Primary Production in Lakes

JACOB VERDUIN

Biology Department, Bowling Green State University, Bowling Green, Ohio

ABSTRACT

The mean photosynthetic rate of lake phytoplankton under optimal light (natural con-
ditions) is about 0.5 micromoles of 02 evolved or CO2 consumed per microliter of organisms
per hour; l-2 micromoles per milligram ash-free dry Ljceight per hour; and 0.2 micromoles
per microgram of chlorophyll per hour. Simple equations are presented for computing
photosynthetic yield per square meter of water surface. Computations for several lakes
yielded values mostly lying between 150-200 millimolcs per square meter per day. A year-
round study under completely natural conditions in western Lake Erie showed writer
yields of about 40 millimoles per square meter per day and summer maxima of about 300.
The annual curve followed the solar radiation curve closely. It is estimated that 28 days
are required for a gram of phytoplankton (ash-free dry weight) to produce an additional
gram of glucose available for reproduction; 80 per centJ of the photosynthate produced is
required to support phytoplankton respiration.

Three quantities, related to primary pro-
duction in lakes, are measured by aquatic*
biologists: (1) the volume of autotrophic
organisms, (2) the ash-free dry weight of
suspended particles (organic seston), and
(3) the concentration of chlorophyll-eac*h
of these expressed per unit volume of water.
These are measures of standing (arop, and
they represent roughly quantitative esti-
mates of the populat,ion of autotrophic or-
ganisms upon whi& primary producation is
based.
Some biologists have studied photosynthc-
sis under approximately natural cwndit,ions
and have related the ohscrvcd Oz-prodwtioll
or COZ-consumption lo one or mow of thcl
quantities named above. l“ron1 sucah data
it is possible to compute ~ncan photos?;n-
t)hetic yields per unit of standing crop. ‘I‘abl~
1 present’s a summary of suc*h yields w~m-
puted from the literat,urc. These data rrlprc-
sent total or true photosynthesis, i.e. the
experiments incaluded darkened hot tles to
take account of the C‘C)s cwntrihutccl by rw-
piration, or of the 02 so (Y~IISUII~C~. Riley’s
(I 941) data arc inc~ludcd lwur~~~ t holy per-
mit computation iI all t hrw c*atcgoriw.
and because they shop\- I hat the rates for
marine phytoplankton NO in gc~~ral agrw-
rnent with those for f’rwh IIX~W phytoplank-
t,on. The data in t)his tahlrl suggest that
t!hc mean photosynthctich ratct of a nlic~rolitcl
of aut atrophic organisms, under optimal
light’ (natural conditions) amounts to about
0.5 mic*romoles of 02 evolved or CO2 con-
sumed per hour (6 fig C). The rate per
milligram of ash-free dry weight is about 1
or %, ( 12-24 pg C) and the rate per micro-
grant of chlorophyll is about 0.2 micromoles
of ( J3 (2.4 pg C). These values represent
~~un~crous t,ests and can he regarded as
fairly reliable means, but results from in-
dividllal test,s vary widely. The data repre-
wit rlpilimnion collections. Manning and
.Juday ( 1941) state that hypolimnion samples
exhibit yields which are less than one-third
so high.
A’I smvey of the literature she\\-s that
there is a rough inverse correlation between
the dr@h of the epilimnion (or total depth
ill llnstratified lakes) and the standing crop
of auf obrophic plants (,Manning and Juday
19-l1, Rwnak 1919, Verduin 1954). Thus
lakw \vith an epilimnion layer of the order
of CHH~meter are likely t,o have standing
cbropx of t,hc order of 50 microlitcrs of plant
volu~nc: per liter of water, 15 milligrams ash-
t’rcc dry w4ght, and :30micrograms of chloro-
phyll. Ilakes having an epilimnion depth
of’ thrb order of 10 meters are likely to have
st atldillg (*raps about one-tenth so high.
rI’hc%~~ values arc intended to indicatNc
ordcr~ of magnitude only. Large \-:triations
iii stallding crop arc obsci*\-cd, ot’ ~~oi~rsc,
86 JACOB VERDUIX

