g ir‘"“-—w“-mM-W.0MW‘ i
‘D

THE INFLUENCE OF A POWER PLANrS
COOLING WATER EFFLUENT ON THE -
PHYTOPLANKTON POPULATIONS AND _

CORRESPONDING PRIMARY PRODUCTIVITY.
NEAR THE WESTERN SHORE OE -
LAKE ERIE

Thais for the Degree Of M. S,
QEQHIGAN STATE LENWERSITY
THOMAS. VERE KREH

£923 '

ABSTRACT

THE INFLUENCE OF A POWER PLANT'S COOLING WATER
EFFLUENT ON THE PHYTOPLANKTON POPULATIONS
AND CORRESPONDING PRIMARY PRODUCTIVITY NEAR
THE WESTERN SHORE OF LAKE ERIE

By

Thomas Vern Kreh

Phytoplankton populations near the western shore of Lake Erie were
studied at one-month intervals from 16 April to 12 November 1971 in
conjunction with the first year operation of a steam-electric generating
facility. Estimates of phytoplankton numbers, volumes and species
composition were made from samples collected in cooling water sources
which included the Raisin River and Lake Erie (three stations) as well
as the plant discharge canal. Gross primary productivity (GPP) was
examined approximately twice per month using the light-dark bottle
oxygen method from 21 May 1971 to 14 June 1972. Also experiments
were conducted to determine the effects of transferring water samples
in various parts of the cooling system on GPP.

During the study period, algal concentrations near the Monroe fossil
fuel plant varied from 1,791 X 103 counts/liter to 23,289 X 103 counts/
liter. There was no statistically significant difference between pre-
operational densities (1970) and post-Operational densities (1971) for
the spring, summer and fall growing seasons. In addition, no yearly
statistical differences in total algal cell volume were evident among
the three seasons.

Among the three major algal classes, diatoms were most abundant in
April, May and November; green algae predominated in June and July; and

blue—green algae dominated in August. No differences in class volumes

Thomas Vern Kreh

could be detected for 1970 and 1971 except in the case of higher summer
diatom volumes during 1971.

A total of 184 phytoplankton species were observed during the study
period in 1971. Mere species comprised a significant percentage of the
total volume in 1971. The river and discharge stations consistently
contained higher numbers of species and greater species diversity than
lake stations.

At the lake station, higher GPP at all measured depths below the
surface occurred in 1971 compared to 1970, independent of plant Operation.
This was probably caused by greater light penetration in 1971. Average
midday hourly GPP was highest at the discharge site and lowest in the
river. Community respiration was lowest at the lake station and highest
at the river station. Ratios of daily GPP and community respiration were
lower at the discharge site during summer and fall 1971 relative to 1970
ratios. Cooling water use appeared to directly affect these ratios.

Cooling system experiments indicated that stimulation of intake GPP
occurred at ambient temperatures of 16 C (A t - 9 C) but at intake
temperatures of 26 C (A t - 9 C), inhibition of GPP resulted. In both
experiments, cooling water returned to the lake environment exhibited a
GPP similar to the lake GPP which indicated rapid community recovery.
Community respiration of intake water was generally higher at temperatures
of the discharge canal but decreased when incubation of discharge water
occurred at the lake station.

GPP was slightly stimulated in the discharge canal and community
respiration was stimulated even more so. Most of the community respira-

tion appears to be heterotrophic rather than of phytoplanktonic origin

Thomas Vern Kreh

because algal biomass accumulations occured in discharge water comcomi-
tant with higher than expected respiration. Apparently the cooling
system acted to stimulate GPP and even more so the heterotrophic consump-
tion of organic detritus so the net effect on the lake was to decrease

organic input without effectively altering GPP in the lake proper.

THE INFLUENCE OF A POWER PLANT'S COOLING WATER
EFFLUENT ON THE PHYTOPLANKTON POPULATIONS
AND CORRESPONDING PRIMARY PRODUCTIVITY NEAR

THE WESTERN SHORE OF LAKE ERIE

By

Thomas Vern Kreh

A THESIS

Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of
MASTER OF SCIENCE

Department of Fisheries and Wildlife

1973

I wish to
less problems
the preparatic
tributions pm
as Iy other g
Murphy,

To all I
ening discus
and J. D. Lu
also like tc
Wish t0 thaI
Faker and C

Supper
“a? of a c,
indebted f
UniveTSity
Scienee FE

And «
lupin,ti
an deep};

friends 7

ACKNOWLEDGEMENTS

I wish to thank Dr. R. A. Cole for his special attention to the end-
less problems on my project and for his criticisms and suggestions during
the preparation of the manuscript. I particularly appreciate the con-
tributions provided by my major professor, Dr. E. W. Roelofs, as well
as my other guidance committee members, Dr. C. D. McNabb and Dr. P. G.
Murphy.

To all my fellow graduate students I will always recall the enlight-
ening discussions in which we engaged, and especially to T. J. Edwards
and J. D. Lu for their willing assistance in the field work. I would
also like to thank J. Ashley and R. Freeman for their field support. I
wish to thank K. Hunter for diatom slide examinations as well as T.

Ecker and C. Annett for providing chemical data.

Support was made possible through the Detroit Edison Company by
way of a contract with the Institute of water Research to whom I am
indebted for the study opportunity. Partial support of the Michigan State
University computer laboratory use time was made possible by the National
Science Foundation.

And to my wife, Kathleen, for her constant encouragement and
inspiration for her ceaseless perseverance in typing the manuscript, I
am deeply indebted. Appreciation is also extended to my relatives and

friends for their interest and concern.

ii

 

INTRODUCTION
DESCRIPTION '
MATERIALS Ah"

Field E
Data A:

RESULTS .

