A M . ZOOLOCIST, 8:31-42 18C8).

Aquatic Primary Production

CHARLES R. GOLDMAN

Department of Zoology and Institute of Ecology, University of California, Davis

95616
SYNOPSIS. The ecosystem concept has been particularly useful and extensively employed in the
study of aquatic primary productivity. The flow of energy through the system is an attractive
area of investigation when it involves some process, but has a more restricted value when units
of biomass are simply converted to calories. Although we are able to measure primary produc-

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tivity in terms of the carbon fixed, we are not yet able to measure the actual change in the
oxidative state of the newly fixed carbon. The fate of photosynthate as food for higher trophic
levels is therefore dependent upon a considerable array of biological and environmental varia-
bles. Primary productivity is considered in terms of its evolution from measures of standing crop
and yield, which have been gradually replaced by measures of rate of carbon uptake or oxygen
production, or by measure of nutrient loss, or by change of COa in the environment. Data from
five lakes are used to illustrate the evolutionary thread of eutrophication and the great range in
primary productivity to be expected on the basis of either unit volume or unit surface area at
different trophic states. Light and nutrients are important in limiting primary productivity, and
are contributing factors to the great variability which one may encounter within a given lake.
Only with a sounder understanding of productivity at the base of the food-chain can we have
any real hope of controlling the productivity of aquatic environments for the benefit of man.

The concept of "primary productivity" ment must be studied concomitantly with

is one aspect of the more general concept the biological. This was a major step for-
of the "ecosystem." Judging from the his- ward, but with the spectacular availability
torical development of ecological theory, of new research tools and methods for
ecosystems as conceptual divisions have measuring vast arrays of biological and
been most easily studied in the form of physical parameters, the researcher now
standing bodies of water. Eighty years ago finds himself in a new and sometimes seri-
Forbes (1887) published his classic "The ous dilemma. What is to be measured, why
Lake as a Microcosm," in which he dis- should it be measured, and what is it likely
played remarkable insight into the inter- to add to our understanding of the ecosys-
dependence of the plants and animals in a tem? In the following I give, and some-
lake. If he appeared less aware of the great times illustrate, my impressions of the di-
importance to his microcosm of such abiotic rection that primary productivity studies
factors as erosion and domestic pollution, have taken, what progress has been made
it is probable that in 1887 they were con- along the way, and what some of the op-
siderably less visible than they are today. portunities are for gaining a better under-
The ecosystem concept continued to devel- standing of the first level of the aquatic
op through the work of other aquatic biolo- food-chain.
gists such as Thienemann (1925), and The object of conducting a study of the
eventually gained maturity in papers by primary productivity of a particular aquat-
Hutchinson (1941) and Lindeman (1942). ic environment is to obtain quantitative
The last author considered productivity information about the amount of energy
the focal point of activity within his dy- available to support what Ohle (1956) has
namic ecosystem. called the "bioactivity" of the system. This
One result of this historical development is considered to be the system's capacity
is that the productivities of aquatic systems for the formation of potential energy and
have been more intensively examined and the subsequent reconversion of this to ki-
more carefully measured than terrestrial netic energy on a unit volume or area basis.
ones. A further result was the realization In terms of Lindeman's trophic-dynamic
that the physical and chemical environ- model, one measures productivity in order
31
32 CHARLES R. GOLDMAN

to compute efficiency of energy transfer ly, we are barely in a position to list some
within the system. In this kind of analysis, of the factors which may be significant.
some value derived from the measured pri- Among these factors are those relative to
mary productivity is used to express the the immediate history of the organisms, the
input of energy into the herbivore level. past and present condition of the environ-
Lindeman (1942) developed a model in ment, and the trend of growth in both
which he introduced the concept of energy populations. Not only do the primary pro-
flow, illustrating it with data collected in ducers themselves have rather different
natural ecosystems. It was consistent with abilities to utilize the available light en-
his purpose to express the data as energy ergy, but the consumers of this trophic level