TABLE 1.
Ph?ytoplanktorr photosy,l thesis undrI, rate in the euphotic zone to the rate under
optimal light optimal light. Estimates of standing crop
To indicate variation observed routinely among irl- are abundant in the literature. Light
dividual samples write X + 3 after eachvalue in the
table. To convert to micrograms COz multipI) penetration data are less abundant, but if
by 44, O2 by 32, carbohydrate by 30, carbon by 12. Secc*hi disc readings are included a con-
siderable amount of information is avail-
Micromoles CO% absorbed
or 02 evolved per able. To make Secchi disc readings approxi-
Reference mattlly convertible to euphotic zone depths,
plit/hr mg/dry- pg chloro-
wt/hr phyll/hr simultaneous determinaGons with the Secchi
disc&and submarine photometer were made
0.8 0.22 hlanning k Juday (1941) during August 1955 by the crews of two
0.22 Gessner (1949)
0.5 1.6 Verduin* (1956) vessels assigned to the synoptic survey of
2.0 *Jackson & McFadden Lake Erie. The data are plotted in Figure
(1954) 1. The line, drawn by inspection through
0.6 1.3 0.31 Riley* (1941) the scatter diagram, suggests that an ap-
* Average of year-round research under natural proximate estimate of the depth of euphotic
conditions. The other data represent summer con- zone can be obtained by multiplying Secchi
ditions. A11 are in tempcraie zone. disc readings by 5. The scatter in Figure 1
indicaat’es that the relationship is not pre-
caise. Different conversion factors appear
in t’he literature; Riley (1941) used a factor
of :<; from the data of Bursche (1955) a
factor of about 2.5 can be computed; Raw-
son (1950) suggests that the factor decreases
as transparency increases, and he lists a
factor of 4.3 when the Secchi disc reading is
about 1 meter.
The ratio of mean photosynthesis in the
euphot)ica zone to the rate under optimal
light is about 0.65. This value has been
determined from numerous experiments in
western Lake Erie (Verduin 1956) and can
he (lorived from Figure 12 of Manning and
,Juday (1941).
The following equations are now pre-
s 0 seni ~1 for computing approximate daily
FIG. 1. Relation of Secchi disc reading (SI), photosynthesis per square meter of water
meters) to depth associated with one per cent of (Kl).
surface light (EZ, meters). An approiimate csti-
mate of euphotic zone is obtained by multiplying I’,, = P, X Y, X D1 X 0.65 (1)
Secchi disc reading by 5.
Where I’, is mean phytoplankton crop per
from time-to-t’ime in a given lake and among cbubic*meter in the euphotic zone, Y, is daily
different lakes of similar epilimnion depth. photosynthesis per unit standing crop under
The data in Table 1 (‘an be used to pro- optimal light (Table l), and D1 is the depth
vide estimates of photosynthesis in a given of the euphotic zone. This equation applies
lake if the following quantities are known: to lakes whose euphotic zone is confined to
(1) the standing crop, expressed as plant the epilimnion, or to unstratified lakes show-
volume, organic seston, or (Ahlorophyll, (2) ing 1 per cent or less of surface light on the
the depth associated I\-ith one per cent of bottom.
surface light, which marks the lower limit B~ause photosynt,hetic yields in the
of the euphotic zone (Ruttner 1953: 127), hypolimnion are so much smaller than in the
and (3) the ratio of Nolan photosynthetic epilimnion (Manning and Juday 1941) a
modificcl 0(111:11i01r is l)iUosoi~to(1 for I:lkos
whose euphotic* zone (1st~~llclsD c.ollsic~c~1,al)l(~
distance into t h(l hypolinlllioll, or for 1111-
stratified lakes showing light intensities
considerably ill (\xwss of I p(‘r wnt of s11r-
face light’ on the bot,tom :
Y,, = r;\ x Y,, x I)2 (‘-‘I
Where F’i is mean phSvvtoplanktjoll cwp ill
the epilimnion, I’, is as above, and 112 is
depth to midthcrmoc~line, or to botStom in
unstratified lakes. 111 this equation it is
assumed that the entire stallding crop in the
epilimnion is exposed to optimal light, and
that! the contribution of the hypolirnllion is
negligible. The first assumption is over-
optimistic, and the se(Aond is unduly pessi-
mistic. They, therefore, tend to counteract,
each other, yielding a simple tquation which
is likely to provide a fair approsimatioll of
the photosynthesis yield.
To test the usefulness of these equations
they have been used to wmputc IF0 ill sewn
Wisconsin lakes studied l)y lIarming and
,Juday (1941). The yields from ecluations
(1) and (2) are compared in Table 2 with
yields computed by 1lanning and ,Juday,
using a det,ailed method (estimating light
intensity and photosynthetic yield at l-
meter intervals in the vertical column, and
at hourly intervals throughout, the day)
which was admittedly more refined than the
data warranted. In applying equaGons
(1) and (2) I used the standing crop of
chlorophyll determined by ALfanning and
Juday , and the hourly photosyntShet,ic+
yield reported by them (Table I, of this
paper), and assumed that there were 12
hours of daylight during \vhicah the photo-
synthetic rate was maintained. The rough-
est approximat ioll in this type wmputation
is the failure to (wrrwfj for dim light when the
~1x11is low, or obscwred by clouds. Stee-
mann Xelsen has sho\vll (I 95-I : 3 17) that
marine photosynthesis (PFl ni?) is not,
linearly proportional to light intellsity, hut
that an intensity cwrrwponding to one-
third of full sunlight \vill support, a photo-
