THE

COMPOSITION
IN THE

OF PARTICULATE
WESTERN
NORTH

ORGANIC
ATLANTIC1

MATTER

David W. Menzel and John H. Ryther

Woods

Hole Oceanographic

Institution,

Woods

Hole,

Massachusetts

ABSTMCT

The composition of particulate

matter in the western North Atlantic is examined to dctcrmined relationships
bctwecn carbon, nitrogen, phosphorus, and chlorophyll.
The regrcssions of phosphorus or chlorophyll
on nitrogen or carbon, when extrapolated
toward the
origin, indicate appreciable but variable amounts of nitrogen and carbon in the absence of
phosphorus and chlorophyll,
The regression of phosphorus on chlorophyll,
on the other
These observations
indicate that chlorophyll
and
hand, has its intercept
at the origin,
phosphorus are decomposed or mincralizcd
at csscntially the same rate, while carbon and
nitrogen are more refractory.
The relationship
bctwccn these paramctcrs and the distribution of detritus is discussed.

decomposed or denatured following death

of the organisms (Yentsch 1962). PhosIt has become apparent that the microphorus also is quickly liberated from the
scopic particulate matter of the sea consists dead material
(Harvey
1960), whereas
of a mixture of living microorganisms and
nitrogen and carbon are both relatively
dead detrital material, the relative propormore refractory to decomposition.
tions of which may vary widely. The chcmiMethods for the collection of the particucal composition of both fractions appears to late matter, normally microfiltration,
prcEle highly variable, at least in terms of those elude the possibility of mechanical separaelements or compounds that have been inand dead fractions.
tion into living
vestigated in the laboratory. The combinaFluorcsccnt
microscopy
using acridine
tj.on of high light intensity and nutrient dcorange (Ferguson Wood 1955), which difpletion (as occurs in surface waters, particuferentially stains living and dead material,
larly in the tropics) produces algal cultures
can be used with some success, but the
that are relatively poor in chlorophyll, nitrotechnique dots not lend itself readily to
gen, and phosphorus. Low light intensities
rapid, quantitative
analysis.
and a rich supply of nutrients ( typical of
A method is needed in quantitative ecolconditions near the bottom of the cuphotic
ogy for the routine measurement or cstimazone or near the surface in tcmperatc or
tion of the biomass of living phytoplankton.
boreal winter conditions ) produce cultures
For the reasons stated above, simple chemiwith high contents of pigment, nitrogen,
cal analyses for any single clement or comand phosphorus. The reported variability
pouncl
cannot provide a sufficiently reliable
of these constituents may be as much as an
index
of
this parameter. The possibility
order of magnitude or more (K&chum and
remains that the measurement of several of
Redfield
1949; Kuenzler
and Ketchum
the chemical constituents of the particulate
1.962) .
matter from a given sample, or a series of
The composition of the nonliving organic
samples from a given homogeneous envimatter, or detritus, depends primarily upon
ronment,
and a study of their interrelationits age and the resulting stage of its dccomships may prove to be a more satisfactory
position or mineralization.
In the euphotic
approach to the problem. The present rezone, phytoplankton
pigments are quickly
-port shows the relationships between carbon, nitrogen, phosphorus, and chlorophyll
l Contribution
No. 1334 from the Woods Hole
Oceanographic
Institution,
supported
by U.S.
in the marine particulate matter collected
Atomic Energy Commission Contracts No. AT(30-1)
during two cruises of the Woods Hole
2646 and AT(30-1)1918
and Office of Naval ReOceanographic Institutions research vessel
starch Contract Nonr-2196.
179
INTRODUCTION

180

DAVlD

W.

MENZEL

AND

Chain. The first cruise was made in January 1962 and extended in a rhumb line
from 41 N lat, 7149 W long, to 37 N lat,
6848 W long (Chain Cruise 22), and the
second in April 1962 from 4230 N lat to
20 N lat along 65 W long (Chain Cruise
25).
Since, as will be shown later, detrital
levels vary considerably
in space, thus
influencing the apparent ratio of one element to another, it was necessary to attempt
corrections for these levels before insight
could be gained into the composition of
living plant cells themselves. The approach
used here is to plot regressions of one
element on another. This technique requires sufficient variability
in the various
parameters for the slope of the line to be
determined. Sufficient variation was present within individual
stations in April to
permit this approach with a degree of reliability.
However, the data from January
did not permit station-by-station
examination because the vast number of values were
consistently low. Only a number of higher
values scattered throughout the section permitted an estimate of the slope of the line.
Data from south of the Gulf Stream could
not be evaluated by either method since
the quantity of particulate matter was uniformly low and homogeneous. The following discussion, therefore, is confined to
stations north of the Gulf Stream. Except
where noted, all samples were taken from
the upper 100 m.
METHODS