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units (calories). Subsequent workers have are not bomb calorimeters and have very
continued to probe the concept of energy different abilities to utilize the calories that
flow. However, advances in biochemistry, they consume.
physiology, and ecology require an elabora- Disregarding for the moment this prob-
tion of the model of energy flow to the lem of how best to present data, let us con-
point where data relating it to the natural sider how much of primary production is
world are extremely difficult to acquire. available to the rest of the food-chain. Per-
Energy is interesting to talk about when haps the best way to answer this is to con-
the discussion is referred to some process. sider how much is not available. The com-
In an imaginary world in which the func- munity of phytoplankton has a definite
tional units are discrete trophic levels, it is respiratory need which obviously must be
not only possible but stimulating to de- met before any energy can be passed on to
scribe the flow of energy through the sys- higher trophic levels. The magnitude of
tem. But when the functional units of the this need is impossible to determine with
system being investigated are conceived of samples of natural phytoplankton because
as macromolecules there seems to be only of the presence of bacteria. However, a
limited merit in translating what amount minimum figure of 10% seems acceptable
to rate measurements of biomass accumu- from studies using the oxygen light-and-
lation into energy units. dark bottle (e.g., Patten, 1966). In such a
As an illustration, the widely used C14 study the difference between gross produc-
method for estimating primary productiv- tivity and net productivity represents the
ity measures the physical incorporation of rate of community respiration, and ranges
carbon atoms into living cells. Neither this from 10% of gross productivity to several
method nor any other practical technique hundred percent. An appreciable part of
adequately measures the change in oxida- this respiration is due to phytoplankton,
tive state of the carbon fixed. Moreover, and there seems to be no reason to make
the subsequent ecological role of newly the assumption that algal respiration re-
fixed carbon is even less amenable to quan- mains at some fixed percentage of gross
titative study. Photosynthate may be used photosynthesis while community respira-
in numerous ways by the photosynthetic tion fluctuates widely in response to chang-
organism. The subsequent ecological role ing conditions. Bunt (1965) measured oxy-
of the photosynthate depends on which of gen consumption of bacteria-free algal cul-
these possibilities is realized. Even if we tures with a mass spectrometer and re-
could specify completely the fate of newly corded values between 20 and 100% of
fixed carbon in producer organisms, we gross photosynthesis. The results were in-
would be unable to describe its transfer to fluenced by quality of the light and tem-
heterotrophic organisms in any but the perature. The number of factors likely to
most general terms. In particular, we are influence the ratio of algal respiration to
still not in a position to describe the energy photosynthesis is large and any figure arbi-
conversions which take place when a popu- trarily taken to represent this ratio is not
lation of natural zooplankton grazes a natu- apt to be very reliable.
ral population of phytoplankton. Present- Besides requiring a portion of their auto-
 AQUATIC PRIMARY PRODUCTION

trophic production for respiration, phyto- of energy from phytoplankton to herbi-

plankton communities must also reserve a vorous zooplankton, and there is evidence
portion for the maintenance of community that the amount of energy involved, may
structure. This "information cost" has sometimes be of the same order of magni-
never been measured directly but there is tude as that recovered in paniculate form
some indirect evidence that it must be paid. in routine C14 productivity studies (Fogg
When an aquatic system is altered with the and Watt, 1965).
aim of increasing the production of fish, Another factor that should be considered
zooplankton and fish production may in- in studies of energy flow is the role of
crease disproportionately more than does allochthonous material in supporting the