synthetic yield amounting to two-thirds of
the rate at) full sunlight. And I have show)
(Verduin 1956) that, phot’osynthetic yields
on cloudy days were cwnsiderably higher
Yield by Manning
and Juday (mmol)
Yield from rquation
Cl) or (2) (mmol)
17 89 (1)
53 56 (1)
57 58 (2)
57 72 (1)
83 78 (2)
87 96 (2)
146 325 (2)
than one would predict on the basis of ob-
wrvcd light intensities. The approxima-
tion, t hcrefore, is not so rough as would at
first appear, and Table 2 shows close agree-
n1cnt ) in most cases, between the yields
computed from cquatJon (1) or (2) and those
reported by Manning and Juday. It seems
likely that these simple equations provide
as reliablt an estimate of the photosynthetic
yield per square meter of water as does the
more tedious metShod outlined by Manning
and ,Juday.
It should be emphasized that the least
rt~liabltl factor in these equations is Y,,
the phot’osynt,het,ic rate per unit of standing
crop. I believe that significant differences
can be established between rates for dif-
ferent lakes. Manning and Juday did not
study this aspect of the problem. The low
value of photosynthetic rate (per mg ash-
free dry weight) for Wisconsin la.kes (Table
1) as compared to Lake Erie, Pymatuning
Reservoir, and Georges Bank, is evidence
that such differences exist.
Pennak’s publication (1949) provides the
data needed for yield computations using
equation (I) or (2). In Table 3 these yield
estimates arc presented. Pennak’s st’and-
ing cbrops are expressed as organic seston,
a photosynthetic rate of 1.4 micromoles per
mg per hour was used to compute the yields,
and Secchi disc readings were multiplied
by 5 to provide euphotic zone dept’hs. The
actual determination of Y, and of euphotic
zone depths in Colorado lakes might result
in a considerable revision of the values in
Table :3. Computations for western Lake
Erie, Sandusky Bay, and some roadside
ditches in Kansas are included. The most
 III If‘igure 2 the monthly 11~~~11 photosy11-
t hctich yields per square meter per day for
\\.aters in the island region of west em Lake
.!I1 yields \vere computed using :L v:il ue of 1.4 Ij:rie are graphed against time of year.
millimoles per gram ash-free dry weight per hour
For the I’S Yield per unit of phitoplanktbn crop, These estimates are based on observed pH
and assuming a 1%hour da>, Whew I), is given changes in the bay near the IT. 1‘. Stone
cclllat ion (1) whew 1)~ is given Ilaboratory between early morning and lat’e
rIsect. afternoon (Verdnin 1956), and t’hey repre-
Depth
sent’ photosynthetic rates measured under
Organic
Euphotic
zone
to mid-
thermo-
c*ompMely natural conditions. ;I line has
Seston Pho~;xt~esis
I.ake been drawn through the highest values ob-
(%5’
lh)
served at various seasons of the year. The
values lying below this line (February and
i2llf?IE! 1.9 7 145 .ipril) were associated wit’h high turhidities
Base Iline 1.7 6 170 which severely limited photosynthesis. The
Haydens I.8 5 152 calear waters which are usually encountered
ISeasleJ 2.4 -I 162
Gaynor 13.6 3 450 a fen- miles to the northwest’, or east, of the
noulder 4.7 5 255 island region (Verduin 1954) almost cer-
Kossler 1.2 6 122 t’ainly supported yields similar t,o those ob-
(Pennak 1949) served when the currents were accomodat-
Erie 2.8 5 153
(Verduin unpubl .) ing enough to move clear water into the bay.
Sandusky 13 1 .-4 200 The curve in Figure 2 bears a st,riking
Bay (UcQuate unput~l.) similarit’y to the curve for radiant energy
Kansas 15 0 .6 151
ditches (Rntelaff 1952) flux in this latitude (Verber 1955) which
has been drawn in as a broken line. It is
significant’ t’hat maximal yields were ob-
served in .June, a month which has been
-5 c*onsistenbly associat,ed with minimal diat’om
-4 carops (Chandler 1944, Chandler and Weeks
-3 *5 1945, Verduin 1951).
Y I’nder completely natural conditions
/ , -2
/ \ CO2 consumptSion during daylight hours is
\ \ I
‘0°: / y-y
. l very nearly equal to COZ evolution during
-0
0 ’ ’
JFMAMJJASOND
’ b * 0 ’ ’ ’
darkness (Verduin 1956). Therefore the
Y ONTH ratio of photosynthesis by the aut’otrophic
Frc. 2. Photosynthesis (I’S, mmol/m2/d:t~~) community to respiration by the tot!al com-
llnder completely natural conditions in western munity must be close to unity. It is
1,ake Erie, graphed against time of J.esr. The yield
is proportional to the solar radiation cIIrv(~ dificultj to distinguish between the respira-
(broken line, hundreds of g.c:\l lcm2/dny), tory contribution of autotrophic organisms
and that of the other members of the com-
interesting feature ill ‘1’at)lo :! is the uni- munity. 111 vigorous laboratory caultures
formity of yield estimates. 1Iost lakes shon algal respiration usually represents about
yields between 150 and 200 nGllimoles (1 .8 10 per cent of maximal photosynthesis
and 2.4 grams (‘) per square meter per day. (Ryther 1954), and when diatom popula-
(iaynor Lake, in Colorado, shows a (~)n- tions from Lake Erie, concent’rated using
siderably higher yield ; consequently that 90. 25 bolting cloth, were studied (Verduin
would be a most interesting one in \vhich to 1956) the mean ratio of respiration to
det’ermine act’ual photosynthetic yields per maximal photosynthesis was 0.125. If
unit of organic seston and ac*tl~:tl depths of this value is regarded as representative of
euphotic zone. the allt,otrophic community an estimate of

