent fruit plantings are more closely
spaced. Many growers have been unable to control tree size adequately by
means of size-controlling rootstocks, chemical growth retardants, spur-
type strains, or dormant pruning. ll-ee crowding hinders orchard opera-
tions, while productivity and fruit quality decline as tree interiors become
heavily shaded. The practice of summer pruning is now being reevaluated
as a possible method of controlling tree growth, inducing flower bud
formation, and improving fruit quality in more intensive orchard systems.
Mika (1986) reviewed the effects of pruning on fruit tree physiology.
Our review was undertaken to summarize the recent information on
summer pruning. Possible explanations for varying results are discussed
and areas of future research needs are identified.
B. Terminology
Pruning may be defined as the removal of plant parts to maintain a
desirable form by controlling the direction and amount of growth. Fruit
growers prune to shape a tree and to regulate the bearing of a tree so it
produces annual crops of quality fruit (%key 1964). The two types of
dormant pruning cuts are ( 1) heading, which is the removal of a portion of
a shoot or branch, and ( 2 ) thinning, which is the removal of a shoot or
branch a t its point of origin.
9. SUMMER PRUNING OF APPLE AND PEACH TREES 353
The term summerpruning is ambiguous and implies only that pruning

is performed while leaves are on the tree, without regard to the type or
severity of pruning being performed. Several types of summer pruning
cuts have been described:
Pinching was described by Quintency in 1719 (cited by Blake

1917) as early-season breaking of young vigorous shoots with
fingers, leaving two or three nodes a t the shoot base.
Heading refers to cutting current season’s shoots back, usually to
4-6 mature leaves.
Stubbing refers to cutting into 2-, 3-, or 4-year-old wood or to the
first spur or fruit.
Watersprout removal is the elimination of watersprouts, usually
before the base of the shoot has lignified.
Thinning refers to making several thinning cuts in the tree tops
during late summer to improve light penetration.
The Lorette system of pruning was described by Tukey (1964) and

consists of heading shoots to 6-10 mm when they are about 30 cm long
and 8 mm in diameter. The process is repeated a t approximately monthly
intervals through the summer.
In an effort to minimize labor costs, many commercial fruit growers
have recently developed methods of mechanical pruning that have been
referred to as mowing, shearing, topping, and hedging. In this review the
term topping will refer to nonselective mechanical heading of tree tops.
The terms shearing, mowing, and hedging refer to the nonselective
mechanical heading of shoots a t the tops and sides of trees, which are
usually grown in hedgerow systems. Summerpruning will refer to selec-
tive pruning cuts (heading and thinning) made by hand while leaves are
on the trees.
11. PEACH
A. General
Blake (1916, 1917) initiated one of the first summer pruning experi-
ments with peach in the United States in 1912. The practice proved no
more advantageous than dormant pruning, and there was little interest in
summer pruning of mature peach trees until the late 1950s, when Harris
and Brown ( 1958; Brown and Harris 1958) studied postharvest pruning
of early-season peach cultivars in California. At about the same time,
some progressive California peach growers began to top trees mechanically
after harvest (Anon. 1962; Christ 1979);this practice is currently used by

some growers in the mid-Atlantic region, either before or following har-
vest or both. Mechanical shearing of peach hedgerows has become a
standard practice to control tree size and shape (Emerson and Hayden
1975; Hayden and Emerson 1975, 1976,1979,1984; Phillips and Weaver
1975; Chalmers et al. 1981; Horton 1985; Walsh 1985). Summer topping
and shearing of peach are commonly said to have many advantages over
dormant pruning, including control of tree size and shape, improved light
distribution in the tree canopy, advanced fruit maturity, compressed
harvest period, improved fruit size and color, improved flower bud cold
hardiness, reduced pruning expenses, and suppressed tree vigor. How-
ever, recent data have not supported all these claims (Marini 1985,1986),
and other results (Walsh 1985; Hayden and Emerson 1984) must be
viewed as preliminary because long-term studies are ongoing.
B. Vegetative Growth
Detailed growth measurements of mature peach trees following sum-
mer pruning and shearing are limited, probably because collection of
such data is quite time consuming. A commonly used measure of overall
tree size and vigor is trunk diameter or trunk enlargement. Summer
heading of container-grown peach trees reduced the diameter increase of
trunks within 20 days after treatment and the reduction was positively
related to pruning severity (Rom and Ferree 1985). Final shoot diameter
of container-grown trees was similar whether the trees were headed after
60 or 90 days of growth (Rom and Ferree 1985); however, diameter of
current season shoots on mature trees was reduced more by pruning in
July than later in the season (Rom and Ferree 1984).Summer pruning had
little effect on trunk growth of mature trees (Daniel1 1973). n u n k size of
hedgerow trees also tended to be greater as shearing was delayed later in
the season ( Walsh 1985). Topping of mature ‘Loring’ trees in June, July,
or June plus July did not influence trunk enlargement, whereas July
topping of mature ‘Sunqueen’trees reduced trunk enlargement compared
to dormant pruning (Marini 1985). The ‘Loring’ and ‘Sunqueen’ trees
were grown on the same farm and were treated similarly, yet results were
conflicting. Peach cultivars appear to respond differently to similar prun-
ing treatments.
A more important measure of vegetative growth may be shoot length,
because tree crowding is related to excessive shoot extension. When
measured a t the end of the growing season, summer pruned and topped
trees were smaller and had shorter shoots than dormant pruned trees.
However, following dormant pruning, all trees appeared similar in size
with shoots of similar length (Marini 1985). The year following treat-
ment, summer pruned or topped trees usually had as much or more shoot
extension as dormant pruned trees. Shoots on young field-grown trees
(Blake 1917; Marini 1985) and container-grown trees (Rom and Ferree
1985) tended to grow several weeks later following summer pruning,
compared to unpruned or dormant pruned controls.
The vegetative responses to summer pruning seem to vary with tree
vigor, cultivar, time, and type of pruning. Summer pruning and shearing
generally tend to reduce tree size, but no more than a similar type of
dormant treatment. Secondary growth (trunk and branch enlargement)
and dry weight accumulation were usually reduced during the season of
summer pruning, but shoot growth was not suppressed the season after
treatment. The bulk of research data indicates that pruning in the
summer time is no more effective than a similar type of dormant pruning
for controlling tree size and growth.
C. n e e Physiology
Summer pruning may temporarily elevate net photosynthesis ( P n )and
transpiration (Tr) rates of leaves on container-grown peach trees. Rom
and Ferree ( 1985)found that Pn and Tr of leaves on container-grown trees
were increased within 3 days after treatment and remained at elevated
levels for 24 days when pruned 60 days after growth began. When
subsequent regrowth was pruned 30 days after the original pruning, a
second cycle of increased Pn and Tr was apparent after 10 days and was
maintained for 24 days. Net photosynthesis rate of leaves 3 nodes below
the pruning cut (on subsequent regrowth) and leaves 7 nodes below the
pruning cut (on the main shoot)had similar responses. At the conclusion
of the experiment, summer pruned trees had fewer leavedtree, less total
leaf area/tree, and leaves with smaller average size and lower specific leaf
weight compared to nonpruned control trees. Effects on leaves were
greatest from late-season pruning (Rom and Ferree 1985).
Partitioning of dry matter and carbohydrates have been studied only
in container-grown peach trees (Rorn and Ferree 1985). In general, dry
weights of all tree parts (leaves, shoots, and roots) a t the end of the season
were suppressed by summer pruning when compared to a nonpruned
control. Dry weight suppression was greatest when pruning was delayed
until later in the season. Pruning treatments generally did not alter the
root :shoot ratio. Distribution of water-soluble carbohydrates in various
plant tissues was not altered by summer pruning, whereas early summer
pruning (60 days after growth began) reduced root starch. Pruning 90
days after growth began increased total root carbohydrate content (Rorn
and Ferree 1985).
D. Light Penetration
n e e growth and fruit production are dependent on a tree’s ability to
intercept and utilize sunlight. Critical light levels have not yet been
established for development of peach flower buds and fruit, but cultural
practices that enhance light distribution in peach trees would probably
improve production of quality fruit. One of the major reasons for summer
pruning has been to improve light penetration into the tree canopy, but
the effectiveness of the practice appears to vary with time and type of
pruning as well as with the orchard system being studied. Light penetra-
tion of three hedgerow systems was increased an average of only 8% by
shearing the sides and top; the greatest increase in light levels was in the
south side of the tree (Rom et al. 1984). Kappel et al. (1983) compared
light levels in four hedgerow systems before and after summer shearing.
Six days after shearing, before regrowth, light levels were increased in the
top meter of the trees by approximately 5570,but light was not increased
in the lower portion of the trees. A month after shearing, after consider-
able regrowth, light levels were still greater than preshearing levels but
lower than immediately after shearing. Unfortunately, there were no
nonsheared controls for comparison in either of these studies.
Summer topping of mature open-center trees in June and/or July more
than doubled light levels in the interior portion of the tree for the
remainder of the season, but light levels at the tree periphery were not
enhanced (Marini 1985). Summer pruning also dramatically improved
light penetration into centers of young vigorously growing trees, whereas
summer topping had very little effect on light penetration (Marini 1985).
Continual shearing or topping of vigorously growing trees seems to result
in the formation of dense shoot growth a t the tree tops, which absorbs a
high percentage of light. This emphasizes the importance of careful
dormant pruning, or use of a shearing practice that makes cuts at
different levels, such as slotting saw pruning (Cain 1971; Ferree 1976)to
prevent the development of an extremely dense layer of branches, which
could limit light penetration.
E. Fruit Production
Most reports concerning peach yields following summer shearing have
involved hedgerow plantings. Postharvest shearing (September)of hedge-
rows resulted in greater yields than shearing in May through August
(Walsh 1985). Fruit size was reduced by preharvest shearing and the
reduction seemed to be cumulative; size progressively decreased with
each additional year of shearing. Unfortunately, a dormant pruned con-
trol was not included in the experiment. Compared to dormant pruning,
June shearing each year (when shoots were 20-25 cm long) increased yield
9-2370 for 4-year-old trees, depending on cultivar (Hayden and Emerson

1984). Shearing in July, August, or June and July reduced yield, but
average fruit size was unaffected by treatment. Summer topping of
mature open-center ‘Sunqueen’ trees in July ( 3 weeks before harvest)
reduced yield 1870,while summer pruning (thinning cuts) reduced yield
6% compared to dormant pruning. Reduced yield of summer topped trees
resulted from a combination of fewer fruit per tree as well as smaller fruit.
However, summer topping mature ‘Loring’ trees in June, July, or June
and July did not affect yield compared to dormant pruning (Marini 1985).
The limited information available indicates that summer pruning or
topping has minimal influence on peach fruit color, firmness, or soluble
solids (Johnson and Coston 1985; Marini 1985). Summer topping and
pruning have not advanced fruit maturity nor condensed the harvest
period (Marini 1985; Walsh 1985),but may actually delay fruit maturity
on young trees (Blake 1917). The influence of summer shearing and
topping on yield and fruit quality seems to vary with cultivar and time of
treatment. Published data do not conclusively indicate that summer
shearing is preferable to winter shearing in various orchard systems.
More research is needed to compare shearing or topping during the
summer with a similar treatment performed during the winter.
E Cold Hardiness and n e e Longevity

Time of pruning long has been shown to influence peach tree productiv-
ity and longevity (Gourley and Howlett 1941; Daniel1 1975). Reports
concerning cold hardiness following summer pruning are both sparse and
conflicting. Theoretically, summer pruning might either enhance or reduce
the cold hardiness of fruit trees. Cold hardiness of peach shoots was
related to carbohydrate levels (Flore et al. 1983). Improved light penetra-
tion following summer pruning may improve the carbohydrate status of
shoots and flower buds in center portions of the tree. However, pruning or
shearing during the summer also stimulated late-season growth, which
may delay the acclimation process (Blake 1917; Kappel et al. 1983). The
loss of foliage may also result in reduced carbohydrate levels throughout
the tree, which may adversely affect cold hardiness.
Walsh ( 1985)recently reported midwinter tree damage a t the southern
end of hedgerows sheared in August, but not on trees sheared in June or
September. Hayden and Emerson ( 1979)reported reduced tree mortality
in hedgerow plots sheared in June and July compared to dormant pruned
open-center trees; they suggested that reduced winter injury may have
resulted from shading of trunks in hedgerows oriented north-south,
rather than from shearing itself. Winter injury of 1-year-oldshoots exposed
to -27.8OC in the field was not influenced by summer shearing of hedge-
rows in July, August, or July and August (Rom and Ferree 1984). Sum-
mer topping and summer pruning had little influence on the onset or
duration of vegetative bud rest, or flower bud cold hardiness of mature
open-center trees over a 3-year period (Marini 1986).
The objective of several recent studies has been to determine the
susceptibility of peach trees to diseases following pruning at various
times of the year. Spring and summer pruning wounds were susceptible to
fungal gummosis incited by Botryosphaeriu dothidea (Moug. ex. Fr.)Ces
and de Not. for nearly a week after wounding, but wounds were most
susceptible following spring pruning (Reilly and Okie 1984). Pruning in
winter, early spring, or summer had no effect on the degree of Cytospora
infection on mature trees, but there was less dieback and wood discolora-
tion below the summer pruning cuts than winter or spring pruning cuts
(Wilson et al. 1984). Pruning cuts that left a raised collar of tissue a t the
branch junction resulted in the least infection, while cuts that resulted in
stubs had the greatest dieback (Wilson et al. 1984).Observations in New
Jersey indicated that trees topped or pruned in July or August were no
less susceptible to Cytospora infection than trees pruned in late winter,
and Cytospora infection was often observed in small stubs resulting from
summer topping in the top of trees (R. P. Marini, unpublished data).
Although data vary slightly with time and type of pruning, as well as
geographic location, summer pruning generally seems to be no more
effective for preventing winter injury or disease infection than late
winter pruning.
G. Summer Pruning in Intensive Orchards

The concept of intensive peach orchards to produce relatively large
crops in young orchards is currently being tested. Most high-density
orchard systems have been designed for mechanical pruning and harvesting
to minimize the need for skilled labor. Various types of summer pruning
have been utilized to eliminate vigorous, nonproductive, upright shoots
and allow adequate light penetration for the production of quality fruit.
The llitura system uses a rigid trellis on which the main tree limbs are
held in an angled position (Chalmers et al. 1978).Horton ( 1985)described
a Y-shaped system without trellis support. Both systems require preharvest
mechanical shearing to allow light for proper fruit development. The
designers of peach meadow orchards utilized extremely high tree popula-
tions (5000-10,000 trees/ha) and used summer pruning to maintain the
small tree size needed for the intensive orchard. The system was designed
for a combine harvester that separated fruit from the cut canopy. Early-
maturing cultivars were able to produce flower buds on the resulting
regrowth in areas with long growing seasons (Erez 1982). Young and
Crocker (1982)developed a peach orchard system that is a compromise

between the traditional open-center orchard and the meadow orchard.
n e e s were planted a t fairly high densities (1667-5000 treedha), but were
trained to an open vase shape. After harvest, trees were severely topped to
0.75 m above ground to leave the lower portion of scaffold limbs plus a few
side shoots. Summer topped trees grew later into the fall and had less
peach rust-induced early defoliation than dormant pruned controls. Sum-
mer topped trees in the intensive orchard produced greater yields and
larger fruit than traditionally spaced and pruned trees.
H. Commercial Applications of Summer Pruning

Decreasing availability of skilled labor, increasing labor costs for prun-
ing and harvesting, and the need for early returns, especially where peach
tree short life is serious, have stimulated interest in alternative peach
orchard systems. Such systems involve high tree populations, either as
hedgerows or meadow orchards, and utilize mechanical pruning and
sometimes mechanical harvesting. Summer pruning appears to be required
to maintain the tree size and shape to allow light penetration, spray
penetration, and movement of orchard equipment within the plantings.
In areas with long growing seasons, systems utilizing pruning during or
after harvest to allow subsequent growth of fruiting shoots for the
following season appear commercially feasible, although economic stud-
ies are not yet published.
Hedgerow systems requiring summer shearing have been studied in
northern areas of the United States for more than a decade. Such systems
provide the opportunity for rather large yields early in the life of the
orchard. Unfortunately, the cold winter climate of some of these areas is
not conducive to regular cropping of peaches. It is difficult to determine
the feasibility of these systems in areas that may not be profitable for
peach production. The hedgerow system, involving summer shearing,
has often been compared to traditional low-density, vase-shaped trees
that were dormant pruned. Walsh (1985) has attempted to eliminate
selective pruning altogether by shearing hedgerows a t various times
during the summer. Yields after 3 cropping years of such systems have
averaged no more than 12,000kg/ha, which is not particularly good when
the tree population of 667 trees/ha is considered. Leuty and Pree (1980)
found that standard peach orchards had lower yields than summer sheared
hedgerow systems during the first 3 years of production; however after 7
fruiting years, cumulative yields were similar for all systems. Packouts
have not been reported for the fruit, but average fruit size was reduced by
0.95 cm for a t least one season by preharvest shearing (Walsh 1985).
Therefore, it is difficult to determine if a hedgerow system with no
selective dormant pruning is a feasible alternative to traditional plantings.

Additional research is needed to determine if summer shearing is more
economical than dormant shearing in hedgerow systems.
Summer topping and summer pruning of mature vase-shaped peach
trees does not appear to offer any advantages over traditional dormant
pruning. Summer topping does allow better light penetration into the
tree, but fruit quality and cold hardiness of flower buds were not improved.
Summer topping reduced dormant pruning costs nearly 2570, but also
reduced the crop value 12970.Over a 2-year period, summer topping was
less profitable than dormant pruning mature ‘Sunqueen’trees, but results
may vary for other cultivars (Marini and Rossi 1985). Postharvest top-
ping may be a profitable practice that would limit tree size and reduce
pruning costs without adversely affecting fruit size. Summer topping of
young, nonbearing trees did not seem practical because dense growth in
the tops of trees caused shading of the tree interior. Summer pruning,
involving thinning cuts, may be advantageous in training young nonbearing
trees (Blake 1916).
111. APPLE
A. Vegetative Growth
For well over a century, pomologists in the United States have pro-
moted the idea that summer pruning suppresses apple tree vigor in
comparison to dormant pruning (Saunders 1863; Heinicke 1975; Hayden
and Emerson 1975; Utermark 1977; Carlson 1979, 1982; Stembridge
1979). In most of these reports, a seemingly logical hypothesis was
discussed: when new growth develops in the spring, the energy for such
growth comes from reserves in the tree from the previous year. After
vegetative growth ceases, the canopy of leaves not only provides energy
for fruit growth but also replaces the reserves within the permanent parts
of the tree. By removal of significant portions of the leaf surface prior to
the replacement of tree reserves, tree vigor in the next year is decreased
compared to dormant pruning. In a lengthy review in 1952, Gardner et al.
concluded that “summer pruning may have a dwarfing or an invigorating
influence (as compared with a corresponding winter pruning), depending
on its severity, kind, the stage of development of the plant and on
environmental conditions -particularly nutrient supply, soil moisture,
and light.” Since that time many reports have been published relating to
summer pruning effects on tree vigor. Conclusions vary for several rea-
sons, including the control treatment( s ) utilized, the measure(s ) of tree
vigor being evaluated, and when growth measurements were made. These
complications are in addition to other factors such as time and severity of

pruning, overall tree vigor, cultivar, and rootstock.
Regardless of the control used, summer pruning has often been shown
to depress trunk growth (Alderman and Auchter 1916; Greene and Lord
1983; Marini and Barden 1982a,e; Myers and Ferree 1983c; Preston and
Perring 1974; Sako and Laurinen 1982). In contrast, other reports indi-
cate no effect of summer pruning on trunk growth (Emerson and Hayden
1984; Ferree 1984; Thylor and Ferree 1984). Lord et al. (1979b) found a
decrease in some experiments but no effect in others.
Arrested or decreased rates of root growth have also been reported
(Thylor and Ferree 1981; Marini and Barden 1982e; Myers and Ferree
1983a; Bubenheim and Barden 1984). These effects of summer pruning
are not surprising. Considerable root growth occurs in the late summer
and fall when shoots and fruits are no longer major sinks for photosyn-
thates. The importance of leaves to the fall flush of root growth was
shown by Heinicke (1936).
Although conclusions vary in regard to the short-term effects of sum-
mer pruning on shoot growth, most can be reconciled. Several reports
indicate that shoots were shorter on summer pruned trees than controls
(Ferree and Stang 1980; Myers and Ferree 1983a,c; Eiylor and Ferree
1984). In each of these studies the controls were unpruned or the trees
were lightly dormant pruned, leaving measured limbs unpruned. It is
apparent, therefore, that the reported responses are to pruning in the
summer but do not directly deal with the hypothesis that summer
pruning is more dwarfing than dormant pruning. From these studies, we
can conclude that summer pruning suppresses shoot growth but we can
make no comparisons between summer and dormant pruning.
In a direct comparison of summer and dormant pruning, Marini and
Barden (1982a) found that total shoot growth per headed shoot on
August pruned trees was equal or greater than that of dormant pruned
controls. This result was supported by later work in which growth responses
were similar to three pruning severities applied during the summer and
winter (Barden et al. 1983).
Summer pruning has a minimal effect on growth in subsequent years.
Whether using young trees in containers or trees in the orchard, there is
general agreement that summer pruning is not devitalizing or more
dwarfing than dormant pruning (Lord et al. 197913; Myers and Ferree
1983b,c, 1984; Thylor and Ferree 1984; Greene and Lord 1983; Marini and
Barden 1982a,e; Barden and Marini 1984).
B. n e e Physiology
Summer pruning effects on various physiological processes have been
studied in both container-grown and orchard trees.
In container-tree studies both net photosynthesis ( P n )and transpira-

tion ( T r )were temporarily higher in basal leaves of summer pruned than
unpruned control trees (Thylor and Ferree 1984).Summer pruning delayed
the normal seasonal decline in Pn of basal leaves but had little impact on
dark respiration (Rd)or Tr (Marini and Barden 1982c; Myers and Ferree
1983a). In field studies, summer pruning was found to increase Pn, Rd,
and Tr of shoot leaves (Marini and Barden 1982~). This effect was greater
for interior than peripheral leaves. The temporary increase in Pn can
hardly be expected to compensate for the leaf area removed. There would,
therefore, be a sizable (depending on pruning time and severity) reduc-
tion in the production of photosynthate.
Using I4Cwith container-grown trees, Marini and Barden ( 1983)found
that photosynthates moved largely to the roots of both control and
summer pruned trees 2 days after summer pruning. At 21 days after
pruning, however, 14C-labeledphotosynthates moved primarily to regrowth
in pruned trees while movement in the controls was still to the roots.
These data indicate that differences in growth response to summer prun-
ing could occur depending on when the summer pruning is done. The
regrowth that results from summer pruning decreases markedly as the
pruning is delayed from June to August (Aselage and Carlson 1977;
Carlson 1979; Ferree 1984; Maggs 1965; Miller 1982).One could reach the
conclusion that pruning in June or July might depress trunk and root
growth more than would August pruning due to the lack of regrowth in
August. The confounding factor, however, is that the leaves on regrowth
may ultimately contribute to the reserves of the tree although at the
outset they were sinks. This is supported by the data of Bubenheim and
Barden ( 1984),who reported that autumn root growth was suppressed by
July and August pruning but not by that done in June.
The suppressed growth of trunk and roots has been frequently cited as
evidence of depleted reserves in the tree, which should lead to decreased
vigor in the subsequent year. However, there are very few published data
showing decreased shoot length in the year after summer pruning. A
tenable hypothesis is that apple trees characteristically store more reserves
than are required for normal growth. Priestley (1960) reported that the
overall level of carbohydrate did not limit growth. Thylor and Ferree
( 1986)found little if any effect of summer pruning on carbohydrate levels
in the basal stem and roots of containerized trees or the spurs or roots of
orchard trees.
Further insight was provided from a recent experiment where container-
grown trees were pruned during the dormant period or summer and trees
from each treatment were harvested monthly or bimonthly (Bubenheim
and Barden 1984). Fkgardless of when it was done, pruning invigorated
shoot growth. June pruning stimulated growth during the next year as
did dormant pruning. The invigoration from July or August pruning