Comun
E

Com

TABLE OF CONTENTS

Page
INTRODUCTION . . . . . . . . . . . . . . . . . . . . . 1
DESCRIPTION OF THE STUDY AREA . . . . . . . . . . . . . 5
MATERIALS AND METHODS . . . . . . . . . . . . . . . . . 12
Field and Laboratory Procedures . . . . . . . . . 12
Data Analysis . . . . . . . . . . . . . . . . . . 16
RESULTS . . . . . . . . . . . . . . . . . . . . . . . . 18
Community Structure . . . . . . . . . . . . . . 18
PhytOplankton Abundance . . . . . . . . . . . 18
Total Numbers . . . . . . . . . . . . . 18
Total Volumes . . . . . . . . . . . . . 18
Total Carbon Content . . . . . . . . . . 21
Class Abundance . . . . . . . . . . . . . . 21
Species Composition by Class . . . . . . . . 23
Blue-green Algae . . . . . . . . . . . . 23
Green Algae . . . . . . . . . . . . . . 23
Diatom . . . . . . . . . . . . . . . . . 24
Other Species . . . . . . . . . . . . . 25
Variety of Species . . . . . . . . . . . . . 25
Total Numbers of Species. . . 25
Annual Differences in Species Abundance. 32
Species Diversity and Equitability . . . . . 34
Seasonal and Yearly Statistical Comparisons . 36
Community Metabolism . . . . . . . . . . . . . . . 38
Effects of Depth . . . . . . . . . . . . . . 38
Spatial and Temporal Effects . . . . . . . . 4O
Midday Productivity . . . . . . . . . . 40
Midday Respiration . . . . . . . . . . . 4O
Diurnal Gross Primary Productivity . . . 42
Daily and Seasonal Gross Primary
Productivity . . . . . . . . . . . . 42
GPP/R ratios . . . . . . . . . . . . . . 45
Source of Power Plant Effects . . . . . . . . 47
DISCUSSION . . . . . . . . . . . . . . . . . . . . . . 54
Changes in Community Structure . . . . . . . . . 54
Annual Variation in Gross Primary Productivity . . 54
Influence of Power Plant Operation on Intake
Community Metabolism . . . . . . . . . . . . . 57
LITERATURE CITED . . . . . . . . . . . . . . . . . . . 62
APPENDICES . . . . . . . . . . . . . . . . . . . . . . 65

LIST OF TABLES

Table Page

1 Mean annual percent composition of domi-
nant species examined from May to November
1970 (Marcus, 1972; X < 12 of the total
algal cell volume) . . . . . . . . . . . . . . 26

2 Phytoplankton species enumerated from April
to November, 1971, with their mean annual
percent composition by station (X < 1% of the

total cell volume) . . . . . . . . . . . . . . 27-31
3 Number of distinctly different species accumu-

lated temporally and spatially with the mean

number of species at a station in parentheses 33
4 Results of a two-way analysis of variance

testing various parameters by the year (Y)

and station (S) factors with the significance
probability of the F-statistic based on the
interaction mean square (Y x S) . . . . . . . 37

5 Mean gross primary productivity (GPP) and
community respiration rates (R) estimated
during the midday interval in MgC/m2 lhr
(X i 1 S. E. ). . . . . . . . . . . . . . . . . 41

6 Mean daily and seasonal gross primary
productivity during 1971 and 1972 (gC/mzlday
and season) compared to 1970 means (Marcus,
1972) . . . . . . . . . . . . . . . . . . . . 44

7 Ratios of gross primary productivity/community
respiration at the sampling stations (based on
gC/m2 /day) with seasonal and yearly averages
for the years 1970 and 1971 . . . . . . . . . 46

8 Comparative cooling system gross primary
productivity (GPP) and respiration (R) in
the upper cubic meter on two dates (X i
l S.E.) . . . . . . . . . . . . . . . . . . . 50-52

9 Comparison of various parameter that could
potentially influence phytoplankton populations
and primary productivity in the study area during
1970 and 1971 (GPP and R means are for the year
period). . . . . . . . . . . . . . . . . . . . 55

iv

Table

 

 

 

 

10 Mean observed and expected values for para-
meters associated with the Monroe Power Plant
operation in 1971 . . . . . . . . . . . . . .

APPENDICES

Al water temperature elevation in the cooling sys—
tem at the one meter depth, Raisin River flow,
and pumping rate at the Monroe Power Plant site
recorded at the time of plankton collections and
primary productivity studies . . . . . . . . .

A2 Periodic total residual chlorine concentrations
(ppm) condensed from plant operational reports
Of 1971 mid 1972 O O O O O O O O O O O O O 0

A3 Physical parameters recorded at the time of
primary productivity studies . . . . . . . . .

A4 Measured daily solar radiation in conjunction
with primary productivity studies of 1971
“d 1972 O O I O O O O O O O O O I C O O O 0

A5 Meteorological information pertaining to
plankton and productivity studies (1971-72)

A6 Mean total algal numbers (1971) (X 103/1iter)

A7 Mean total algal volume (1971) (X 10-4 ml/
liter) 0 O O O O O I O O O O 0-. I I O O O 0

A8 Mean individual phytoplanktonic volume (1971)
in p O O O O O O C C O O O O O O O O O O O 0

A9 Mean volume of blue-green algae (1971) (X 10—6
ml/liter) O O O O O O I O O O O O O I O O O O 0

A10 Volumetric dynamics of Aphanizomenon flos-aquae
(x 10-5 Ill/liter) o o o o o o o o o o o o o o o

All Volumggric dynamics of Anacystis incerta
(x 10 Ill/liter) o o o o o o o o o o o o o o 0

A12 Volumetric dynamics of Anacystis thermalis
(x 10-5 nllliter) o o o o o o o o o o o o o o 0

A13 Volumetric dynamics of Anacystis cyanae
(XlO’Sml/liter)...............

A14 Mean volume of green algae (1971) (X 10--6

ml/liter) . . . . . . . . . . . . . . . . . . .

Page

58

65

66

67

68

69—70

72

73

74

75

76

76

77

78

 

lable

A15

A16

A17

A18

A19

A20

A21

A22

A23

A24

A25

9&6

A27

A28
A29
A30

AL31

Vol‘
(X

Vol
10"

Table

A15

A16

A17

A18

A19

A20

A21

A22

A23

A24

A25

A26

A27

A28
A29
A30

A31

Volumetric dynamics of Mougeotia elegantula
ml/liter) O O O O O O O O

(x 10'5

10-5 ml/liter). . .