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the production of phytoplankton (McCon- energy requirements of consumer organ-
nell, 1965; Goldman, 1967). A possible in- isms. This material is included in Ohle's
terpretation of this observation is that a (1956) definition of bioactivity. No natural
large amount of primary production is un- aquatic ecosystem is entirely closed. Poten-
available to grazers but remains with the tial energy enters in the form of organic
phytoplankton community as "informa- solutes and debris. Parsons and Strickland
tion" necessary for the maintenance or de- (1962) and Sorokin (1965) have discussed
velopment of community structure. Grazers the role of bacteria in incorporating or-
remove production which is in excess of ganic solutes into paniculate matter. More-
this threshold level. If the magnitude of over, Sorokin has shown that this type of
the information cost is high relative to pri- production of paniculate matter can be the
mary production and is not itself a fixed most important as far as producing food
percentage of primary production, then a for filter-feeding crustaceans is concerned.
small increase in the rate of growth of the Organic matter, both paniculate and dis-
phytoplankton will provide a relatively solved, may also appear in the water as a
larger increase in the food supply of grazers result of chemosynthesis. This mode of
and eventually the fish which consume formation should be considered primary
them. production, but is not usually measured.
There are additional difficulties that Its magnitude is probably not great in
must be met in the course of fitting meas- many systems, but Sorokin has found it to
urements of primary productivity to the be very important in the Rybinsk reservoir
trophic-dynamic model. It is now known and in the Black Sea.
that a highly variable, yet often significant,
portion of the photosynthate produced dur- EVALUATION OF METHODS
ing a C14 light-and-dark-bottle experiment Before continuing with a discussion of
is not retained by the producer organisms, primary productivity and succession in
but moves into the environment in soluble lakes it is important to turn our attention
form (Fogg and Watt, 1965; Watt, 1966; to methods of measuring primary produc-
Hellebust, 1965). It is impossible to meas- tivity, for much of the current research in
ure the absolute magnitude of such excre- aquatic primary productivity is still di-
tion by a community of natural plankton rected toward achieving more accurate
because the excreta can serve as a substrate measurements or toward greater under-
for bacterial growth (Hobbie and Wright, standing of the methods already in use.
1965) and thus find their way back to par- Methods of measuring primary productiv-
ticulate or inorganic form during the incu- ity are reviewed and discussed in greater
bation period. To be sure, the material so detail in Ryther (1956), Odum (1956),
formed is rightly considered primary pro- Strickland (1960), Doty (1963), Goldman
duction, and it ultimately serves as a (1963), and Strickland and Parsons (1965).
source of energy for organisms at the higher Although some methods of measuring pri-
trophic levels. The pathway along which mary productivity have preceded the great
this energy flows, however, is quite different development in instrumentation in the last
from that usually modeled for the transfer two decades, they have been greatly acceler-
34 CHARLES R. GOLDMAN

ated by the use of modern instruments, method is of limited use because changes
radioactive isotopes, and computers for in standing crop of phytoplankton reflect
analysis of data. only the net effect of many biological and
physical events. For example, standing
Standing Crop and Yield crop may be greatly diminished by preda-
The standing-crop method has been used tion and water movements, while at the
by aquatic biologists for the longest time same time photosynthetic rates may remain
and is still used for estimating the produc- high. The measurement of plant pigments,
tivity of rooted plants (Westlake, 1965), also a standing-crop measurement, is fre-
quently used (Richards, 1952). Some prob-

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and for periphyton growth on slides. Pri-
mary productivity is the conversion of in- lems in the estimation of chlorophyll a are
organic matter into organic matter. Stand- discussed by Tailing and Driver (1963).
ing crop is that part of the production
UPTAKE OF NUTRIENTS
which is physically present in the system,
and does not include what is lost in respira- An early attempt at measuring the rate
tion. Findenegg (1965) has made useful of production in aquatic ecosystems was
comparisons between the standing crop and made by measuring the materials removed
productivity. Biologists in the early part in a given aquatic system and calculating
of this century were, for the most part, how much biological production would be
content to look at biological systems at required to absorb this amount. In the
rest. To set the whole scheme of biological North Sea, the decreases in CO2 and phos-
interactions in motion took several more phate were used by Atkins (1922, 1923),
decades. Preoccupation with measuring and Cooper (1938) considered decreases in
standing crop dominated the interests of oxygen and nitrate. Steele (1956) has esti-
both freshwater and marine scientists dur- mated the annual cycle of plant produc-
ing the long period when nets were yield- tion in a northern portion of the North
ing a myriad of exciting new plankton or- Sea (Fladen Ground) by considering
ganisms to be described. Standing-crop changes in the inorganic phosphate in re-
measures were most frequently reported in lation to vertical mixing of the water mass.
terms of numbers of organisms. Recogni- Phosphorus is considered by many to be a
tion of the lack of precision in equating particularly difficult element to use because
large phytoplankters to smaller forms led it is often taken up and stored by organ-
to the more precise practice of converting isms well in excess of their requirements
counts to biomass or to volumes, and even- for optimum growth (Gerloff and Skoog,
tually thought was given to production in 1954). These methods were, of course, in-
terms of yield of organisms. This intro- direct and were influenced to an uncertain
duced the time factor, and fisheries biolo- extent by the interactions of both plants
gists, like farmers, expressed yields in terms and animals. Still they were of real impor-
of tons of fish produced per acre of water tance in directing the field towards attempt-
surface per year. ing to achieve more precise measurements
Productivity is defined as the rate at of a dynamic system.
which biological production occurs. Thus Methods of Measuring Oxygen and COt
an operational definition of primary pro-
ductivity is the rate at which photosyn- In moderately to highly productive aquat-
thesis occurs. An early attempt at estimat- ic environments it is feasible to measure
ing phytoplankton productivity was made short-term fluctuations in dissolved oxygen
by Lohman (1908) from changes in the directly in the environment. As a first ap-
standing crop. Estimates of phytoplankton proximation, it is assumed that as each
productivity are still made today from peri- mole of carbon dioxide is reduced in pho-
odic measurements of standing crop {e.g., tosynthesis a mole of oxygen is released
Maciolek and Kennedy, 1964), but the into the water. Thus, if suitable correc-
 AQUATIC PRIMARY PRODUCTION 35