(when little regrowth occurs) could be overlooked because it was not
manifested for several months-but it did occur. This could be an espe-
cially serious problem in short-term experiments in which growth is not
determined during subsequent seasons.
C. Light Penetration
One of the benefits of summer pruning is an increase in photosynthetic
photon flux density ( P P F D )within the tree canopy. The vital importance
of adequate PPFD is well documented for flower bud formation (Auchter
et al. 1926; Cain 1973; Lakso 1980), fruit set (Doud and Ferree 1980;
Jackson and Palmer 1977), fruit color (Doud and Ferree 1980; Heinicke
1966), fruit size (Heinicke 1966; Seeley et al. 1980), and soluble solids
(Doud and Ferree 1980; Heinicke 1966).
Porpiglia and Barden (1981) showed that immediately after pruning
shoots to 2 to 3 leaves in August, PPFD was increased both at the
periphery and within the canopy. However, in the year following this
summer pruning, PPFD was lower in summer pruned than control trees,
which were less severely headed in the dormant season. PPFD through-
out the canopy was increased for the remainder of the season by summer
pruning in mid-August (Marini and Barden 1982b). However, in the
following year, PPFD was lower within summer pruned trees than in
comparably dormant pruned controls. Morgan et al. ( 1984)studied effects
of summer pruning as a supplement to dormant pruning and found
increased PPFD penetration in summer pruned trees. Likewise, Thylor
and Ferree (1984) found that summer pruned trees were more open in
September and October.
The limited number of studies in which light penetration effects of
summer pruning were studied are in general agreement. Summer pruning
increases light levels within the tree canopy during the season of pruning
but may decrease levels during the year after treatment.
D. Yield
In several studies, summer pruning has been found to decrease yields;
in some cases no effect was found. Crowe and Brown ( 1975),Engel ( 1974),
Mika et al. (1983),Sako and Laurinen (1982),Stiles (1980,1981),and Van
Der Boon (1980) all reported decreased yields resulting from summer
pruning. Emerson and Hayden ( 1981,1984)reported that winter pruning
supplemented with summer hedging decreased yields compared to winter
pruning alone or winter or summer hedging. In a study involving summer
vs. dormant pruning, Barden and Marini ( 1984)found no effect on either
total number or weight of fruit per tree over a 4-year period. Thylor and
364 RICHARD P. MARINI AND J O H N A. BARDEN
Ferree (1984)reported a decrease in yield per tree from summer pruning

but no effect on yield expressed on a canopy volume basis.
The lack of reports that show an increase in yield as a result of summer
pruning are particularly interesting when compared to the proposed
benefits of summer pruning. Heinicke ( 19751, Stembridge (1977a,b, 1979),
Carlson ( 1979),and Utermark ( 1977)proposed two major benefits to be
derived from summer pruning: (1)buds break at the base of pruned
shoots and develop into fruiting spurs, and ( 2 ) increased light levels
within the tree canopy as a result of summer pruning strengthen spurs
and increase flower bud formation.
Although there have been a few reports of increased flowering following
summer pruning (Stembridge 1977b),there are others where no effect was
found (Greene and Lord 1983). Mika et al. (1983) found that summer
pruning decreased flower bud formation.
One of the proposed benefits of summer pruning is enhanced flower
bud development due to improved light penetration, and possibly a
redistribution of growth substances (Stembridge 1977b). Pruning in late
June and early July increased flower bud formation more effectively than
late pruning (Bunemann and Struklec 1980). The majority of research
data indicates no improvement in flowering or fruit set the year following
summer pruning compared to dormant pruning (Ferree and Stang 1980;
Greene and Lord 1983; Myers and Ferree 1983c; n y l o r and Ferree 1984).
In most of these studies, summer pruning was performed rather late
(greater than 90 days after bloom, which is after flower bud initiation).
Increased light levels late in the season appear to be of questionable
benefit to flower bud development. However, early summer pruning to
improve flowering may reduce fruit size.
We have also observed vigorous shoot growth the year following treat-
ment from spurs on 2- and 3-year-old wood within 25 cm below summer
heading cuts (Marini and Barden 1982a). Sometimes individual spurs
bloomed within a few weeks after August heading, especially on ‘Golden
Delicious’(unpublished data). These two responses would reduce slightly
the amount of bloom the following year. Summer heading of terminal-
bearing cultivars has also reduced flowering the following year because
flowers are normally produced on 1-year-oldwood (Ferree and Stang 1980).
Failure to confirm the anticipated effects of increased PPFD on yield
does not seem to be related to fruit set. Ferree and Stang (1980)found
some increases in fruit set, but Greene and Lord (1983) and Thylor and
Ferree ( 1984) found no effect.
E. Fruit Size
Published data relating time of pruning to fruit size are rather limited.
There are, however, several reports indicating decreased fruit size resulting
from summer pruning. Stiles (1980, 1981), Marini and Barden (1982d),
Myers and Ferree (1983b),Greene and Lord (1983),and Taylor and Ferree
(1984) all reported adverse effects of manual summer pruning on fruit
size. Myers and Ferree (198313)found that the suppression in fruit size due
to summer pruning was greatest from early July pruning and was
progressively less from August and September pruning. Ferree (1984)
found that summer hedging decreased fruit size when compared to either
hedging or hand pruning during the dormant season. Perring (1978)
found a 28% increase in fruit size from late summer pruning and Taylor
and Ferree (1984)found an increase in one year following a decrease the
previous year. These are the only reports seen by the authors that have
shown positive effects of summer pruning on fruit size. In some cases,
little or no effect of summer pruning on fruit size was reported (Preston
and Perring 1974; Lord and Greene 1982).
By removing sizable quantities of leaf area during the summer, it
should not be unexpected that fruit size can be suppressed. Whether or
not the effect is measurable could depend not only upon the cultivar,
growing season, and crop load, but particularly the timing and severity of
the summer pruning. On the basis of published results, fruit size suppres-
sion is one of the potential negative effects of summer pruning (Ferree
1984; Barden and Marini 1984; Myers and Ferree 1983b; Taylor and
Ferree 1984). Under most conditions smaller fruit may lead to both
lowered total yields and reduced returns per bushel.
F. Fruit Quality
Among the many effects of summer pruning of apple trees are various
aspects of fruit quality. The quality factors that are well documented
include red color, soluble solids, firmness, bitter pit incidence, flesh
calcium levels, and water core.
1. Fruit Color. Since red coloration of apple fruits is directly depend-

ent upon light, it would be logical to expect that increased PPFD as the
result of summer pruning would increase red color of apples. Several
studies have reported such increases (Vincent 1917; Seeley et al. 1980;
Lord and Greene 1982; Marini and Barden 1982d; Morgan et al. 1984;
Preston and Perring 1974; Stiles 1980; Taylor and Ferree 1984). A few
reports (Myers and Ferree 198313; Miller 1982)have found either no effect
or inconsistent results. As a result of very severe summer pruning in
Georgia, Miller (1982) found an increase in sunburned fruit.
With poor coloring cultivars or under marginal coloring conditions,
summer pruning may provide beneficial effects; with excellent coloring
strains under good coloring conditions, little or no benefit may occur.
2. Soluble Solids. One of the more serious undesirable effects of

summer pruning is a depression of soluble solids in the fruit. Marini and
Barden (1982d), Myers and Ferree (1983131, Greene and Lord (1983),
Thylor and Ferree ( 19841, and Morgan et al. ( 1984)have all reported lower
soluble solids following summer pruning. This response has not been
universal, as Miller (1982), Lord and Greene (1982), and Ferree (1984)
reported no effect of summer pruning on soluble solids. Worth noting,
however, is that we have seen no reports that indicate a positive effect of
summer pruning on fruit soluble solids. I t should not be unexpected for
fruit soluble solids to be reduced by pruning during the final 5-10 weeks
before harvest because of the reduced leaf/fruit ratio. Obviously, the
degree of this effect could vary markedly with the timing and severity of
the pruning, and particularly the proximity of the fruit to the pruning cut.
3. Firmness. Several researchers have evaluated summer pruning
effects on flesh firmness and the lack of an effect is consistent (Miller
1982; Lord and Greene 1982; Myers and Ferree 198313; Greene and Lord
1983; Thylor and Ferree 1984; Marini and Barden 1982d).
4. Bitter Pit. Several published reports have shown a decrease in the
incidence of bitter pit as the result of summer pruning (Borsboom 1976;
Preston and Perring 1974; Van Der Boon 1980; Marini and Barden
1982d).Other researchers (Engel 1974;Greene and Lord 1983;Schumacher
et al. 1974) found no effect. Myers and Ferree (1983b)reported a sizable
reduction in the incidence of cork spot, another calcium-related disorder,
from summer pruning.
Since shoot growth competes with fruit for available calcium, summer
pruning might have the potential to reduce this competition and thereby
increase calcium levels in the fruit. Lord et al. (1979a) and Preston and
Perring ( 1974)reported increased fruit calcium levels after summer prun-
ing. Lord and Greene (1982)and Marini and Barden (1982d)reported no
increase in fruit calcium. The results are inconsistent as to both calcium
levels and the incidence of bitter pit. Shoot growth is most competitive
for calcium during active shoot growth. Most summer pruning has been
done in July or August, when shoot growth had already stopped. There-
fore, shoot removal in July or August may have less effect on fruit cal-
cium levels than would pruning in June when the resulting regrowth may
compete with the fruit for calcium. It hardly seems likely that summer
pruning can be depended upon to provide a consistent increase in cal-
cium and a decrease in the incidence of bitter pit.
5. Water Core. Relatively few researchers have reported on effects of
summer pruning on water core. Myers and Ferree ( 198313)and Marini and
Barden (1982d) found reduced water core levels in summer pruned trees
and hypothesized that increased fruit :leaf ratios may have suppressed
the severity of water core.
Conflicting results of summer pruning on fruit size and soluble solids
may be explained, to a large extent, by the type and severity of pruning,
as well as crop load. Fruit size and soluble solids of apple (Greene and
Lord 1983)and peach (Marini 1985)were reduced by severe heading cuts
near the fruit, but not by thinning cuts a t a greater distance from the
fruit. Adequate leaf area near the fruit appears to be important for fruit
development. ‘Stayman’ apples on June-headed trees with regrowth had
smaller fruit, but greater soluble solids than August-headed trees with
no regrowth (Marini and Barden 1982d). An unpublished preliminary
study with ‘Stayman’trees also indicated competition between fruit and
storage organs for late-season photosynthate. Fruit soluble solids were
greatest on scaffold branches that were girdled near the trunk of control
trees, whereas soluble solids were similar for fruit on girdled and non-
girdled branches of August-headed trees. Therefore, it seems that severe
summer pruning reduces photosynthate translocation to both fruit and
storage organs, since trunk enlargement is also suppressed following
summer heading.
IV. SUMMARY
Summer pruning is generally thought to suppress tree growth by

lowering the photosynthetic capacity of the tree, thereby reducing the
carbohydrate reserves (Ferree 1979; Stembridge 1979),which are required
for spring growth (Priestley 1960). Although carbon partitioning in
mature trees has not been studied following summer pruning, there is
indirect evidence that carbohydrate reserves are reduced by the treat-
ment. l h n k (Lord et al. 1979b; Marini and Barden 1982a; Marini 1985)
and root (‘hylor and Ferree 1981; Rom and Ferree 1985; Marini and
Barden 1982a) growth have been suppressed the year of pruning, possi-
bly due to reduced supplies of photosynthate or growth regulators from
the tree top during the late summer. 14Cstudies have shown that the roots
and trunk are major sinks for photosynthate produced late in the season
(Hansen 1967; Priestley 1962; Quinlan 1969),and late-season root ( Priestley
1964; Heinicke 1936) and trunk (Kozlowski 1962) growth have been
associated with an accumulation of assimilate from the foliage. In addi-
tion to possibly limited carbohydrate reserves in the trunk and roots,
growth the following season might be expected to be suppressed by the
restricted root system following summer pruning ( Kramer and Kozlowski
1979). However, shoot growth the year following summer pruning has
rarely been suppressed when compared to nonpruned or dormant pruned
control trees (Lord et al. 1979b; 'bylor and Ferree 1984; Marini and
Barden 1982s; Marini 1985).
The size of the root system relative to that of the shoot system may
influence shoot growth. A small root system should theoretically sup-
press shoot growth by limiting the supply of water, nutrients, and hor-
mones to the plant top. Maggs (1964)hypothesized that leaves and roots
may normally function below their maximum potential because halving
the leaf surface or root system did not reduce the growth increment by
50%. Kramer and Kozlowski ( 1979)suggested that each plant species has
a characteristic root : shoot ratio. If the ratio is drastically altered, physi-
ological changes occur that lead to compensatory growth and tend to
bring the root :shoot ratio back into its characteristic balance. Data from
summer pruning of container-grown apple trees support this concept.
Dry weight of root systems and root :shoot ratios of summer pruned
trees were less than that of nonpruned trees a t the end of the growing
season. However, shoot growth and root :shoot ratios were similar for all
treatments a t the end of the following season (Bubenheim and Barden
1984; Marini and Barden 1982f).
There may be at least two possible explanations for the lack of shoot
growth control the year following summer pruning. One explanation is
that total carbohydrate reserves may not be adequately reduced by sum-
mer pruning. Rom and Ferree (1985) reported that the percentages of
water-soluble sugars, hydrolyzed starch, and total extracted carbohy-
drate on a dry weight basis from leaves, shoots, and roots of container-
grown peach trees were not significantly reduced by summer pruning.
A second explanation is that only a small quantity of carbohydrate
reserve may be necessary for adequate spring growth. The importance of
reserve carbohydrate for spring growth of mature trees has not been
studied in detail, but there have been several reports for container-grown
trees. Quinlan (1969) detected reserve 14C in all new growth the spring
following late-summer I4C fixation, and Hansen (1971) estimated that
about 50-70% of the carbohydrate requirements of early-spring growth
were supplied from reserves rather than from current photosynthate.
Priestley (1962) reported that only about one-third of the extractable
carbohydrate supply in young apple trees was depleted during spring
growth, and Hansen and Grauslund ( 1973)reported that less than 25% of
the reserve carbohydrate was used in spring growth. Hansen (1967)
reported that 75% of the I4C fixed by young trees in October was lost,
probably to respiration, by spring. About 15% of the 14C originally fixed
was recovered in shoots and leaves the following June. Priestley (1964)
attempted to restrict carbohydrate accumulation in young apple trees by
stem girdling in September. Girdling successfully restricted winter root
growth, but there was no effect of extractable carbohydrate per unit bark
tissue, below the girdle, or on growth the following spring. Priestley

suggested that nongirdled trees may have utilized the surplus reserves
for winter root growth. We found that mature apple trees that were
summer pruned for 3 consecutive years had 12% less total nonstructural
cabohydrates in the bark tissues in November than dormant pruned
trees. However, at bloom time the summer pruned trees had 9%more total
nonstructural carbohydrates than dormant pruned trees (unpublished
data). The excess carbohydrate in control trees may have been used for
winter respiration or root growth.
These studies indicate that relatively low levels of reserve carbohydrate
are adequate for normal spring growth. There may be a critical minimum
level of reserve required for spring growth, above which no suppression of
growth is observable. Summer pruning, as normally performed, may not
have suppressed reserves below that minimum level. There is still a need
to determine the importance of reserve carbohydrate for spring growth
and fruit set. At this time, however, it does not seem feasible to control
shoot growth of fruit trees by altering carbohydrate levels in the tree
during late summer.
One of the problems in pruning studies is selection of appropriate
controls. Controls used in various studies have been unpruned (Ferree
and Stang 1980; Myers and Ferree 1983c; Lord et al. 1979b; Morgan et al.
1984),lightly dormant pruned (Myers and Ferree 1983a,b, 1984; mylor
and Ferree 1984,1986; Greene and Lord 1983),and comparably pruned in
the dormant period (Marini and Barden 1982a,b; Stiles 1980, 1981). If
the objective is to compare dormant and summer pruning, it is very
difficult if not impossible to make valid comparisons of shoot length
unless comparable pruning is done and the only variable is time of
pruning. For example, if shoots are headed to 3 to 4 leaves in June, July, or
August, is it valid to compare these shoots (with or without regrowth) to
an unpruned control or even a lightly pruned dormant control? Such
comparisons may lead one to conclude that summer pruning suppresses
shoot growth. However, unless there is a comparable dormant pruned
control, one can only evaluate the tree’s response to summer pruning. As
noted by Marini and Barden (1982a), at the end of the growing season,
summer pruned trees had shorter shoots and were smaller than the
not-yet-pruned controls. However, after the controls were dormant pruned,
trees were similar in appearance. A definitive study would be one where
the treatments included a nonpruned control, a set of trees pruned a
certain way during the summer, plus another set of trees pruned identi-
cally during the dormant period.
The published research results from summer pruning studies often
have not supported many of the long-held views on the practice. The
major advantage of summer pruning may be that pruning can be spread
out during the year whenever labor is available. Summer pruning may
have a place in training young fruit trees, and may also be used to improve
fruit color for poor coloring cultivars grown in climates where color
development is less than ideal. However, summer pruning must be used
with caution to avoid reduced fruit size and soluble solids, which may
reduce crop value. Thinning cuts, which are made some distance from the
fruit, have little effect on fruit size or soluble solids. One of the major
proposed advantages of summer pruning is to suppress tree vigor. After
reviewing the literature, it is our opinion that summer pruning is no
more effective than dormant pruning for suppressing shoot growth and
limiting tree size.
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ROM, C. R., D. C. FERREE, and G. A. CAHOON. 1984. The influence of three
tree training systems within hedgerows on light distribution, cropping, and efficiency
of ‘Redhaven’and ‘Redskin’peaches. Ohio Agric. R e s . Deu. Ctr. R e s . Circ. 283:49-53.
SAKO, J., and E. LAURINEN. 1982. Effect of summer pruning on the growth
and yield of apple trees. A n n . Agric. Fenniae 21:8-12.
SAUNDERS, W. 1863. “Report of the Commissioner of Agriculture for the Year
1863,” p. 552. USGPO, Washington, D.C.
SCHUMACHER, R., F. FANKHAUSER, and W. STADLER. 1974. The influ-
ence of summer pruning and calcium chloride on physiological disorders in the
cultivar Maigold. Schweir. Z. O b s t . Weinbau 110(24):654-657.( H o r t i c . A b s t r .
45:3725; 1975. )
SEELEY, E. J., W. C. MICKE, and R. KAMMERECK. 1980. ‘Delicious’ apple
fruit size and quality as influenced by radiant flux density in the immediate growing
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STEMBRIDGE, G. E. 1977a. Our experience with high density peach plantings.
Compact Fruit D e e 10:48-52.
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10
Orchard Floor Vegetation Management
E. J. Hogue and G. H . Neilsen
Agriculture Canada, Research Station
Summerland, British Columbia, Canada VOH 1ZO
I. Introduction 377
11. Major Orchard Floor Management Systems 379
111. Effects on the P e e and Crop 382
A . Vigor 382
B . Leaf Nutrient Content 390
C . Fruit Yield 395
D . Nutrient Content a n d Quality of Fruit 398
E . Root Development 402
I v. Effects on the Soil 405
A . Organic Matter 405
B . Nutrients 407
C . pH 410
D . Moisture and Other Soil Physical Properties 411
E . Temperature 415
I. INTRODUCTION
Orchard floor vegetation management before the twentieth century is

not well documented. The wide row spacing and high scaffold branches
found in an English pear orchard planted before 1760 (Janick 1986)were
probably typical of early orchards designed to accommodate grazing. We
suspect that orchard soil management was not a major preoccupation in
Europe until after the transition of fruit growing from small, low-density
orchards to larger, more intensive plantings where higher yields and
better fruit quality became important concerns. In such orchards grazing
under and around trees was no longer practical. In New York, sod
orchards began to be tilled toward the end of the nineteenth century and
cover crops came into use during the 1890s (Slate 1976). One of the
earliest experiments examining the effect of sod in an orchard was
started in 1902 in Massachusetts (Faust 1979). Also in 1902, a Missouri
trial compared straw mulching with “other common methods of [orchard]
soil management” ( Shaw and Southwick 1936). Experimentation with
377
378 E. J. HOGUE AND G. H. NEILSEN
clean cultivation in New York orchards had actually begun 3 years earlier
(Faust 1979).
Prior to World War I, most Europeans orchards were cultivated to
control vegetation (Robinson 1983).However, while clean cultivation was
believed to offer advantages of increased growth, and was widely adopted
in the early days of orcharding, various developments began to reverse
this trend. For example, in Britain the use of grass increased in the 1920s
with the advent of mowers developed to cut grass on airplane landing
strips (Robinson 1983).Also, a comprehensive research program at East
Malling Research Station showed that a frequently mowed grass and
clover sward, in most cases, was a superior form of soil management for
English orchards because it maintained or increased organic matter, and
thus improved soil structure and water penetration (Greenham 1953).
In the fruit-growing area of British Columbia, grass sod was adopted
as a standard practice commencing around 1935 and paralleled the
changeover from furrow to sprinkler irrigation (Fisher 1976). Perceived
advantages included reduced soil erosion, reduced winter injury to trees,
and improved fruit color.
During the 1950s and early 1960s the advent of broad spectrum
herbicides enabled efficient and economical control of vegetation around
the base of trees and permitted a reevaluation of the desirability of this
vegetation. This development coincided with the increased use of dwarfing
rootstocks and closer tree spacing, which made it especially necessary to
examine the influence of orchard floor vegetation management on tree
growth, tree productivity, and fruit quality.
A number of other reviews have focused in whole or in part on aspects
of orchard floor management. The effects on the distribution and effectiveness
of root growth was reviewed by Atkinson (1980),the chemical composi-
tion of the fruit by Perring (1984b),general fruit quality by Proebsting
( 1972), and soil properties by Rogers and Greenham ( 1948), Greenham
(19531, and Haynes (1980/1981). Specific vegetation management sys-
tems including mulching (Jacks et al. 1955)and the effects of herbicides
in orchards (Robinson 1974; Atkinson and Herbert 1979; Stott et al.
1979; Atkinson 1985)have also been reviewed. Only a few reviewers have
attempted to assess the effects of several orchard floor vegetation man-
agement systems on both the tree and soil (Haynes 1980; Atkinson and
White 1981).
This review examines the four major orchard vegetation management
systems: permanent orchard floor vegetation, mulching, mechanical cul-
tivation, and herbicides. We have summarized the effects of each system
on both the soil and the crop: changes in the chemical and physical
properties of the soil, and the effects on tree growth and nutrition and
fruit yield and quality.
10. ORCHARD FLOOR VEGETATION MANAGEMENT 379
11. MAJOR ORCHARD FLOOR MANAGEMENT SYSTEMS
1. Permanent Orchard Floor Vegetation. The use of a permanent

