Title Page
 Table of Contents
 Species of algae and their distribution...
 Distribution of aquatic insects...
 Productivity theory

Group Title: Productivity of Florida springs. NR 163-106 (NONR 580-02) Report to Biology Branch, Office of Naval Research; progress 1953-1957
Title: Productivity of Florida springs
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00073979/00003
 Material Information
Title: Productivity of Florida springs NR 163-106 (NONR 580-02) Report to Biology Branch, Office of Naval Research; progress 1953-1957
Physical Description: 4 v. : illus. ; 28 cm.
Language: English
Creator: University of Florida -- Dept. of Biology
United States -- Office on Naval Research
Publisher: s.n.
Place of Publication: Gainesville
Subject: Freshwater biology -- Florida   ( lcsh )
Springs -- Florida   ( lcsh )
Genre: non-fiction   ( marcgt )
General Note: Frequency varies.
Funding: This collection includes items related to Florida's environments, ecosystems, and species. It includes the subcollections of Florida Cooperative Fish and Wildlife Research Unit project documents, the Sea Grant technical series, the Florida Geological Survey series, the Coastal Engineering Department series, the Howard T. Odum Center for Wetland technical reports, and other entities devoted to the study and preservation of Florida's natural resources.
 Record Information
Bibliographic ID: UF00073979
Volume ID: VID00003
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved, Board of Trustees of the University of Florida
Resource Identifier: aleph - 001871216
oclc - 21432514
notis - AJU6211

Table of Contents
    Title Page
        Title Page
    Table of Contents
        Page 1
        Page 2
        Page 3
        Page 4
        Community matabolism of Silver Springs
            Page 4
            Page 5
            Page 6
            Page 7
            Page 8
            Page 9
            Page 10
            Page 11
    Species of algae and their distribution in Florida springs
        Page 12
        Page 13
        Page 14
        Page 15
        Page 16
        Page 17
        Page 18
    Distribution of aquatic insects in relation to environmental gradients
        Page 19
        Page 20
        Page 21
        Page 22
        Page 23
    Productivity theory
        Page 24
        Page 25
        Page 26
Full Text

Pp L., ,

Annual V.
*-ann nrv

Wit:.fh OQ~: kvwn V. '
.0 .3~~~:Q
13i Xi~:va -'e
r~i:X -.~ i. *1s~

D3 vsrn *itB CWin

I .: )


0 !-, !. I '!I -'- I Yi.

Of El.P

1 10 ', .


Introduction and Abstract

Ao Community metabolism of Silver Springs
Alkalinity, hardness, and chlorides (2nd semi-annual
report, p. 8)
Nitrogen-phosphorus ratio, comparisons between springs
Boron (by H1.T Odum and Bruce Parrish)(in 2nd semi-
annual report, p. 6-7)
Light intensity
Photosynthetic efficiency
Further measurements of production and respiration
Bacteria (by H.T. Odum and Osilio 1alindo) in 2nd
semi-annual progress report, pa 15314o
Pyramids of weight and number
Downstream losses from community production
Long nnCco Stability
Balance sheet for silver Springs

Bo The species of algae and their distribution in Florida
Springs (by Larry wrhitford, N.C. State Uollege)

Ca Distribution of aquatic insects in relation to
environmental gradients (by Wo'. Sloan)

D, Productivity theory

Productivity measurements and atmospheric diffusion in
Timds regulator; the optimum efficiency for maximum
power in physical and biological systems (by IIo To
Odum and R, 0. Pinkerton, Dept. of Chemical Engo)

p @


Prepared by: Howard To Odum with sections written by WC. Sloan and

NR: 163-106


ANNUAL RATEs 1nw (4 y, ars)

CONTRACTOR: T.;t., of Biology, University of Florida Gainesville
(with Biology Branch, Office of Naval Research)

FRINCIPLE INVESTIGATl,: T, ..- ,d To Odum (In the course of the original
application this ;:.-r.j became administratively listed under We Co Allee,
Head of the i:l'.'L~ii Dept,)

Collaborat,:i. s durinye current period:
John H.o .,.'.., i j..p-o o C .L .:.y ""
Larry A.o hitt E~d, NoCo State College, fall time during summer 1953
Richard CGo Phl.2:;:t.,; .r' Di;-. of Chemical Engineering

E~~iam C. Sloan, i~'- 1, 1952 to present
Osilie C,..... Jnaea 1953, financed by Dept. of Biology
Bruce Parrish, June 1953, financed by Dept. of Biology
.ade Hampton, F. to May, 1953


Objectives: A :.'d;:l of the basic factors controlling -individual,
population, and community productivity by an analysis of the unique
conditions uappliod by outflows from selected constant temperature


ao Dul in rent r2Fe 22Ej d (Jan. 1, 1953 to Deo 31, 1953)

By means of new methods it has been possible to measure theover-
all community metabolism as well as the standing state comImunity in Silver
Springs. Photosynthetic rates have been determined by downstrem:- gradient
methods, transplantation growth plots, and bell jars diurnally and annually
Respiration rates have been estimated with bell jarso The doiwntream losses
of particulate, and (.. -:-.i.':.d organic matter have been found to balance
the excess of photosynthesis over respirationo The community has thus been
demonstrated to be in a constant ~~z:l-,l.;ature steady state sat somewhat comparable
to a climax on land Nitrate, phosphate, and major chemical constituents
are essentially constant There is an approximate constancy of standing
crop of organisms 71*t11-i.g;,; the production rates in summer are three times
those in the winter. s-,.- evidence :':.;gect.. that there are photeperiodio

changes in reproduction rates in spite of constant temperature
Rates of protein synthesis estimated from nitrate uptake down-
stream agree (1) with photosynthetic quotients obtained from
carbon dioxide and oxygen uptake downstream and (2) with the
nitrogen content of the conmunityo The overall annual production
of 50,000 lbs./acre is the greatest productivity we know of on
land or seao huch high figures seem reasonable with the flow
of high nutrient, rram water, and high light intensity over a
dense periphyton community Theoretical concepts of stady state
thermodynamics have been applied to show that self maintaining
ope systems tend to adjust to high power and low efficiency
output. The 3% photosynthetic efficiency observed in Silver
Springs is in agreement with this principle Pyramids of weight
and pyramids of number have been determined including bacteria
These pyramids are aindlrllc.: to some in the literature, The
contribution of an acre of a fertile stream annually is readily
inferred from data obta.iin.' on downstream increase of bacteria,
chlorophyll, and organic matter.

In other springs Mr. Sloan has related stability of insect
populations to chlorinity and to gradients of stability of
environr-ental factors, Dro Lo A, .hitford during the summer
made an.. ecological r?'- taxonomic study of the distribution
of algae in 26 coo.t;rating springs. From his lists and from
analytical data on the chemostatic water in these springs one
can infer culture conditions necessary for many species

1Yhere consistent with accuracy ~:.-tL~ry t-, prove a point,
some of the ia..A;ui accuracy that might be obtained is sacrificed
in any one technique to p-..it the study of all aspects of the

be Since start of. ro- at
'his contract was begun June 1, 1952. In the first 6 months
which preceded the current report period, the oxygen gradient and
transplant proO.;' ::. .vlty methods were developed. Ia)s were
constructed and .c -,'.~r.tive chemical and biological data were
obtained for 40 i'...r.oti.' springs, A miscellaneous exploratory
experiments were conducted as reported in the first progress
report, Experiments begun earlier in the springs on chlorinity
control of marine invasion were completed, Intensive work on
Silver Springs was 'eguno

Immediate: prr.:Ps production rates of higher trophic levels:
ao periphyton animals by glass slide growth and
-ieE:.i;on rates on Saoittaria blades
b, fish by tagging, visual racingj of colored tagsa
and recapture with gig; growth in cages
Construct a laboratory periphytonflow-system, algal
culture as a stream model
Complete nitrogen picture with total nitrogen analyses;
check some nitrate patterns.