tions can be made for the diffusion of oxy- enough for measurable changes to occur,
gen across the air-water interface this meth- but not long enough for depletion of nu-
od can be used to estimate the net rate at trients or bacterial growth on the bottle
which the community is incorporating CO2, surface. Respiration can be estimated by
without enclosing it in a bottle (Odum, pairing each "light bottle" sample with a
1956). If measurements are made at regu- "dark bottle" from which light is excluded.
lar intervals over a 24-hr period, the aver- Putter (1924) and Gaarder and Gran
age hourly decrease in oxygen during (1927) were apparently the first to use this
hours of darkness can be determined. It is method. They recorded the oxygen in the
then assumed that respiration removes this initial sample and then, after incubation,

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amount of oxygen each hour throughout the change in oxygen that occurred in the
the day, making it possible to estimate the light and the dark bottles. From this they
gross rate at which the community incor- calculated net and gross productivity.
porates COo. There is an analogous meth- In methods dependent on measurements
od in which fluctuations in CO2 are re- of oxygen one has to make some assump-
corded. tion about the photosynthetic quotient
The pH meter, which was thought for a (moles O2 liberated/moles CO2 incorpo-
while to be the panacea for the afflictions rated). For the photosynthesis of carbo-
of aquatic biologists, was quickly brought hydrate the ratio is unity. For the synthesis
to bear on the problem of measuring pho- of an algal cell, however, the expected ratio
tosynthesis. As carbon dioxide is removed is higher, and presumably varies with the
from an aquatic system during photosyn- physiological state of the algae and the nu-
thesis, the pH rises. This shift has been trients available (Myers and Cramer, 1948;
used to estimate both photosynthesis and Ryther, 1956).
respiration. The sea and some fresh waters Oxygen methods in general have rather
are too buffered against changes in pH to poor sensitivity and are of no use if the
make this method useful in all environ- gross incorporation of inorganic carbon
ments, but it has been employed with suc- during the test period is less than about
cess in lakes (Verduin, 1956), and for con- 20 mgC/m3. In many of the less produc-
tinuously monitoring the growth of cul- tive aquatic environments several days may
tures (Beyers, 1965). CO2 may also be meas- be required for this much photosynthesis to
ured by standard volumetric or gasometric occur. During this time bacteria may de-
techniques. velop on the insides of the container, in-
Although CO2 and O2 can be measured validating the results.
with relative precision, the overall preci- Photosynthetic rates can also be meas-
sion of productivity measurements made by ured in light and dark bottles by determin-
these techniques is not generally great be- ing the amount of radioactive carbon fixed
cause of uncertainties in the corrections for in paniculate form after a short incubation
diffusion, water movements, or extended with Na2C14O3 (Steemann Nielsen, 1952).
enclosure time. Some of the oxygen pro- Sensitivities with this method are much
duced may not be immediately released greater, and much shorter periods of incu-
from higher aquatic plants, thus causing a bation are possible. By adjusting the speci-
lag period in the evolution of oxygen. The fic activity of the Na2C14O3 inoculums it is
big advantage of this method relative to possible to obtain easily measurable
the more sensitive C14 method is the esti- amounts of C14 in particulate form after
mate one gets of community respiration. only 2 hr. Unlike the oxygen method, how-
ever, the result with the dark bottle does
Light-and-Dark-Bottle Methods not provide an estimate of community res-
Some of the uncertainties just mentioned piration and for this reason the C14 meth-
can be reduced by making the measure- od gives the ecologist somewhat less to
ments on samples which have been en- work with.
closed in glass bottles for a time just long Because the C14 method is so sensitive
36 CHARLES R. GOLDMAN

and rapid, it has been widely used. One ities will be less than the actual rates.
outcome of its popularity has been a great These comments on methods have been
deal of scrutiny of the method itself. After presented as a caution to those ecologists
15 years of use, however, it is still not clear who might be tempted to rely too heavily
whether the C14 method estimates gross on published values of primary productiv-
productivity, net productivity, or some- ity in generalizing about the transfer of
thing in between. One of the latest papers energy through aquatic food-webs. In this
on the subject (Bunt, 1965) concludes that regard, it is significant that those of us who
the results most closely estimate net pro- are measuring primary productivity are
ductivity, but also suggests that the prob- still spending a disproportionate amount