orchard floor vegetation is likely the most common orchard soil manage-
ment system. B s t s of other systems often consider this the standard or
control treatment and it has been referred to as a sod, sward, grass, or
grassed-down treatment, with considerable variation in the species com-
prising the orchard floor plant cover. The growth of this vegetation is
generally controlled by mowing but other control treatments such as the
use of mulch in the tree row or a narrow herbicided strip is frequently
combined with permanent floor vegetation. The recent major advances in
selecting grass species and cultivars most suitable for interplanting in
horticultural crops have been reviewed by Butler (1986) and will not be
discussed here.
2. Mulching. A wide variety of inorganic and organic mulching mate-
rials have been used with tree fruits (Yhble 10.1).Although some mulching
treatments have covered the whole of the orchard floor, most have been
restricted to near the tree. The combination of mulching in the row with
sod between the rows has been referred to as sod mulching. Mulching has
also been combined with other soil management treatments as illustrated
by Robinson and O’Kennedy (1978), who used straw, and Lord et al.
(1968),who used hay and black polyethylene in combination with herbi-
cides. The effects of mulching on soil properties beneath a wide variety of
crops has been reviewed by Jacks et al. (1955).This review is confined to
its use in temperate-zone tree fruit production.
3. Cultivation. Cultivation is the mechanical movement of soil to
improve conditions for crop growth and the term is used interchangeably
with tillage. It was the traditional method of maintaining a loose soil
structure and controlling weed growth in commercial orchards during
much of this century (Woodbury et al. 1917; Bailey 1920). The extent of
cultivation can range from frequent cultivations throughout the year to
various minimum cultivation systems such as with the use of annual
cover crops. In the discussion to follow, an effort has been made to
distinguish between the effects of various cultivation systems. Little
attention has been given to the establishment and the effects of the
various summer and winter cover crops, subjects covered in detail else-
where (Childers 1983).
4. Herbicides. The introduction in the 1950s of chemicals capable, at
low rates, of controlling plant growth provided another means for the
orchardist to control the orchard floor vegetation. In low-density plantings
only the area under the tree drip line is treated. In higher density orchards
Table 10.1. Selected References Indicating Type and Method of Mulching in Experiments in Temperate n e e Fruit Orchards
in Various Geographical Areas
~~~
Mulch Crop Method and rate of application Geographical area Reference
Organic mulches
Straw mulch Apple, peach Not indicated Indiana Baker (1941, 1943)
Sour cherry -0.1 m depth beneath tree to drip line Michigan Kenworthy ( 1954)
Apple - 17.9 MT/ha in 1.2 X 1.2 m square UK White and Holloway ( 1967)
around tree, renewed annually a t rate
of 13.4 MT/ha
0.1-0.2 m depth over orchard floor Germany Weller (1969)
Apple, peach 0.15 m depth, 3 m wide strip in tree row, Australia Baxter (1970)
renewed every second year
Straw mulch plus Apple Barley straw a t 17.5 MT/ha every second Ireland Gormley et al. (1973 a , b ) ,
herbicide year plus 250 kg/ha calcium ammonium Robinson and O’Kennedy (1978)
nitrate in year 1 to avoid nitrogen
deficiency
Cereal straw Peach 0.1 m depth, 26.9 MT/ha eventually to Australia Cockroft (1966),
cover whole orchard floor Cockroft and Wallbrink ( 19661,
Cockroft and Tisdall(l974)
Wheat straw -0.1 m depth (sufficient to prevent grass Ohio Havis and Gourley (1937)
growth) beneath the trees
70-90 k g per tree per year within tree Ohio Wander and Gourley ( 1943)
drip line
Depth sufficient to control weed growth, Connecticut Judkins and Rollins ( 1943),
renewed also to achieve this, under trees Judkins and Wander ( 1945)
180-360 k g per tree in 5 y r period likely New York Boynton et al. (1952),
within drip line of tree Boynton and Anderson ( 1956)
Meadow hay Apple Renewed annually to maintain a 0.15m New Hampshire Latimer and Percival ( 1944)
depth around trees to just beyond the
tree drip line
Alfalfa hay Sour cherry -0.1 m depth, renewed annually beneath Michigan Kenworthy (1954)
tree to drip line
Alfalfa hay Apple 0.3 m per year depth within the tree New Mexico Sullivan and Enzie ( 1972)
drip line
Strawy manure Apple, peach 1.3 m square around trees Indiana Baker (1941,1943)
Sawdust Apple Not indicated Indiana Baker ( 1943)

0.12m depth, reapplied in year 2 Australia Baxter ( 1977)
Renewed annually t o -0.15 m depth New Hampshire Latimer and Percival ( 1944,1947)
around trees to just beyond drip line
Sour cherry -0.1 m depth beneath tree to drip line Michigan Kenworthy ( 1954)
Seaweed Apple Renewed annually to achieve -0.15 m New Hampshire Latimer and Percival(l944, 1947)
depth around trees to just beyond
drip line
Inorganic mulches
Glass wool cinders Apple To cover surface ( ? ) Indiana Baker (1943)
Laminated plastic Apple To cover ground surface between tree rows Israel Moreshet et al. (1975)
vacuum coated during growing season, removed a t
with alloyed harvest
aluminum
Black polyethylene Apple, peach 1.3 m width, centered on tree Australia Baxter (1977)
plastic
Apple 1.1 m width centered on tree Norway Mage ( 1982)
3.0m wide strip centered on the tree row B.C., Canada Neilsen and Hogue ( 1985)
it is more convenient to apply the chemicals in a continuous band in the

tree row. The grassed-alley, herbicide strip system was shown to be a good
orchard floor management option in the 1960s (von Engel 1968)and has
been in common use since. The system of maintaining a completely clean
orchard floor with herbicides (overall herbicide), introduced shortly there-
after (Robinson 1974), has gained some acceptance in Europe, particu-
larly in high-density plantings. In North America, however, the grassed-alley,
herbicide strip remains the most common orchard floor management
system (Crabtree and Westwood 1976). The literature on the herbicides
suitable for use in orchards is extensive. A partial list is presented in
n b l e 10.2 to indicate the evolution in choice of chemicals. This choice
today varies with type of crop species, climate, soil, cultural practices,
availability, and registration requirements of different areas.
111. EFFECTS ON THE TREE AND CROP
A. Vigor
1. Permanent Orchard Vegetation. The extreme debilitating effect of
complete sod on fruit tree vigor was recognized very early in the history of
commercial fruit growing (Bedford and Pickering 1914; Howard 1924)
and these observations gave much of the impetus to the extensive use of
cultivation in orchards in the first half of the twentieth century. Subse-
quent experimental measurements involving variables such as tree girth,
number and size of leaves, annual extension growth, pruning weights,
internode lengths, and root and top dry weight were generally poorer for
fruit trees grown in association with a complete sod cover. Although
many of these observations were made on apple trees, a grass-induced
vigor reduction has been observed for most fruit trees including peach
(Baxter 1970; Lord and Vlach 1973; Welker and Glenn 1985) and pear
(Raese 1977). Newly planted trees are especially sensitive to vigor reduc-
tion in the first years of growth (White and Holloway 1967; Bould et al.
1972; Robinson and O’Kennedy 1978)but mature fruiting trees also show
reduced vigor when grown in association with sod (Rogers et al. 1948;
Greenham 1965; Haynes 1981a; Neilsen et al. 1984).There are studies to
indicate that dwarfing rootstocks of apples such as M.2 (Dancer 1963)
and especially M.26 (Whiteand Holloway 1967)are very sensitive to sod
competition. The reduced tree growth has been attributed to excessive
competition by sod for nitrogen (Bould and Jarrett 1962) and moisture
(Dancer 1963) or both (Bould et al. 1972; Haynes 1981a).
Despite observations of possible vigor reduction related to sod-induced
soil moisture stress, few measurements have been made of leaf water
potential or conductance for fruit trees under permanent orchard vegeta-
Table 10.2. Selected References Demonstrating the Evolution of Herbicide Use in Temperate n e e Fruit Orchards
Rate(s )
Herbicide(s )" ( k g ha ') Crop( s ) Controlh Toxicity' Geographical area Reference
DNBP + fuel oil 3 Apple nursery F N New York Curtis (1952)

Monuron 2.8 Sour cherry E N Wisconsin Gilbert et al. (1959)
CIPC 22 P N
Mon. + CIPC 2.8 + 22 E N
Monuron 5.6 Apple F Wisconsin Holm et al. (1959)
11.2 E N
Diuron 11.2 E N
Dalapon 8.7 F N N.S., Canada Leefe and Longley ( 1960)
Dalapon + 2,4-D 8.7 + 1.1 E N
Amitrole 5.6 F N
Amitrole 11.2 E N
Simazine 18 Peach, sour cherry E S Michigan Larsen and Ries ( 1960)
9 E N
18 Apple, pear E N
4.5 F N
Diuron 5.6 Apple P S Washington Benson and Degman ( 1961)
11.2 G M
22.4 E H
Simazine 5.6 P S
11.2 F M
22.4 G M
continued
Rate(s )
Herbicide(s)" (kg ha ' 1 Crop(s ) Controlh Toxicity' Geographical area Reference
Diuron + amitrole 9.0 + 5.6 Young apple E H Ontario, Canada Saidak and Rutherford (1963)
4.5 + 5.6 G N
Simazine + amitrole 4.5 + 5.6 G N
Simazine + dalapon 4.5 + 11.2 G N
Simazine 2.8 Sour cherry G N Wisconsin Gilbert et al. ( 1964)
5.6 E N
11.2 E S
Diuron 2.8 G N
Isocil 2.2 Apple E S Delaware Fisher (1965)
4.4 E H
Bromacil 1.1 E S
2.2 E H
Amitrole 5.6 P N Oregon Mellenthin et al. (1966)
Simazine 4.5 P N
Amitrole + simazine 5.6 + 4.5 Apple G N
Diuron 4.5 F N
Amitrole + diuron 5.6 + 4.5 E N
Simazine 3.4 Plum P N California Lange et al. ( 1969)
6.8 P- E N
A trazine 4.5 P S
Prometryn 4.5 P- E S
Diuron 6.8 E N
Bromacil 4.5 E N
Isocil 4.5 E N
Terbacil 4.5 E N
Amitrole 4 Apple N Victoria, Australia Baxter and Newman ( 1971 )
Amitrole + simazine 4 + 2.1 N
Paraquat + diquat 1 + 0.6 N
Dichlobenil 4.5 Peach G N Georgia Daniel1 and Hardcastle (1972)
6.7 E N
Simazine 4.5 G N
Nitralin 4.5 G N
Terbacil 2.8 E N
Paraquatd 0.6 E N
Dichlobenil 6.7 Apple F N Quebec, Canada Lareau and Hogue (1976)
9.0 G N
Terbacil 2.2 G N
4.5 G N
Metribuzin 2.2 G N
4.5 E N
Glyphosate 2.2 Peach G-P N Florida Arnold and Aldrich (1979)
Glyph. + napropamide 2.2 + 4.5 Pecan G-F N
Naprop. + terbacil 4.5 + 1.1
Paraquatd 0.6 G-F N
Diuron 4.4 Apple F N Ontario, Canada Heeney et al. ( 1981)
Simazine 4.4 F N
Terbacil 4.4 E N
Dichlobenil 8.8 G N
Oryzalin 9.0 Peach G N West Virginia Welker (1984)
Oryzalin + diuron 4.5 + 2.2 E N
Oryzalin + simazine 4.5 + 2.2 E N
Diuron 2.2 G N
Simazine 2.2 F N
"Herbicide common names according to Hopen and McLane ( 1984); 'Toxicity: N, none; S, slight; M, moderate; H, high.
refer to same for chemical names and authorities. dSeveral applications during season.
'YO control: P, 0-50; F, 50-70; G, 70-90; E, 90-100.
tion. Such techniques have been demonstrated in other circumstances by

Jones and Cumming (1984)and Brun et al. (1985).
The plant interaction known as allelopathy has received increasing
attention as an additional factor affecting crop growth. However, despite
a wealth of published information on this subject (Rice 1979; Putnam and
Duke 1978), there is still little evidence that allelopathy is important in
the field (Newman 1982). While there is some evidence supporting its
occurrence in a variety of orchard floor cover crop or weed species (Skroch
and Shribbs 1986),Shribbs et al. ( 1986)found that the competitiveness of
four ground cover species with ‘Golden Delicious’ apple seedlings in the
greenhouse was related to their biomass production and not to allelopathy.
Furthermore, the results of a field study with young apple trees impli-
cated competition for nitrogen as the major factor of interaction between
the trees and both grass and broadleaf ground covers (Shribbs and
Skroch 1986b).
The amount of nitrogen fertilizer necessary to satisfy grass growth and
positively affect tree vigor has not been definitively established. How-
ever, it is likely to be in excess of 200 kg/ha in most orchards, as a
number of published studies have indicated an inability to offset vigor
reduction in the short term a t annual rates of nitrogen fertilization
ranging from 60-120 kg/ha over 3 years (Dancer 1963), 120-160 kg/ha
over 4 years (Greenham 19651, up to 180 kg/ha under irrigation for 3 years
(Neilsen et al. 1984) and as high as 270 kg/ha for 3 years (Rogers et al.
1948).In a 5-year study under irrigation, Mason ( 1964)found no increased
extension growth for mature trees of five major British Columbia apple
cultivars fertilized at rates of nitrogen ranging from 30 to 140 kg/ha. In
this study, nitrogen was applied in the fall in a narrow band a t the drip
line of the trees. However, ‘Topred Delicious’ trees in sod were similarly
band fertilized in the first 5 years and showed a growth response to
nitrogen fertilization over a range varying from about 2.6 to 10.3 kg/ha x
number of years of apple tree age (Miller 19831. This was attributed to
adequate moisture supplies in the soil although there was no assessment
of nitrogen mineralization in this fine-textured clay.
There have also been studies of methods to avoid the reduction in vigor
associated with sod while maintaining a complete ground cover. Bould
and Jarrett ( 1962)established relatively pure stands of a number of cover
crops and found wild white clover (5!Fifoliumrepens L.) and natural
swards to exhibit less reduction of tree vigor than timothy (Phleum
pratense L. ) and ryegrass (Loliumperenne L. 1, apparently as a result of
less competition for nitrogen. Clover had maintained its dominance after
4 years but its density had decreased to less than half the original values,
especially on plots fertilized a t twice the regular rate (70 kg/ha per year).
Use of maleic hydrazide and 2,4-D to suppress but not eliminate natural
sward growth still resulted in significant vigor reductions for ‘Golden
Delicious’ and ‘Cox’sOrange Pippin’ ( Stinchcombe and Stott 1983).
Cover crop species can also influence the vigor of fruit trees in ways not
related directly to competition for moisture or nutrients. Orchard floor
vegetation, for example, can influence the populations of disease-causing
microorganisms and nematodes (Norris 1986). The potential benefit of
vegetation for suppression of pathogens suggested by some has not been
established unequivocably. On the other hand, available evidence indi-
cates that orchard floor vegetation can increase the difficulty of disease
management by hosting various pathogens (Norris 1986).The extensive
literature on this topic is the subject of other reviews (e.g., Thresh 19811.
Permanent orchard vegetation, with or without “weed” species, also
influences tree vigor through its effect on insect and mite populations.
The interaction has been reported variously to be beneficial, detrimental,
or of no consequence, and a reasonable conclusion is that no general state-
ment can be made on the effect of orchard plant species on pest manage-
ment. Each situation must be judged on its own merit (Norris 1986).
The picture on vertebrate pests, especially voles, emerges much more
clearly with population increases and damage to fruit trees related to the
cover and food sources supplied by orchard floor vegetation (Byers 1984).
2. Mulching. In general, mulching increases the growth and vigor of

fruit trees (Haynes 1980).Improved fruit tree growth has been attributed
to a number of factors associated with the use of mulch. In Australia,
increased tree circumference of peach was associated with the use of a
straw mulch that improved structure, soil moisture, and soil rooting
depth for trees growing in shallow soils overlying a massive clay subsoil
in the Goulburn Vdey (Cockroft 1966;Cockroft and Tisdalll974). Improved
initial growth of ‘Cox’sOrange Pippin’ apples on M.26 rootstock without
irrigation in Southern England was attributed to less moisture stress
under straw mulch relative to complete sod or herbicide strip treatments
(Whiteand Holloway 1967). Of 11treatments, the best first-year growth
of ‘Smoothee Golden Delicious’ on seedling rootstock was obtained with
straw mulch in nonirrigated plots in North Carolina (Shribbs and Skroch
1986a). This study strongly indicates the importance of soil moisture for
tree establishment. By the end of the third year, although trunk diame-
ters were still highest in the mulched plots, the relative growth rate of the
trunk was equaled in plots that were herbicided, cultivated, or covered
with common ragweed (Ambrosia artemisiifolia L.).
Boynton and Anderson (1956)working in New York state concluded
that growth increases of ‘McIntosh’apple trees associated with the use of
hay mulch were likely a consequence of addition of nitrogen from the

mulch material and were similar to growth responses to inorganic nitro-
gen fertilizer in their 5-year factorial experiment. Reduced tree growth
might have been expected in this experiment if a mulching material with
a low C :N ratio without compensating nitrogen fertilizer had been used,
as demonstrated for sawdust mulch by Latimer and Percival ( 19471.
Mulching has a range of other benefits and specialty uses. For exam-
ple, Shaulis (1946) measured increased initial peach tree vigor and sur-
vival in Pennsylvania orchards planted on sloping and eroded land. More
vigorous pear and apple trees and increased soil temperatures under
black plastic have been reported in Norway, where soil temperatures are
more likely to limit tree growth (Mage 1982; Frimanslund 1984). There
are also reports suggesting that mulching young trees reduces the haz-
ards of replanting old orchards (Kenworthy 1954).Although much research
indicates short-term benefit to tree vigor, long-term growth can also be
increased as indicated in a 20-year study by Toenjes ( 1941)in which trees
grew larger with the sod-mulch system compared to cultivation-cover
crop soil management. Sustained tree vigor can, however, present a major
problem in high-density plantings where trees can become too large for
their original planting densities.
3. Cultivation. Cultivation also has generally increased vigor of fruit

trees when compared to permanent orchard floor vegetation (Dancer
1963; Leefe and Longley 1960; Lord and Vlach 1973; Goode and Hyrycz
1976; Haynes 1981a; Mage 1982; Miller 1983; Miller and Glenn 1985).
However, there are also reports of no difference (Neilsen and Hogue 1985)
or of a reduction in growth of cultivated trees compared to trees under a
grassed surface (Rogers et al. 1948; Robinson and O’Kennedy 1978)
perhaps because of the surface root damage caused by tillage. Tillage has
been frequently combined with cover cropping, and Woodbury et al.
(1917),for example, found that apple trees grown under clean cultivation
followed by winter cover cropping made 45% greater average yearly
increase in trunk diameter than trees in grass. Similar benefits of using
the tillage-cover crop system have been reported by others (Judkins and
Riollins 1943;Dancer 1963;Judkins and Wander 1945; Olney and Armstrong
1942). However, long-term benefits to the vigor of trees from cultivation
may not be measurable. Olney and Armstrong (1942), reported that as
peach trees under grass got older and well established, their vigor increased,
especially on sloped land, while under cultivation there was greater soil
erosion and a progressive decline in growth.
Relative to mulch, cultivated apple trees can have less vigor when a
high water table restricts rooting zone depth (Butijn and Schuurman
1957).Similar vigor has been measured when trees were irrigated remov-
ing the potential benefit of the soil moisture-conserving properties of an
inorganic mulch (Neilsen and Hogue 1985). However, Proebsting ( 1958)
found the vigor of irrigated peaches to be less under cultivation than
under an organic mulch.
Pees under cultivation have often been found to have comparable vigor
to those under herbicides (Fisher 1965; Gilbert et al. 1959; Mage 1982)
likely because of the similar effects that these two treatments have on
availability of moisture and nitrogen. For newly planted peach, in addi-
tion to a comparable growth response, these treatments provided
significantly increased freezing resistance of bark and wood tissue of
dormant scions (Marriage and Quamme 1980).
4. Herbicides. The elimination of competing species with herbicides

has been shown to increase growth, usually measured as a trunk diameter
increase, particularly of young fruit trees (Gilbert et al. 1959: Larsen and
Ries 1960; Leefe and Longley 1960; Gilbert et al. 1964; White and
Holloway 1967; Lange et ai. 1969; Baxter and Newman 1971; Lord and
Vlach 1973; Stott 1976; Arnold and Aldrich 1980; Atkinson and Lipecki
1980; Atkinson and White 1981; Welker and Glenn 1985).
The annual increase in tree size generally reflects the degree of weed
control (Lange et al. 1969; Skroch et al. 1971; Welker 1984). However,
Mellenthin et al. (1966) found that tree growth could be related to the
degree of weed control maintained in one location but not in another where
the primary competition, Agropyron repens L., was less prevalent.
Welker and Glenn (1985) found increased growth of newly planted
peach trees when the vegetation-free area around the trees increased from
0.36 to 13.0 m2. This supports English results with apple, where overall
herbicides provided better growth and yields than the grassed-alley,
herbicide strip system (Atkinson and Lipecki 1980). However, the com-
parison of growth of trees under herbicide treatments has generally been
made with a nonweeded or grassed control. When herbicide treatments
were compared with hand-weeding, Gilbert et al. (1959) found similar
growth of sour cherry in a 4-year trial. However, Ries et al. (1963) for
peach and Raese et al. (1974)for pear obtained more growth on simazine-
and amitrole-treated trees than on hand-weeded or plastic-mulched trees;
Daniel1 and Hardcastle (1972) obtained more growth from peach with
chemical weed control than from cultivating; and with ‘Golden
Delicious’/M .26 apples growing in Ireland, Robinson and O’Kennedy
( 1978)obtained more vigor in overall herbicide plots than with any of five
other soil management treatments.
Where fruit tree growth was not related to degree of weed control,
interacting factors such as additional soil moisture provided by mulching

(Lord et al. 1968),irrigation (Crabtree and Westwood 1976),or herbicide
toxicity (Skroch 1970; Lord and Greene 1975)were found to affect growth.
Although herbicides have become a valuable tool in removing compet-
ing vegetation in the orchard, these chemicals can easily reduce tree vigor
because of toxicity to the crop. Considerable testing has been carried out
in the field (e.g. Holm et al. 1959; Benson and Degman 1961; Saidak and
Rutherford 1963)and in the greenhouse (Luckwill and Casely 1966; Lange
and Crane 1967) to measure better the tolerance of fruit crops to herbi-
cides, as well as the interactive influence of other factors such as root-
stocks, scion cultivars, soil types, and organic matter (Lange et al. 1969).
Although simazine is probably the most widely used residual chemical
for preemergence vegetation control in fruit crops (Atkinson 1985), as