pllytoplmkt 011 I~c~~)l’o(l~l~~tio~~
(311 1)~ 1n:1do
for western I,akc~ I<%0 as folio\\-s: l’hot o-
synthesis under optimal light is 1 .(i milli-
moles per gram ash-free dry weight per horn
(Table l), or
1.6 X 0.65 X 12
_-.-___- = 6.2 mn~ol/g/day.
2 >
The factor of 2 is introduced because the
euphotic zone represents about one-half of
the total water mass (Verduin 1954).
Respiration of the aut ot rophic communit,y
amounts to about (1.6 X 0.125 X 24) = 5.0
mmol/g/day. The photosynthate avail-
able for reproduction amounts to about 1.2
mmol/g/day, or approximately 20 per cent
of t’he total photosynthetic yield. This
estimat’e is similar to Riley’s (1941) Table
VI, from which a mean total plankton pro-
duction representing 15 per cent of total
gross production can be computed. Con-
vert’ing to glucose oiw obtains (1.2 X 30) =
36 mg/g/day. Thus it, would require
1 would provide even a roughly quantitative
___ = 28 days for a gram of ash-free dry
(0.036)
weight to produce an additional gram of
glucose available for reproducfion. This
estimate of regeileration time agrees in
general with IGnscle’s turnover caoefficicnt
(0.83 per month) and Grimm’s observation
of annual diatom yield (8-10 times maximal
st,anding crop) as reported by Ruttner
(1953: 149-151).
Of t,he three quantities used as a basis for
comput’ing photosynthetic4 yield (Table 1))
organic seston would seem to be the least
reliable estimate of autotlrophic* organisms
because important fractions of non-auto-
trophic organisms and of detritus are in-
eluded. The \rariatioll :unong individual
samples, ho\vCi’er, is high regardless of the
base used, and is only slightly higher in the
organic seston-based rates than in the ot)hers.
(See Riley’s 1941 (*orrelations between t’hese
bases and oxygen production, for example.)
It is noteworthy that the average rates com-
puted from Riley’s \I-ork are about) 50 per
cent higher than those from the work of
Manning and ,Juday ( 194 1) in both t’he ash-
free dry w-eight,- and the c*hlorophyll-based
89
c~olllllllls. ‘I‘h(l differences, t horf~for(~, tloes
11ot rofltlcbt a higher fraction of non-auto-
trophic mat)erial in the Wisconsin organic
seston. If the curve in Figure 2 is used to
c*omputc an average photosynt,het ic yield
(per &/day) a value of about 140 mmol is
obtaincd. This agrees closely with bhe value
of 153, for Lake Erie, in Table 3. The latter
is based on centrifuged organic sest’on de-
terminations and an assumed photosynthetic
yield of 1.4 mmol/g/hr. This comparison
is particularly significant because Figure 2
represents CO2 change under completely
natural caonditions and is not based on esti-
mates of phytoplankton volume, organic
seston, or chlorophyll. These observations
suggest that organic seston values (means
f ram numerous determinations) provide a
fairl) reliable index of photosynthetic
capacity in spite of the large variations in
ratio of autotrophic to non-autotrophic ma-
terial which must be present among individ-
ual samples. However, a method which
dist8inction between autotrophic organisms,
non-autotrophic organisms, and detritus
would caontribute much to aquatic biology
It should be emphasized that the produc-
tion estimates in t)his paper represent rough
approximations. They should provide a
mobive (not a substitute) for more detailed
research in this field. The large variation
in photosynthetic yield among individual
samples (Ryther 1954, Verduin 1956) makes
the study of numerous samples mandatory
regardless of the precision achieved in in-
dividual determinaGons. Consequently, a
time-consuming method which provides a
high degree of precision may be less valua-
ble than a more rapid method with a larger
scatter which will disappear when many
samples are averaged. Moreover, any de-
t,ailed study of one aspect must be accom-
panied by equally detailed determination of
other aspects. For example, careful deter-
minations of photosynthetic rates will not
improve the area-based yield estimates if
detSerminations of light penetration are not
equally refined. And the actual euphotic
zone depth for a specific community is a
function of that community’s efficiency
under dim light; the roughly approximate
90 .J.‘LC!OIJ VERDUIN