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lem may only be resolvable in terms of a of time on simply finding out more pre-
particular set of experimental conditions. cisely what is being measured.
There are other uncertainties associated
PRIMARY PRODUCTIVITY AND SUCCESSION
with the C14 method already mentioned.
IN LAKES
There is now ample evidence that a por-
tion of the C u fixed during incubation Having briefly discussed methods of
may seep out of the algal cells in the form measurement, I shall consider succession
of water-soluble organic compounds (Fogg in five lakes which exemplify major differ-
and Watt, 1965). This material is part of ences in primary productivity and trophic
the primary production although it follows state. The concept of eutrophic and oligo-
a different path through the food-chain, trophic types of lakes is not a new one. It
presumably being utilized by bacteria was used by Naumann (1919) to indicate
(Hobbie and Wright, 1965). The amount the difference between the more productive
of photosynthate liberated extracellularly lakes of the cultivated lowlands and the
is large enough to be measured with pre- less productive mountain lakes. The troph-
cision and a number of workers, including ic state of five different aquatic environ-
the author, are now routinely including ments is summarized in Figure 1. It should
quantitative studies of extracellular prod-
ucts of photosynthesis as part of the meas-
urements of primary productivity.
A serious technical problem associated
with C14 concerns the calibration of radio-
active sources and of instruments used for
measuring radioactivity. In order to calcu-
late productivity in terms of carbon uptake
it is necessary to know accurately the
amount of C14 initially in the sample, the
amount of C14 added in microcuries, and
the number of microcuries recovered in
participate form by filtering the sample
through a membrane filter. Steemann Niel-
sen (1966) and Goldman (1963, 1968) de-
scribe different approaches to solving these
problems.
Arthur and Rigler (1967) confirm the
warning of Guillard and Wangersky (1958)
that another possible source of error exists rhgo rreso ej dys

in the C14 method. These authors found TROPHIC STATE

evidence that under certain conditions FIG. 1. Profiles of the change in photosynthesis
with depth in five lakes during summer. The gen-
phytoplankton cells are damaged during eral trophic state of these lakes is also indicated in
the filtration, and if steps are not taken to relation to their relative carbon assimilation per
circumvent this loss, calculated productiv- unit of surface area.
 AQUATIC PRIMARY PRODUCTION 17

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HG. 2. Permanently frozen Lake Vanda in the Wright dry Valley, Antarctica (77°32'S lat.
161°33'E long.). Previous shorelines are evident along the right side of the lake. Official U. S.
Navy photograph.

be kept in mind that the general progres- world (Fig. 2). The lake is permanently
sion from an oligotrophic to eutrophic and sealed under 3 to 4 m of very clear ice,
finally to a dystrophic lake is as much a which transmits 14 to 20% of the incident
result of the original basin shape, climate, radiation to the water below. This pro-
and edaphic factors, as it is of geologic age. vides enough light to power the photosyn-
It is unlikely that some shallow lakes ever thesis of a sparse phytoplankton popula-
passed through a stage that could be con- tion to a depth of 60 m (Goldman, et ah,
sidered oligotrophic, and it is just as un- 1967). Lake Vanda can be classified as
likely that the first lake to be considered ultraoligotrophic, since its mean produc-
here (Vanda) will ever become eutrophic. tivity is only about 1 mgC/m 2 /hr (Fig. 1).
Lake Vanda, located in "dry" Wright Lake Tahoe in the Sierra Xevada of
Valley near McMurdo Sound in Antarctica, California and Xevada is an alpine lake
is one of the least productive lakes in the long esteemed for its remarkable clarity.
 CHARLES R. GOLDMAN

depletion in oxygen in its deep water dur-

ing summer stratification and also under
ice cover during late winter.
Clear Lake is an extremely eu trophic
shallow lake, with periodic blooms of a
bluegreen alga (Aphanizomenon) and in-
organic turbidity greatly reducing the
transparency of the water. The photosyn-
thetic zone is thus limited to the upper 4
m (Fig. 1), with a high intensity of pro-
ductivity per unit volume yielding an aver-