little as 0.5 ppm has been shown to be toxic to some fruit seedlings in
sand culture (Lange and Crane 1967) and 2.2 kg/ha applied to the soil
surface in container experiments (Skroch 1970). Sharma et al. (1977)
found that leached soil-appliedsimazine reduced photosynthesis of ‘Golden
Delicious’/M.7 by an average of 22% during the 50-day period of their
greenhouse study but caused no measurable growth inhibition. However,
such a significant effect on photosynthesis must result in sooner or later
to a reduction of tree vigor and productivity, the onset of which can be
gradual and difficult to detect. Hence, there is concern about the leaching
of residual herbicides, particularly photosynthetic inhibitors on coarse
soils, or with repeated use of these chemicals leading to an accumulation
of residues in the surface soils where competition-free tree roots will grow.
Glyphosate, a very effective and widely used chemical for postemergence
control of orchard vegetation, has been shown to be harmful even a t very
low rates to fruit trees when applied to foliage or immature bark (Atkinson
1985). Prunus species trees are especially sensitive.
Thus, even with emphasis on precise and careful application and under
the best application conditions it has been difficult to avoid some leach-
ing and drift of herbicides and the resulting effects on growth, vigor, and
even survival of young fruit trees.
B. Leaf Nutrient Content

1. Permanent Orchard Vegetation. The reduction in vigor of a range
of tree fruits commonly associated with establishment of a complete sod
cover in orchards has long (Bedford and Pickering 1919; Howard 1924)
and frequently been associated with decreases in leaf nitrogen concentra-
tion (e.g., Greenham 1965; Bould et al. 1972; Stott 1976). Reduced leaf
nitrogen concentration was also acknowledged by Rogers et al. (1948),
who advocated maintenance of orchard soil fertility through use of a

frequently mown sod. However, experiments in which leaf nitrogen con-
centration of fruit trees has not been decreased by a frequently mowed
sod cover have been reported for mature trees with a long history of
adequate nitrogen and hence apparently adequate nitrogen reserves (White
and Greenham 1967). It is generally recognized, however, that leaf nitro-
gen concentration of trees growing in sod is more easily affected by
suppression of ground cover than by nitrogen fertilization (Mason 1969a;
Greenham 1976; Neilsen et al. 1984). Although most of the reductions
reported are for mid to late growing season leaf nitrogen concentrations,
Bould and Jarrett (1962) found that sod cover also reduces nitrogen
supply to trees early in the growing season (green-cluster)as indicated by
lower total nitrogen concentration in xylem sap.
Conversely, increased leaf phosphorus concentration has been reported
for apples trees with sod cover (Bould and Jarrett 1962; Greenham 1965;
Haynes 1981a; Neilsen and Hogue 1985; Shribbs and Skroch 1986b).The
higher phosphorus concentration of apple leaves may reflect a restricted
growth of the trees due to low nitrogen levels (Bould and Jarrett 1962).
However, others have not always found a reciprocal relationship between
nitrogen and phosphorus. Bould et al. (1954) found elevated leaf phos-
phorus content when nitrogen levels were adequate. Ferauge ( 1969)found
soil phosphorus to be mobilized by the cover crop, especially if it was deep
rooted. Recently, Atkinson and White ( 1980)suggested that mycorrhizal
infection, which can enhance phosphorus uptake, is transferable from
grass to tree roots. Atkinson ( 1986)has also suggested that the degree of
mycorrhizal infection and root growth in the early spring may be impor-
tant factors affecting phosphorus supply to apples.
Sod can also increase leaf potassium concentration (e.g., Bould and
Jarrett 1962; Greenham 1965; Stott 1976; Haynes 1981a; Neilsen and
Hogue 1985; Shribbs and Skroch 1986b) and reduce leaf scorch symp-
toms typical of potassium deficiency if leaf concentrations are not already
a t adequate or luxury levels (Wallace 1928; Rogers et al. 1948). Results
for calcium and magnesium are more variable, with a few reports of
increased leaf calcium and magnesium (Greenham 1965; Neilsen et al.
1984)or of leaf calcium (Haynes 1981a)resulting from orchard sod cover.
However, decreased leaf magnesium has also been associated with a
sod-induced increase in leaf potassium (Neilsen and Hogue 1985) in a
study where major differences were measured in the relative ability of a
sod to recycle magnesium and potassium. The effect of sod cover on leaf
micronutrient status has rarely been reported despite an early observa-
tion that sod cover reduced iron deficiency symptoms of apple trees in
England (Wallace 1929).
2. Mulching. Boynton and Anderson ( 1956)reported that decomposa-

ble mulch with a favorable C :N ratio was likely to increase leaf nitrogen
concentration of apples since available soil nitrogen supply was increased.
Also, most mulches suppress the competition for nitrogen by under-tree
vegetation. This is most clearly indicated by field experiments with black
plastic mulch that increased leaf nitrogen concentration relative to apples
grown under sod (Mage 1982; Neilsen and Hogue 1985).
Fewer reports have concerned the effects of mulching on leaf phos-
phorus, although Wander and Gourley ( 1943) reported increased levels
in apple trees mulched with wheat straw and receiving no other fertil-
izers. Lower leaf phosphorus concentrations (with elevated leaf nitrogen)
have, however, been reported for black plastic mulching ( Neilsen and
Hogue 1985),apparently as a consequence of year-long removal of under-
tree vegetation.
Increased leaf potassium concentration is one of the earliest (Baker
1941) and most frequently recognized consequences of mulching with
organic materials, although often the increases represent luxury uptake
(Wander and Gourley 1943; White and Holloway 1967; Baxter 1970).
Improved potassium nutrition has been attributed to the addition of
significant amounts of potassium with the mulch as well as to improved
potassium uptake following improved root growth or improved soil mois-
ture conditions (Baker 1943). The latter two factors were apparently not
significant under irrigation in British Columbia, where leaf potassium
decreased under black plastic mulch in a similar manner to that observed
under other ground cover suppression treatments (Neilsen and Hogue
1985). The increase in leaf potassium has frequently had an adverse effect
on leaf calcium and magnesium concentration (Wander and Gourley
1943; Baxter 1970).This is consistent with observations that magnesium
deficiency in apple was more common under straw mulch than grassed
plots (Butijn and Schuurman 1957).
There has been little assessment of the effects of organic or inor-
ganic mulches on micronutrient nutrition, although Wander and Gourley
( 1943) observed increased leaf boron levels of apples grown with a wheat
straw mulch.
3. Cultivation. Howard (1924)found that cultivation, compared to a
grassed soil surface, greatly improved leaf size and color of a wide variety
of fruit crops, a response he attributed to increased availability of soil
nitrogen. Greenham (1965)found that in cultivated plots sufficient nitro-
gen mineralized from the soil to maintain apple trees for 32 years in a
satisfactory condition without added nitrogen as indicated by leaf color
and nitrogen concentration; and Mason ( 1969a)showed that leaf nitrogen
concentrations of 30-year-old‘McIntosh’trees increased with the amount
of tillage. Hill (1962)found that peach trees under sod required twice as
much nitrogen (45.4 g per year of tree age) to provide leaf nitrogen levels
comparable to trees under cultivation. Thus, cultivation is generally
reported to increase leaf nitrogen in the short term when compared to
grass (Ljones 1958; Dancer 1963; Bould et al. 1972; Haynes 1981a;
Miller and Glenn 1985; Neilsen and Hogue 1985). However, Bould et al.
( 1972) found that although grass clearly depressed leaf nitrogen in the
initial years, cultivation did not consistently increase leaf nitrogen in
later years.
In contrast, leaf phosphorus and potassium have been found to be lower
in trees under cultivation than in trees under grass (Bould et al. 1972;
Neilsen and Hogue 1985; Shribbs and Skroch 1986b) or under mulch
(Baker 1941; Wander and Gourley 1943). Greenham (1965) found that
tillage decreased leaf phosphorus compared to permanent vegetation only
when trees received no added nitrogen, while leaf calcium was decreased
under tillage with or without added nitrogen. Wander and Gourley (1943)
found that tillage increased leaf calcium compared to mulching.
4. Herbicides. Proebsting ( 1958)found that leaf nitrogen concentra-
tion of established peaches in herbicided plots varied considerably in a
3-year comparison of six soil management treatments. In the first year,
leaf nitrogen was second highest of all treatments (2.60%)while in the
third year, leaf nitrogen was the lowest (2.41%).However, a t no time was
leaf nitrogen altered sufficiently to affect yield or growth adversely. Miller
and Glenn ( 1985)showed that leaf nitrogen of four apple cultivars receiv-
ing five rates of nitrogen was higher in trees in herbicided plots compared
to grassed plots only a t the lowest, or half the recommended rate of
nitrogen application. Welker and Glenn (1985) found leaf nitrogen of
young peach trees to be positively correlated with the size of the area
maintained free of vegetation with herbicides. Neilsen et al. ( 1984)obtained
increased leaf nitrogen from established ‘Golden Delicious’ apple with
both early and year-long weed control, and Johnson ( 1980)increased leaf
nitrogen from established ‘Cox Orange Pippin’/M.2 by imposing herbi-
cide strips of either 0.9 or 1.8 m on previously grassed rows. Atkinson and
White ( 1980),however, found that nitrogen was not consistently higher in
leaves of young or mature apple trees in herbicided plots compared to
trees under grass. Over a 7-year period, differences were small in leaf
nitrogen concentration of ‘Cox’sOrange Pippin’/MM. 106for two extreme
treatments, including 125 kg/ha of nitrogen on overall herbicided plots
and grassed with no added nitrogen. Notwithstanding results such as
these, it appears that in general the herbicide effect on leaf nitrogen
increase is similar to that of other treatments such as tillage and mulching
(Proebsting 1958; Lord and Vlach 1973; Miller and Glenn 1985; Neilsen
and Hogue 1985),which involve removal of competition. Variation of this

effect may relate to variation in soil nitrogen mineralization potential and
initial nitrogen status of the trees.
When Ries et al. (1963) demonstrated that simazine increased leaf
nitrogen of nonbearing apple and peach and leaf nitrogen of bearing apple
trees, it appeared that some herbicides might have a direct effect on fruit
tree nutrition. However, in subsequent trials, repeated applications of
simazine had no effect on leaf nitrogen concentration of ‘Jerseyland’
peach (Lord et al. 1967). Lord et al. ( 1968),also looking for a simazine-
induced effect on nutrient composition of young apple trees, measured no
differences in leaf nutrient levels directly attributable to simazine. Raese
et al. (1974), however, reported that amitrole plus simazine herbicides
resulted in higher nitrogen in ‘d’Anjou’pear leaves. In other trials, Raese
( 1977) suggested that these herbicides could partially substitute for
nitrogen fertilizers on young ‘d’Anjou’ pears. However, Heeney et al.
(1981)also found no effect of repeated herbicide use, including simazine,
on leaf nitrogen concentration of apples. Thus, it appears that, as suggested
for other crops (Tbeedy and Ries 1967),simazine may promote increased
leaf nitrogen only under certain growing conditions.
Atkinson and White ( 1980)found, in a young apple orchard in the first
bearing year, that leaf phosphorus was lower in overall herbicide than in
grass plots not under irrigation. Miller and Glenn (1985)also found leaf
phosphorus to be significantly lower in trees in herbicide plots compared
to grass sod, although peach leaf phosphorus concentrations were not
influenced by the size of the herbicided area around the tree up to 13 m2
(Welker and Glenn 1985). Neilsen and Hogue ( 1985) showed that apple
leaf phosphorus concentration of trees in plots kept clean until mid
summer and left to weed over for the rest of the year was similar to that of
trees in grass, whereas trees in plots kept clean year round were lower in
phosphorus than those in grass. The decrease in leaf phosphorus concen-
trations attributed to removal of vegetation with herbicides was usually
insufficient to cause growth effects.
Vegetation control with herbicides generally caused a reduction of fruit
tree leaf potassium concentration (Atkinson and White 1980; Miller and
Glenn 1985; Neilsen and Hogue 1985),which was occasionally accompa-
nied by an increase in leaf magnesium (Neilsen and Hogue 1985; Miller
and Glenn 1985). However, Lord and Vlach (1973)found that herbicides
did not affect concentrations of peach leaf potassium and magnesium
compared to grass. Similarly, herbicides have seldom influenced levels of
leaf calcium when compared to permanent orchard vegetation (Lord and
Vlach 1973; Heeney et al. 1981; Miller and Glenn 1985). However, Raese
(1980)found that a single application per year of simazine a t 3.6 kg/ha,
which provided poor weed control, increased leaf calcium in ‘Delicious’
apple over trees in grass, and good weed control with paraquat increased
calcium over both treatments. However, the combination of the two
chemicals, providing excellent weed control, failed to increase leaf cal-
cium compared to the weedy check.
C. Fruit Yield
1. Permanent Orchard Vegetation. Many studies have reported signif-
icant yield reductions of apple, pear, and peach under sod management,
which accompanied reduced vigor and leaf nitrogen concentration ( Saidak
and Rutherford 1963;Cockcroft 1966; Baxter 1970; Skroch 1970; Gormley
et al. 1973a; Lord and Vlach 1973; Raese 1977; Stott et al. 1979). Both
number and size of fruit have been affected in the short term, with
long-term yield reduced because of smaller trees. Peach seem especially
sensitive to yield reduction as a consequence of excessive competition by
sod for available nitrogen (Judkins and Wander 1945; Proebsting 1958).
Dwarfing apple rootstocks (e.g., M.9 and M.26) also seem sensitive to
yield reduction (Robinson and O’Kennedy 1978; Neilsen and Hogue
1985).The competitive effects of a sod cover appear early as evidenced by
the failure of young apple trees planted in sod, without nitrogen fertiliza-
tion, to produce an adequate number of flower buds (Latimer and Percival
1947).Even with nitrogen fertilization, bloom development may be poorer
for trees competing with sod (Neilsen and Hogue 1985) with the result
that yield efficiencies (expressed as kilograms harvested fruit per square
centimeter trunk area) are reduced (Miller 1983).As indicated previously
in the discussion about tree vigor, the amount of nitrogen fertilizer
required to overcome yield reduction is not precisely known. Bollard
(1957) found 0.91 kg of ( NH4)zS04per ‘Jonathan’ apple tree per year
insufficient to maintain yield of grassed down trees at the level of trees
remaining under clean cultivation without extra nitrogen. However, in
some studies, slight yield reductions of mature fruiting trees shortly
after grassing down have been more than compensated for by improve-
ments in fruit quality (Rogers et al. 1948).
2. Mulching. In some comparative studies, maximum fruit yields
have been achieved under mulch. For example, Baxter (1970) working
with shallow soils in Australia, measured significantly greater blossom
density per unit of tree circumference, double the yield, and larger fruit
size in the first two fruiting years for mulched, compared to clean culti-
vated peach and apple trees. Under the adverse soil physical conditions of
the Goulburn Valley in Australia, peach trees subjected to the ‘ktura
system of soil management, which includes straw mulching, yielded
three times those of commercial nonmulched trees on similar soil types
(Tisdallet al. 1984).Under similar adverse soil conditions in the Netherlands
(heavy soil with poor drainage), Butijn and Schuurman (1957) found
apple yield and net income to rank first for the straw mulch plots, and in a
long-term trial with ‘d’Anjou’pear in a clay adobe soil, Westwood et al.
(1964)obtained a better yield with a heavy hay or straw mulch than with
tillage. These studies show that straw mulching can be especially impor-
tant in terms of maximizing yield where soil physical conditions are poor.
Boynton and Anderson (1956) found that total apple yield increased
with the amount of hay mulch used in a study where the effects of
mulching were similar to and additive to the effects of nitrogen fertiliza-
tion. Thus, suppression of the competitive influence of orchard ground
cover and direct additions of nutrients are also contributing factors to the
yield increases under mulch. Proebsting (1958)found that highest subse-
quent 5-year fruit yields were measured for mulched peach trees where
tree vigor had been depressed by 3 previous years under orchard grass
sod. Not surprisingly, then, some experiments show that fruit yield
under straw mulching is not significantly greater than herbicide-treated
trees (Baxter 1977) and trees under black plastic mulching do not yield
significantly more than trees with other ground cover suppression treat-
ments (Neilsen and Hogue 1985). Usually, as demonstrated for black
plastic mulches, fruit yields of mulched trees greatly exceed those of
trees grown with full ground cover (Mage 1982). Often increased yields
are preceded by significantly increased flowering ( Frimanslund 19841.
Robinson and O’Kennedy( 1978)identified an example of decreased apple
yield under straw mulch in Ireland and suggested that this was a conse-
quence of reduced temperature and aeration of the soil.
3. Cultivation. Cultivation usually increases early fruit yields corn-
pared to permanent orchard vegetation but not to herbicided plots (Cockroft
1966; Daniel1 and Hardcastle 1972; Lord and Vlach 1973; Crabtree and
Westwood 1976; Mage 1982; Miller 1983; Neilsen and Hogue 1985). How-
ever, reduced yields of cultivated relative to herbicide-treated orchards
have occasionally been reported (Gormley et al. 1982; Misra et al. 1984;
Miller and Glenn 1985), perhaps because of the surface root pruning
effect of tillage (Coker 1959). Palmer and van Haarlem (1944) reported
that clean cultivation, compared to grass, increased yields of apple in the
early years of a 15-year trial in Ontario but that total yields over that
period were similar. Butijn and Schuurman (1957)compared clean culti-
vation, straw mulching, green manuring, and grass sod and found that
apple yielded best under clean cultivation and poorest under grass during
the first 4 years in production. However, over the 17 years of the Netherlands
trial, differences between treatments diminished due to the relative increase
in yield of trees under grass and the decline in yield of trees in clean
cultivated soil. Likewise, Rogers and Raptopoulous ( 1945),using mature
trees of four apple cultivars on M.9, were unable to show differences in
yield between continuously cultivated trees and those under five different
cover crops over a 4-year period. Palmer et al. (1941),comparing several
treatments in a long-term peach trial, obtained the best yields frcjm
tillage plus manure (9000 kg/ha yearly) and the lowest yield from tillage
plus mineral fertilizer (275 kg/ha of 4-8-12 fertilizer yearly), the differ-
ence being attributed to the amount of nitrogen added.
The differences in yield of trees under tillage and those under grass
appears to be due to more than the availability of soil nitrogen. Even
when high rates of nitrogen fertilizer were added to trees under grass,
their yield was not equivalent to that of trees under tillage (Bollard 1957;
Hill 1962; Bould et al. 1972). Proebsting (1958)studied the effect of soil
management systems and added nitrogen, under irrigation, i.e., where
soil moisture was presumably eliminated as a factor. He found that tillage
followed by winter cover crops produced better yielding trees than those
under permanent grass or legume sods. This result was attributed to
increased nitrogen mineralization.
Cultivation can promote late summer growth of fruit trees because of
the generally improved soil nitrogen and moisture conditions. Thus, it is
suspected of reducing cold acclimation of trees with presumed greater
susceptibility to winter injury (Westwood and Bjornstad 1981).
4. Herbicides. Increased fruit tree yield is generally related to the
increased tree size whether tree growth is related to vegetation removal by
herbicides or by some other method (Webster and Brown 1980).Chemical
weed control on 4-year-oldnonbearing pear resulted in markedly increased
vigor, larger trunk diameter, and greater subsequent yield of larger fruit
than trees where weed control had been delayed until trees were 6 years
old (Raese et al. 1974). In a 9-year study, Stinchcombe and Stott (1983)
found that overall herbicide promoted increased tree yield, especially in
the early years, a result consistent with a number of studies (White and
Holloway 1967; Robinson 1974; Stott 1976).However, in later years yield
differences tend to be less pronounced.
Robinson and O’Kennedy ( 1978)compared six methods of soil manage-
ment in a ‘Golden Delicious’/M.26 planting between 1965 and 1976 and
showed that overall herbicides gave higher yields than any other treat-
ment. Reviewing the yield advantages of the overall herbicide system of
apple orchard management, Robinson ( 1983) reported a 21% increase in
yield of ‘Cox’s Orange Pippin’ between 1968 and 1981; Stott (1976)
reported a 41%increase in yield; and Atkinson and White ( 1981) reported
reduced returns due to competition from grasses and annual weeds
within the herbicide strip of 25 and 3470,respectively, in 1975 and 45 and
55%, respectively, in 1976.
Proebsting (1958)found that soil management did not affect yield of
mature peach trees in the first three years of his study but in the fourth
year herbicide increased yields over grass or alfalfa sod, mulch, or cultiva-
tion followed by winter cover crops of rye or vetch. In the fifth year, trees
in herbicided plots outyielded sod.
Similar yields were measured for ‘Bisbee Delicious’ apple trees kept
completely weed free with herbicides year round or only until midsum-
mer, cultivated, or mulched with black plastic (Neilsen and Hogue 1985).
These yields exceeded those under sod. Miller ( 1983)found that nitrogen
source or rate did not affect yield of ‘Topred Delicious’ apple, although
soil management affected yield in one of the two years, both herbicides
and cultivation being higher than sod.
Mage (1982)found that herbicides and cultivation gave similar yields
of apple, both better than permanent grass sod but both inferior to
plastic mulch. Likewise, Misra et al. (1984) obtained greater yield in
15-year-old‘Fkd Delicious’ under hay mulch, or by growing beans around