Long gl~S: Compare production rates of different springs
with the best of t, techniques.
I Co, ..:. production differences with chemical
di.f:'-;.eaiges in the thermostatic chemostats.
T ,'.';^ Sr:.':-'.mrints to demonstrate relative roles
of competition, incidence, time lag factors,
';.:. physical composition on community structure.
Cor.I:.' o;. hu>, activities^ efficiencies, .i:'
community structure of springs with other
"' .l'i steady state communities such as coral
reef, :--o;...cal streams and equatorial estuarieso


A study begun earlier but completed as part of this project:
Odum, 1.T, '.-',l,. Factors controlling marine invasion into
Florida wa :.. u OullQ of marinee Science of the Gulf and
anrifbc:. volo ,ppo1$4-156 o

Sti.u..-lated b: -,-'T.; project although not directly :l''o''ted:
Laessle, A),. I.,O'. The use of root characteristics to
op':r. ,te various ribbon-naEvcoc species of Saittaria from
species of .W-rli .:.;.o Turtox News, volo M3T (1F7 eo 2 ppo

The ,.:,';-. :c;'::. an.uual progress report was d:Tstributed in
July 1953o Beca.-.co of the change in policy to an annual
report ':,..;:: '-' of the 2nd o;id.-annual :...,C; are
at'; -,1h,2 apart of ;L 'j;i report; and referred to within to
avoid d'.Ulirc.: r ".-c. p. per:.. on aspects of i-,i,...',,gs work
were pr":c.-:" .-, at the .i...i-:, A'ao..:. of SE Biologists0 and
Flao Acado of Science meet-ings~




In the 2nd ,,r ,io-.;r-t..! report(po4-5) is reported l1: nitrate
analyses made with the ;.,.:,-. "ous phenol-disulfonic acid method
These analyses r.:" a 5 :.> variability 'l.":i 'a; in the method
but established ;h"' ?."-.A l nitrate level, demonstrated a seasonal
constancy, and a -..,-'l.. 1o'.,..O.loei. uptake statistically

The strychnidine method for determining nitrates was then
adopted with :;.r :i.' .:l 2 ':*:..-,.ibility and reduced costo. The
pattern previously shown has been cor':...:'icJ. The nitrate uptake
Is found to d'.c:."-: .: both at night and during the d;C,"o This new
data is summarized with the old in Table lo Jote the approximate
constancy of the nitrates seasonally

Long gl~S: Compare production rates of different springs
with the best of t, techniques.
I Co, ..:. production differences with chemical
di.f:'-;.eaiges in the thermostatic chemostats.
T ,'.';^ Sr:.':-'.mrints to demonstrate relative roles
of competition, incidence, time lag factors,
';.:. physical composition on community structure.
Cor.I:.' o;. hu>, activities^ efficiencies, .i:'
community structure of springs with other
"' .l'i steady state communities such as coral
reef, :--o;...cal streams and equatorial estuarieso


A study begun earlier but completed as part of this project:
Odum, 1.T, '.-',l,. Factors controlling marine invasion into
Florida wa :.. u OullQ of marinee Science of the Gulf and
anrifbc:. volo ,ppo1$4-156 o

Sti.u..-lated b: -,-'T.; project although not directly :l''o''ted:
Laessle, A),. I.,O'. The use of root characteristics to
op':r. ,te various ribbon-naEvcoc species of Saittaria from
species of .W-rli .:.;.o Turtox News, volo M3T (1F7 eo 2 ppo

The ,.:,';-. :c;'::. an.uual progress report was d:Tstributed in
July 1953o Beca.-.co of the change in policy to an annual
report ':,..;:: '-' of the 2nd o;id.-annual :...,C; are
at'; -,1h,2 apart of ;L 'j;i report; and referred to within to
avoid d'.Ulirc.: r ".-c. p. per:.. on aspects of i-,i,...',,gs work
were pr":c.-:" .-, at the .i...i-:, A'ao..:. of SE Biologists0 and
Flao Acado of Science meet-ings~




In the 2nd ,,r ,io-.;r-t..! report(po4-5) is reported l1: nitrate
analyses made with the ;.,.:,-. "ous phenol-disulfonic acid method
These analyses r.:" a 5 :.> variability 'l.":i 'a; in the method
but established ;h"' ?."-.A l nitrate level, demonstrated a seasonal
constancy, and a -..,-'l.. 1o'.,..O.loei. uptake statistically

The strychnidine method for determining nitrates was then
adopted with :;.r :i.' .:l 2 ':*:..-,.ibility and reduced costo. The
pattern previously shown has been cor':...:'icJ. The nitrate uptake
Is found to d'.c:."-: .: both at night and during the d;C,"o This new
data is summarized with the old in Table lo Jote the approximate
constancy of the nitrates seasonally

Nitrate Analyses in Silver Springs in ppm

Boil: 3/4 mile
Day, time9 sky downstream:
analyses mean analyses mean

Strychnidine Me tho.:
Novo 27, 1953 night (2:30 a.m.) 10 .463 10 4539
Octo 22, 1953 day (2:00 p.m) almost 10 .493 10 .574
Octo 10, 1953 day (1:00 p.m.) clear 10 .401 10 .380

Phenol-disulfonic Acid Method
April 9, 19553 c, noon, clear 10 .453 10 o330
May 8, 1953 night 10 .499 10 o468
May 14, 1955 noon c.r-. 10 .462 10 o440
May 25, 1953 1:30 p.m. few cumulus 5 e40 5 34
Mean TY "TSO

Phospjgt ea

Additional inorganic phosphate determinations were carried out
in Silver Springs. The data in Table 2 shows a small significant
uptake of phosphate between the boil and downstream. (The possibility
that the decrease of phosphate and nitrate is in part due to the
entrance of dl.~:~:.c;. i: (.ji.7.ty water from small side boils is now
being checked,)

Table 2, Dissolved Inorganic Phosphorus in Silver Springs
in ppm

Day, time, sky

April 9, 1953, noon clear
May 14, 1953, noon, clear
May 25, 1953, noon, few cur,,.l.us
Oct. I, 19553 noon, broken clouds
Oct. 10, 19535 noon, clear
Oct. 22s 1953, 2:00 pom., cumulus
Novo 27, 1953, night


Boil 3/4 mile
analyses mean analyses mean
analyses mean analyses mean


.603 '~~9~

-- -- -~- -A -~au r~ u~ lir~~

-I -- --~---- --I----

Table 1o

Nitrogen-phosbhorrs ratio, comparisons between springs

The downstream decrease of nitrate is o057 ppm and the down-
stream decrease of phosphorus is .0026o The uptake ratio is
therefore 21o9/1 by weight if it can be assumed that the down-
stream docrE eso is ..ti:-ely uptakeo The ratio in the water is
a457/00543 or 8r4/o1 Thus relative to the water the nitrogen
decreases twice as fast as phosphorus and thus should limit first.
Full confidence in this result cannot be placed until the
contribution of the side boils is checked and the downstream
organic nitrogen is measured From Table 3 below it is interesting
that Silver Springs has a higher N/P ratio than other springs
analyzed. Thus nitrate-N may be limiting in many springs.