Four liters of seawater were filtered

through Whatman ( GFC ) micropore glass
filters for the determination
of carbon,
nitrogen, and chlorophyll.
To determine
carbon, the pads were rinsed with isotonic
NazS04 solution to remove chlorides and
subsequently analyzed by a modification of
the method of Fox, Issacs, and Corcoran
The sensitivity
was increased
(1952).
somewhat by reducing the strength of the
oxidizing
agent to 4.5 g K2Cr207/liter
I12SOb and reducing the strength of the
titrant by one-half. This method does not
actually determine carbon, but, rather, total
oxidizable material.

JOHN

II.
I

h
i
\
*;

5-

4-

RYTHER
I

I*/

APRIL

9
2

/:

I?
P

3
c,
i-2

2-

%l @.
L

.
*

0:

/
,*,.*

/;/X

I,*/

x:
m

&/,&I

*X
I

X43

l,,

12

16

20

24

26

PARTICULATE
C
pg-of.
/liter

FIG. 1. The relation between particulate carbon

and nitrogen in the euphotic zone, January and
April 1962.

Two different methods were used for the

estimation of nitrogen, Samples from Chain
Cruise 22 were analyzed by a microKjcldahl method and those from Chain
Cruise 25 by the micro-Dumas
method
using the P. H. Coleman Companys automated nitrogen analyzer.
Particulate phosphorus samples were collected by filtering
1 liter of seawater
through 0.8-p membrane filters containing a barrier of MgC03 (Strickland and
Parsons 1960). The barrier was removed
and placed in a digestion tube to which was
added 1.5 ml concentrated H2S04. After
the HzS04 was refluxed for 1 hr, the
samples were cooled, their pH adjusted to
7.5 with I-N NaOH, and the phosphate
determined calorimetrically
by the method
of Murphy and Riley ( 1958).
Chlorophyll was measured by the method
of Richards with Thompson
(1952) as
modified by Creitz and Richards ( 1955).
RESULTS

AND

DISCUSSION

Carbon : nitrogen

relations

The relations between particulate carbon

and nitrogen during January and April 1962
show a difference in the slope of the regressions between the two seasons, indicating
C : N ratios (by atoms) of 12.5 : 1 in January and 5.3 : 1 in April ( Fig. 1). Part of
this divergence may have resulted from

PARTICULATE

ORGANIC

MATTER

IN

TIIE

WESTERN

NORTH

181

ATLANTIC

FIG, 3. The relation between particulate

carbon
and phosphorus in the euphotic zone, January and
April 1962.

where the nutritional demands of the phytoplankton may help to retain this balance.
However, in old water (e.g., below the
euphoticzone
to depths as great as 4,000
m), the C : N ratio may decrease to 2.5 : 1.
The vertical profiles of carbon and nitrogen in deep water illustrate this fact (Fig.
2A, B). Similar conclusions regarding the
relationships between carbon and nitrogen
in deep water may be derived from the
data of Parsons and Strickland ( 1962).
Carbon : phosphorus

FIG. 2. The vertical

distribution
of particulate
carbon, nitrogen, and C : N ratios at A) 38 N lat,
65 W long, and B) 36 N lat, 65 W long, in
April 1962.

different analytical techniques, although rcpeated tests of nitrogen recovery by the

micro-Kjeldahl method indicated a loss not
exceeding 20%. In both cases, however,
th.c intercept of the regression is at or near
the origin, suggesting that carbon and nitrogen probably occur in the same proportions
in all particulate matter sampled at the
same time. It follows that C : N ratios in
the nonliving fraction retained by the filter
are the same as in the living phytoplankton.
This hypothesis appears valid on the basis
of observations made in the euphotic zone,

relations

Phosphorus is rapidly remineralized

or
solubilizcd, as is evident from the fact that
none was present in particulate form below
200 m.
The regression of phosphorus on carbon
for the same group of samples as considered
above shows much greater scatter (Fig. 3),
the cause OFwhich will be discussed below.
However, a linear relationship is still apparent for each cruise. Another difference
is that the regression line does not pass
through the origin but shows an appreciable
amount of carbon with no phosphorus. This
substantiates the contention that phosphorus is more rapidly mineralized from dead
material than either nitrogen or carbon and
implies the existence of a detrital residue
containing C and N but not P.
While the regressions of P on C for all
the April stations show a high degree of
scatter (Fig. 3), this scatter is greatly reduced when each station is plotted independently (Fig. 4). Furthermore, the slope
of the regression for the April data obtained

DAVID

W.