the trees than when the plots were kept clean with paraquat. Conversely,
the yield of peach trees in the first 6 years receiving two applications of
paraquat or one application of paraquat plus simazine was comparable to
that obtained under cultivation or hay mulch, and all were better than
under mowed grass (Lord and Vlach 1973). Over 11 yielding years, four
cultivars of apples on M.4 yielded less under herbicidal treatment with-
out tillage than under tillage alone (Weller 1969). Lord et al. ( 1967) did
not obtain increased yield with their herbicide treatment in peach com-
pared to trees in grass sod because of relatively poor weed control.
Thus, yields of fruit trees were always higher in plots where vegetation
had been controlled with herbicides over those either under a grass sod or
weedy, whether they were under the grassed-alley, herbicide strip, or the
overall clean with herbicides system. However, yields in herbicided plots
appear to be generally the same or slightly higher, and only occasionally
lower than yields under tillage or mulching.
D. Nutrient Content and Quality of Fruit

1. Permanent Orchard Vegetation. Decreased fruit nitrogen and in-
creased fruit potassium and phosphorus concentrations, paralleling the
changes in leaf nutrient composition, have been reported for apples from
trees under sod in the United Kingdom (Wilkinson 1957; Perring 1975;
Stott et al. 1979). Increased fruit phosphorus concentration seems espe-
cially pronounced for trees under sod orchard cover when fertilized with
phosphorus ( Perring 1975), when nitrogen fertilizer rates are low ( Perring
1984b), or when the sod cover involved clover (Atkinson and Herbert
1979). This result is considered important since low fruit phosphorus
concentrations have been associated with increased incidence of low-
temperature breakdown (Johnson et al. 1983). In contrast, fruit flesh
calcium concentration, so important to apple fruit quality ( Perring 1968),
has either been unaffected by orchard vegetation treatments ( Wilkinson

1957)or affected more by fruit size than by any treatment involving sod
orchard floor cover (Perring and Jackson 1975). Understandably then,
elevated flesh calcium concentrations have been reported for apples from
trees in sod where apple size has been reduced (Stott et al. 1979; Neilsen
et al. 1984). However, since calcium concentration of apples can be
effectively increased by calcium sprays, achieving elevated fruit calcium
levels through fruit size (and yield) reduction is clearly unnecessary
(Stott et al. 1979). Little systematic research has been reported on other
fruit nutrients although greater fruit sodium and sulfur concentrations
have been measured when the orchard floor is sod (Wilkinson 1957).
Perring and Holland ( 1985)also demonstrated that yearly fluctuations in
the concentration of many fruit minerals are often larger than differences
due to experimental treatments, with the consequence that any effect of
soil management must be assessed over a number of years.
Decreased fruit nitrogen levels often improve fruit quality. Thus, some
reported consequences of the permanent orchard vegetation manage-
ment system include yellower ‘Golden Delicious’ (Gormley et al. 1973a,
1982; Stott et al. 1979; Neilsen et al. 1984),yellow ‘d’Anjou’pears (Raese
1977)’and improved red fruit coloration for apple cultivars such as ‘Cox’s
Orange Pippin’ (Greenham 1965;Stott et al. 1979)and ‘WorcesterPearmain’
(Rogers et al. 1948). Also, earlier maturity but low yields of highly
colored ‘Elberta’ peach were associated with grass sod (Proebsting 1958).
Other fruit quality improvements reported for apple include reduced fruit
drop (Rogers et al. 1948), relative freedom from bitter pit and storage
disorders (Stott et al. 1979)’firmer fruit a t harvest (Neilsen et al. 1984),
and for pear, higher soluble solids and reduced storage scald and rot
(Raese 1977). One must suspect, however, that many of these differences
could have been achieved by reduced nitrogen fertilizer rates. Further-
more, the improvement in fruit quality often did not compensate for
reduced yields. This suggests further caution in interpreting the effects of
permanent orchard vegetation on fruit quality as does the result of
Johnson et al. (1983)that there were marked seasonal differences in the
effect of overall clover sods on red coloration and storage quality of fruit.
2. Mulching. Little information is available concerning the effects of

mulching on fruit mineral content, in part because much mulching research
was completed prior to recent interest in fruit mineral content. In some
early work, Wander and Gourley (1943) reported reduced calcium, ele-
vated potassium, phosphorus, and magnesium, and unaffected boron
levels in fruit flesh and cores from mulched trees. When organic mulches
act as a source of readily available nitrogen, development of red fruit color
can be reduced as reported for apples grown with hay mulch (Boynton
and Anderson 1956; Latimer and Percival 1944, 1947). With regard to
effects not clearly related to mineral nutrition, straw mulch reduced fruit
quality of ‘Golden Delicious’ apple in Ireland as indicated by reduced
soluble solids, acidity, and color, and this effect was attributed to reduced
soil temperature rather than increased nitrogen (Gormley et al. 1973a).In
Ontario, use of hay mulching as a means of conserving soil moisture was
unsuccessful in prevention of cracking of ‘Italian’prune (Cline and Tehrani
1973).However in a 1-yearstudy in Israel, Moreshet et al. ( 1975)reported
increased percentage of color, size, and sugar content of ‘Orleans’apples
harvested from the bottom third of treated trees when reflectant alumi-
num mulch was used between the rows. The use of reflectant mulch has
since become a common cultural practice in Japan to obtain better fruit
coloration of some apple cultivars.
3. Cultivation. Cultivation generally increases fruit nitrogen concen-

trations compared to grass (Hulme 1956; Perring 1975)whereas phospho-
rus and potassium levels are generally lower (Wander and Gourley 1943;
Wilkinson 1957; Perring 1975). Furthermore, fruit with greater nitrogen
availability should have larger fruit and lower fruit calcium. However,
Perring (1975)found that cultivation had no effect on calcium levels, nor
was there the usual inverse relationship between fruit calcium and size.
He did, however, find that cultivation reduced fruit magnesium levels.
Gormley et al. (1973a)found that ‘Golden Delicious’ fruit quality was
generally better in fruit from cultivated plots compared to fruit from
noncultivated (herbicide)plots. With ‘Cox’sOrange Pippin’, however, the
differences were very small (Gormley et al. 1973b)and the slightly higher
soluble solids in ‘Jonathan’attributable to the cultivation treatment was
not detected by a taste panel. In a follow-up study with the same
plantings, Gormley et al. (1982)concluded that soil management contin-
ued to influence fruit quality after 15 years, but to a lesser extent than in
the early years of the trial. In most seasons in these Irish studies there
was a low negative correlation between yield and soluble solids. Thus,
tillage provided better fruit quality than the higher yielding overall
herbicide treatment.
With 30-year-old ‘McIntosh’ apple, Mason ( 1969a) found that fruit
quality was largely unaffected by cultivation. Cultivation affected only
firmness, which decreased as tillage increased. Tillage had no effect on
fruit grade. With established ‘Delicious’trees, cultivation had no effect on
quality except to cause green fruit flesh (Mason 196913). These effects
reported by Mason appear to be related to increased nitrogen availability
to the fruit.
4. Herbicides. Johnson (1980)imposed herbicide strips of 0.9 and 1.8

m centered on the tree rows in a previously grassed orchard of 15-year-old
‘Cox’s Orange Pippin’/M.2 and obtained significantly higher nitrogen

concentrations compared to fruit from the grassed area. The increased
availability of nitrogen reduced fruit color in all 5 years of the trial.
However, Johnson et al. ( 1983)found that fruit nitrogen concentration of
‘Cox’s Orange Pippin’ over a 6-year period was generally unaffected by
overall herbicide treatment compared to the herbicide strip or white
clover sod management systems when similar rates of nitrogen fertilizer
(75 kg/ha) were applied. Perring ( 1984c) found that nitrogen concentra-
tions in apples from trees in a herbicide strip differed little from fruit
grown under permanent orchard vegetation after the first year, when fruit
nitrogen concentrations were influenced by the nitrogen mineralized from
a recently killed sod. Overall herbicide treatment resulted in significantly
increased fruit nitrogen only in the last 2 years of an 8-year trial. He
suggested that a t this site, soil nitrogen availability had not declined
sufficiently to result in large effects on fruit nitrogen concentration.
Low fruit phosphorus appears to increase the risk of low-temperature
breakdown in England (Johnson et al. 1983)and a reduction in apple fruit
phosphorus concentration is also generally associated with increased use
of herbicides in orchards. For example, Johnson ( 1980)and Johnson and
Johnson (1983)reported lower fruit phosphorus concentration in three of
five years for herbicide strip treatments compared to grass. Although the
fruit phosphorus concentrations were not regarded as critical for storage
quality of these apples, the results indicated an important trend. Also,
Johnson et al. (1983) reported consistently lower fruit phosphorus con-
centration, 1974-1980, in overall herbicide treatments relative to herbi-
cide strip management and Perring (1984a) confirmed a trend toward
higher fruit phosphorus concentration as the width of the herbicide strip
decreased. However, a residual (2-year)increase in fruit phosphorus from
killed grass occurred when converting from herbicide strip to overall
herbicide management. Furthermore, fruit species may differ in this
response with Perring ( 1984b) reporting that in a 5-year investigation,
overall herbicide use decreased phosphorus concentration of ‘Conference’
but not of the interplanted ‘Cornice’pear. In a single year of this study,
phosphorus concentrations of fruit of ‘Merton Glory’ cherry and ‘Victoria’
plum were unaffected by the overall herbicide to herbicide strip conver-
sion. Little attention has been directed tQ fruit phosphorus concentration
outside England.
Use of herbicides does not appear to lead to major changes in fruit
calcium, magnesium, or potassium concentration. Johnson et al. ( 1983)
indicated that over 6 years potassium, calcium, and magnesium concen-
trations in apple fruit were not generally affected by herbicide strips or
overall herbicide soil management systems compared to a clover sod. In a
3-year study with ‘Golden Delicious’ apple, Neilsen et al. ( 1984)compared
permanent vegetation with year-round vegetation removal with herbi-

cides and found decreased fruit potassium concentration in one year and
decreased fruit calcium concentration in two years. Furthermore, the
decreased fruit calcium concentration was associated with increased
average fruit size.
Perring ( 1984c)measured reduced fruit calcium concentrations for the
overall herbicide treatment in the 1year out of 8 when average fruit size
was increased. In the same trial, potassium and magnesium concentra-
tions were not affected by the herbicide treatment, and the normal strong
positive correlation between potassium and magnesium was apparent in
alternate years only. Perring ( 1984b) found that manganese was highest
in 1 year under overall herbicides, intermediate in herbicide strip, and
lowest under permanent orchard vegetation. The direct influence of man-
ganese on fruit was, however, unknown.
Gormley et al. (1973a)found that ‘Golden Delicious’apple from grassed
plots displayed better eating quality than fruit grown in soil maintained
free of vegetation with herbicides. This was attributed to a higher per-
centage of soluble solids resulting from lower yield and a higher leaf :fruit
ratio under grass. Fruit quality differences between the vegetation man-
agement treatments were less marked with ‘Cox’s Orange Pippin’ or
‘Jonathan’ (Gormley et al. 197313).In their follow-up study, Gormley et al.
(1982)found no differences of fruit firmness of ‘Golden Delicious’but in
two years when overall herbicides gave the highest yields, the fruit had
lower soluble solids. In ‘Cox’s Orange Pippin’ there was a negative corre-
lation between yield and soluble solids. They concluded that although the
overall herbicide method is an efficient system of soil management resulting
in increased yields, fruit quality may be reduced in some seasons. Johnson
et al. (1983) compared overall herbicide to herbicide strip and overall
clover for 7 years in an experiment involving annual nitrogen, phospho-
rus, potassium, and magnesium fertilizer applications. Soil management
did not consistently affect size, red color, or susceptibility to bitter pit or
senescent breakdown of ‘Cox’s Orange Pippin’ apple. It was concluded
from this study, which had large fertilizer inputs, that soil management
treatments would not likely be advocated nor condemned on the basis of
fruit mineral composition or storage behavior.
E. Root Development
1. Permanent Orchard Vegetation. Early observations by Howard
(1924)in India indicated that a grass cover restricts the total amount of
root development of apple, forcing tree fruit roots downwards and reduc-
ing markedly the number of active rootlets during the monsoon season.
More recent research has tended to confirm these early observations. Gras
and Tkocm6 (1977)measured lowest root density in the surface 25 cm of

grassed treatments in a long-term soil management trial, and Atkinson
et al. (1978) found 75% of the roots of mature ‘Cox’s Orange Pippin’
apple in the surface soil of the herbicide strip while most of the roots
under grass were below 10 cm. As a consequence of this root distribution
pattern, uptake of 15Nwas always much less from the grassed alleys.
Atkinson (1983) has also observed that sod affected root structure.
llees under sod exhibited more short (lateral) than long (extension)
roots. Also, a higher percentage of the roots beneath an irrigated sod were
infected with vesicular arbuscular mycorrhizae, which can improve phos-
phorus uptake. A more branching rooting habit was also noted by Coker
(1959) on ‘Cox’s Orange Pippin’/M.9 apple growing under permanent
grass relative to trees grown under cultivation. However, the depth to
which roots of fruit trees penetrate is also influenced by other factors,
including subsoil structure and consistency. Thus, in an experiment
where the subsoil was clay, Cockcroft and Wallbrink (1966)found many
pear and peach roots near the surface (8 cm) beneath a white clover sod
compared to a cultivated treatment. However, root density near the soil
surface was even higher with the straw mulch and bare surface (herbi-
cide) treatments.
2. Mulching. Improved shallow and lateral root development has been
a recognized consequence of straw mulching, especially where mulches
act to conserve moisture (Yocum 1937). Early studies revealed more roots
closer to the surface and even extending into the mulch itself (Baker 1943)
and more recent studies involving root counts have indicated greater root
density, especially in the surface 7.5 cm beneath the mulch relative to
cultivated treatments (Cockcroft and Wallbrink 1966) and even in the
surface 30 cm relative to full ground cover or cultivated soil treatments
(Whiteand Holloway 1967).As indicated by Greenham ( 1953),increased
root growth beneath organic mulches is attributed to improved soil
moisture and moderated soil temperatures, which encourage increased
root growth in a medium generally rich in nutrients. Although much less
information is available, black plastic mulching would also appear to
affect fruit tree root growth. Observations of grubbed trees from black
plastic treatments indicate a tendency to multibranching and many small
roots but they demonstrate no clear difference in root vigor (Mage 1982).
Clearly, mulching can significantly increase the effective soil volume of
shallow soils.
3. Cultivation. Root growth can also be altered by mechanical culti-
vation. In tree excavation studies carried out in England, Coker ( 1959)
showed that 16-year-old‘Cox’s Orange Pippin’/M.9 apple growing under
cultivation for 9 years had very few roots near the surface as a result of
repeated pruning by tillage implements. The equivalent trees under grass

had 25-5390 of their fibrous roots in the surface 15 cm. Thus, cultivation
of trees established under grass (or mulch, see above) could result in a
severe check to tree growth as a result of root loss. Cockcroft and
Wallbrink (1966), comparing root distribution of peach and pear under
tillage and nontillage management systems, found that the roots of fruit
trees grew into the surface 8 cm of soil of all but the tillage treatment.
Butijn and Schuurman (1957) in a long-term trial with several apple
cultivars, mostly on M.2 and M.4, found that trees under grass had the
poorest root development but those under tillage were only slightly
better, both having a considerably lower number of roots than trees under
straw mulch or green manure treatments. Mitchell and Black (19681,
however, comparing tillage to grass found no significant treatment effect
in total weight of peach roots. Goode and Hyrycz (1976) grew M.2
rootstocks for 2 years under grass and cultivation with two levels of
nitrogen. After tree excavation they measured fibrous (<1mm) and coarse
(>1mm) roots and found no soil management effect a t the low nitrogen
level. However, at the higher nitrogen level, cultivation reduced root
growth. Furthermore, it appeared that soil applied urea had a significant
depressing effect on the production of fibrous roots.
4. Herbicides. When under-tree vegetation is controlled through the

use of herbicides, tree roots can grow to the surface without competition.
For example, Gras and Tkocmd (1977) reported that the highest root
density was in the surface 5 cm of a soil treated with herbicides for 7
years; similar results have been obtained by other workers (e.g. Atkinson
1977).Atkinson and White ( 1980)observed that 5-year-oldtrees of ‘Cox’s
Orange Pippin’/M.26 had the highest root weight under overall herbicide,
the lowest under permanent orchard vegetation, while herbicide strip
trees were intermediate. This effect was also observed in a trial where the
effect of three management systems on apple root length and weight were
compared (Haynes 1980/81). The differences in tree root distribution
were large after only 3 years, with the herbicide treatment most effectively
encouraging root growth close to the surface.
The most common orchard management system in the world is currently
the grassed alley and herbicided tree row. In this system it is generally
assumed that young trees with the bulk of their roots close to the trunk
have similar root distribution whether the alley is grassed or not. As the
trees age, use of the grassed alley by the roots could be expected.
However, even after 12 years, the uptake of l5No3by 12-year-oldapple
trees from under the grassed alley was small compared with that from the
herbicide strip in the tree row (Atkinson et al. 1980). Even with large
trees, uptake of l5No3from the herbicide strip was much higher than
from the grassed alley (Atkinson and White 1980), indicating that while
trees age and increase in size the bulk of their roots remain in the
vegetation-free zone.
There have been few critical studies of the direct effect of herbicides on
the morphology of roots of trees growing under normal orchard condi-
tions when the effect of weed competition or cultivation damage have
been excluded (Atkinson 1980).
IV.EFFECTS ON THE SOIL
A. Organic Matter
1. Permanent Orchard Vegetation. Increased soil organic matter con-
tent in orchards appears to be a natural consequence of sod cover (Greenham
1953).Although the magnitude of the organic carbon increase varies with
soil type (Simons 1958), the largest increase occurs in the surface soil,
especially the top 5 cm, where almost a 2% increase in organic carbon
relative to mechanical cultivation has been reported after only 3 years
(Haynes and Goh 1980~). In one study, failure to measure a significant
increase in soil organic matter under sod was attributed to the rapid
decomposition of orchard grass residues (Rogers et al. 1948). Decaying
cover crop roots, of course, can increase soil organic matter only to their
respective rooting depth.
The benefits of increased soil organic matter content have been ascribed
to improved physical and chemical properties, including improvements in
nutrient exchange capacity, water absorption, soil structure, and mainte-
nance of a healthy soil flora and fauna (Greenham 1953).
2. Mulching. Addition of organic mulching materials to orchard soil
surfaces also conserves and can even increase the organic matter content
of orchard soils (Havis and Gourley 1937; Greenham 1953; Delver 1980),
especially immediately beneath the mulch (Wander and Gourley 1943).
Haynes (1980) attributed the improved retention of organic matter to
lack of cultivation, lower soil temperatures, and fewer wetting and drying
cycles a s well as to the addition of organic matter. The nature of the
organic mulch can also influence the potential increase in organic matter.
Latimer and Percival (1947) showed that after 4 years the decomposed
organic layer beneath hay mulch exceeded that beneath seaweed mulch
and was much greater than beneath sawdust, which had barely decomposed.
However, exceptions to the general pattern of organic matter increase
do occur as noted by Goode and White (1958a), who found that the
organic content of the soil was barely maintained a t its organic level
despite the addition of large amounts of organic material to the soil
surface. This was attributed to compaction and poor aeration of some of

the mulched materials. Although there have been no studies of the effects
of inorganic mulches of soil organic matter levels, a decline in content
would be anticipated.
3. Cultivation. A decline in soil organic matter content in cultivated
orchard soils was recognized long ago (Woodbury et al. 1917) and has
frequently been documented (Havis and Gourley 1937; Wander and
Gourley 1943; Goode and White 1958a; Haynes and Goh 1 9 8 0 ~ )This .
decline can be rapid. Goode and White (1958a) working in England
reported organic matter content a full percentage point lower than the
grass sward control after only 5 years of light cultivation. The decline in
organic matter content has been particularly pronounced for surface
layers of the soil. Haynes (1980) reported that compared to grassed or
zero-tilled herbicide-treated soils, cultivated soils have lower organic
matter in the surface layers but can have a higher content in the subsoil.
This relationship depended upon the depth of cultivation and soil mix-
ing. Rogers and Raptopoulus (1945, 1946) were able to increase soil
organic matter content of an English orchard, relative to year-round