The very striking differences in the communities in the
different springs in spite of similarities in the chemical
composition of the major elements and similarity of temperature
and light is one of the most fascinating aspects of the whole
project. Constitu.Tilo that might be responsible are nitrate-N
and phosphate and the N/P ratio. There are wide and radical
differences in N/P ratios The adoption of the strychnidine
method has permitted nitrate determinations on the high chlorinity
springs as in some L +.-.: in Table 3o As a beginning to the
comparative study of various springs (which is the main objective
of next years study) some N/P ratios are shown in Table 3S Some
idea of the large community differences can be obtained from
Dro "hitford's algal lists in subsequent sections below

Table 3o N/P ratios in various springs

Spring, date N03-N Inorgo P N/P
pM by weight
Juniper, Deco 4, .040 .024 1.66
!.ekiva (Orange Cos), Deco 4 0062 0132 e47
:nee De Leon (Volusia Col, Dec. 4a .157 .052 3,00
Alexander, Dec. 3 .034 .050 068
Silver Glenn, Deco. 3 .023 022 1o00
Salt, Dedo 3 0060 .012 5o00
Ilomosassa, Nov. 1953 .080 .008 .10
Rock, Deco 6, 1953 .112 .127 000
Silver, mean of 300 analyses .444 .0409 /r
;eekiwachee, June 6. .12 .018 6,7
Orange, June 6. .22 .081 2o7
Sanlando, June 19, .17 .172 .98
blue (Alachua Coo), June 26o .85 .092 9,2
Ichtucknee, June 9, .40 .064 6a3
-UII~IL~C -I---~---__I-UIP( IPI(OIIUL~lrm --rY~.-

Light Intensit,

Measurement of light intensities has just commenced, but some
initial figures enable an estimate to be made of the magnitudeso
Measurements were made with a submarine photometer obtained from
Fred 6chueler, 30 Albemarle Road, Vialtham 54, iasso containing a
"eston photronic cell #8594YRo A diurnal curve of light intensity
at the surface and 8 ft. deep on a clear winter day is given in
Figure I with],ight intensity expressed in microamperes as read.
Approximately/4 foot-candles are equivalent to a microampereo As
shown approximately half of the Incident radiation penetrates to
the photosynthetic surfaces of the community which are mostly
in shallower water than 8 ft. The bed of plants is 2 to 3 ft. thick
and 99% or more of the light reaching the plants is absorbed in
the first 1 1/2 fts. :hen the photometer is inverted at the
water surface no appreciable light intensity is measured so that
99% or more of the light is absorbed Of the light incident on
the water much is deflected by refraction and reflection in
connection with ripples, for there is a strong flicker in the
microampere reading when the wind ripples the surface As is known
in other waters the absorption in the first meter of the very
clear water is not logarithmico Further measurements are in
progress to determine whether or not this water is more transparent
than that in the Sar,'. o Sea. From the point of view of vertical
stratification the largest intensity differences occur in the
first meter. In I ..' w 1 note that the length of day at 8 fto
depth is ohly 3igb'hly h-.ter than at the surface even in the winter

Phos ;i'L:b Eff l

At the sameme tme t'at light was measured in Figure 1, the
productivity was determined by the downstream gradient method of
oxygen measurement as outlined in previous reports and corrected for
simultaneous respiration on bell jar estimates This production
gives an estimate of C % "f Phe incident li~ht in the photo-
synthetic range is taken o:.4 foot candles/ricroampere in Figure 2.
If a calculated intensity is obtained for a clear day f-.sri
Kennedy (1949. Icy,3,. ..tion of daily insolation energy i'ullo of
Amero MIeteorole Soc., vol. 30, pp. 208-213.), an efficiency of
3.2% is obtained. If the annual production of 50,000 lbso/acre is
used with an annual insolation figure from laurwitz and Austin
(1944. Cl'-atology, 2cGraw Hill.), 3.5% is obtained.


In an effort to understand how the greater plant production of
spring and summer is translated into ani-al production without
grois changes in the community, two series of measurements are being
made as indexes to r.- productive activity of two invertebrate
@ populations* 'h.e percent of female shrimps (Paleomonetes) with
S eggs and the number of clumps of Pomacea (Ampultar-a ) eggs on
125 meters of rock and board walls whfTcEhconstitute one border of
the boil area. Although Incomnlete there is data enough in Table
4 to suggest a definite photoperiodic nulse in breeding intensity
although some breeding occurs winter and summer At constant
temperature it is very likely that the stimulus involves light
directly or indirectly.

Table 4, Seasoal. Variation in Egg Counts in 6ilvor S.-. ;ngs

arm!il(BW-<^"^WTUtasseswIaYsO owneverwi--^<^e^^^o^llrsimmenaenu r aM nr w m r a t ,. n, .n .-.mass ran.nwl.w.ief^>-<-


with eggs

"r"^n #
oT.'-7 2'.- lumps
:,:-2 125 meters
of shoreo


Oct. 1
Oct. 15
Oct., 22

Dee 19
Jane 7, 3.:',


-00 -


/OO -

~(#) ,.+

hi I

I t

Time of Day

Figure lo DV. nal ,.areh of LiCrht ci-:~nty in
Doc. 19 "...'3g




Further measurements of production and resiration

The meth.-:.ij for measuring overall productivity by oxygen
g adi.c',;- met l. ,(- :'..-: transplants, and bell jars were outlined
in the first ir- c;:, ..;s report and detailed data is reported
from the early part .'? the current report period in the
2nd semi-annual C..c .t (attached) on ppo 16-21e Since that time
two additional full day curves of oxygen and carbon-dioxide
have been obtained which confirm the previous patterns Some
additional summer data is all that is now needed to show a
detailed picture of the overall community production rates ral2ative
to annual changes in length of day and light intensity.

Another series of 6 black bell jar respiration measurements
were conducted Nov. 'f This timerubber tube was used rather
than copper (The use of copper was potentially a stupid blunder)
for extract ng water from inside the bell jarso As listed in
Table 5 below, these measurements fortunately agree with the
previous measurements on p. 19 of the 2nd progress report The
bell jar measurements were made during the dayo In the first
hour after the jars were lowered over the plants there was a
net oxygen increase This may be accounted for by the suggestion
that oxygen diffu:::. into the water from the tiny visible bubbles
imbedded in the per.i .3:y.ono In the second and third hour the
oxygen de: c.'a:;-.. at about the same rate and it is these rates that
are used as respiration estimates.