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age of about 300 mgC/m 2 /hr during the
FIG. 3. Castle Lake as viewed from the west rim. growing season. Because Clear Lake is
This cirque lake with a surface area of 19.5 hec- shallow, it does not stratify for more than
tares is located in Siskiyou County, California. a few hours at a time during the summer,
T.39N.,R.5W.,S13 at an elevation of 5600 feet.
and the phytoplankton which sinks below
the light zone is continuously returned to
Although it is more productive than Lake it by mixing.
Vanda, it is still oligotrophic. The lake is Cedar Lake lies near Castle Lake in the
characterized by a deep euphotic zone, with Klamath Mountains (Fig. 4). Its shallow
photosynthesis occurring in the phytoplank- basin is nearly filled with sediment and it
ton (Fig. 1) and attached plants to a depth is nearing the end of its existence as a lake.
of about 100 m. Although the production Numerous scars of similar lakes found in
under a unit of surface area is not small, the area presage Cedar Lake's future. Ter-
the intensity of productivity per unit of restrial plants are already invading the
volume is extremely low. Lake Tahoe's low lake, and higher aquatic plants reach the
fertility (as inferred from its productivity surface in many places. The photosyn-
per unit volume) is the result of a restricted thesis beneath a unit of surface area
watershed, whose granitic rocks provide a amounts to only about 6.0 mgC/m 2 /hr
minimum of nutrient salts. This situation during the growing season as the lake is
is rapidly being altered by human activity now only about 4 m in depth (Fig. 1).
in the Tahoe basin. The cultural eutrophi- Cedar Lake may be considered a dystrophic
cation of the lake is accelerated by sewage lake. Some lakes of this type pass to a bog
disposal in the basin and by the exposure condition before extinction. In others,
of mineral soils through road building and their shallow basins may go completely dry
other construction. Since Lake Tahoe's
water is saturated with oxygen all the way
down, the decomposition of dead plankton
sinking slowly towards the bottom is essen-
tially complete. This means that nutrients
are returned to the system and because of
a water retention time of over 600 years the
increase in fertility will be cumulative.
Castle Lake, located at an elevation of
5600 feet in the Klamath Mountains of
northern California, shows some of the
characteristics of Lake Tahoe as well as
those of more productive environments
(Fig. 3). It is, therefore, best classified as
mesotrophic. Although the lake has a mean FIG. 4. Cedar Lake, Siskiyou County, California.
The lake has a maximum depth of just under 4 m
productivity of about 70 mgC/m 2 /hr (Fig. and the invasion of the lake basin by rooted plants
1) during the growing season, it shows a is well underway.
 AQUATIC PRIMARY PRODUCTION 39

of photosynthesis per hour are all based on

4-hr experiments made at mid-day during
AUGUST 17, 1957
summer. Had the experiments been per-
0 <
WEATHER CLOUDY
formed during a different time of day, the
WITH RAIN
results would have appeared quite differ-
ent. A diurnal study of photosynthesis
with depth is therefore particularly in-
structive because it reflects this variation
in photosynthesis as the light pattern
changes throughout the day (Fig. 6). If

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02 04 06 08 10 12 14 16 18 20 22
z HOURS
photosynthesis-depth curves such as those
s shown in Figure 6 are integrated, we can
or plot the diurnal course of photosynthesis
beneath a unit of surface area. This has
been done for six diurnal studies at Castle
UJ Lake, California (Fig. 7). The light curve
5.0 AUGUST 22,1957 is included to show the very close relation-
WEATHER CLEAR, ship between photosynthesis and light dur-
BRIGHT ing the months of June through September.
2.5 The consistent symmetry of these curves
enables an investigator to make rather ac-
curate conversions of short term experi-
ments to a day's productivity on the basis
04 06 08 10 12 14 16 J8 20 22 of measured light.
HOURS Theoretical curves have been developed
FIG. 5. Average phatosynthetic carbon assimilation by Tailing (1965), Vollenweider (1965),
with variation dn solar radiation at Brooks Lake,
Alaska. The primary productivity on a cloudy day
is compared with' that occurring on a bright day.
Values are from the average o£ two stations run
simultaneously. (Goldman, 1960a)