cultivation, by turning in a range of spring- and autumn-sown cover
crops. Making a different comparison, however, Havis and Gourley ( 1937)
found lower organic matter content associated with cultivated orchard
soils relative to sod or mulch even though a system of incorporating
annual cover crops was used in the cultivated orchards.
Although a decline in soil organic matter content has so frequently
been reported for cultivated orchard soils, there is little direct evidence
relating the decline in soil organic matter content to tree growth prob-
lems. In fact, Greenham ( 1955)indicated that for a fine sandy loam soil a t
East Malling no harmful decline in organic matter had resulted from 20
years of cultivation. In this study, leaf fall and root dieback were consid-
ered significant sources of orchard soil organic matter.
4. Herbicides. Atkinson et al. (1980)measured lower organic matter
in the tree row where herbicides had been used for 6 years than in the
grassed alley of a semimature orchard. After killing the grass in the alley
with a herbicide, the organic matter remained constant for 3 years. In his
review, Haynes (1980) indicates that the use of herbicides in no-till
management may preserve temporarily the original distribution of organic
matter in the soil profile, as opposed to tillage, but after 3-5 years the
organic matter concentration is usually lower than under grass sod. It is
generally higher than in soil under continuous tillage, probably because
the natural compaction of the surface provides conditions less favorable
for mineralization. However, soil organic matter can be expected to decrease
with continous herbicide use due to less plant material being returned to
the soil.
B. Nutrients
1. Permanent Orchard Vegetation. Haynes and Goh (1980a) esti-
mated the nitrogen return to the orchard floor via sod clippings to be
approximately 400-600 kg/ha per year. As a consequence, total soil
nitrogen under sod was significantly higher than under overall herbicide
and clean cultivation. However, despite the likelihood of increased organi-
cally bound nutrients such as nitrogen beneath sod, the readily available
N03-N averaged only 1-2 ppm and NH4-N was near zero during the
growing season in a long-term NPK fertilizer trial in England (White and
Greenham 1967). These values were low relative to similarly fertilized,
cultivated plots. Thus, although a sod cover supplies high amounts of
potentially available nitrogen, it also uses large amounts of the available
nitrogen as it is mineralized.
Extractable soil phosphorus does not appear to be increased by sod
relative to cultivation (Greenham 1965)or mulching (Latimer and Percival
1947)although Deist et al. ( 1973)found that phosphorus levels deeper in
the soil were increased by cover crops. Potassium returns are high from
sod clippings with annual estimates over a 3-year period ranging from 321
or 608 kg/ha for short- and long-grass treatments in New Zealand (Haynes
and Goh 1980a) and 217 kg/ha in irrigated British Columbia orchards
(Neilsen and Hogue 1985). Over the 6 years of these two studies, calcium
and magnesium returns to the orchard soil averaged 22 and 7%, respec-
tively, of potassium returns, an indication of the relative potential for
accumulation of these three nutrients beneath sod. The few comparative
measures of exchangeable soil bases made under sod in orchards suggest
significantly greater exchangeable calcium and magnesium but not potas-
sium, compared to orchards whether herbicided or tilled. This result is
attributed to less leaching of calcium and magnesium from sod treat-
ments (Haynes and Goh 1980b; Neilsen and Stevenson 1983).
2. Mulching. The potential to add significant amounts of nitrogen by
using organic mulches is well known (Harley et al. 1951). Weeks et al.
(1950) reported increased total soil nitrogen in the surface 15 cm in an
apple orchard measured 10 years after completion of 20 years of mulching
with low-grade hay. Thus, such fertility changes may have long-term
significance to orchard nutrition (Greenham 1953). In the short term,
however, the availability of N03-N beneath mulches will depend upon the
C :N ratio of the mulching material. Haynes ( 1980)suggested that mulches
having C : N ratios less than 30 :1will release nitrogen to the soil, while a t
ratios greater than 30: 1 available nitrogen in the soil will decrease.
Kenworthy ( 1954), working with cherries, found that sawdust mulches
with low C :N ratios required 5-6 years of decomposition before nitrogen
release exceeded that immobilized by the soil flora.
Available soil phosphorus has generally increased in orchard soils upon
additions of decomposable mulches (Goode and White 1958a; ntkey and

Schoff 1963), especially seaweed (Latimer and Percival 1947). After 12
years of mowed-grassmulching on top of a herbicide strip, water-extractable
phosphorus averaged 19 mg/liter beneath the mulch in comparison to 3
mg/liter beneath sod between the tree rows (Delver 1980).
Increased soil potassium availability was one of the early reported
consequences of mulching with hay and straw (Wander and Gourley
1938; Latimer and Percival 1947; Boynton et al. 1952) and this finding
has more recently been reconfirmed (Delver 1980; Shribbs and Skroch
1986b). In part, this reflects the large amounts of potassium added by
many mulching materials, although there are also indications of improved
potassium availability, possibly because of reduced potassium fixation
near the soil surface. For example, Wander and Gourley ( 1945)attributed
the rapid ( 4 months) penetration of fertilizer nitrogen to a 20 cm depth to
its application with a straw or hay mulch.
Soil calcium and magnesium concentrations have been reported to
increase ( n t r k and Partridge 19471, decrease (formagnesium) (Goodeand
White 1958a),or be unaffected (Stephenson and Schuster 1945; Boynton
and Anderson 1956)beneath organic mulching materials. It is likely, as
suggested by ntrk and Partridge ( 1947), that the original nutrient con-
tents of the soil and the mulching material are important considerations
in interpreting such apparently contradictory results. With the exception
of an increase in hot-water-soluble boron reported beneath hay mulch by
Boynton and Anderson (1956), there has been little assessment of the
effects of organic mulches upon the availability of other nutrients.
Also, there has only been limited assessment of the fate of soil nutri-
ents under inert mulches, although such studies might provide informa-
tion on leaching of soil nutrients or the mineralization potential of
residual soil organic matter.
3. Cultivation. Cultivation often increases the availability of soil

N03-N, especially relative to full ground cover (Woodbury et al. 1917;
Judkins and Rollins 1943; White and Greenham 1967; Haynes 1980).
Furthermore, Dickson ( 1937) found reduced soil N03-N concentration
under minimum cultivation (undertaken until mid-May ) compared to
normal cultivation until August. This indicates that the extent and
timing of weed or cover crop regrowth can influence seasonal N03-N
trends under cultivation. Seasonal variation in N03-N leaching or deni-
trification are also important. Miller and Glenn (1985)found that late in
the dormant season (March, West Virginia) no N03-N accumulated in the
surface 30 cm of the soil in either cultivated or sod plots, provided
near-normal (for tree growth) nitrogen fertilization rates were used. Although
accelerated mineralization of organic matter and greater N03-N availabil-
ity are generally agreed consequences of cultivation (Haynes 1980),total

soil nitrogen content tends to decline under cultivation (Haynesand Goh
1980~).
The effects of cultivation on other soil nutrient levels are unclear. For
example, soil phosphorus values have been reported to decrease (Goode
and White 1958a), to increase (Schuricht et al. 1983), or to be not
significantly different (Deist et al. 1973) under cultivation relative to
permanent orchard vegetation. Similar contradictory findings could be
reported for exchangeable calcium, magnesium, and potassium, suggesting
that cultivation is unlikely to be a major factor affecting the soil concen-
tration of these nutrients.
4. Herbicides. Haynes and Goh ( 1 9 8 0 ~found) that during summer

and autumn, N03-N in the surface layer of herbicided soil was notably
higher than in grassed plots. They also showed that N03-N concentra-
tions in the topsoil fluctuated seasonally in both herbicided and grassed
soil with peak concentrations during midsummer of 21 pg/cm3 in the
former vs. 10pg/cm3in the latter. Atkinson et al. (1980)found soil N03-N
concentrations in overall herbicided plots to be higher, especially in the
alleys, than in grassed plots a t both 0-15 and 15-30 cm depths. However,
there was no indication when the sampling was carried out.
Haynes and Goh ( 1 9 8 0 ~determined
) that extractable and total phos-
phorus, and to a lesser extent exchangeable phosphorus, accumulated a t
the surface of untilled herbicided plots, probably from the applied fertiliz-
ers. Atkinson and White ( 1980)measured consistently greater phospho-
rus concentrations in the soil of herbicided strips than under the grassed
alley and, in a more detailed examination (1976) of the distribution of
phosphorus, they found that phosphorus concentration was much higher
in the row than in the alley of an orchard in the grassed-alley, herbicide
strip management system for 9 years. In plots where the above system
was practiced for 6 years, and then converted to either overall herbicide or
overall grass, the distribution of phosphorus remained very similar after
3 years.
Potassium accumulated in the surface soil of herbicided plots when
compared to cultivated plots (Haynes and Goh 198Oc) and the higher
potassium levels appeared to be due to more restricted downward move-
ment of applied potassium. Other studies have shown that potassium
levels do not vary from the herbicided row to the grassed alley as do
phosphorus levels (Atkinson and White 1976).
Compared to grassed plots, surface soil (0-5 cm depth) in herbicided
plots was much lower in exchangeable calcium and magnesium (Haynes
and Goh 1980~).This is undoubtedly related to the cycling of these
cations by orchard floor vegetation since it is known that vegetation
reduces leaching of these cations from the surface soil. In this same trial,
the surface 5 cm of the herbicided plots maintained a higher level of
exchangeable calcium than the same layer of cultivated soil, a result of
the higher organic matter of the herbicided soil. However, Haynes and
Goh (1980a)in another trial found similar exchangeable calcium concen-
trations in herbicided and cultivated soil, both treatments having lower
calcium than grassed soil. More recently, Miller and Glenn (1985)found
that over 4 years, soil calcium levels were not affected in the 0-30 cm
profile of the herbicide strip. This may be related to depth of sampling,
since Haynes and Goh (1980a) found no significant differences in base
saturation of soil in different treatments below the 15 cm soil depth.
C. pH
1. Permanent Orchard Vegetation. Most comparison studies have
indicated higher soil pH under grass than when orchard floor vegetation
is removed (Atkinson and Herbert 1979; Haynes and Goh 1980b; Neilsen
and Stevenson 1983). However, it is unlikely that the existence of sod can
prevent the inevitable pH decline associated with the continued use of
acidifying nitrogen fertilizers in orchards. This is evidenced by a 14-year
study a t Efford Horticulture Station in Hampshire, England, by Goode
and Higgs (1977)in which soil pH declined from 6.3 to 4.4 for grass plots
fertilized with (NH4)2S04but to 4.1 in the surface 15 cm of similarly
fertilized herbicide-treated plots.
2. Mulching. Although Haynes ( 1981b)suggested that in-row mulching
with grass clippings as practiced in Western Europe and the United
States may result in higher soil pH due to transfer of calcium and
magnesium into this area of the orchard, there is little consistent research
data to support this suggestion. At best, small relative increases in soil
pH (Kenworthy 1954)and exchange capacity and base saturation (Reuther
1941)have been reported after hay and straw mulching. More frequently,
mulches have not affected soil pH (e.g., lbkey and Schoff 1963).
3. Cultivation. Soil pH can decline in cultivated soils relative to
orchards with permanent under-tree vegetation (Schuricht et al. 1983).
This has been attributed to increased leaching of calcium and magnesium
from the surface layers (Haynesand Goh 1980b).Frequently, however, the
pH decline has been similar in magnitude to that observed from other
ground cover suppression treatments (Deist et al. 1973). In some stud-
ies (Wander and Gourley 1943; Goode and White 1958a),no measurable
pH decline occurred in cultivated soils. Incorporating lime before plant-
ing is an effective way to correct acidity problems that have developed
in orchards.
4. Herbicides. Long-term herbicide use in orchards has often been

associated with a decrease in soil pH. Robinson (1974) reported that 5
years of simazine application resulted in a soil pH of 4.4 relative to a
cultivated soil pH of 6.6 in the same planting. Other workers have found
smaller reductions in soil pH attributable to herbicide treatments over
similar periods of time (Atkinson and White 1980; Haynes and Goh
1980b; Miller and Glenn 1985).Acidity increased throughout the 0-30 cm
soil profile of a closely spaced planting after 5 years of overall herbicide
(Atkinson et al. 1980).pH was 6.2 a t the lowest apple tree density of 2.4 m
tree spacing but 7.1 a t a tree spacing of 0.3 m, indicating the possible
influence of root density on cation leaching. Lipecki et al. (1985) in a
detailed look at soil pH a t three different depths across the interrow of
orchards under three different managements found the lowest pH along
the rows and the highest in the tracks of the orchard machinery in all
management systems. The overall herbicided soil had lower pH at all
three depths than the soil under completely grassed or grassed-alley,
herbicide strip management. The pH pattern was similar a t all depths
but differences decreased with soil depth. Thus, although there is appre-
ciable variation in the rate of pH change in different locations, undoubt-
edly attributable to variation in soil physical properties, fertilizer
applications, and moisture regime, there is a consistent reduction of soil
pH with the use of herbicides in orchards.
D. Moisture and Other Soil Physical Properties

1. Permanent Orchard Vegetation. Few detailed measurements have
been made of soil moisture conditions under sod in orchards. From such
measurements Rogers et al. ( 1948)concluded that soil under permanent
grass “began losing water 15-30 days earlier in the spring than did most
cultivated soil.” The implications of this observation for early-season tree
growth and nutrient uptake have not been pursued. An actively growing
sod also requires a proportion of the water throughout the growing
season, and in nonirrigated areas this competition for water can become
critical (Tbkey and Schoff 1963). The extent of soil moisture depletion
varies with the vigor, rooting depth, and frequency of mowing of the
orchard sod. Shallow-rooted grasses such as Kentucky bluegrass (Poa
pratensis L.) or annual bluegrass (Poa annua L.) deplete less moisture
from the whole orchard soil profile than deep-rooted sods comprised of
Ladino clover (nifolium repens L.) or S.23 perennial ryegrass (Lolium
perenne L.) (Toenjes et al. 1956; Goode 1956). Toenjes et al. (1956)
suggested mowing about mid-June to conserve moisture under grass
sods early in the summer, while repeated mowings of new growth were
suggested to reduce water use by sod during the rest of the summer. More
41 2 E. J. HOGUE AND G. H. NEILSEN
recent measurements have confirmed the existence of high water deficits

in the soil around fruit trees and that sod competition modified the
pattern of water depletion with more water being used from deeper in the
soil profile (Atkinson and White 1976).
On the other hand, sod improves other soil physical properties as
reflected by decreased soil bulk density and increased soil porosity (At-
kinson and Herbert 1979). Decreased bulk density means an increase in
total pore space available for root growth and an increase in water-holding
capacity. Such changes have been associated with increased earthworm
populations (Haynes 1980/81) and with lower soil penetrometer values
under grass (Atkinson et al. 1980) except where a dense stiff root mat
exists (Rogers et al. 1948).Such soil properties are likely to reduce surface
moisture runoff and erosion and, in fact, the soil and moisture conserva-
tion benefits of sod in orchards have long been recognized in North
American extension bulletins concerned with soil conservation (e.g.,
Bregger and Brown 1945; Anthony et al. 1948).
2. Mulching. Conservation of soil moisture has long been considered

one of the most significant advantages of mulching fruit trees. For
example, Baxter ( 1970), using sporadic shallow gravimetric sampling
and tensiometers, compared a straw mulch system with a weed-free
herbicide treatment and found greater available water under the mulch.
Dancer (1964)used gypsum blocks a t shallow soil depths and found the
highest percentage moisture under a strawy manure mulch relative to a
cultivated bare surface especially during an intense May-June drying
period. In small plots without fruit trees, Goode and White ( 1958a)found
that, in contrast to sward and clean cultivation treatments, soil beneath
wheat straw mulch remained close to field capacity throughout four
summer seasons. The general improvement in soil moisture status is
likely a consequence of both improved infiltration capacity and reduced
evaporation (Greenham 19531. However, countering these benefits is the
possibility that reflected heat from mulches could increase transpiration
of the mulched trees (Waggoner et al. 1960).
Increased infiltration might increase leaching ( n r k and Partidge 1947)
but reduction in surface runoff and soil erosion in mulched orchards
(Haynes 1980) suggests a net soil and moisture conservation benefit.
Very likely, the type of mulch will influence the degree of soil moisture
conservation. For example, absorbent mulches such as peat (Greenham
1953) or sawdust ( n r k and Partridge 1947) have been found to reduce
rainfall penetration, especially from light rains, while seaweed mulch
(Latimer and Percival 1947) has dried and shrunk after application,
thereby losing some of its moisture conservation characteristics. Although
few measurements have been made concerning soil moisture beneath
10. ORCHARD FLOOR VEGETATION MANAGEMENT 41 3
inert mulches, the significant reduction in evaporation that is likely to

occur suggests that soil moisture content can be higher beneath black
plastic mulches (Mage 1982).
The moisture-conserving properties of mulches may sometimes be
undesirable. For example, Black ( 1963)measured a potentially undesir-
able increase in percentage moisture and degree of saturated pore space
beneath a straw mulch on a silty clay soil susceptible to waterlogging
in winter.
Improvements in soil physical properties such as porosity and develop-
ment and stability of a desirable soil structure have frequently been
reported for orchard soils mulched with organic materials (Haynes 1980),
especially when compared to cultivation (Goode and White 1958b). The
improvements in soil structure have been attributed to a regular supply of
organic matter, especially in fine-textured soils (Delver 1980),to protec-
tion of the soil surface from raindrop impact (Greenham 1953), and to
retardation of soil surface slaking and sealing (Haynes 1980). In Aus-
tralia, regular additions of a straw mulch have been used as an integral
part of the n t u r a soil management system to maintain a stable, open soil
structure with rapid water infiltration, good aeration, and a high root
concentration in soils with physical properties that normally limit crop
growth (Tisdall et al. 1984).
Although mulches generally improve soil physical properties, Black
(1963)reported a higher mortality of young peach trees growing under
a mulch of wheat straw, apparently as a consequence of the development
of an impermeable layer 15 cm beneath the mulch. There are few data
concerning the potential for soil structural deterioration over time be-
neath inorganic mulches where organic materials would not be added at the
soil surface.
3. Cultivation. Fkmoval of weed and cover crop competition by culti-

vation increases soil moisture content relative to permanent orchard
vegetation (Goode and White 1958a; Dancer 1964). However, moisture
content of cultivated soils may not differ significantly from mulched soils
(Goode and White 1958a) and may be less than herbicide-treated soils
(Robinson 1974).Also, the moisture content of cultivated soils may show
considerable seasonal variation depending upon the time of cultivation
and cover crop or weed regrowth. For example, Black and Mitchell ( 1970)
measured higher water loss throughout a 67.5 cm orchard soil profile
under trashy cultivation compared to clean cultivation or herbicide appli-
cation. This was attributed to an invasion of summer weeds after a single
spring plowing and discing. Palmer and van Haarlem (1944)also report
that frequent and deep cultivation can lead to drier soils due to evapora-
tion from newly exposed soils.
Compared to other soil management systems, cultivation frequently

results in a deterioration of other soil physical properties (Butijn and
Schuurman 1957; Cockcroft and Tisdall 1974). Increased bulk density
has been reported relative to sod (Robinson 1974) and straw mulch
(Reuther 1941).Furthermore, decreased stability of soil aggregates (Goode
and White 1958a) and decreased macroporosity and oxygen diffusion
rate in fine-textured silt soils are reported consequences of soil cultivation.
Other dangers of completely clean cultivation included a rapid deteriora-
tion of soil structure and the possible development of a hardpan layer
restricting tree root growth immediately below the zone of tillage (Greenham
1953). Haynes (1980) also distinguishes between short- and long-term
consequences of soil cultivation with immediate consequences of cultiva-
tion often a decrease in soil bulk density and porosity, whereas over the
long term, soil aggregation deteriorates resulting in increased suscepti-
bility of soils to crusting and impeded drainage. However, the conse-
quences of physical deterioration of orchard soils on fruit trees are less
clear. In one study, Deist et al. (1973) found a significant compaction of
soils in a clean cultivated orchard but concluded that the values obtained
were still well above critical values and that regular tillage of such a
gravelly soil was unlikely to seriously compact the soil.
Decreases in soil infiltration capacity (Havis and Gourley 1937) and
saturated hydraulic conductivity of cultivated orchard soils ( Haynes
1980/1981) can parallel the decline in quality of soil physical properties