Volumetric dynamics of Pediastrum simplex (X

10-5 ml/liter). . .

Mean diatom volume (1971) (X 10—

Volumetric dynamics
(x 10-5 ml/liter) .

Volumetric dynamics
10-5 m1/liter). . .

Volumetric dynamics
(x 10-5 m1/liter) .

Volumetric dynamics
(x 10-5 ml/liter) .

Volumetric dynamics
(x 10“5 ml/liter) .

Volumetric dynamics
(X 10'5 ml/liter) .

Volumetric dynamics
(x 10-5 m1/1iter) .

Volumetric dynamics
10'5 ml/liter). . .

Volumegric dynamics
(X 10 ml/liter) .

Mean number of species at each station in 1971
Mean phytoplanktonic diversity (1971) . . . . .

Mean phytoplanktonic equitability (1971)

 

 

of Coscinodiscus radiatus

6 m1/liter) .

Volumetric dynamics of Pediastrum duplex (X

 

of Melosira granulata (X

 

of Stephanodiscus astraea

 

of Stephanodiscus tennis

 

of Fragilaria capucina

 

of Fragilaria crotonensis

 

of Tabellaria fenestrata

 

of Cryptomonas ovata (X

 

of Ceratium hirundinella

 

Page

79

80

80

81

82

82

83

83

84

84

85

85

86
87

88

Analysis of variance of factorial design comparing

year, station and month factors . . . . . . . .

vi

90-92

Figure No .

 

10

Figure No.

1

10

LIST OF FIGURES

Page

Map of the study area in relation to
western Lake Erie . . . . . . . . . . . . 6

Map of the study area including positions
of the sampling stations . . . . . . . . 7

Mean water temperatures at the 1 m depth
recorded in conjunction with phyt0plankton
callections O O O O O O O I O O O I O O O 10

Location of ig_situ sites in the study

area used for the cooling system experi-

ments of 18 August 1971 and 14 October

1971 . . . . . . . . . . . . . . . . . . 15

Mean total algal numbers, total algal cell

volume and total cell carbon in 1971 by

station (total numbers ; total volume

-----; total carbon __,__) . . . . . . . 19-20

Mean percent composition of algal classes
on a volumetric basis for the study period
of 16 April to 12 November 1971 . . . . . 22

Mean phytoplanktonic diversity and

equitability by station on each sampling

date (hatched boxes—-diversity; clear
boxes--equitability) . . . . . . . . . . 35

Depth profile of average gross primary
productivity measured during the midday

interval for the years 1970 ( ) and

1971 ( ----- ) at the three stations in the

study area (X i l S.E.) . . . . . . . . . 39

Average gross primary productivity estimated
during the diurnal period on two study dates
(lake . ; discharge —---; river ) 43

Schematic diagram of control and transfer
light-dark bottle combinations used to

measure effects of the power plant cooling

system on Ig_situ gross primary productivity

and respiration . . . . . . . . . . . . . 49

vii

INTRODUCTION

This paper focuses on the effects of a steam-electric generating
facility's heated effluent on phytoplankton populations and corresponding
primary productivity in waters near the western shore of Lake Erie. The
post-Operational data presented in this report is compared to pre-operational
data gathered during 1970 studies (Marcus, 1972). This report is one in
a series of terrestrial and aquatic impact studies embarked upon to
assess the influence of a fossil-fuel plant on the physical, chemical,
and biological limnology of the study area.

In recent accounts, western Lake Erie has been described as highly
enriched and photosynthetically very productive. Davis (1964) reported
a consistent increase in phytoplankton numbers off Cleveland, Ohio,
between 1920 and 1963 and associated this change with increased nutrient
additions. Although no comparable data has been reported for the western
basin, some reports have suggested that standing crops have also increased
there during the past century. Abundances of phytoplankton are presently
concentrated enough to cause periodic nuisances (FWPCA, 1968) at beaches,
marinas, etc. Recent (1969) summer beach samples from the western shore
of Lake Erie averaged 4,000 counts of algae/ml of water (Mich. DNR, 1970).
Further alteration of the phytoplankton community may occur at least
locally in the vicinity of a large steam-electric station. Particularly
in eutrophied environments the potential changes may aggravate nuisance
algal growths. Given a mandmade temperature increase, biologists have

predicted potential changes in: (1) species composition, (2) average cell

2

size, (3) biomass, (4) species diversity, (5) gross primary productivity
(GPP), and (6) respiration (R).

All biological processes involve complex temperature-dependent
physical and chemical activities. Photosynthetic rates are controlled
by two physiological processes, i.e.: photochemical and thermochemical
(Giese, 1962; cited in Hoar, 1966). Light absorption by photosynthetic
pigments (photochemical) is limited at lower temperatures while enzymatic
reactions (thermochemical) are controlled at relatively high temperatures
when the chlorOphyll mechanisms are saturated with light energy. Within
the algal species' tolerance limits, metabolic rates nearly double with
temperature increases of 10 C (the Q10 value). Patrick (1969) has pro—
posed that as upper limits of temperature tolerance for an algal species
are approached, cellular respiration will comprise most of the metabolism.
With increases in temperature, the respiration by the whole community
(R) should increase more than gross primary productivity (GPP) because
respiration is a characteristic of all life whereas GPP is only a function
of plants.

A considerable body of data is now accumulating on thermal effects
in once-through cooling systems; particularly in marine environments.
Generally, thermal wasting stimulates primary productivity until some
upper ambient temperature is reached. The point at which temperature rise
becomes inhibitory depends at least on the thermal history of the environ-
ment, the change in temperature across the condenser and the kinds of
organisms present in that aquatic system. Narinner and Brehmer (1966)
examined temperature changes (A t) as related to primary productivity.
At ambient water temperatures of 10 C and in the presence of a A t of
5.4 C, carbon-14 uptake rates increased over controls but at ambient

temperatures of 15 C and given the same A t, photosynthetic rates

 

éecreased. Morg
uptake (0‘14) 0‘
uptake was inhi‘
of 8 C). An es
when transferri
15 C) was repor
oxygen evolutiI
compared to di
Changes 1

and diversity
induced in Mr
it is assumed
mlEther of spE
0f heated efj

the trend in

b°th intake
and reportec'

year. More

in 10th 3?
factor. Pa
Creek arti:
above Vere
Other an“
algae in I
those ’50“
having ma

Hon
“hen the

This may

3

decreased. Morgan and Stross (1969) reported that stimulation of carbon
uptake (C-l4) occurred at or below ambient temperatures of 16 C but
uptake was inhibited at temperatures of 20 C or greater (given a A t

of 8 C). An estimate of the recovery ability of enclosed communities
when transferring post-condenser water to pre-condenser conditions (A t =
15 C) was reported by Suchanek (1971). Essentially no differences in
oxygen evolution occurred between discharge water incubated at the intake
compared to discharge water incubated in the discharge canal.