MENZEL

AND

JOHN

I-I. RYTHER

PARTICUL ATE NITROGENS a - a f. /lifer

"0

12

16

20

24

28

2-4

PART/CU! ATE CARBON

P9 - at/
FIG. 6. The relation between particulate
carbon,
by individual
stations. For location, see Fig. 4.

within these environments, it retained the

same relative proportions of the four abovementioned major constituents.
The regressions of either phosphorus or
chlorophyll
on either carbon or nitrogen,
when extrapolated to the origin, indicate the
presence of appreciable
but variable
amounts of carbon and nitrogen in the absence of both chlorophyll and phosphorus.
However, the rcgrcssions of nitrogen on
carbon and of chlorophyll on phosphorus
both pass through the origin. In addition,
nitrogen and carbon, but not phosphorus or
chlorophyll,
are found in the particulate

nitrogen,

12

16

20

24

28

lifer

and chlorophyll

in April

1962, considered

matter below the euphotic zone. These

observations, plus evidence in the literature
dealing with culture experiments, lead to
the conclusion that chlorophyll and phosphorus are quickly decomposed, mineralizcd, or denatured following the death of
phytoplankton, leaving a detrital residue of
nitrogen and carbon which slowly changes
in relative proportion, with nitrogen being
somewhat more refractory than carbon.
The fact that the regressions of chlorophyll or phosphorus on carbon or nitrogen
extrapolate to positive values of carbon and
nitrogen with no phosphorus or chlorophyll

186

DAVID

W.

MENZEL

populations by pigment analyses. III. A note

on the use of Millipore
mcmbranc filters in
the estimation of plankton pigments.
J. Marinc Rcs., 14: 211-216.
FEIUXJS~N
WOOD, E. J.
1955.
Fluorescent
microscopy in marine biology.
J. Conseil, Conscil Perm. Intern. Exploration
Mcr, 21: 6-7.
Fox, D. L., J. D. ISSACS, AND E. F. CORCORAN.
1952. Marine lcptopcl, its recovery, mcasurcment, and distribution.
J. Marinc Res., 11:

29-46.
H. W.
1960. The chemistry
and fertility of sea waters.
Cambridge
University
Press, London.
240 p.
KETCHUM,
B. H., AND A. C. REDFIELD.
1949.
Some physical and chemical characteristics
of
algae grown in mass culture.
J. Cellular Comp.
Physiol., 33 : 281-300.
KUENZLER,
E. J., AND B. EI. KETCIIIJM.
1962.
Rate of phosphorus uptake by PhneodactyZum
tricornutum.
Biol. Bull., 123: 134-145.
MURPIIY,
J., AND J. P. RILEY.
1958. Single solution method for the determination
of soluble
HARVEY,

AND

JOHN

H. RYTHER

phosphate in sea water. J. Marine Biol. Assoc.

U.K., 37: 9-14.
PAHSONS, T. R., AND J. D. H. ST~UCKLAND.
1962.
Oceanic detritus.
Scicncc, 136: 313-314.
RICHARDS,
I?. A., WITII
T. G. THOMPSON.
1952.
The estimation and characterization
of plankton populations
by pigment analysts.
II. A
spectrophotometric
method for the estimation
of plankton pigments.
J. Marine Res., 11:
156-172.
STEELE, J. H., AND I. E. BAIRD.
1961. Relations
between primary production,
chlorophyll
and
Limnol. Occanog., 6: 68particulate carbon.
78.
STRICKLAND,
J. D. H., AND T. R. PARSONS.
1960.
A manual of seawater analysis.
Bull. Fisheries
Rcs. Board Can., 125: l-185.
Marine plankton, p, 771YENTSCH, C. S. 1962.
797. In R. A. Lcwin, [cd.], Physiology
and
biochemistry
of algae. Academic Press, New
York.
-,
AND R. F. VACCARO.
1958. Phytoplankton nitrogen in the oceans. Limnol. Oceanog.,
3 : 443-448.