Table 5o 1'urther Bell Jar Respiration Measurements

Time .la:.o Or;. Oxygen decrease lbso/acre/yr
oxygen measure *-..J. p-:-' hro per jar glucose
in minutes ppm

108 .84 22,400.
95 .61 16,200.
106 044 11,720.
90 .34 9,060.
114 A46 12,240.
92 .52 135840.
mean: 14,240
*If 1 ppm 02 g .66 ppm dry weight instead of
1 ppm 02 094 ppm glucose, mean respiration is 10,000 Ibs/acre

The diurnal curves of oxygen and carbon dioxide at the down-
stream station on Jano 7, i..'153 again showed the lag between
oxygen release and carbon-dioxide uptakeo The oxygen curve follows
the light curve with a rapid return to night values at sundown
whereas the carbon-dioxide uptake reaches a peak later than the
oxygen and does not return to night values until several hours after
dark Thus the timing of light and dark reactions of photo-
synthesis show up even in this overall community respirometoy
Peculiarly a plhtosy.~c~hetio quotient greater than 1 was obtained


Pyrads of weigjLt and number

quantitative estates of the main com-n"unity components
were made and anal r.:-.1 in the first part of this report period
and are :,..l.,vi on 10-15 of the att:-.x. 2nd oG--r a
report,, Included in this are estimates of bact'ia. Further
estimates of ... i..1pi .:,.',p are 'n.,.p;m.d:

Downstream losses from .- i, it;r pi':..:.,lt:l.on
A major Lp.?.. of work ;]il3 fall has been an estimation of the
downstream Ir. ,:, of ..' :T.u':.c;. As indicated in the balance sheet
section below thcse eocatiates 'idicate that the overall metabolism
estimates now :"~.'- ..y :.J c.-': Estimates of downstream loss
include: BOD '.1.,1,.:, 1:Ci.n:irin'te organic matter, bacteria
chlorophyll(by il'.U. s'-' l:.:h.all, Flao State Univo), particulate
organic matter, and gross plant clumps 'The BOD values were
inconsistent and thus the permanganate method was adopted.
These data wre s': ., in Table 6. It is quite clear that
there is a v.-'.r- snall downstream increase in content of organic
matter which appear i.,.. .i. fiflicantly i.a.l. per liter until one
multiplies by, the large water disc,.1'r,;o It is then apparent that
the steady downstream loss of cr:.in-Vi matter mainly as inviable
dissolved substances I?.ia microscopic particulate organic matter
is half of the }j ...;..;...1 of the 3/4 mile headwater region
(bee Table 7). Just as in the terrestrial community, possibly
the coral rf.., or the lak E t e community is adjusted to produce
considerable excess orgai.c matter over that needed to balance

Tal.,.3 6. .-1.. .s 3/ mile co ..w iv ". .. .

Mlea.uri.-nt, :.c. ..,C1 Boil 3/4 mile
.. ............ ...... ...... -'.. *.. .'.. '~.~.'.r..s~. ji .'iJ .. ... i, ,i,, .n
BactcriA-- (IoInrici 1; :,-1) 10 99/ec 10 988o/ee

Chlorophyll(centrl L7",r- and extract 10 liters, by Ucli. '.. marshall)
July 3, l'.. 1 o0 m/ 1 o43mg/T
Dissolved Organic Uatter (Permanganate method, value, in ppm 02)
Oct0 1, l .a 5 3 o54 3 110O
Oct. 10, 1953 10 .69 10 .81
,ct. 22, ,105 5 .57 5 o62
Jean: "*-3553

5 day TOD (values in ppm 02 used up)
Octo 1, 1953 5 15 5 -o04
Oct. 10, 1953 5 o.134 5 *o04

Particulate organic matter (plankton net concentrate of 84 liters)
July 2, 195 1 .11 mg/l1

Algal cells ( 84 liter concentrate) 1 2.2/ce 1 67o/eo

Clumps of SagitS...i.:.- ('. weight) 3 296 -ms/


Ln o s" r..o st.. lity

In 1861 Loc._~ 'O: vis:i-l Silver Springs and wro~-e a paper
describing the cs :.cal effects observed and some aspects of the
community (Lcc.o i:-, Jo 1861o On the optical pheno:.,n" ,
presented by tlh., Silver Spring in Marion Co, PFla., Anr..-'.r Jo of
Scie, 2nd Series, volo 31, pp0 1--12.) His description of the
dominant plants as *c.,:. gr.ass and ~o.:.;-like plants" with
,.lado 3"-2: fto .,. .- tha.: that the -~s.!-:nui.ty in its gross
aspects is in :.. .: .~ tion as it was in 1861o

Balance sheet fo,"- Silver

The estimates of dissolved organic matter lost downstream
in Table R account Fo-r' the oxe.-o of production over respiration
Thus the pro .1:.I, disc:.c~-;.1 in the 2nd progress report (po 22)
seems solvec. ;''- annual balance sheet is given in Table 7o
Some further data l.'. be obta ned to check these estinateso
Perhaps for flr,:t :t ..! the oie;all community -,.?abolism of
a steady state natural community has been determined. It is
both exciting and sobervn:'; to know that the community does not
change radically so that data now oLi-ain-d can be repeated by
anyone late:'. or ,I-'.,;:" the ".: -is for .urt;he-r work without
repit.tione --l'; .. one forgets to measure tocda7. can
be mau. -;-...: or .: y. This reproducibility should
permit rapid i.-. <:es : -9 ble in the successive studies
in famous local such as L. "a.nt0;,,-, Lro indiicr,'eg and
Linsley pondo

Table 79 A;,:! 1l Balance Sheet For Silver Springs


Photoa~3 t.L~~. ".i -'. o..::.ed for ;ojuJp: .ration 456950o
,- ith o,, "''::- t ,. ,p-"? in tl. -.:50 00 t..

Lespiration 15,100.

Downstream 3.on
Clumps 590,
Particulate 13,320,
Dissolve, 15,000.

una- couint.Ka, .1o l65000*

Protein p'-Aul:: I: ion 20,800.
LrC 0 0G6-1, 0

Bo The Species of Al.-:;-. and heir Distributio n in Florida Springs
by "'.ry Ao x h 'r, ep.-.r..nt of Biology
N. CG State College, Raleigh

The Jw... '. *,: ;. :: ..- of ', o of the te-d.c;al Florida spring r .'l ru
are two -.c:t::. of -' :.an 4 :2n and i-l: : or six species of .[3s'n.,
P~.7. ., -.. a .. -, .. .... ', bl..- r;.-. al is by f: ." the most
...o... ... ._-. to a .--.th of 30 f.jct in the 1jo.l-, .t,
the i .nCIr., and is -. '..':; in the runs. A hit a spo is more ::;n..ns,
in areas of '. .:. ..11 _. deep in the boils-aTn masses of v'OagIt-i.
P.ctu.i;: fos extensive mats and clumps on sand, ooze and rock, and
C:i.-,' .:s: T''-.:; occur in and on all Iypasi of aquatic flowering plants. On
it and on the .. '"; of -.''",-. Cf,_.;! plental Achn,t~ lance
--;'. -. (." '. ....: '. ,'... *. :,T ^ 71^:;i 1.-', r.. n.' and ;-y}i-.- I.: 'T. ;.i &4(ld
.' v-- .. 3 '. ,, .; ..; .. -'i^o ;^,- to o fi e ::T' ,i.a.Li,.g
ZzYJ ujUO I .31n-j fiv Jpu.
together -%, a. .. others account for,, i. is estimated, over one h.-l the
organlc matter '." "" :?. , ,'; ,irgs, -ry leaf, stem, Plo't:va:':.i.i,
filaments, i:., or wood :.;:r.rc':- -%mcd to the sunlight for a few US as
a coj3.,$ple cover of *.1' 3c.1.;,, iUTjii-a: a few weeks the layer is c~-:c'al cells
thick. Light is the most important v. r'.3Able,. The under side of leaves has
only 10.=--1 of the number -.. on the ur.e-r surface. Surfaces within a
plant mass 3:":;-, 2:;' have much --.3, and although the water is exceptionally
<.ci, PI: .'?.c are .".". loes in itr.:,, :o.n.T (over 20 ..Iot).,

There .' ".:.2..t';,, r -.-r. in F ;..j dominant -:i'o of b-L., pool and
rano The shallows of the [.p. "and the c.-gon of the run have more filamen-
tous diatoms and ,':. ;:., because they are not washed away as rapidly
by the c:rr.:.., but the .:.. crop is .. ttlo greater especially in beds
of flw .r p ,,.. ,-. :. omie .:rt in the doriia:ntB toward the fila.
m'3ntout.i.'irto;.;.2; : .-. ;-..', Melosira and Nl.t3hia and the green algae Rhizo.
cloniu and .. .