Changes in community structure as measured by species composition
and diversity indices are also useful indicators of the degree of stress
induced in phytoplankton populations subjected to condenser passage.

It is assumed that an unstressed environment will support the largest
number of species. Few field studies have dealt with direct influences
of heated effluents on community composition. Gebelein (1971) followed
the trend in the number of phytOplanktonic diatom species collected in
both intake and discharge samples at Northport, New York (A t = 15 C)

and reported no significant differences between stations throughout the
year. More studies have emphasized thermal effects on periphyton communities
in lotic systems with the temperature appearing to be the controlling
factor. Patrick, st 31. (1969) found that diatom species of White Clay
Creek artificially exposed in §i£g_to temperatures averaging 34.2 C or
above were eventually replaced by blue—green algae in a few days time.
Other authors have described the species composition of thermally adapted
algae in hot springs but these exceptional environments (60 C) are unlike
those found in the vicinity of power stations releasing heated effluents
having maximum temperatures of 35-40 C.

Morphological changes, such as decreased cell size, may be effected
when the upper temperature limit for a species is exceeded (Patrick, 1969).

This may be particularly important to algae taxonomy in a given study area.

4

Generally, the question to be addressed is whether the intermittent
operation of a 3,200 megawatt (MW) fossil fuel plant has influenced the
phytOplankton pOpulations in a stimulatory or inhibitory manner. But
specifically the following hypotheses will be tested and are discussed
herein: (1) numbers of algae will increase, (2) cell biomass (volume)
will increase, (3) species composition will change from diatoms to green
and blue-green algae, (4) the number of species will decrease, (5) the
species diversity will decrease, (6) gross primary productivity will
increase, (7) community respiration will increase even more than gross

primary productivity, and therefore, (8) GPP/R ratios will decrease.

DESCRIPTION OF THE STUDY AREA

Previous reports have described the study area in detail (Cole,
1972; Marcus, 1972; Nalepa, 1972; Parkhurst, 1971) regarding the physical,
chemical and biological character determined in a comprehensive survey at
the Monroe Power Plant site. The study area has a variety of habitats
that could be affected by the discharge of cooling water. The power
plant is situated on the filled-over, former Raisin River delta on
western Lake Erie near Monroe, Michigan (Figures 1 and 2). The intake of
the once-through cooling system is designed to take all river water
available along with supplementary lake water which is drawn through
the old river mouth. A dredged discharge channel, 2.5 km long, 150 m
wide and having a mean water depth of 7 m, was constructed in 1970 to
receive the cooling water once it passes through the condensers.

The Raisin River receives municipal wastes and reprocessed paper
wastes from the city of Monroe less than 2 km upstream from the intake
for the cooling system. A yearly dredging schedule is maintained in the
last 2 km of river to remove organic sludge buildups and permit the pass-
age of large vessels into a small harbor near Monroe. An industrial
plating operation discharges wastewater 1 km above the plant intake where
it is mixed with river water. All of these river disturbances cause a
distinct difference in the quality of the water between the river and
the lake. The quality of water masses in the western basin of Lake Erie
are probably most influenced by inflows from the Detroit River (95%) and

the Maumee River (22). Wind-induced currents most frequently move

   

DETROIT RIVER

Figure 1. Map of the study area in relation to western Lake Erie.

a):

 

Monro.

   
        

/{
Raisin
lUvor

VVostorn
Lake

LoIPl' one
on o E".

Bay

6m

/ Ilkml

é Prov-lung Current

0 Plankton
P Prlmsry Productlvlty

06

Figure 2. Map of the study area including positions of the sampling
stations.

8

northward along the west shore and through the study area. This pro-
duces uniform vertical mixing of various parameters in the study area.

The Monroe Power Plant first began Operating on May 3, 1971, but
intermittent Operation of Unit I (the plant is composed of 4 separate
units) was a common occurrence in 1971. Ranges in electrical output
from 0 to 830 MW of electricity were primarily a function of the frequency
of repairs. 0n five study dates in 1971 and 1972, the plant was not
generating electricity but cooling water was continuously pumped despite
the fluctuations in electrical output. Each unit has a set of three
intake pumps which are activated as electrical units are put into
operation. Temperature changes across the condenser during the study
period varied from 0 to 10 C while mechanical effects in the cooling
system remained the same except on the last few sampling dates when the
pumping rate was doubled from 23 m3/sec to 46 m3/sec (Appendix A1). A
A t of 10 C was approximately the highest temperature change and this
corresponds to maximum electrical output. The discharge canal was not
subject to flowing conditions in 1970 as it was in 1971. Exchange with
lake water in 1970 occurred only through wind-generated mixing and
no river water entered the discharge canal. The water pumped through the
cooling system takes about seven minutes moving at 2 m/sec with a con-
denser contact time of approximately 12 seconds.

Chlorine is introduced upstream from the condensers to control bac-
terial and other growth. Chlorine is applied for one-half hour periods
twice daily during winter, spring and fall and four times daily in the
summer. The desired residual concentration is less than 0.5 mg/liter total
residual chlorine at the head of the discharge canal. Total residual
chlorine, as indicated by plant records, is normally less than 0.50

mg/liter but has been as high as 0.75 mg/liter (Appendix A2).