during summer, and their flora and fauna

become those of vernal ponds.
In examining some aspects of the pro-
ductivity of these five lakes, the variation
in both the intensity of photosynthesis and
the depth to which it occurs is evident (Fig.
1). The great importance of the total avail-
able light can scarcely be overemphasized.
This was first made apparent to the author
during studies of primary productivity and
limiting factors in three oligotrophic lakes
of the Alaskan Peninsula, where weather
conditions imposed severe light limitations 0.0 2.0 4.0 0.0 2.0 4.0
on the phytoplankton productivity (Gold- Mg C • rrr'-hour" 1
man, 1960a). The average photosynthesis CASTLE LAKE DIURNAL
(mgC/m3/day) on both a cloudy and a Vlll-l-61
bright day was essentially proportional to FIG. 6. Diurnal photosynthesis-depth curves at a
the available light energy (Fig. 5). single station in Castle Lake, California. Measure-
ments were made under different light conditions
In Figure 1, photosynthesis with depth occurring during the four time intervals noted on
has been presented for five lakes of greatly 1 August 1961. For the light curve for this day see
differing trophic character. These values Figure 7.
40 CHARLES R. GOLDMAN

response, with nitrogen of particular im-

portance. Trace elements, either singly or
in combination, have been found to stimu-
late photosynthesis in a variety of lakes. In
general, some component of the phyto-
plankton will respond positively to almost
any addition of nutrients, but the com-
munity as a whole will tend to share some
common deficiencies. Justus Von Liebig
did not intend to apply his law of the mini-

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mum as rigidly as some have interpreted
it, and we can best envision limitation of
nutrients from the standpoint of the bal-
ance and interactions of the whole nutri-
ent medium with the community of organ-
isms present at any given time. Much
about the nutrient requirements of phyto-
0400 0800 1200 1600 2000 0400 0600 1200 1600 2000
plankton can be gleaned from the excel-
TIME (HOURS) lent treatise of Hutchinson (1967).
FIG. 7. Integral diurnal productivity curves (brok- It must be borne in mind that the pri-
en lines) and photosynthetically active radiation
(solid lines) based on measurements made at Castle mary productivity of a given lake may vary
Lake, California. All curves were developed from greatly from place to place, and measure-
the integration of vertical productivity profiles such ments made at any one location may not
as the one constructed for 1 August 1961 shown in provide a very good estimate for the lake as
Figure 6. a whole. This problem has been explored
and Rodhe (1965) to express the integral in reservoirs by Sorokin (1959), and in
photosynthesis. These give rather good Brooks Lake, Alaska, by the author. In the
estimates if there is not too great an asym- rather sterile Brooks Lake the productivity
metry in the distribution of the phyto- per unit volume and per unit of surface
plankton population and if the light curve area increased as one approached the major
decreases uniformly with depth. This pro- tributaries providing the lake's nutrients.
cedure has very limited usefulness, how- Variability in productivity beneath a
ever, in a lake like Clear Lake where the unit of surface area is particularly evident
percentage of light transmitted per meter in Lake Tahoe, where attached algae are
varies greatly in the euphotic zone (Fig. already becoming a nuisance in the shal-
8), or in a lake such as Naknek, Alaska, low water, and transparency is often mark-
which receives volcanic ash from tributary
streams and has a strongly asymmetrical 1/
_-^ •o

curve of photosynthesis with depth (Gold- as -

man, 1960a). <

Nutrient limiting factors have been re- 10

V
-

viewed by Lund (1965) and examined by S'5 -

—.
the author in a number of lakes (Gold-
DEPTH

man, 1960o, b; 1964; 1965). In Brooks - -~ " -

Lake, Alaska, a sequence of the most limit- 2.S -

ing factors ranged from magnesium in the /

spring, through nitrogen in the summer, to 30 -

\
phosphorus in the fall (Goldman, 1960a). 1 I t 1 t i 1 1
In Castle Lake, potassium, sulfur, and the 4 8 12 16 20
tTRMSUlSSIOK/IIETER
24 28 32 36 <

trace element, molybdenum, were found to FIG. 8. Variation in the percentage of light trans-
be the most limiting. In Lake Tahoe, iron mission with depth in Clear Lake, California. Val-
and nitrogen gave greatest photosynthetic ues computed for each successive depth interval.
 AQUATIC PRIMARY PRODUCTION 41