The :.; :....: ...; I to. :3 a wide variation in chcfic-.,- c :-1-.: .ic.. Ap-
plaronty very 3 .1 7 .-:":.-: ."- 'r.. prevents abundant growth of most of I.,am
in the boil, but in the presence of a few ppm of cxyg-n there can be a wide
variation in mi ... .-:i, "; ,, .e, c-l.ci~Ei, chlorides, total dissolved solids
and hardness ;...:,'; any marked change in the flora. Growth of Plectonema
is a7l .:r'T- where there is only a r',:; parts of chloride per mil on n~;ii
:ih'or there is P -..' :" ppm. Most of the dominant diatoms grow well at
chloride i~nCr:, ~:.::; -,;' I.; :. 7y in the oligohaline range (up to 600 ppm)o

The ap.r.:: ;. Tr.it. absence i 'Ict.:rn.:c in Manatee :-.:: .' and its
abundance in the very :-..1 F- L-i, .; siy cannot .be -" na un
less it is to 7 ':'y; de."' f? n.' intohe f...nr spring. SiIrvL,-.,'.y the
factors which '?.JA.; the growth of H drodt in abundance in a few sprWi.gs
and apparently prevent its growth o era are unknown The same is ta
of the genera Vaucheria and Dichotomosi hn which are present and abundant
in only a few !a:T, .-..l-genus ora should be mentioned here also.
As a guns it should be P:l.Og among e osinants even in the .s16 springs
Since it is a very '.".- genus, however, with species adapted to a wide
range of conditions it is not !;.kod as a dominant I',ocrtu" no one species is
abundant and wit~t.c d in the springs. Apparently one p..io0 is present
in Silver spring, another in 1. 3.c.tll% and still others in saline waters. An


accurate knowledge of the species and their distribution might help determine
* some of the important factors bearing on algal distribution, Concentration
of chlorides, and speed of current may be important here, The great diminution
in amount of the species of C0ldoora in Wakulla run between July Ii and
August 1, 1953, cannot as yet explained

The quieter water at the c.dgcs of the spring pool and along the edges
of the run support a much more varied flora than that of the boil and run
channel. As pointed out* above, there is no great variation in the species
of dominant or their total abundance but the total number of species is much
greater, RhisocloniUn is abundant only here, as are certain species of Spirogra
and other filamentous algae. The unattached filamentous diatoms and some
single-celled and colonial green and blue-green algae are much more abundant
or sometimes present only here,

There is no true plankton in the springs or their runs. There is, however,
a considerable pseudoplankton or tychoplankton composed of microorganisms con-
stantly being detached and carried down the runs. (See Odum, Second Semi-
annual Report p. 22). That a true plankton flora would develop in the spring
waters under pond or lake conditions is proved both by the floras of nearly
lake waters with similar heirc.-il composition ard by a few cases where spring
waters are impounded. An cxca~ itd boat basin about 150 feet 1ong off Silver
Springs run contains many phytoplankton species in such genera as Endorina
Dictyosphaerium, Anki odesmus, Kirchnerielle Micratiniumo and Ped isrumo
Very small eaibaym ents alon'hior runs, and *evn the' water in submOrged oats
indicate that a true, and rich, plankton flora would develop under empoundmento

The dominant 2.:,o of the springs and their runs constitute what secms
definitely a perraneant vernal floral TJidespread collections from more than
ten springs at all seasons SRicn.-to almost no seasonal changes in dominance
There is possibly a a.llr::." on:a upward in winter of shade species (see below)
but this cannot, as yet, be definitely confirmed. Such genera as Microspora
Stigeocloniumredogonium and narrow-celled s e.es of Mou eotia whcSl" I' r
abundant only in spr g in most habitats are abundant n the springs the year
round. No sexual reproduction in the common green algae has been observed at
any season in boil, pool or swift runo

Certain species of al., in the springs seem definitely to be shade
plants, since they are found only in deep boils or in masses of vegetation
and since temperature is not a variable here. All the fresh-water red algae
such as Audouinella, C-osnroSon and Thorea belong here. Thorea has been
collection is ~4e Spr~~ngi at a-deptI of 30 feet or more Ahithr
Xenococcus and possibly a few diatoms are also shade formso

The rapidity with which the dominant algae attach themselves to leaves,
stems, and other swr.-fac'x: is amazing Planted glass and wood strips have
several hundred diatoms per sq. ma at the end of five days. Observations
of young and older leaf surfaces indicate that attachment is as rapid here,
or more so. In about 15 days there is an average of 100% coverage one cell
thick Within a month there is a tangle of epiphytes several cells thick
The type of attachment surface is important and there is also a fairly definite
order of attachment, On smooth surfaces such as glass or plastic strips,
* certain algae and Naias leaves Cocconeis lcentula and Achnanthes lanceolata
are the pioneers n aves, especl of eria and"
three are pioneers with more Cocconeis than th others


Next, Xenococcus and Pseudoulvella attach in considerable numbers
but apparently are soon covered and greatly reduced in numbers, G on
lo ng.ce, Go oisphrum and uSlna ulna attach over the pioneer spe
and all remain in lrge Mmbers Triom eon Among these the unattached
filamentous diatoms become abundant The chief ones are iFr aria co~nstrue
Fo ca inat Nealosira itaJ ca cad M. Lanuata Several oti~ --atoms and
sever lgreenit e;Z"6rF1p7..lly species of L coconiua Microsporag and
Oedogonium make up a minor part of the pe- ati toget her with a
few less abundant blue-green ?..LI,
This permanent mat continuously loses cells to the flowing water but
growth keeps it relatively uniform in thickness and composition. Deeper in
the masses of vegetation and on the under side of leaves Cocconeis lacentula
is the chief diatom and Xenococcus and AmphitliM are press n income a e
The total flora in the a areas is onlyabout 5 to 15% of that on the top
side of leaves or in better lighted areas

Some study has been made of the types of algae attaching to other
surfaces than plants in the springs Bare wood is the most common such
material, A rather wide variety of species are found on wood, S dra
is the commonest diatom along with Stigeocloni um, hisoolonium, r
certain species of SgpRol a and edoonium in the gen algae, andHt
Langba, and Chroococcus among the bluegrieens
Painted wood seems to havo a much smaller variety of species. Apparently
the chief ones are blue-greens such as Schizothrix that form tough geSatinous
layers over the paint surface, In this matthe diatom Rholodia is sometimes
abundant and a number of green algae such as I---d E~Iido attach
to it,

Iron surfaces seem to favor greea algae although the diatom S_ S
Sa attaches directly to iron, The genera Rhizoclonium and Stigoc S m
are most abundant together with the blueqggreexw Um and Anabana,
Epiphytic diaoams on the algae attached to the iron are abundanto
A copper screen submerged in Silver Springs pool for several months had
a surprisingly large amount of algae on ito All were green algae. The fila-
mentous species, MiMros ra stJagrm formed a mat in which StlaOoctytia
shroeteri Scenedesmweus o' b S diamoh um and So biua were a anto
Tere wre no aae or epip Iae present, Apparently even in
running water copper inhibits the attachment or growth of most species