9

Water temperature in the study area increased steadily at all stations
from April until July, 1971, and then declined gradually into November,
1971. Inshore stations1 had higher water temperatures than the lake
stations (Figure 3). The river usually had a 1-2 C greater temperature
than the lake water at the l m depth. The discharge exhibited wide
variations in temperature relative to intake waters during the study
period (Figure 3; Appendix Al) because of intermittent plant operation.
The change in surface water temperature within the discharge canal was
minor (:_2 C). Heated water from the discharge canal never reached any
of the three lake stations sampled for phytOplankton parameters. For
the 1971 study period, lake stations never warmed above 26.5 C similar
to 1970 temperatures. River temperatures in 1971 were similar to 1970
regimes. Maximum surface temperatures of 25 C were reported for August
by Marcus (1972) at the discharge site compared to 34 C on 30 June 1971
(Appendix A3).

Secchi transparency readings were greatest at the lake station,
lowest in the river and intermediate in the discharge canal (Appendix A3),
but measurements were rarely greater than 1 m at any of the three
stations. With the exception of 29 February, 1971, the average reading
for the lake station was 0.8 m, 0.5 m for the river and 0.5 m for the
discharge canal. Sestonic solids were highest at the inshore sites and
varied with wind velocities.

Daily solar radiation was nearly constant from 15 April to 18 August
1971; thereafter, lower radiations reached the lake during 1971 (Appendix

A4). Meterological data indicated a corresponding reduction in the

 

1Inshore stations indicates the river (9) and the discharge canal (8)
in combination.

10

.mcowuooaaoo
couxcmaaoumnm nuwa oowuocsmcoo CH vowuooou Somme a H ocu um monoumuomaou nouws coo: .m osswfim

2km: whin—

NT—— 9.0— 570 open 975 2.0 Oflen OTC
l. P r b P IF D

b

rm
to
wu—
we.

so“

(a) sunivuadnai

 

 

.I . I 8.... .3

III 32¢ /\ run

I I I 092.030

 

11

percent possible sunshine for the same period (Appendix A5). Daily
radiation recorded in the spring of 1972 was like spring and summer
radiation in 1971.

Total alkalinity was usually highest in the river; ranging from
120-175 mg/liter CaCOS. The lake water ranged from 90-132 mg/liter,
while the discharge had intermediate concentrations of 114-160 mg/liter.
Lake water had characteristically higher pH ranges (8.1 - 8.8) than
inshore sites (river and discharge canal) with the river most often
exhibiting the lowest pH (6.9 - 7.8).

Water at inshore stations contained consistently higher concentra-
tions of total carbon than the lake and concentrations in the river
usually exceed concentrations in the discharge. Particulate organic
carbon usually comprised not more than 372 of the total carbon. Concen-
trations of total carbon varied between 22 and 64 mg/liter.

Total phosphorus fluctuated without trend at all stations. Con-
centrations of total phosphorus varied between 0.07 mg/liter and 0.32
mg/liter in the study area. The lowest concentrations occurred in the
lake and highest concentrations occurred in the river.

Mean total nitrogen fluctuated more widely than carbon or phosphorus,
with the lowest concentrations (0.6 to 1.2 mg/liter) determined on 17
September 1971 and highest concentrations in April, 1971 (0.5 to 2.7
mg/liter). Inshore stations averaged higher concentrations of nitrogen

than the lake sites.

MATERIALS AND METHODS

Field and Laboratory_Procedures

Population sampling was completed at four-week intervals from 16
April to 12 November 1971. Water for chemical analyses was obtained at
the same time. Standard primary productivity measurements were made
approximately every 2 weeks, weather permitting, at three stations during
the interval of 15 April 1971 to 14 June 1972.

The stations chosen for phytoplankton analyses (Figure 2) coincided
with those selected by Marcus (1972). Sites 1, 3 and 6 are located in
the lake proper; station 8 in the discharge canal and station 9 in the
river (Figure 2). Each site had two substations where two depths were
sampled, i.e.: 0.5 m and 2.5 m. All offshore stations had buoy markers
to orient the investigator. Inshore stations were located from shoreline
reference points. Water was retrieved using an 8.1 liter VanDorn water
sampler (Wildco Co., Saginaw, Mi.) and a subsample was placed in a 500
m1 polyethylene bottle. A formalin preservative was used (37% formaldehyde)
in a 1:25 solution to arrest biological activity.

Laboratory treatment involved using the membrane filter technique
first described by McNabb (1960) and adopted by American Public Health
Association (1971). Between 10 and 40 ml of water were filtered through
a 0.45 p "millipore" filter which was placed on a 2 x 3 glass slide and
thinly coated with immersion 011. After "cleaning", the filters were
subsequently examined at 200x or 450x using a dark phase microscOpe.
Species frequencies in 30 random fields were converted to estimated numbers]

liter by a modified equation applied to this study:
12

l3

no/liter - d x 10}2

(quadrant area in uz) (m1 filtered)
where the d value or theoretical density corresponds to a given frequency.
A “EQEEEW is considered as a cell, colony or filament. Live and preserved
wet mounts were qualitatively examined prior to filter enumeration.

Diatoms were identified separately from other algae by performing
proportional counts on permanent mounts prepared according to Weber (1971).
Total counts of centric or pennate diatoms recorded on the millipore
filters were assigned to diatom species on the basis of their relative
frequency observed in a permanent "hyrax" mount. Admittedly, the densities
are only gross estimates of the existing numbers per unit volume of water.

Phytoplankton taxonomy was based on keys of Hustedt (1930), Patrick
and Reimer (1966), Prescott (1962), Taft (1945), Taft and Taft (1971),
Tiffany (1934) and Weber (1971). Taxonomic revisions of blue-green algae
were applied according to Drouet (1968) and Drouet and Daily (1956).

Volumetric estimates were derived from random measurements of cell
volume and total counts/liter for each species during the study period.

The mean species volume was calculated using average dimensions of the
regular geometric configuration most closely resembling the cell. Mean
cell size was obtained from the quotient of total algal cell volume and
total counts/liter.