edly reduced near streams which drain dis- Beyers, R. J. 1965. The pattern of photosynthesis
turbed watersheds. In July, 1962, the pro- and respiration in laboratory microecosystems, p.
61-74. In C. R. Goldman, fed.], Primary produc-
ductivity of Lake Tahoe showed great in- tivity in aquatic environments. University of
crease near areas of high nutrient inflow California Press, Berkeley.
(Goldman and Carter, 1965). This condi- Cooper, L. H. N. 1938. Phosphate in the English
tion was even more evident in the summer Channel. J. Marine Biol. Assoc. U. K. 23:181-195.
of 1967 when Crystal Bay at the north end Doty, M. S. [ed.] 1963. Primary productivity meas-
of the lake, and the southern end of the urement, marine and freshwater. U. S. Atomic
Energy Commission Report TID 7633.
lake showed different periods of high pro- Findenegg, I. 1965. Relationship between standing
ductivity. This variability in productivity

Downloaded from https://academic.oup.com/icb/article/8/1/31/239570 by guest on 23 October 2024

crop and primary productivity, p. 271-289. In
may be influenced by discharge of sewage C. R. Goldman, [ed], Primary productivity in
and land disturbance. Were it not for the aquatic environments. University of California
great volume of the lake (155 km3), it Press, Berkeley.
Fogg, G. E., and W. D. Watt. 1965. The kinetics of
would already be showing more severe release of extracellular products of photosyn-
signs of eutrophication. thesis by phytoplankton, p. 175-186. In C. R.
In the foregoing, I have attempted to Goldman, [ed.], Primary productivity in aquatic
give my impressions of aquatic primary environments. University of California Press,
Berkeley.
productivity, treating the subject both as Fortes, S. A. 1887. The lake as a microcosm. Re-
a research task and as a body of informa- published 1925. Illinois Nat. Hist. Surv. Bull. 15:
tion to be interpreted. I believe that bio- 537-550.
logical productivity can no longer be con- ~ Gaarder, T., and H. H. Gran. 1927. Investigations
sidered a matter of simple academic inter- of the production of plankton in the Oslo Fjord.
J. Cons. Int. Explor. Mer 42:1-48.
est, but must be recognized as a problem Gerloff, G. C, and F. Skoog. 1954. Cell contents of
of unquestioned importance for survival. nitrogen and phosphorus as a measure of their
The productivity and harvest of most of availability for growth of Microcystis aeruginosa.
the world's terrestrial and aquatic environ- Ecology 35:348-353.
ments must be increased if the world popu- Goldman, C. R. 1960a. Primary productivity and
lation is to have any real hope of having limiting factors in three lakes of the Alaska
Peninsula. Ecol. Monographs 30:207-230.
enough to eat. This increase is not pos- Goldman, C. R. 19606. Molybdenum as a factor
sible unless we gain a much better under- limiting primary productivity in Castle Lake,
standing of both aquatic and terrestrial California. Science 132:1016-1017.
productivity. Only with a sounder under- "• Goldman, C. R. 1963. The measurement of primary
standing of the processes which control productivity and limiting factors in freshwater
with carbon-14, p. 103-113. In M. S. Doty, [ed],
productivity at the level of the primary Primary productivity measurement, marine and
producers can we have any real hope of freshwater. U. S. Atomic Energy Commission Re-
understanding the intricate pathways by port TID 7633.
which energy moves and biomass accumu- Goldman, C. R. 1964. Primary productivity and
lates in various links of the food-chain. micronutrient limiting factors in some North
American and New Zealand Lakes. Verh. Intern.
Verein. Limnol. 15:365-374.
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124. ments. University of California Press, Berkeley.
Atkins, W. R. G. 1922. Hydrogen ion concentration Goldman, C. R. 1967. Mol)bdenum as an essential
of sea water in its biological relation. J. Marine micronutrient and useful water mass-marker in
Biol. Assoc. U.K. 12:717-771. Castle Lake, California, p. 229-238. In H. L. Gol-
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and respiration in a marine diatom with the eliminating serious errors in the measurement of
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207:1373-1375. Mer 32 (1): (in press).
 42 CHARLES R. GOLDMAN

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some marine flagellates. Limnol. Oceanog. 10: using C". Fiz. Rast. 6:125-133.
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