and result in increased surface runoff. The resultant increase in erosion
from cultivated plots has been associated with a decline in vigor of peach
(Olney and Armstrong 1942)and was one of the main reasons for recom-
mendations to abandon clean cultivation for peach in Ontario (Palmer
and van Haarlem 1944). Several soil- and water-conserving systems of
cultivation have been developed for orchards, and these include contour
and strip cultivation (Breggerand Brown 1945),trashy cultivation (Shaulis
1946),and annual cover crops in combination with minimum early spring
cultivation (Anthony et al. 1948; Greenham 1953). Although annual
cover crops can be effective in reducing autumn and winter soil and water
loss, Li et al. (1942)in a 15-year study found that cultivation in fall and
spring still resulted in decreased soil aggregate stability, hydraulic con-
ductivity and infiltration, and increased compaction, runoff, and erosion
relative to orchards managed with permanent vegetation cover.
4. Herbicides. Other than general statements about herbicide-treated

orchard soils having a lower soil moisture deficit than grassed soils
(Haynes 1980),very little has been written about the effect of herbicides
compared to other management systems.
Atkinson and Herbert ( 1979)reported bulk density of herbicide-treated
soil to be about 8% higher than grassed or cultivated soils. Long-term

herbicide use also affected pore size distribution but the proportion of
large pores (>15 pm) was not necessarily reduced. Across a range of soil
types and tree fruit crops, Atkinson and White (1980) found that soil
bulk densities were approximately 9% higher under herbicide than under
grass or cultivation. A reduction in the total pore volume could occur by
reducing pores within a limited range of pore sizes or across the entire
range, depending on the soil (Atkinson and White 1980).
E. Temperature
1. Permanent Orchard Vegetation. Although soil temperature varia-
tion a t 10 and 25 cm depth under sod has generally been intermediate
(several degrees Celsius cooler in the summer and warmer in the winter)
relative to year-round cultivation or straw mulch (Weller 1969) there are
few data relating soil temperature to fruit tree performance.
Soil temperature may influence fruit trees indirectly by its relationship
to air temperature. Although summer air temperatures in orchards with
sod have been observed to be cooler than in orchards with tilled, bare soil,
differences have been recorded mainly in spring during radiation frosts.
Such frosts occur on clear nights when unchecked soil radiation cools the
air nearest the soil below freezing. Since orchards with permanent vegeta-
tion have cooler spring soil temperatures than tilled, herbicided, or
plastic-mulched soil (Neilsen et al. 1986), the amount of heat to be
radiated is less. In addition, grass or other vegetation has a larger surface
area for radiation to the air above and loses heat rapidly (Haynes 1980).
Skroch and Shribbs (1986) cite instances when temperatures in the
orchard during radiation cooling have been 0.5°-1.00C, even 2OC lower in
grassed than in tilled orchards. Hamer ( 1975)reported that the recorded
minimum temperature during a radiation night was 3.3OC cooler over a
heavily grassed soil than over a bare soil and recommended that grass in
orchards be cut short during blossom time. Krezdorn and Martsolf ( 1984)
indicated that maintenance of weed-free, moist, compact soil furnished
the best reservoir of heat for cool nights and reduced frost damage
to orchards.
2. Mulching. Organic mulches generally insulate the orchard soil
and as a consequence lessen orchard soil temperature variability, reduc-
ing daily and annual temperature extremes (Greenham 1953). Thus,
mean soil temperatures beneath mulch in summer are frequently lower
(Gormley et al. 1973a) especially on days with high incident solar radia-
tion (Dancer 1963). Mean monthly temperatures a t 10 cm depth below a
10-20 cm thick straw cover have frequently been lo-2OC less than those
beneath bare soil in the summer months while during the winter similarly
measured temperatures could be l 0 C higher under mulch relative to bare

soil (Weller 1969).
Although early research by 'hkey and Schoff ( 1963)indicated summer
soil temperatures were reduced under all mulches, including nondecom-
posable ones, more recent measurements in Norway by Mage ( 1982)have
indicated that mean daily summer season soil temperatures were about
3OC higher under black plastic mulching, while Neilsen et al. (1986)in
British Columbia measured a 33.890 increase in degree days above 10°C
a t 20 cm depth during the growing season over 4 years. However in the
same study, mean monthly soil temperatures were often lo-2OC lower
during periods of radiation cooling during the winter.
Hammond and Seeley (1978)used various plastic mulches in conjunc-
tion with insulation plus heating and cooling equipment to create a wide
range of soil temperatures ( lo-4.5' to 18O-26OC)for sweet cherry, peach,
and apple trees in early spring. Anthesis was neither advanced nor
delayed, except in frozen soil, by any of the treatments, indicating that
ambient temperature and not root temperature plays the major role in
bud development.
Mulch-induced soil temperature changes will be important if they are
near critical thresholds. For example, in Ireland where summer tempera-
tures are cooler than in many fruit-growing regions, lower mean soil
temperatures were recorded in mulched plots and this was deemed to
affect 'Golden Delicious' fruit quality negatively (Gormley et al. 1973a).
The reduction in heat transfer from soil under mulch to air during cool
spring periods could increase the risk of frost damage a t certain times of
the year as suggested by Haynes (1980), who cited German research
indicating that daily fluctuations of air temperature were greatest above
straw mulch. Similar risks of extreme soil and air temperature fluctua-
tions associated with black plastic mulching need assessment in fruit
growing areas where winter damage to fruit trees can be a concern.
3. Cultivation. The few comparative studies of soil temperatures
under cultivation indicate that daily summer temperatures tend to exceed
those under sward or mulch (Dancer 1964), averaging lo-3OC higher
than straw mulch a t 10 and 25 cm depths (Weller 1969). In the winter,
corresponding daily soil temperatures were often l 0 C colder for clean
tilled surfaces. In Norway, soil temperatures during summer a t 20 cm
depth were similar to grass and herbicide weed control but lower than
temperatures beneath black plastic mulches (Mage 1982).
4. Herbicides. Summer soil temperature differences a t 5 cm depth
under permanent sod vs. under herbicided surfaces were found to be as
great as 5O-6OC higher in the herbicide plots (Atkinsonand White 1981).
Cockcroft and Wallbrink (1966)reported that temperatures in the top 7.5
10. ORCHARD FLOOR VEGETATION MANAGEMENT 41 7
cm of herbicide-treated bare soil in Australia often exceed 32OC during

sunny summer days compared to no more than 24OC under straw mulch.
However, Mage (1982)in Norway found no difference in soil temperatures
at 20 cm from May to August under herbicided, grass or tilled plots, but
all were 2.3O-3.6OC lower than under a black plastic mulch.
V. CONCLUDING REMARKS
There are major differences in the effects of the four orchard floor
vegetation management options examined on the crop and the soil (Thble
10.3.).Although changes in the quality of the orchard soil environment
may be of long-term importance, most orchardists are more concerned
with improvements in growth and production of the orchard crop. Thus,
with the crop in mind, inorganic mulch, herbicide strip, overall herbicide,
and organic mulch soil management systems score well (Thble 10.3). The
inorganic mulch system has the highest rating since it is the only system
to result in improved growth, yield, and nutrient content of the tree crop,
while presenting no chance of mechanical or chemical injury and no real
hazard from vole damage.
Organic and inorganic mulching, herbicide strip, and permanent orchard
vegetation management systems offer an overall benefit when the quality
of the soil environment is the main consideration (lkble 10.3). The
highest rating for the organic mulch system is a consequence of increased
soil organic matter, increased availability of phosphorus and potassium,
increased soil moisture content, and improved soil physical properties,
which can reduce excess water runoff and erosion in the tree fruit growing
areas where this can be a serious problem. Changes in soil pH can also be
moderated, which would be desirable where near-optimum pH conditions
already exist.
Although important changes in soil properties have been measured
under various soil management systems, it is also clear that optimum
ranges for most soil properties are yet to be identified for tree fruits. Thus,
it is difficult to judge whether improved conditions have resulted from
many soil management-induced changes. This is particularly true for
changes in soil organic matter content, soil nutrient availability (except
for nitrogen), many soil physical properties such as bulk density and
porosity as well as alterations in the soil temperature regime. Because of
the perennial nature of the tree fruit crop, other soil problems such as
acidification may not become apparent until replanting occurs in older
orchard sites.
Other factors such as allelopathy, pest management, and orchard
temperatures have not been discussed sufficiently in this review to pro-
Table 10.3. Overall Summary of the Effects of Various Orchard Floor Vegetation Management Systems on the Crop and Soil
in Temperate n e e Fruit Orchards
Orchard floor vegetation management systems
Mulch Cultivation Herbicide

Factors Permanent
affected by orchard With Overall
floor management vegetation Organic Inorganic Continuous cover crop orchard floor Strip
Crop
h ++ +i + + ++ +i'
Vigor"
Leaf nutrient:
N __ Od + ++ + ++ ++
++ ++ + 0 0 - t
PK
~~
Yield ++ ++ + + +++' ++
- - 0
Fruit quality + 0 0 0
~ ~ ~ ~
Crop safety' ++ ++ ++
~~
Vole damage 0 + i i t t
Crop rating -2 +6 +8 +3 +3 +6 +I
Soil
Organic matter ++ ++ 0 0 0
Fertility:
N __ Od + ++ + ++ ++
PK ++ ++ 0 0 0 0
+ + 0 - 0 __ -
PH
Moisture __ ++ ++ ++ +
Erosion ++ ++ ++ + __ +
Soil rating +3 +9 +4 -2 +1 -1 +3
Total ratings +l +15 +12 +1 +4 +5 +10
“Controlled vigor considered as desirable.

b++, very beneficial; +, beneficial; 0, no appreciable effect; -, detrimental; --, very detrimental.
‘Variations could be expected depending on row width.
dWill depend on the C :N ratio of the mulches.
‘A triple plus given to account for the clear yield advantage shown in some European research.
/Includes potential for mechanical injury, root pruning, herbicide toxicity.
vide an evaluation of how they are influexed by soil and cover crop
management. The importance of these possible interrelationships, how-
ever, must be kept in mind. Allelopathy and integrated pest management
are receiving ever increasing attention and their interaction with orchard
floor management should become clear very soon. There seems to be,
however, little attention devoted to orchard temperature in this regard.
The effect of management systems on vole populations in orchards, also
not discussed, has been well covered recently (Byers 1984) however, and an
evaluation of vole importance in different systems was possible ('hble 10.3).
Comparative economic costs were not taken into consideration but are
likely to be very important to the grower in choosing an orchard floor
management system. Although permanent orchard vegetation over the
entire orchard floor may be the cheapest system to maintain, the cost of
poor establishment, low vigor, and low yields is likely unacceptable. The
high costs and various disadvantages of mechanical cultivation appear to
be limiting its use to preplant soil cultivation where it may have great
value, for example, in incorporating lime materials to adjust soil pH.
Although costs of materials and labor are likely to vary considerably
among fruit-growing areas, the cost and availability of organic and
inorganic materials may well be a major limitation to their widespread
use. The cost of various herbicide materials can also vary, but Atkinson
(1985)concludes that the maximum cost/benefit is achieved with the use
of glyphosate in tree fruit plantings. Use of herbicide strips rather than
overall herbicide will result in further economies.
Thus, although organic mulching in the tree row with grassed alley
appears to be the best orchard floor management system ('hble 10.3),
economic considerations may make the use of an inorganic mulch or herbi-
cides a more appropriate choice. Use of inorganic mulches in vegetable
and small fruit production is widespread and its adoption by the tree fruit
industry may be hastened by the increasing concern about pesticides. In
the meantime, the grassed-alley, herbicide strip system is both sufficiently
economical and beneficial to the crop and soil to represent a good choice
in management systems.
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Subject Index
A E
Abscisic acid, rose senescence, Environment
66 ginseng, 211-26
Abscission greenhouse management, 32-38
rose flower, 63-64 Ethylene, rose senescence, 65-66
rose leaf, 63-64
Acclimatization, micropropagation,
F
278-81,316-17
Filbert, in uitro culture, 313-14
Almond, in uitro culture, 313
Flowering
Anatomy and morphology, ginseng,
honey bee pollination, 239-43
198-201
rose, 60-66
Apple Fruit crops
in uitro, 319-21
chlorosis, 161-65
summer pruning, 351-75
honey bee pollination, 244-50, 254-56
in uitro culture, 273-349
B orchard floor management, 377-430
Boron, pine bark media, 119-22 summer pruning, 351-75
C G
Calcium Genetics and breeding
container growing, 84-85 ginseng, 197-98
pine bark media, 116-17 in uitro techniques, 318-24
Chelates, 169-71 Germplasm preservation, in uitro, 324-25
Chestnut, in uitro culture, 311-12 Ginseng, 187-236
Chlorosis, iron deficiency induced, Grafting, rose, 56-57
133-86 Grape, chlorosis, 165-66
Citrus Greenhouse and greenhouse crops,
chlorosis, 166-68 energy efficiency, 1-52
honey bee pollination, 247-48 Growth regulators. See Growth
Container production, nursery substances
crops, 75-101 Growth substances
Copper, pine bark media, 122-23 ginseng, 226
Cytokinin, rose senescence, 66 rose, 53-73
431
432 SUBTECT INDEX
H Phosphorus
Hazelnut. See Filbert container growing, 82-84
Honey bee, 237-72 pine bark media, 112-13
Photosynthesis, ginseng, 223-26
I Physiology
I n uitro propagation, 57-58, 273-349 ginseng, 211-13
nuts, 273-349 rose, 3-53
rose, 57-58 summer pruning, 351-75
temperate fruits, 273-349 Pigmentation, rose, 64-65
Iron Pine bark, potting media, 103-31
deficiency chlorosis, 133-86 Pistachio, in uitro culture, 315
pine bark media, 123 Pollination
ginseng, 201-2
M honey bee, 237-72
Magnesium Potassium
container growing, 84-85 container growing, 84
pine bark media, 117-19 pine bark media, 113-14
Manganese, pine bark media, 123-24 Propagation
Media, pine bark, 103-31 ginseng, 206-9
Micronutrients rose, 54-58
container growing, 85-87 Pruning
pine bark media, 119-24 apple, 351-75
Micropropagation. See I n uitro peach, 351-75
propagation Prunus, in uitro culture, 322
Mycorrhizae, container growing, 93
R
N Rejuvenation, rose, 59-60
Nitrogen Root, rose, 57
container growing, 80-82 Rose, growth substances, 3-53
pine bark media, 108-12
Nursery crops, nutrition, 75-101 S
Nut crops Seed, rose propagation, 54-55
honey bee pollination, 250-51 Senescence, rose, 65-66
in uitro culture, 273-349 Soil
Nutrition orchard floor management, 377-430
container nursery crops, 75-101 testing, 88-90
ginseng, 209-11 Storage, rose plants, 58-59
pine bark media, 103-31
T
0 Tissue, nutrient analysis, 90
Orchard floor management, 377-430
V
Ornamental plants, chlorosis, 168-69
Vegetable crops, honey bee pollination,
251-54
P
Virus elimination, 318
Peach, summer pruning, 351-75
Pear, in uitro culture, 321 W
Pecan, in uitro culture, 314-15 Walnut, in uitro culture, 312
Pest control, ginseng, 227-29 Weed control, ginseng, 228-29
PH
container growing, 87-88 Z
pine bark media, 114-17 Zinc, pine bark media, 124
Cumulative Subject Index
A ginseng, 9:198-201
Abscisic acid kiwifruit, 6:13-50
dormancy, 7:275-77 navel orange, 8:132-33
rose senescence, 9:66 orchid, 5: 28 1-83
stress, 4:249-50 pecan flower, 8:217-55
Abscission petal senescence, 1:212-16
anatomy and histochemistry, pollution injury, 8:15
1:172-203 Angiosperms, embryogenesis,
flower and petals, 3:104-7 1:1-78
regulation, 7:415-16 Anthurium, fertilization, 5:334-35
rose, 9:63-64 Antitranspirants, 7:334
Acclimatization Apical meristem, cryopreservation,
foliage plants, 6:119-54 6~357-72
herbaceous plants, 6:379-95 Apple
micropropagation, 9:278-8 1,316-17 alternate bearing, 4: 136-37
Actinidia, 6:4-12 CA storage, 1:303-6
Adzuki bean, genetics, 2:373 chemical thinning, 1:270-300
Agaricus, 6:85-118 fertilization, 1:105
Agrobacterium tumefaciens, 3:34 fire blight control, 1:423-74
Air pollution, 8: 1-42 flower induction, 4:174-203
Almond, in uitro culture, 9:313 in uitro, 5:241-43; 9:319-21
Alocasia, 8:46, 57. See also Aroids, light, 2:240-48
edible nitrogen metabolism, 4:204-46
Alternate bearing replant disease, 2:3
chemical thinning, 1:285-89 root distribution, 2:453-56
fruit crops, 4:128-73 stock-scion relationships, 3:315-75
pistachio, 3:387-88 summer pruning, 9:351-75
Aluminum, deficiency and toxicity, in watercore, 6:189-251
fruits and nuts, 2:154 yield, 1:397-424
Amorphophallus, 8:46, 57. See also Apricot, CA storage, 1:309
Aroids, edible Aroids, edible, 8:43-99
Anatomy and morphology Arsenic, deficiency and toxicity, in fruits
embryogenesis, 1:4-21, 35-40 and nuts, 2:154
fruit abscission, 1:172-203 Artichoke, CA storage, 1:349-50
fruit storage, 1:314 Asexual embryogenesis, 1:l-78;
433
434 CUMULATIVE SUBIECT INDEX
2 ~268-310;3:214-314; 7: 163-68, Calcium

171-73, 176-77, 184, 185-87, cell wall, 5:203-5
187-88, 189 container growing, 9:84-85
Asparagus deficiency and toxicity, in fruits and
CA storage, 1:350-51 nuts, 2:148-49
fluid drilling of seed, 3:21 foliar application, 6:328-29
Auxin, and dormancy, 7:273-74 nutrition, 5:322-23
Avocado, flowering, 8:257-89 pine bark media, 9:116-17
Azalea, fertilization, 5:335-37 tipburn, disorder, 4:50-57
Carbohydrate
B
metabolism, 7:69-108
Babaco, in uitro culture, 7:178
partitioning, 7:69-108
Bacteria
Carbon dioxide, enrichment, 7:345-98,
ice nucleating, 7:210-12
544-45
pathogens of bean, 3:28-58
Carnation, fertilization, 1 : l O O ; 5:341-45
tree short life, 2:46-47
Carrot
wilt of bean, 3:46-47
CA storage, 1:362-66
Bacteriocides, fire blight, 1:450-59
fluid drilling of seed, 3:13-14
Bacteriophage, fire blight control,
Caryophyllaceae, in uitro, 5:237-39
1:449-50
CA storage. See Controlled-atmosphere
Banana
storage
Cauliflower, CA storage, 1:359-62
fertilization, 1:105
Celeriac, CA storage, 1:366-67
in uitro culture, 7:178-80
Celery
Bean
fluid drilling of seed, 3: 14
fluid drilling of seed, 3:21
Cell culture, 3:214-314
resistance to bacterial pathogens,
Cell wall hydrolases, 5: 169-219
3~28-58
Chelates, 9:169-71
Bedding plants, fertilization, 1:99-100;
Cherry, CA storage, 1:308
5:337-41
Chestnut
Beet
blight, 8:281-336
CA storage, 1:353
in uitro culture, 9:311-12
fluid drilling of seed, 3:18-19
Chicory, CA storage, 1:379
Begonia (Rieger),fertilization, 1:104
Chilling
Biennial bearing. See Alternate bearing
injury, 4:260-61
Bird damage, 6:277-78
pistachio, 3:388-89
Boron
Chlorine
deficiency and toxicity symptoms in
deficiency and toxicity symptoms in
fruits and nuts, 2:151-52
fruits and nuts, 2:153
foliar application, 6:328
nutrition, 5:239
nutrition, 5:327-28
Chlorosis, iron deficiency induced,
pine bark media, 9:119-22
9~133-86
Brassicaceae, in uitro, 5:232-35
Chrysanthemum fertilization, 1:loo-101;
Breeding. See Genetics and breeding
5~345-52
Broccoli, CA storage, 1:354-55
Citrus
Brussels sprouts, CA storage, 1:355
alternate bearing, 4:141-44
Bulb. See ' h l i p
asexual embryogenesis, 7: 163-68
C CA storage, 1:312-13
Cabbage chlorosis, 9: 166-68
CA storage, 1:355-59 cold hardiness, 7:201-38
fertilization, 1:117-18 fertilization, ]:I05
CUMULATIVE SUBJECT INDEX 435
honey bee pollination, 9:247-48 floral promoter, 4:112-13

in uitro culture, 7:161-70 grape root, 5:150, 153-56
navel orange, 8:129-79 lettuce tipburn, 4:57-58
nitrogen metabolism, 8:181 rose senescence, 9:66
rootstock, 1:237-69
Cloche (tunnel),7:356-57 D
Coconut palm Date palm
asexual embryo genesis, 7:184 asexual embryogenesis, 7: 185-87
in uitro culture, 7:183-85 in uitro culture, 7:185-87
Cold hardiness, 2:33-34 Daylength. See Photoperiod
citrus, 7:201-38 Deer, 6:274-75
herbaceous plants, 6:373-417 Deficiency symptoms, in fruit and nut
injury, 2:26-27 crops, 2:145-54
nutrition, 3:144-71 ‘Delicious’ apple, 1:397-424
pruning, 8:356-57 Disease
Colocasia, 45, 55-56. See also Aroids and air pollution, 8:25
Common blight, of bean, 3:45-46 aroids, 8:67-69
Compositae, in uitro, 5:235-37 bacterial, of bean, 3:28-58
Container production, nursery crops, control by virus, 3:399-403
9~75-101 controlled-atmosphere storage,
Controlled environment agriculture 3:412-61
7:534-545. See also Greenhouse hydroponic crops, 7:530-34
and greenhouse crops; Hydroponic lettuce, 2:187-97
culture mycorrhizal fungi, 3:182-85
Controlled-atmosphere storage. See also root, 5:29-31
individual fruits and vegetables. stress, 4:261-62
flowers, 3:98 tulip, 5:63, 92
fruit quality, 8:lOl-27 Disorder, watercore, 6:189-251
fruits, 1:301-36; 4:259-60 Dormancy, 2:27-30
pathogens, 3:412-61 release in fruit trees, 7:239-300
seeds, 2:134-35 tulip, 5:93
tulip, 5:105 Drip irrigation, 4:l-48
vegetable quality, 8:lOl-27 Drought resistance, 4:250-51
vegetables, 1:337-94; 4:259-60 Dwarfing
Copper apple, 3:315-75
deficiency and toxicity symptoms in by virus, 3:404-5
fruits and nuts, 2:153
foliar application, 6:329-30 E
nutrition, 5:326-27 Easter lily, fertilization, 5:352-55
pine bark media, 9:122-23 Embryogenesis. See Asexual
Corynebacterium flaccumfaciens, 3:33,46 embryogenesis
Cowpea, genetics, 2:317-48 Endothia parasitica, 8:291-336
Cranberry, fertilization, 1:106 Energy efficiency, in greenhouses
Cryopreservation, apical meristems, 1~141-71;9: 1-52
6:357-72 Environment
Cryphonec tria parasitica. See Endo t hia air pollution, 8:20-22
paras it ica controlled for agriculture, 7:534-45
Crytosperma, 8:47, 58. See also Aroids, for energy efficiency, 1:141-71;
edible 9:l-52
Cucumber, CA storage, 1:367-68 embryogenesis, 1:22,43-44
Cytokinin fruit set, 1:411-12
dormancy, 7:272-73 ginseng, 9:211-26
436 CUMULATIVE SUBJECT INDEX
greenhouse management, 9:32-38 control, 4:159-60

navel orange, 8:138-40 honey bee pollination, 9:239-43
nutrient film technique, 5:13-26 induction, 4:174-203; 254-56
Erwinia initiation, 4:152-53
amylouora, 1:423-74 in uitro, 4:106-27
lathyri, 3:34 kiwifruit, 6:21-35
Essential elements, 5:318-30 orchid, 5:297-300
foliar nutrition, 6:287-55 pecan, 8:217-55
pine bark media, 9:103-31 phase change, 7: 109-55
soil testing, 7:l-68 photoperiod, 4 :66- 105
Ethylene pistachio, 3:378-87
CA storage, 1:317-19, 348 postharvest physiology, 1:204-36;
dormancy, 7:277-79 3:59-143