Primary productivity was measured at three stations (Figure 2) from
water samples obtained with a 4.1 PVC VanDorn water bottle. Duplicate
clear and darkened pyrex borosilicate glass bottles were filled with
water obtained at the surface, 0.5 m, 1.5 m, and 2.5 m and placed in_§i£2_
at the corresponding depth of collection. One initial water sample was
taken for each set of four exposure bottles to determine the original oxygen
concentration. Suspension systems consisted of an anchored styrofoam

float, clipped iron rods, and hanger rOpes.

l4

Incubations were performed as near the midday period as possible
and spaced over 3-4 hours. All i2_§i£g_intervals were between the times
of 0900 and 1600 hours with the majority of times between 1000 to 1400
hours. Sample fixation and titrametric determination followed using the
modified Winkler method (A.P.H.A., 1971). Gross primary productivity and
respiration values were obtained by oxygen changes (Strickland, 1960).

Comparative cooling system experiments were executed on 18 August and
14 October 1971 in order to discern the effect of plant operation on
photosynthetic activities of phytoplankton communities. Surface water,
delivered to two light and two dark bottles, was collected at five stations
(Figure 4). The combined influences of heat and mechanical disruption were
incorporated into the design. Control and transported water was placed
at prescribed locations to test the cooling system effects at: (1) river,
(2) intake, (3) lake, (4) upper discharge canal and (5) mouth of the dis-
charge canal (plume). Changes in oxygen concentrations within the 300 m1
bottles were used to describe stimulatory or inhibitory effects.

Diurnal gross primary productivity was estimated at the surface and
0.5 m depth on 26 August 1971 and 1 June 1972. Four incubation periods
comprised the first diurnal study and three were used in the second study.
Midpoints of the time interval were used to position the data points.

Secchi transparency data were obtained using a 20 cm disc (Wildco
Co., Saginaw, Mi.). A YSI model 51A combination oxygen-thermistor meter
was used for temperature measurements. Diel pH ranges were based on a
standardized Instrumentation Laboratories porto-matic pH meter, model 175.
Total alkalinity measurements were performed as described in A.P.H.A. (1971).

Daily solar radiation was measured during the study period by two
instruments: (1) Eppley pyrheliometer (Eppley Lab., Inc., Newport, R.I.),

beginning 15 April 1971 and ending 29 February 1972; (2) Belfort

15

IAISIN "VII

 

 

CIIIK

WESTERN
LAKE
”N ERIE

(13Knu

Figure 4. Location of i2_situ sites in the study area used for the
cooling system experiments of 18 August 1971 and 14
October 1971.

l6

pyrheliometer (Belfort Instrument Co., Baltimore, Md.), from 12 May to
14 June 1972. A polar planimeter was used to convert from area to total
langleys (g cal/cm?) per day.

Meteorological information was condensed from U.S. Dept. of Commerce
records taken at the Detroit City Airport (Appendix A5). River discharge

was obtained from U.S.G.S. guaging station records.

Data Analyses

 

Raw data were transposed to a computer program summarizing para-
meters from the four replicates for each station. Average values were
compiled for number of counts/liter, percent of total counts, average
estimated volume per species and percent of total cell volume for each
species. That information was the basis for determining the relative
rarity or commonness of species.

A two-way analysis of variance was applied to test for vertical
(depth) and horizontal (station) differences in the numbers of observed
species, numerical diversity, numerical equitability, total phytoplankton
numbers, total cell volume, blue-green algal volume (CyanOphyceae),
green algal volume (ChlorOphyceae), diatom algal volume (Bacillariophyceae).
Preliminary comparisons indicated no significant differences between
depths, therefore, all replicates were pooled for testing station differences.
Multiple comparison tests using Tukey's method (Steele and Torrie, 1960)
were applied to estimated means for each station using an a = .05 level
of significance. Logarithmic transformations on total numbers, total
volumes, and all class volumes were used to reduce heterogeneity among
variances. Tukey's value was based on the error mean square produced in
the factorial analysis. Range tests were completed using the appropriate

mean, whether arithmetic or geometric, but arithmetic means are presented

17

in all cases. Using formulae from Strathmann (1967) total carbon con-
tent was estimated from the phytoplanktonic volume.

The Shannon-Wiener diversity index was used to depict phytoplankton
diversity (Pielou, 1969). Equitability (the ratio of observed diversity
to maximum diversity for the same number of species) is calculated from

the formula: e - ZE_where R is the sample diversity and S refers to
the number of samplgd species (MacArthur, 1965). Equitability measures
the relative evenness of species abundances. Diversity is determined
both by equitability and the number of species.

Gross primary productivity and respiration estimates on a volumetric
basis were adjusted to areal units by averaging oxygen production among
all depths and applying a depth factor for the respective sites. Oxygen

content was transformed to carbon uptake by using the photosynthetic

ratio of 0.312 units of carbon fixed/unit oxygen evolved (Westlake, 1969).

RESULTS

Communitygstructure

 

Phytoplankton Abundance

Total Numbers: Estimated total number of counts/liter in the study

 

area fluctuated from as low as 1,791 X 103 on 16 April to as high as
23,289 X 103 on 16 August (Figure 5, Appendix A6), but no vertical
differences were ever ascertained. The river and discharge canal pro-
duced the highest densities on all occasions except one when the river
abundance was less than that at one of the lake stations. The lake
stations consistently contained lesser (p < .05) algal concentrations
than the inshore sites. The north lake station usually yielded higher
densities than the two southerly stations and approached the summer num-
bers recorded at the inshore sites. The southernmost lake station on

all dates except one was among the significantly lowest group of stations

in the multiple range tests for total numbers/liter.

Total Volumes: Figure 5 indicates that algal volumes consistently

 

show temporal shifts at each station in conjunction with algal numbers

but again no vertical differences occurred. Volumetric estimates were

more variable than numerical estimates of abundance so the capacity to
differentiate stations was reduced. The discharge canal was less frequently
among the sites with the greatest volume than when numbers were compared,
but the south lake station was again most frequently among the stations

with the least volumetric abundance (Appendix A7). Shifts in ranks are
undoubtedly due to differences in the mean size of the algal cells at

the different sites. The mean cell size of algae was frequently lowest

18

 

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tosses; 32.2.... .38 15:63:35.... in» ..3..\.o§. 32.5.... .38

Mean total algal numbers, total algal cell volume and total

cell carbon in 1971 by station (total numbers .