Changes in the algal flora down run are not marked where conditions
are not modified by such things as shady banks, ingress of surface water,
water from other springs, or sea-water intrusion. After the first mile
Silver Springs run baccz.nc- narrow and the banks heav-iy wooded It also
receives some brown surface water down runo Consequently about 2 2/2 miles
from the boil flowering plants largely disappear probably due to reduced light
Mats of Vaucheria with some filamentous blue-green algae, and a few of the
usually domininit cdiatoms, are abundant in the shallows. The deeper channel
has relatively little plant life

Wakulla run apparently receives some saline water from springs along
it, with consequent modification of its flora toward the oligohaline type


discussed below, Weekiwachee run soon receives considerable brown surface
water and later sea-water from the Gulf. The diatoms at first dominant
are replaced by others (e iaeq turga and Sedra radians)o Then as
the run nears the Gulf a species of Rivularia becomes very abundant, and
certain salt-tolerating diataos such as osira borreri, du p.
St h n sodCiscus b 10 Witscia and Cylotela memghiniana
become abundant

In the slightly salty (oligohaline) springs, and in the rura as
they near the Gulf, the increase in salinity is first marked by the presence
of the above-mentioned diatoms and not by the dropping out of typically
fresh-water species. The only excepbon seems to be Plectonema wollei.
It disappears when the chloride level exceeds 200 ppm It- is surpsng
to find, near the Gulf, typically fresh-water genera like S ~ro a
Oedogoniumn and uotifa, aboi-ndant, and in healthy condition, along with
typically marine genera uch a and rPoloi annia
When such species as .Clad-or~ aminS Enteromo ha plumosa and
Cyclotella meneghniana ar cornoon or abundant in the boil along with the
usual frshwater do iants it definitely indicates an oligohaline condition
Two. definitely salhnr springs have been studied; Salt Springs (chlorides
2,400 ppm) and Warm S )alt .; n chloridess 9,300 ppm)o The first has several
species of diatoms comoin to abundant which are dominant in the fresh-water
springs These are _CG' .:c .:;.i, ,. Gom-.'nema 1n~ icepS and Achnanthes
lanceolata. The c&i-i': '.-,n 'Alt Spzni s are Enteromogha gumosa
Cocconeis eFntlcae Fc:vl. :'.. flbellata Nitsehia 3radox a)ndI ira
I The alg16 fol1a i1i2;t's it should epj ced on the border between
oligohaline and bra-.:'.c. o

Warm Salt Spring, li~e the other springs, has a dominant algal flora
of blue-greens and dci t:i.i3c, bu t it does not have a single important species
in common with any freshs.water spring studied. Both high temperature (86F)
and salinity are probably important, along with low dissolved oxygen. Dominant
here are the blue greens Spirui3? l Oscillatoria sp., Chroococcus
limneticus; a single &:.c nTga, Ch onea two diatormsneT
eulensteinii and Nitschia -andcrshe ensi3 er important algae, especal
in the run, are l~ucn species), Mitschia linearis, and Cocconeis
Collections from three ",suljur" springs have been studied Sulfates
are mkown to be high in two of them and all are characterized by a hydrogen
sulfide odor and a very lowy oxr.ien tension (Oo0 to 0.7 ppm)o One is oligohaline.
In all cases the dominant .Clor i is filamentous blue-green algae, chiefly
species of Phormidium and Lj . A few diatoms, notably the ubiquitous
Sn a ulna, are present, As the water moves down the run, it becomes better
oxygenatedand probably loses some of the dissolved sulfur compounds due to
the action of the abundant sulfur bacteria. Within a short distance a species
of S iro a becomes aaimdant, at first at the surface, and later on the bottom
of Tae run The blu-r.. .;:.' ::i 'ltc characteristic of the boil begin to drop
off and are replaced by larger species of filamentous blue=greens chiefly
Oscillatoria sp. and other species of Phormidium, The tolerant diatoms,
oimaphoa' giegps and Eunotia pectiaii coe in, Within a half mile (in
Beacher Spring run 7the fora is quite s im ar to that of the quieter portions
of a typical rmun The algae in deeper water, however, are still largely
blue-greens. At the surface filamentous blue-greens are the chief dominant
along with Cocconeis lacentul and GomphoMa lonicepo It is worth noting


that the species PI.eonrr-i I.rallei has not been collected in these springs
nor their sruns. /flLor l.r-Z or oxygenation the entirely typical springs
flora should certainly demwlopo

Apparently onlyly fairly M.i salinity prevents any spring from
a rich algal flora in its lower run. These studies tend to confirm
ones as to the high productivity of Florida Springs.


Detailed lists have been prepared for 26 contrasting springs. Some
sample lists follows

Number of Taxa (genera, species, varieties, forms) Identified
from all Springs Areas

CGihlor-oc:i .." (green algae) 53
Cyanophyease (blue-green algae) 32
Bacilla.iophyceae (diatonm) 66
Rhodopbytceae (red algae) 4
other groups of algae

Itaxs species of filamentous green algae cannot be identified
except when fruitingo Probably a total of 65 taxa or more present
Imany species of blue7green algae yet remain to be identified.
The total will approximate 50 taxa or more
Only a few spsc:i.: S of diatoms remain unidentified. Probably
75 taxa are present
Estimated total taxa inhabiting springs studied 190-200

List of Algae in a Sulfur Spring and Pool
(Beacher Spring)

Spirulina s a) ,.*.l-
Aphanocapsa spo
Synedra sp.
Navicula sp.
Spirulina goiontii
Chroocoecus tur:'- d:a?
Aphanothece spo
Gleocystis gigas?

Idst of Algae in a Salt Spring
(Warm Salt Spring)

Spirulina ? platensis
Amphora ci l.n:rtc.':r i
Chara honicoari. iii
Oscillatoria sp.
Nitschia gandorsheimiensis
Spirulina tenuissima
Aphanotheee nidulans


Aphanocapsa stagnina
Chroococcus spo
Phormidtim sp.
Arphora coffacformai
Amphora proteus
Achnanthes exigua
Navicula i':;io.;
Cocconeis p,:nh.ic.lus
Iyngbya spo
Synechocoecus elongatus
Amphiprora pO ldx~'r:
Phormidiu~ tenue?
Euglena sp.

List of A. ln.3 Typical of Most Springs and their Upper Run

The list is in i:prmo;- nt order of abundance, by volume of species,
The first six species constitute 80=90% of the total average volume

Cocconeis placentula
Syn.~'3a: '..i;.~ I (& vars.)
Gomphonema longiceps (& vars.)
Plectonem a o?..-.i
Acl'-.'n.i la3nceolate varo elliptica
Goi po.:;o: ,E. sphaeiophorum
Amphithrix spo
Fragilaria construens
Cladophora sppo
Spirogyra sppo
Rhisoclonium spp.
Nitsehia ;,.,il!1:.i.
Cypibella sppo
SStigeoclonium. ppo
Oedogonium sppo
Pseudoulvella sp.
Xenococcus sp,
Lyngbya sppo
Chara zeylanica (& fors,)
Vaucheria asp
Hydrodictyon reticulatum
Mougeotia spp,
Microspora appo
Scenedesmus appo
Melosira spp.


\, /i


young -.

Figure 2 :
Diagram of iepii ';tio algae on top
and bottom s~.':.- of Sagittaria
leaf (Silver Springs).