“1 per c~iit, of siii*fm+c~ light ” ikllist lw atl- In 3 previoiis publication (1’twluiii 19X)
justcd to fit specific* wnniunit iw ;iiicl (*Ii- I demonstrated that less than half of the
nutcs. Whcrl suc*h r(~filwrlwlt s arc at- photJosynthesis observed under natural con-
t,empted the choice of tool used for light ditions clan be accounted for by the phyto-
intensity measurement h3c~o1nc~simportant. plankton carops detected by examination in
I have pointed out (Verduin 1950) that t,he Sedgwicak-Rafter cell. Figure 2 further
photronic cells like t’hose in the photometers emphasizes t’hat fact,. Lake Erie, like many
used at the Stone Laboratory arc relatively temperate zone lakes, exhibits vernal and
insensitive to non-visible radiation, and that autumnal diatom maxima with wint,er and
the photosynthetic process also is insensitive summer minima. The annual curve for
to non-visible radiation. Icor primary pro- photosynthesis under natural conditions
duction studies, therefore, t,his type of is not himodal, but shows a summer maxi-
photometer has advantages over the pyrhe- mum at, a time when the diatom crops are at
liometer t,ype, because the latter measures a minimum. It seems likely that other
the non-visible rays which represent about lakes which exhibit bimodal phytoplankton
one-half of surface radiation but are in- abundarwe curves will reveal similar uni-
effective in photosynthesis und are rapidly modal productivity curves. The organisms
screened out by water. responsible for this unexplained photosyn-
This difference in instruments is responsi- thetic act)ivity (their identity, size, biomass,
ble for Rawson’s (1950) conclusion that the metabolic activity, etc.) present one of the
ratio between euphotic* zone depth and major problems in primary production to-
Secchi disc reading decreases with irwreasing day.
transparency. In a private communica-
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 PRIMARY PRODUCTIOS IN LAKES 91

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