Figure 5.

; total

).

volume ------; total carbon

20

 

0

Mn

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cow 0...)
86.. "fl“
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31 n a. m a. ml m w. w
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..3=\.9x.:3§... .38

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Figure 5 (con't.)

21

in the river and discharge canal and grestest among the lake station
(Appendix A8).

Total Carbon Content: Conversion of total volume to carbon content

 

(mg/liter) depends on the algal species present in greatest abundance

and therefore will vary differently than variation in total volume when
the species composition changes. The estimated algal carbon concentration
remained below 7 mg/liter at all stations except on 16 August when it _a
reached 8.4 and 9.6 in the river and canal, respectively (Figure 5). r]

The south lake station produced lower concentrations of carbon

compared to the other lake stations. As expected, this appears to be
more closely allied to volumetric variation than to total number of ‘J

individuals per liter.

Class Abundances

 

A total of six algal classes were represented in the enumeration of
western Lake Erie phytoplankton during 1971. Three classes comprised the
major percentage of algae volumetrically, i.e.: diatoms (Bacillariophyceae),
blue-greens (Cyanophyceae) and greens (ChlorOphyceae). Two dinoflagellates,
one Cryptophycean and five Euglenophytan species constituted the remaining

species in 1971. Cryptomonas ovata and Ceratium hirundinella were two

 

species which provided a notable portion of the total algal volume from
June to September. Otherwise, primary volumetric contributions were
from.the three classes previously noted (Figure 6).

Mean volumetric composition fluctuated expectedly by station but more
so temporally (Figure 6). Although there was considerable overlap in
dominance as seasons changed, generally diatoms were most abundant (> 50%
of volume) in the cooler months of the year 1971, i.e.: April, May and

November. Green algae replaced diatoms as the dominant class in June and

22
PERCENTAGE

 

 

 

A

I

J

’ 1

A

s l

O

N

A

I

J

J I 3

A

' I

o I

N I

‘ fl

1

z
:r
I-’ 59.
In I;
g. I +-
m

 

a..."

20¢)

Sau’hb"

 

 

0...! Classes logic-”bye... lulu-dept Chloropb com. Cymmophycomo

 

Figure 6. Mean percent composition of algal classes on a volumetric basis
for the study period of 16 April to 12 November, 1971.

23
July. Blue-green algae predominated in August, green algae and diatoms

dominated in September while greens and blue-green algae dominated in

October.

Species Composition by Class

 

Blue-ggeen Algae: Strong monthly variation in blue-green algal

 

volumes is apparent (Appendix A9). Highest standing crops existed in

the discharge canal (90,836 X 10'-6

ml/liter) during August when 90% of

the algal volume was composed of blue-green algae. But at that time,
temperatures in the discharge canal barely exceeded river or lake tempera-
tures (Figure 3). This extraordinary abundance was not caused by a temp-
erature change. A resurgence of cyanophyceans occurred in October when

lake stations had larger concentrations than inshore areas. Four blue-

green algae exhibited significant volumes during 1971, Aphanizomenon

 

flos-aquae, Anacystis incerta, Anacystis thermalis and Anacystis cyanae

 

(Appendices A10 to A13). Of those four, A, flos-aquae could be considered

 

the most ubiquitous species, occurring at all stations on seven dates
with a maximum concentration observed during the August "bloom" in
the discharge canal. A, incerta also reached its greatest abundance
during that time. Both species were volumetrically most abundant at
inshore stations on 16 August 1971. High standing crops of A, thermalis
were recorded at the inshore sites for the three summer dates. A, cyanae
occurred in scattered samples but also attained significant volume on
16 August, especially inshore.

Green Algae: Peak volumes of green algae occurred during June to
October with the river and discharge canal consistently containing the
largest volumes of greens on those dates (Appendix A14). One filamentous

green alga, Mogggotia elegantula, flourished in the fall period first in

 

24
the canal and river, then somewhat later in the lake (Appendix A15). Two

relatively large coenocytic algae, Pediastrum duplex and P, sipplex,

 

produced scattered but relatively great volumes in 1971 (Appendix A16 and
A17). P, duplex reached a maximal concentration on 17 June while P,
simplex peaked on 16 August. The latter species had a slightly larger
mean colonial volume than P, duplex but never became as abundant as P,
duplex.

Diatoms: Maximum diatom volumes occurred in the spring and fall
months, but the summer levels never decreased below the minimum concen-
trations that any of the other classes experienced (Appendix A18). Seven

species of diatoms were considered dominants during 1971. Coscinodiscus

 

radiatus exhibited no consistent pattern of seasonal fluctuation in
abundance but its largest volume was recorded on 17 September. Inshore
samples contained lower volumes of this species than the lake sites

(Appendix A19). Temporal variation of Melosira granulata followed a

 

sequence of high spring volumes, low summer and slightly higher fall
volumes. No evident spatial difference appeared to exist for this species

(Appendix A20). The centric diatom, Stephanodiscus astraea, predominated

 

in May samples at all stations (Appendix A21). It usually was higher in
abundance volumetrically at the river and discharge stations. Stephanodiscus
tenuis also fluctuated but was most prevalent in spring samples (Appendix

A22). Fragilaria capucina accentuated itself in the fall chiefly at the

 

lake stations (Appendix A23). During the spring, Fragilaria crotonensis
proliferated in larger volumes at the lake stations than inshore (Appendix
A24). Tabellaria fenestrata was most abundant in the spring, particularly
in May. Its volume was characteristically lower inshore than offshore

(Appendix A25).

25

Other Species: A cryptophycean, Cryptomonas ovata, played a minor

 

 

role volumetrically but was relatively numerous when it occurred. Peak
volumes in May occurred at the northern part of the lake study area but
otherwise showed little spatial variation (Appendix A26). A dinoflagellate,

Ceratium hirundinella, occurred in greatest quantities in the lake at the

 

time of an August algal bloom that was otherwise dominated by blue-green
algae (Appendix A27).