1. Coomla <
2o Aohnanthes
3, Gomphonema
4, Synedra
5o Xenooooous
60 Pseudoulvella 14 /
7, Lyngbya
80 Soenedesmu3
9o Cymbella .
l0, Fragilaria
11o Microspora
12o Stigeoolonium
13o Melos 14o Amphithrix


In the previous semi-annual progress reports, patterns of
insect distribution in Homosassa and Weekiwaohee Springs were
discussed. Variance analysis of this distribution showed that
population density as well as species number increases with
distance downstream from the spring boils. The decrease in
species number as the estuaries are approached was also noted,
It was suggested that this differential distribution is cor-
related with environmental gradients. Graphs of these gradients
based on additional data are shown below.

Figure o3 Dissolved oxygen in parts per million Maximum-
solid line, range dotted line0 Maximum repre-
sents the mean value of DO samples taken through-
out the year within two hours of the suns zenith,
Huliout is the maximum minus the minimum values, all
of which were taken before dawn. Each locus on the
curves represents a collecting station which is
designated by an encircled number,

a) Dissolved oxygen in Homosassa Springs

DO in

p pm/

Boil Distance in miles Estuary

b) Dissolved oxygen in Weekiwaohee Springs

DO in

I- -T- -__- ___ ___

Bol Distance in miles E----t

Boil Distance in miles Estuary

Figure 4 Chloride values in parts per thousand, Maximums.
solid line range-.dotted line, Maximum is the
mean value for chloride samples collected at
high tide,f Range means the difference between
high und low tide values and thus shows the tidal
;' ,\.. t1;,

a) Chloride concent.:-,tions in Homosassa Springs


Distance in miles Estuary

b) Chloride ,- c,-.:e-n.rAtions in Weekiwachee Springs

Cl in
ppt .}



SpeOies Distribution
The total number of species per station is shown below $.n
figure o TabWI 3 shows the number of species per order for
each station and total number of species for each spring system.
No seasonal variation in species composition has been noted al-
though not all of the data have been consolidated

Cl in


^*v W


Figure 5, Number of species plotted against distance down run.

a) Homosassa Springs

Noo of


b) Weekiwaohee Springs

Noo of

Boil Distanoe in miles Estuary

Table 8

Number of species per order found at each
tion and total number of species for each




station number
Boil 12 3 4 5 6 7 8

4 55232110
9 9 9 71 3 0 0 0
5 77454 331
4 9785542 0
1 1 1 00 0 0 0 0
3 77053211
1 020 000000

Total for river-s .I4

station number
Boil 1 2 3 4

2 1011 4 1
1 7 15 10 2
6 9 15 3 1
4 7 8 5 3
0 1 1 1 0
3 7 4 1 1
1 4 3 0 0
S- 4 o0 Cv
Total for river 74

It is seen that curves for both maximum oxygen and oxygen
range seem to resemble the species curves fairly well; at least
in the upper parts of the rivers It seems unlikely however~
that oxygen in excess of 6 or 7 ppm could affect the insect dis-
tribution al'BouP'.', the low DO in the boil areas proper may be
limiting. Investit.r.tions of other lotic systeras have shown that
large n.m'>..rs of s:jeoiea are able to exist in situations whose
DO content never s. .: ..s 6 or 7 ppmo
It apgr.;". fairly ~.. :-i that the increase in chloride con-
tent in the lower .r :'"1 .-:.rrg to control insect distribution
It is difficult to det. &.:'o!o, however, whether the total ohlor=
ide content ',.tio or .'.ll. effect is more IrJo5;.t.. in this
respect, In .-..:.".-' it seems that chloride concentrations in
excess of 0,5 p;.- may serve as a limiting factor In waters
containing more that 1,O ppt chloride typical brackish water
forms are most common Some of these are Enallaa durum (Hagen)9
3T hoQg l~mbeus. (8. .!.:' ), iand the brine fly -~ ET dra sp

The hypottesls that spring boil ,nr,: '. ... are stable was pro-
posed in pr. Springs coa"uSri this :.:''.ral pattern The oxygen range curves
seen in fL ,u.e ,'. .I'.'.' Leotions of the stability of the en=
vironment; these show the boil regions to be the most stable
parts of the systems T': extrw%.ly small seasonal changes in
temperature in these re:...ion; as seen below also indicate sta-
bility in '- .' :, .

Boil 1/2 way down Estuary
Homosassa 1 8 8
Weekiwachee 1 5 6

Seasonal run.-: of "-..,..-rature in degrees centigrade

Evidence of 1.-,-; -*i stability of insect populations is
shown by the fact that the numbers of individuals per sample
collected during d i'-;'*.*r,: t seasons in the boil areas of both
rivers do not d.':,.?: siEnifioantly. This is shown in the table
.Tiabl 9o i..'-. -rais of variance of number of individuals
collecv,:,d in boil ar-2.o

Stations cot..p.-.-.:A j'nte e..~.:ps insects/S sweeps p value

Homosassa April 25 3,8 A05
Homosassa NovT 25 10O0

Homosassa April 25 3o8 >005
Weekiwachee Jiuly 25 7o4

Homosassa Novo 25 l0o0 >3, 0
Weekiwachee July 25 704

However, a significant dt... CiVence is found in the numbers of
individuals per sample collected at different ,ea=oo.s. down=
If those species having the best chance for survival are
those whose optima :"'. within the small range of environmental
fluctuat'ic(. .'. ":bi',;:. ,.- the boil l ,.:a, ., the dif'ca,,o c in
numbers of species bet-; .; boil and downstream areas may be,
at least in part, a fu.noton of this environmental stability,
Ordinarily, the red ution ~' sncies variety by l,-i.Lting
factors results in lurge populations of the few surviving
species since t:-;.: arie now relieved of a certain amount of
competition and predatio' p..c:.. .,eo This is not the ease in
boll areas w.::,.. smell populati.'',.v of the existing few species
is the rule r,0....;:' .. .tion limiting factors were diBs
cussed in the ..'. ': -- ,... : : port and include available food
and predation An at .:..;i:: will be made to evaluate the Import-
ance of these ': the 1. report

1' : C l j 24 o

Productivity .v ,c....,i and atmospheri diffusion in streams

Heretorio-, where the L ..:i.ait method of measuring productivity
has been applied, atmospheric diffusion has been assumed to be
constant during both Ilgl': and 1...:yo Because the gradient flow
method pYoml ,i'o be of Lr-...,1 applicability to springs, to
coral reefs (C,-,.: : ICs and T.So Austin 1949 Organic
productivity of an atoll, Transo Pmero Geophysics6 UL.-:To1., volo
20 (2), pa :15-." "o) to Cetuaries, and to st.at:raI,a it is necessary
to ':n:.i:7 k.- the ,.T, 1 ;' day lb.':ng '7 in at:i.,.o.phJ rice C. ..'J.o.l.:
as a further Irefl .....,t.o It may be expected that the rate of
diffusion of o ;,:..-;, into or out of a flowing system in the course
of a 24 hour cycle will n -:.poend primarily on the oxygen concentration
in the flow It should be possible to determine the function
relating the rate .," dif-.l:.I;.on of oxygen per area to oxygen
concentration for a particular flow system so that thereafter
a correction r;i.,>; be .;-pl.T.od to production measurements in the
same place

Considr;:'k an ..,-;.:0 square unit a.,.: of stream community
Let P be the rate of photosynthesis, R the rate of respiration,
D the rate of di .-.;..; : of o;0:jcu into the stream and 0 the
oxygen concentration. 'n.