flower longevity, 3:66-75 pruning, 8:359-62
kiwifruit respiration, 6:47-48 regulation in floriculture, 7:416-24
rose senescence, 9:65-66 rose, 9:60-66
senescence, 1:204-36; 3:59-143
F sugars, 4:114
Fertilization and fertilizer tulip, 5:57-59
anthurium, 5:334-35 Fluid drilling, 3:l-58
azalea, 5:335-37 Foliage plants
bedding plants, 5:337-41 acclimatization, 6:119-54
carnation, 5:341-45 fertilization, 1:102-3; 5:367-80
chrysanthemum, 5:345-52 Foliar nutrition, 6:287-355
controlled release, 1:79-139; 5:347-48 Frost
Easter lily, 5:352-55 apple fruit set, 1:407-8
foliage plants, 5:367-80 citrus, 7:201-38
foliar, 6:287-355 Fruit
geranium, 5:355-57 abscission, 1:172-203
greenhouse crops, 5:317-403 CA storage and quality, 8:lOl-27
lettuce, 2:175 development in pistachio, 3:382-91
nitrogen, 2:401-4 diseases in CA storage, 3:412-61
orchid, 5:357-58 kiwifruit, 6:35-48
poinsettia, 5:358-60 navel orange, 8:129-79
rose, 5:361-63 quality and pruning, 8:365-67
snapdragon, 5:363-64 ripening, 5:190-205
soil testing, 7:l-68 set, 1:397-424; 4:153-54
trickle irrigation, 4:28-31 in navel oranges, 8:140-42
tulip, 5:364-66 size and thinning, 1:293-94; 4:161
Fig, ripening, 4:258-59 softening, 5:109-219
Filbert, in uitro culture, 9:313-14 tomato parthenocarpy, 6:65-84
Fire blight, 1:423-74 Fruit crops
Floricultural crops alternate bearing, 4:128-73
fertilization, 1:98-104 avocado flowering, 8:257-89
growth regulation, 7:399-481 CA storage, 1:301-36
postharvest physiology and senes- diseases, 3:412-61
cence, 1:204-36; 3:59-143 chlorosis, 9:161-65
Florigen, 4:94-98 citrus cold hardiness, 7:201-38
Flower and flowering dormancy release, 7:239-300
alternate bearing, 4: 149 fertilization, 1: l o 4 4
avocado, 8:257-89 foliar nutrition, 6:287-355
honey bee pollination, 9:244-50, Gibberellin

254-56 dormancy, 7:270-71
in uitro culture, 7:157-200; 9:273-349 floral promoter, 4: 114
kiwifruit, 6:l-64 grape root, 5:150-51
navel orange, 8:129-79 Ginseng, 9: 187-236
nutritional ranges, 2:143-64 Girdling, 4:251-52
orange, navel, 8:129-79 Grafting
orchard floor management, 9:377-430 phase change, 7:136-37, 141-42
pecan flowering, 8:217-55 rose, 9:56-57
pruning, 8:339-80 Grape
roots, 2:453-57 CA storage, 1:308
short life and replant problem, 2:l-116 chlorosis, 9:165-66
summer pruning, 9:351-75 root, 5:127-68
water status, 7:301-44 Greenhouse and greenhouse crops
Fungi carbon dioxide, 7:357-60,544-45
mushroom, 6:85-118 energy efficiency, 1:141-71; 9:l-52
mycorrhizal, 3: 172-213 growth substances, 7:399-481
pathogens in postharvest storage nutrition and fertilization,
3~412-61 5~317-403
Fungicide, and apple fruit set, 1:416 Growth regulators. See Growth
substances
G Growth substances, 2:60-66
Garlic, CA storage, 1:375 apple dwarfing, 3:315-75
Genetics and breeding apple fruit set, 1:417
aroids, 8:72-75 apple thinning, 1:270-300
bean, bacterial resistance, 3 :28-58 CA storage in vegetables, 1:346-48
chestnut blight resistance, 8:313-21 cell cultures, 3:214-314
citrus cold hardiness, 7:221-23 cold hardiness 7:223-25
embryogenesis, 1:23 dormancy, 7:270-79
fire blight resistance, 1:435-36 embryogenesis, 1:41-43; 2:277-81
flower longevity, 1:208-9 floriculture, 7 :399-481
ginseng, 9:197-98 flower induction, 4:190-95
in uitro techniques, 9:318-24 ginseng, 9:226
lettuce, 2:185-87 in uitro flowering, 4:112-15
mushroom, 6: 100-111 meristem and shoot-tip culture,
navel orange, 8:150-56 5:221-27
nitrogen nutrition, 2:410-11 navel oranges, 8:146-47
plant regeneration, 3:278-83 petal senescence, 3:76-78
pollution insensitivity, 8:18-19 phase change, 7:137-38; 142-43
tomato parthenocarpy, 6:69-70 rose, 9:53-73
tree short life, 2:66-70
Vigna, 2:311-94 H
Genetic variation Halo blight of beans, 3:44-45
alternate bearing, 4: 146-50 Hardiness, 4:250-51
photoperiodic response, 4:82 Harvest
pollution injury, 8:16-19 flower stage, 1:211-12
Geranium, fertilization, 5:355-57 index, 7:72-74
Germination, seed, 2:117-41, 173-74 lettuce, 2:176-81
Germplasm preservation Hazelnut. See Filbert
cryopreservation, 6:357-72 Herbaceous plants, subzero stress,
in uitro, 5:261-64; 9:324-25 6:373-417
Histochemistry L
flower induction, 4:177-79 Lamps, for plant growth, 2:514-31
fruit abscission, 1:172-203 Leaves, flower induction, 4: 188-89
Histology, flower induction, 4: 179-84 Leek
Honey bee, 9:237-72 CA storage, 1:375
Horseradish, CA storage, 1:368 fertilization, 1:118
Hydrolases, 5: 169-219 Leguminosae, in uitro, 5:227-29
Hydroponic culture, 5:l-44; 7:483-558 Lemon, rootstock, 1:244-46. See also
Hypovirulence, in Endothia parasitica, Citrus
8~299-310 Lettuce
I fertilization, 1:118
Ice-nucleating bacteria, 7:210-12 fluid drilling of seed, 3:14-17
Insects industry, 2:164-207
aroids, 8:65-66 tipburn, 4:49-65
avocado pollination, 8:275-77 Light
hydroponic crops, 7:530-34 fertilization, greenhouse crops, 5:330-31
lettuce, 2:197-98 fruit set, 1:412-13
tree short life, 2:52 nitrogen nutrition, 2:406-7
tulip, 5:63, 92 orchards, 2:208-67
I n vitro photoperiod, 4:66-105
cold acclimation, 6:382 plant growth, 2:491-537
cryopreservation, 6:357-72
embryogenesis, 1:l-78; 2:268-310 M
flowering, 4:106-27 Magnesium
phase change, 7: 144-45 container growing, 9:84-85
propagation, 3:214-3 14; 5 :22 1-77 ; deficiency and toxicity symptoms in
7~157-200;9:57-58, 273-349 fruits and nuts, 2:148
Iron foliar application, 6:331
deficiency and toxicity, in fruits and nutrition, 5:323
nuts, 2:150 pine bark media, 9:117-19
deficiency chlorosis, 9:133-86 Mandarin, rootstock, 1:250-52
foliar application, 6:330 Manganese
nutrition, 5:324-25 deficiency and toxicity symptoms in
pine bark media, 9:123 fruits and nuts, 2:150-51
Irrigation foliar application, 6:331
drip or trickle, 4:l-48 nutrition, 5:235-326
fruit trees, 7:331-32 pine bark media, 9:123-24
grape root growth, 5:140-41 Mango
lettuce industry, 2:175 alternate bearing, 4: 145-46
navel orange, 8:161-62 asexual embryogenesis, 7 :171-73
root growth, 2:464-65 CA storage, 1:313
J in vitro culture, 7:171-73
Juvenility, 4: 111-12 Media
pecan, 8:245-47 fertilization, greenhouse crops, 5:333
tulip, 5:62-63 pine bark, 9:103-31
woody plants, 7:109-55 Meristem culture, 5:221-77
Metabolism
K flower, 1:219-23
Kale, fluid drilling of seed, 3:21 nitrogen in citrus, 8:181-215
Kiwifruit (botany),6:l-64 seed, 2:117-41
CUMULATIVE SUBJECTINDEX 439
Micronutrients Nursery crops

container growing, 9:85-87 fertilization, 1:106-12
pine bark media, 9:119-24 nutrition, 9:75-101
Micropropagation. See also I n uitro, Nut crops
propagation chestnut blight, 8:291-336
nuts, 9:273-349 fertilization, 1:lo6
rose, 9:57-58 honey bee pollination, 9:250-51
temperate fruits, 9:273-349 in uitro culture, 9:273-349
tropical fruits and palms, nutritional ranges, 2:143-64
7: 157-200 pistachio culture, 3:376-96
Microtus. See Vole Nutrient
Moisture, and seed storage, 2:125-32 concentration in fruit and nut crops,
Molybdenum nutrition, 5:328-29 2:154-62
Monocot, in uitro, 5:253-57 film technique, 5:l-44
Morphology foliar-applied, 6:287-355
navel orange, 8:132-33 media, for asexual embryogenesis,
orchid, 5:283-86 2:273-81
pecan flowering, 8:217-43 for organogenesis, 3:214-314
Moth bean, genetics, 2:373-74 plant and tissue analysis, 7:30-56
Mung bean, genetics, 2:348-64 solutions, 7:524-30
Mushroom uptake, in trickle irrigation, 4:30-31
CA storage, 1:371-72 Nutrition ( h u m a n )
spawn, 6:85-118 aroids, 8:79-84
Muskmelon, fertilization, 1 :118-19 CA storage, 8:lOl-27
Mycoplasma-like organisms, tree short Nutrition ( p l a n t )
life, 2:50-51 air pollution, 8:22-23, 26
Mycorrhizae, container growing, cold hardiness, 3:144-71
9:93 container nursery crops, 9:75-101
Mycorrhizal fungi, 3:172-213 embryogenesis, 1:40-41
grape root, 5:145-46 fire blight, 1:438-41
foliar, 6:287-355
N fruit and n u t crops, 2:143-64
Navel orange, 8:129-79 ginseng, 9:209-11
Nectarine, CA storage, 1:309-10 greenhouse crops, 5:317-403
Nematodes mycorrhizal fungi, 3:185-91
aroids, 8:66 navel orange, 8: 162-66
lettuce, 2:197-98 nitrogen in apple, 4:204-46
tree short life, 2:49-50 nutrient film techniques, 5:18-21,
NFT. See Nutrient, film technique 31-53
Nitrogen pine bark media, 9:103-31
CA storage, 8:116-17 slow-release fertilizers, 1:79-139
container growing, 9:80-82
deficiency and toxicity symptoms in 0
fruits and nuts, 2:146 Oil palm
in embryogenesis, 2:273-75 asexual embryogenesis, 7:187-88
foliar application, 6:332 in vitro culture, 7: 187-88
metabolism in apple, 4:204-46 Okra, CA storage, 1:372-73
in citrus, 8:181-215 Olive, alternate bearing, 4:140-41
nutrition, 2:395, 423; 5:319-20 Onion
pine bark media, 9:108-12 CA storage, 1:373-75
trickle irrigation, 4:29-30 fluid drilling of seed, 3:17-18
Orange. See also Citrus fertilization, 1:119

alternate bearing, 4:143-44 fluid drilling in seed, 3:20
sour, rootstock, 1:242-44 Persimmon
sweet, rootstock, 1:252-53 CA storage, 1:314
trifoliate, rootstock, 1:247-50 quality, 4:259
Orchard and orchard systems Pest control
floor management, 9:377-430 fire blight, 1:423-74
light, 2:208-67 ginseng, 9:227-29
root growth, 2:469-70 hydroponics, 7:530-34
water, 7:301-44 Pesticide and fire blight, 1:450-61
Orchid Pests, vertebrate, 6:253-85
fertilization, 5:357-58 PH
p hysi ol ogy, 5 :2 79-3 15 container growing, 9:87-88
Organogenesis, 3:214-314. See also I n fertilization greenhouse crops,
uitro. Tissue culture 5~332-33
Ornamental plants pine mark media, 9:114-17
chlorosis, 9: 168-69 soil testing, 7:8-12; 19-23
fertilization, 1:98-104, 106-16 Phase change, 7:109-55
foliage acclimatization, 6:119-54 Phosphorus
container growing, 9:82-84
P deficiency and toxicity, in fruits and
Papaya nuts, 2:146-47
asexual embryogenesis, 7: 176-77 nutrition, 5:320-21
CA storage, 1:314 pine bark media, 9:112-13
in uitro culture, 7:175-78 trickle irrigation, 4:30
Parsley Photoperiod, 4:66-105, 116-17
CA storage, 1:375 Photosynthesis
fluid drilling of seed, 3:13-14 efficiency, 7:71-72
Parsnip, fluid drilling of seed, 3: 13-14 ginseng, 9:223-26
Parthenocarpy, tomato, 6:65-84 light, 2:237-38
Passion fruit, in vitro culture, 7:180-87 Physiology
Pathogen elimination, in uitro, carbohydrate metabolism,
5~257-61 7 :69-108
Peach citrus cold hardiness, 7:201-38
CA storage, 1:309-10 cut flower, 1:204-36; 3:59-143
short life, 2:4 dormancy, 7:239-300
summer pruning, 9:351-75 embryogenesis, 1:2 1-23 ; 2:268-3 10
Peach palm. See Pejibaye flowering, 4:106-27
Pear ginseng, 9:211-13
CA storage, 1:306-8 juvenility, 7:109-55
decline, 2: 11 nutritional quality and CA storage,
fire blight control, 1:423-74 8 : 118-20
in uitro, 9:321 orchid, 5:279-315
root distribution, 2:456 pollution injury, 8: 12-16
short life, 2:6 pruning, 8:339-80
Pecan root pruning, 6:158-71
alternate bearing, 4: 139-40 rose, 9:3-53
fertilization, 1:106 seed, 2:117-41
in uitro culture, 9:314-15 subzero stress, 6:373-417
Pecan flowering, 8:217-55 summer pruning, 9:351-75
Pejibaye, in vitro culture, 7:189 tomato parthenocarpy, 6:71-74
Pepper (Capsicum) tulip, 5:45-125
CA storage, 1:375-76 watercore, 6:189-251
CUMULATIVE SUBJECTINDEX 441
Phytotoxins, 2:53-56 ginseng, 9:206-9

Pigmentation orchid, 5:291-97
flower, 1:216-19 rose, 9:54-58
rose, 9:64-65 tropical fruit, palms 7: 157-200
Pinching, by chemicals, 7:453-61 Protected crops, carbon dioxide, 7:345-98
Pineapple Pruning, 4:161, 8:339-80
CA storage, 1:314 apple, 9:351-75
in uitro culture, 7:181-82 apple training, 1:414
Pine bark, potting media, 9:103-31 chemical, 7:453-61
Pistachio fire blight, 1:441-42
alternate bearing, 4:137-39 light interception, 2:250-51
culture, 3:376-93 peach, 9:351-75
in uitro culture, 9:315 phase change, 7: 143-44
Plantain, in uitro culture, 7:178-80 root, 6:155-88
Plant protection, short life, 2:79-84 summer, 9:351-75
Plum, CA storage, 1:309 Prunus
Poinsettia, fertilization, 1:103-4; 5:358-60 in uitro, 5:243-44; 9:322
Pollination root distribution, 2:456
apple, 1:402-4 Pseudomonas
avocado, 8:272-83 phaseolicola, 3:32-33, 39, 44-45
embryogenesis, 1:21-22 solanacearum, 3:33
fruit set, 4:153-54 syringae, 3:33,40; 7:210-12
ginseng, 9:201-2
honey bee, 9:237-72 R
kiwifruit, 6:32-35 Rabbit, 6:275-76
navel orange, 8:145-46 Radish, fertilization, 1:121
orchid, 5:300-302 Rejuvenation
protection, 7 :463-64 rose, 9:59-60
Pollution; 8:l-42 woody plants, 7:109-55
Postharvest physiology Replant problem, deciduous fruit trees,
aroids, 8:84-86 2:l-116
CA storage and quality, 8:lOl-27 Respiration
cut flower, 1:204-36; 3:59-143 fruit in CA storage, 1:315-16
foliage plants, 6:119-54 kiwifruit, 6:47-48
fruit, 1:301-36 vegetables in CA storage, 1:341-46
lettuce, 2:181-85 Rhizobium, 3:34, 41
navel orange, 8:166-72 Rice bean, genetics, 2:375-76
pathogens, 3:412-61 Root
seed, 2:117-41 diseases, 5:29-31
vegetables, 1:337-94 environment, nutrient film technique,
Potassium 5~13-26
container growing, 9:84 grape, 5:127-68
deficiency and toxicity, in fruits and pruning, 6:155-88
nuts, 2:147-48 rose, 9:57
foliar application, 6:331-32 tree crops, 2:424-90
nutrition, 5:321-22 Rootstocks
pine bark media, 9:113-14 alternate bearing, 4: 148
trickle irrigation, 4:29 apple, 1:405-7
Potato citrus, 1:237-69
CA storage, 1:376-78 fire blight, 1:432-35
fertilization, 1:120-21 light interception, 2:249-50
Propagation. See also I n vitro navel orange, 8:156-61
floricultural crops, 7:461-62 root systems, 2:471-74
442 CUMULATIVE SUBIECT INDEX
stress, 4:253-54 Stress

tree short life, 2:70-75 benefits of, 4:247-71
Rosaceae, i n uitro, 5:239-48 climatic, 4: 150-51
Rose on plants, 2:34-37
fertilization, 1:104; 5:361-63 protection, 7 :463-66
growth substances, 9:3-53 subzero temperature, 6:373-4 17
in uitro, 5:244-48 Sugar
allocation, 7:74-94
S flowering, 4:114
Salinity, 4:22-27 Sugar beet, fluid drilling of seed,
air pollution, 8:25-26 3:18-19
Scoring, and fruit set, 1:416-17 Sulfur
Seed deficiency and toxicity, in fruits and
abortion, 1:293-94 nuts, 2:154
on flower induction, 4: 190-95 nutrition, 5:323-24
fluid drilling, 3:l-58 Sweet potato, fertilization, 1:121
kiwifruit, 6:48-50 Symptoms, deficiency and toxicity, in
lettuce, 2:166-74 fruits and nuts, 2:145-54
rose propagation, 9:54-55
vegetable, 3:l-58 T
viability and storage, 2:117-41 Taro. See Aroids
Senescence Temperature
cut flower, 1:204-36; 3:59-143 apple fruit set, 1:408-11
rose, 955-66 CA storage of vegetables, 1:340-41
Sensory quality, CA storage, 8:lOl-27 cryopreservation, 6:357-72
Shoot-tip culture, 5:221-77. See also fertilization, greenhouse crops,
Micropropagation 5~331-32
Short life problem, fruit crops, 2:l-116 fire blight forecasting, 1:456-59
Small fruit, CA storage, 1:308 interaction with photoperiod, 4:80-81
Snapdragon fertilization, 5 :363-64 navel orange, 8:142
Sodium, deficiency and toxicity, in nutrient film technique, 5:21-24
fruits and nuts, 2:153-54 plant growth, 2:36-37
Soil seed storage, 2:132-33
grape root growth, 5:141-44 subzero stress, 6:373-417
management and root growth, 2:465-69 Thinning, apple, 1:270-300
orchard floor management, 9:377-430 Tipburn, in lettuce, 4:49-65
plant relations, trickle irrigation, Tissue
4: 18-21 culture, 1:l-78; 2:268-310; 3:214-314
stress, 4:151-52 dwarfing, 3:347-48
testing, 7:l-68; 9:88-90 nutrient analysis, 7:52-56; 9:90
Soilless culture, 5:l-44 Tissue culture, aroids, 8:75-78
Solanaceae, in uitro, 5:229-32 Tomato
Somatic embryogenesis. See Asexual CA storage, 1:380-86
embryogenesis fertilization, 1:121-23
Storage fluid drilling of seed, 3:19-20
cut flower, 3:96-100 parthenocarpy, 6:65-84
rose plants, 9:58-59 Toxicity symptoms, in fruit and nut
seed, 2:117-41 crops, 2:145-54
Strawberry 'Ikansport, cut flowers, 3:lOO-104
fertilization, 1:106 'Ikee decline, 2:l-116
in uitro, 5:239-41 'Ikickle irrigation, 4: 1-48
nlip W
fertilization, 5:364-66 Walnut, in uitro culture, 9:312
physiology, 5:45-125 Water
n n n e l (cloche),7:356-57 cut flower relations, 3:61-66
n r f g r a s s , fertilization, 1:112-17 fertilization, greenhouse crops, 5:332
'hrnip, fertilization, 1:123-24 fruit trees, 7:301-44
light in orchards, 2:248-49
U trickle irrigation, 4: 1-48
Urd bean, genetics, 2:364-73 Watercore, 6:189-251
Urea, foliar application, 6:332 Watermelon, fertilization, 1:124
Weed control, ginseng, 9:228-29
V Weeds
Vase solutions, 3:82-95 and lettuce research, 2:198
Vegetable crops virus, 3:403
aroids, 8:43-99 Woodchuck. 6~276-77
diseases, 3:412-61 A
and quality, 8:lOl-27 Xanthomonas phaseoli, 3:29-32, 41,
fertilization, 1:117-24 45-46
fluid drilling of seeds, 3:l-58 Xanthosoma, 8:45-46,56-57. See also
honey bee pollination, 9:251-54 Aroids, edible
hydroponics, 7:483-558
mushroom spawn, 6235-118 Y
tomato parthenocarpy, 6:65-84 Yield determinants, 7:70-74; 97-99
Vernalization, 4: 117
Vertebrate pests, 6:253-85 Z
Vigna, genetics, 2:311-94 Zinc
Virus deficiency and toxicity, in fruits and
benefits in horticulture, 3:394-411 nuts, 2:151
elimination, 7:157-200; 9:318 foliar application, 6:332, 336
tree short life, 2:50-51 nutrition, 5:326
Vole, 6:254-74 pine bark media, 9:124
Cumulative Contributor Index
Aldwinckle, H. S., 1:423 El-Goorani, M. A., 3:412

Asokan, M. P., 8:43 Esan, E. B., 1:l
Atkinson, D., 2:424 Faust, M., 2:vii, 142; 4:174; 6:287
Aung, L. H., 5:45 Ferguson, A. R., 6:l
Bailey, W. G., 9:187 Ferree, D. C., 6:155
Baird, L. A. M., 1:172 Fery, R. L., 2:311
Barden, J . A., 9:351 Flick, C. E., 3:214
Barker, A. V., 2:411 Geisler, D., 6:155
Bass, L. N., 2:117 George, W. L., Jr., 6:25
Beer, S. V., 1:423 Goldschmidt, E. E., 4:128
Benschop, M., 5:45 Graves, C. J., 5:l
Blanpied, G. D., 7:xi Gray, D., 3:l
Buban, T., 4:174 Grierson, W., 4: 247
Byers, R. E., 6:253 Griffen, G. J., 8:291
Caldas, L. S., 2:568 Grodzinski, B., 7:345
Campbell, L. E., 2:524 Hackett, W. P., 7:109
Carter, J. V., 3:144 Halevy, A. H., 1:204; 3:59
Cathey, H. M., 2:524 Hendrix, J. W., 3:172
Chin, C. K., 5:221 Hogue, E. J., 9:377
Cohen, M., 3:394 Huber, D. J., 5:169
Collier, G. F., 4:49 Hutchinson, J . F., 9:273
Collins, W. L., 7:483 Isenberg, F. M. R., 1:337
Conover, C. A., 5:317; 6:119 Iwakiri, B. T., 3:376
Coyne, D. P., 3:28 Jackson, J . E., 2:208
Crane, J. C., 3:376 Janick, J., 1:ix; 8:xi
Daie, J., 7:69 Jensen, M. H., 7:483
Davenport, T. L., 8:257 Joiner, J . N., 5:317
Davies, F. S., 8:129 Jones, H. G., 7:301
DeGrandi-Hoffman, G., 9:237 Jones, J . B., Jr., 7 : l
De Hertogh, A. A., 5:45 Kang, S.-M., 4:204
Dennis, F. G., Jr., 1:395 Kato, T., 8:181
Doud, S. L., 2 : l Kawada, K., 4:247
Dunavent, M. G., 9:103 Kierman, J., 3:172
Eans, D. A., 3:214 Kofranek, A. M., 8:xi
Elfving, D. C., 4:l Korcak, R. F., 9:133
444
CUMULATIVE CONTRIBUTOR INDEX 445
Krezdorn, A. H., 1:vii San Antonio, J . P., 6:85

Lakso, A. N., 7:301 Saure, M. C., 7:239
Larsen, R. P., 9:xi Schneider, G. W., 3:315
Larson, R. A,, 7:399 Schuster, M. L., 3:28
Li, P. H . , 6:373 Scorza, R., 4:106
Litz, R. E., 7:157 Scott, J. W., 6:25
Lockard, R. G., 3:315 Sharp, W. R., 2:268; 3:214
Loescher, W. H., 6:198 Shear, C. B., 2:142
Lorenz, 0. A , , 1:79 Sheehan, T. J., 5:279
Maraffa, S. B., 2:268 Smock, R. M., 1:301
Marini, R. P., 9:351 Sommer, N. F., 3:412
Marlow, G. C., 6:189 Sondahl, M. R., 2:268
Maronek, D. M., 3:172 Soule, J., 4:247
Mayak, S., 1:204; 3:59 Sparks, D., 8:217
Maynard, D. N., 1:79 Splittstoesser, W. E . , 6:25
Mika, A , , 8:339 Srinivasan, C., 7:157
Mills, H. A., 2:411; 9:103 Stevens, M. A , , 4:vii
Molnar, J. M., 9:l Styer, D. J., 5:221
Monk, G. J., 9:l Swietlik, D., 6:287
Monselise, S. P., 4:128 Syvertsen, J. P., 7:301
Moore, G . A , , 7:157 Tibbitts, T. W., 4:49
Mor, Y., 9:53 Tisserat, B., 1:l
Murashige, T., 1 : l Titus, J. S., 4:204
Neilsen, G. H., 9:377 Webster, B. D., 1:172
Niemiera, A. X., 9:75 Weichmann, J., 8:101
Ogden, R. J . , 9:103 Wetzstein, H . Y., 8:217
O’Hair, S. K., 8:43 Whitaker, T. W., 2:164
Ormrod, D. P., 8:l White, J. W., 1:141
Pellett, H. M., 3:144 Williams, M. W., 1:270
Pokorny, F. A., 9:103 Wittwer, S. H., 6:xi
Poole, R. T., 5:317; 6:119 Wright, R. D., 9:75
Porter, M. A., 7:345 Wutscher, H. K., 1:237
Proctor, J . T. A., 9:187 Yadava, U. L., 2:l
Richards, D., 5:127 Yelenosky, G . , 7:201
Ryder, E. J . , 2:164; 3:vii Zieslin, N., 9:53
Sakai, A , , 6:357 Zimmerman, R. H., 5:vii; 9:273
Salisbury, F. B., 4:66

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HORTICULTURAL REVIEWS

Editorial Board, Volume 9

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Van Nostrand Reinhold

2 Plant Growth Regulators in Rose Plants

3 Nutrition of Container-Grown Woody Nursery Crops

VII. Timing of Nutrient Applications 90

4 Elemental Status of Pine Bark-Based Potting Media

5 Iron Deficiency Chlorosis

6 Ginseng: Industry, Botany, and Culture

7 The Honey Bee Pollination Component of Horticultural

IV. Concluding Remarks 259

8 Tissue Culture of Temperate Fruit and Nut n e e s

9 Summer Pruning of Apple and Peach n e e s

10 Orchard Floor Vegetation Management

W. G. BAILEY. Department of Geography, Simon Fraser University,

G. H. NEILSEN. Agriculture Canada, Research Station, Summerland,

'Present address: Division of Agriculture, Arizona State University, Tempe, Ari-

lack thereof, when trying to study vegetative efficiency of a fruit tree

directly with beauty in the fruits, vegetables, flowers, and landscapes of

In temperate climates, controlled-environment horticulture has become

In response, researchers and commercial greenhouse operators were

11. STRUCTURAL MODIFICATIONS

UV-resistant ), flat fiberglass (25-mil), flat fiberglass (premium-grade,

(Durolite f ) materials transmitted 76-8270, while rigid PVC (Novaroof)

energy savings. Goldsberry found that fiberglass-reinforcedplastic ( FRP )

repeated by extending polyethylene across the house at gutter height,

its unsurpassed light transmittance. In the United States, double-film

between two layers of polyethylene in the greenhouse roof at night and

spaces. Hay and Goldsberry ( 1978)conducted a two-yeartest with single-

( N P V )economic analysis to nighttime heat load reduction data for three

111. ALTERNATIVE ENERGY SOURCES

mented thermal mass, expanded daily temperature ranges, and a highly

Table 1.1. Total Tkansmission Factor: Short-Term Simulation Results

Solar shed Control house

Week Experimental Simulatedb Experimental Simulatedb

Oct 8-14 1.58 1.62 (1.73) 1.11 1.18 (1.26)

=From Lau et al. ( 1984).

Rotz et al. (1979) undertook a computer simulation study of energy

Table 1.2. Computer-Simulated Energy Use in Heating a 4000-m2 Commercial

Glasshouse Double polyethylene

Fuel Energy Energy

Insulation and solar heat

"Glass is replaced by double acrylic material.

2000 4000 6000 8000 Collector area ( 1 1 2 )

Percentage of total energy supplied

Edward Island. In Israel, a plastic greenhouse was built over an artificial

polyethylene shed-type rock storage greenhouse that was not equipped

Inlet dUC1 Charginp Inlet ducl Oischarginy

water distillation as well. Daily production of fresh water in excess of 2

screens to reduce heat loads by 30% or more (Manning et al. 1981).The

heated greenhouses maintained a t 18'C daytime and 16'C nighttime

Waste heat can be obtained from nonindustrial sources. In West Germany,

small-diameter pipes lying on the surface of the soil. Energy savings of

IV. ENVIRONMENTAL MANAGEMENT

A. Root Zone Heating

the minimum nighttime air temperature was lowered below normally

In bedding plant trays, media temperature varied as much as 10°C,

B. Alternative Heating Methods

Many of these devices can be installed in a greenhouse range leading to