dO/dt a P o R D

If the dmir.'.-o, .: "--.,:.:; .:',,i,':me and nighttime oxygen concentrations
is very mail then bocth ha n.. Dr c.n be assumed consmta.-: as ohas
been done ho~'-- .., ; in the ap.ii:l Ao 'n the diff.'r'.; be to.on
day and night vc.1l;::;. of >O/dt is the productivity(uncorrected).
dO/dt can always be evaluated by measurements at two stations on
a flowing -:'t.;-; at an interval of time between upaL:.:e,:;
inul.icraT,.- and dovmst;ream ii'.. m.'ei, ~iti1 to permit the water to

If '-y;:;.'i content is alternately .?.y ml.,-:.-Psaturated and under-
saturated the c conOentration is .:.n:ntJi.';d in u ,r11'.-'iu~aU
with the atmospe1'- each day., ^t this time D can be ,::*,;ur.::;d
zero and the xc,.f; of the equation evaluated using other', ;ieasurementso

In springs or in the majority of streams which do not become
supersaturated it is necessary to evaluate Di for a given
section of cstr.:'-r .., f th i-.cl.londr nt "' .-i~':cr,,; -.;, of R and P with
black bottle c::,'c:riments etc. Fb~ silver Springs at night at
the downstream sta ::.. d6/dt is .5 ppm, which amounts to 68 mg/
sqo ft/hro R-cp,.:-iation bn d on lo.1.1 jA'v- is 18 mg/sq. fto,/hr
-substitution in the above equation gives

Di a 68 0O-18 W 86 mg/asq ft/ hr when the oxygen
concentration is ,.0 ppm and
the temperature is 72 dego Fo
This calculation of course i.l ...ts fish respiration awli..*...*,
magnitude has yet to be determined


The diffusion rate into Silver Springs during the day as the
values at the downstream station reach 5 ppm may be expected to be
less than at night when on.y.on concentration is at 3 ppm, Using
night values of di:.' :'.lt as has been done in production
measurements thus '..,'....'estimates the production The maximum
error that ':.:d .ult j .- 1':-.!yi-,: that b .o,! diffusion in Look plr.co
during the day r-.,.: -i;s t: 1j,i)00J lbs./aereo The error is
probably T.,"...ui; half this

Another kind of loss involving oxygen bubbles in the shallow.
water on :>iw;;.,..r 0:.J's -.1 ch rise to the surface without dissolving
has been v l C.V. r 1r o: -... i. ino t yet '.. .i -,

Timrnda ,~re. .-.l.:'. j The o.timum efficiency for maximum
awwer ..;.;, in. ,,. al and biolorical
by II T.o "C,,: tri.l r0 C,'. Pinke-ton (Dept. of Chem. rng.)

Loe"OU:;Fe the ;: ...,' .:v.. of t111,iil.mo:yn.rios does not indicate
the mnagnittidi,.- of .;.-:ts of entropy increase, a general theoretical
expression has been deri' ;0 with the use of concepts of steady
state therlyl~'7i. -~ ,ics to -:i.;., ly a"'-tji:.:. W,',od regulator" to
mathh "times a.,".j ., Our proposition is that natural systems
tend to operate at that efficiency which produces a maximum
power output, which is leas than the maximum efficiency An
expression icat3J .l ;-.. 'iolency .r:1r ;, :.-~,1miL power was developed
which was aplld.'-d to: (a) AtIi:,o<.'s r ':.!in.r,, (b) a water wheel
turning a gri; ,-; ..- : (3) one b~, ..t k !, :,.1 .n.. another 7..Ittery,
(4) a thern n.-: .-u -,r.,. ; Cn.n ,.,ctrci mai.;., (5) a ther-mal
diffusion cne,~h'n,, (6) the metabolism of an organism, (7) food
capture by an oi... ,:-,. its Raintenance, (8) photosynthesis,
(9) a self sustaining- climax community, (10) growth and mainte-
rina7o of a civilization. The essence of the general i'r,;vation,
,.n.. the -'1))ll. .cation to a :i:';r.,i ; state .1..',:,j.;rity .like Silver
Springs is extrace'-. i. '-., the ri,:....-.i. t and summarized below.

A driving "i *. '.'. coii. to a dri ,...* ou can be G ..:.
at one extreme of no work :..,ir Ing acco.;:-i.l!.i.:d in the dri,:r,;
process or at another extreme of so great a strain that the system
is stopped T.'.e.; extremes, the first with 0 efficiency and the
2nd with 100% efficiency produce a zero useful power output"
If power output in the form of growth and maintenance is at a
premium in the survival of y-stci.ain nature, they may be expected
to be adjusted for an ,p;--,..;!u efficiency that is not maximum The
expression for efficiency in terms of T,.:,.xs.mum power adjustment is
derived below:


T dS/dt a JlX1 .f J2X2 1 774. z2

J1 a (j + cf2)X1 cfX2 /553

J2 -cf.X- 1 cX2
J _Cf: CXX

f g Xix1 when J-: 0 When X 0 J2 a X

Jg A when '. 1 and Js g 0

If R x/f

E Pg2/P
at rmfar-X.mrny, -,- :.:;- .: ; (' q ..2...i,,.' ). R a 1/ and

E 1
2(1k 2 ./rAf2
The ,;it-.; -I ns .. c :. the '. Il'.1!i', ,
J is the output f .. is the '...ja.. "for_ the hQI f te output
process s; '- is the 1 ~::f'' .u:.n; Xp is the input the~ c:.;, mii l /
"force"; 'is the 1 ., iv .i":.n value of Jg when ;y; is 0-
c is the :eire" -. "vvity iv : the value of J n X isj O f i
the factor of !,'. -..'t onality re~'i:':7. X1 and Xp. E is the
output off:! ;.-. r P] is the useful power input; I is the useful
power output. t is i~.-'h tlTgc T is the absolute to,, rature; and
S is the siCt.,:r,,, (.o. methods n-..:> see Denbigh, ie e, '5;..
Ti'lieri.i l/..;:.. i .. '.,, S /- r state "~ :A' :U T.-"2.-n, 103 ppo)

Hlow let us apply this t, an ecological st,-,adc' state aonmunity
like Silver Springs vi.-'e the *:'.;.3 Oic driving ".u. '-.-"' is the
absorption of r.'ir.tion .I. the Sn.ct .c u i.' the maintenance
and self rep3 a.:,. of the comunmityo If all tho power output
is replacement, the :;: :c..,:.-,,' e c .:.:..,:. and thus the m,:.s..: .-,.m size
that can be m!.,i': ."1 in s:hc id,; state occurs when the thermodynamic
force ratio adji. :; ....-~: (h) is 50% One can vi..ia.:-. that a
community not in climax would have a net growth output and thus
would increase in (C ..;,:.Ity until its ...ni. ..ai replac:.r-t
requirement bE:l,:n.-..:'1 i- th and dissipative rates at t 50%
Communities which did not follow this scheme might be expected
not to have equal survival value8 It is thus suggested that the
largest size system that can be maintained in steady state on a
given potential S .:, .., source is one whose replacement costs
approach 50/: of trhe ; i.; This result suggests that there
is a definite ra'.. to '.-. c---ected between standing .1p: and
productivity(as c-u .17.:'ca obse-vc,. ) that corresponds to that
rate of entropy ine..'. .t results in u~:i..'r.,i! po_,': 2.9 Simlar
ratios might be c-T.-.'.: wi)ira.y variant types of comrMunritiese

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