Comparing volumetrically dominant species common to the 1970 and
1971 study periods (Tables 1 and 2) suggests that most of those species
did not comprise nearly the same average yearly percentage of the total
cell volume. Also, there appears to be no relation between yearly dif-
ferences in discharge canal environment and the change in species dominance.
Fewer species were volumetrically dominant (: 1%) in 1970 than 1971;

those important in 1971 but not in 1970 were P, capucina, P, crotonensis,

 

.1, fenestrata, M, elegantula, P, simplex and g, hirundinella (Table 2).

 

 

 

‘P, duplex is the only species which exhibited a higher percent composition
during 1971. Of the species in Table 2, eight averaged a lower percent
of the total cell volume in 1971 relative to 1970. This implies that

percentages of the total cell volume were more evenly distributed among

more species in 1971 samples than in those of 1970.

Variety of Species

 

Total Number of Species: Table 2 illustrates that in 8 months of

 

water collection, 184 phytoplankton species were observed. Of that
number, diatoms were represented by 80 species; green algae, 79 species;
blue-green algae, 17 species; euglenoid algae, 5 species, dinoflagellates,

2 species; and cryptophyceans by 1 species.

26

Table 1. Mean annual percent volumetric composition of dominant species
examined from May to November, 1970 (Marcus, 1972; X < 1% of the
total algal cell volume).

 

Percent by Station

 

 

 

 

 

Species (16) 1 3 6 8 9
Coscinodiscus radiatus 1.8 2.2 3.4 X X
Cyclotella meneghiniana X 1.1 X 2.6 7.1
Melosira granulata 13.7 8.5 12.5 15.3 8.2
Stephanodiscus alpinus X 1.2 X X 3.0
S, astraea 7.4 2.3 3.4 5.0 3.9
S, binderanus 13.9 21.6 14.5 9.8 32.1
S, niagarae 1.2 1.1 6.6 4.1 X
S, tenuis 1.2 X X X 3.4
Anacystis cyanae 13.0 17.7 16.7 13.8 12.3
A, delicatissima (S, incerta) 10.5 10.7 8.8 13.8 3.0
Aphanizomenon flos-gguae 19.3 12.6 12.2 11.5 5.9
Oscillatoria sp. (Microcoleus vaginatus) 6.7 7.4 10.2 10.3 4.2
Pediastrum duplex 2.1 3.9 3.5 4.7 3.2

 

é, elachista var. conferta (A, incerta) 1.6 1.4 2.4 2.8 2.0

 

27

Table 2. Phytoplankton species enumerated from April to November, 1971
with their mean annual percent composition by station (X < 1%
of the total cell volume).

 

Station
Bacillariophyceae 1 3 6 8 9

 

Achnanthes lanceolata (Breb.) Grun. X X X

A, minutissima Kutz. X

Amphiprora ornata Bailey

Amphora ovalis (Kuta.) Kutz.

A, ovalis var. pediculus (Kutz.) V.H.

Asterionella formosa Hass.

Cocconeis disculus (Schum.) Cleve.

mark Ehr.

Coscinodiscus radiatus Ehrenberg 10.7 10

Cyclotella bodanica Eulenst.

Q, kuetzingiana Thwaites

g, meneghiniana Kutz.

Q, ocellata Pant.

E, stelligera (Cleve and Grun.) V. H.

Cymatopleura solea (Breb. and Godey)
Wm. Smith

Cymbella affinis Kutz.

Diatoma hiemale (Lyngbye) Heiberg

Q, tenue var. elongatum Lyngb. X X

2, vulgare Bory. X

Epithemia sorex Kutz. X

Fragilaria brevistriata Grun. X X X X

Fragilarig capucina Desm. 2.5 2.8 2.8 1.1

.E. construens (Ehr.) Grun. X

F. crotenensis Kitton 4.0 1.6 4.8 1 0

E, pinnata Ehr.
Gomphonema olivaceum (Lyngb.) Kutz.

Q, parvulum (Kutz.) Kutz

Gygosigma kutzingii (Grun.) Cleve.
Q, scalproideg (Rabh.) Cleve.

S, spegcerii (Quek.) Griff and

Renfr.

.flgngggghig amphioxys (Ehr.) Grun. X X
Melgsira gmbigug (Grun.) O. Mull. X X
M. W (Ehr.) Ralfs 3
M. 1515351191 0. Mull 1.

smalls Patr. var. .canalis
contents Grun.

 

 

 

xxxx

 

 

xxxxx
x x
x x
x x
xxx

 

6 14.6 6.9

 

l—‘CD
#0

 

 

 

xxxxx
xxxxx°
xxxxx
xxxxx
xxxx-

 

 

x
x
xx

 

xxxx
x
xxxxxxx

1"
b

xxx
x
xxxx
xx xxx°
x xxxxx-

x

xx
xxxxx-

E
x
x

.exigua (Gregory) 0. Mull. X X X
(Ehr.) Kutz. X

E

H

E“
><><a:x

magazmmn
C)
'1
.5
xxx

5
§

28
Table 2 (con't.)

Bacillariophyceae l 3 6 8 9

 

mutica Kutz. X

reinhardtii (Grun.) Grun.

tripunctata (0.F. Mull.) Bery. X X X
Nitzschia acicularis (Kutz.) Wm.
Smith

ppphibia Grun.

angustata (Wm. Smith) Grun.
apiculata (Greg.) Grun.
denticula Grn.

dissipata (Kutz.) Grun.
filiformis (Wm. Smith) Schutt
fonticola Crun.

_, gracilis Hantzsch

Nitzschia hol§§t1c§_ Hust.

S, hungarica Grun. X
. linearis Wm. Smith

. lorenziana Grun.

. palea (Kutz.) Wm. Smith
_, sigma (Kutz.) Wm. Smith
Pinnularig_borealis Ehr.
Rhoicosphenia curvg£§_ (Kutz.) Grun. X
Rhopalodia giphg_ (Ehr.) 0. Mull.
Stephanodiscus alpinis Hust.

. astraea (Ehr.) Grun. 2
binderanus (Kutz.) Krieger

duping (Fricke) Hust.

slums. Ehr.

ggngig_ Hust. 1.5 1.2
X

 

uzuznz

xxx

xx
x
x

xx

xxxx

lzlzlzlffilzlzlz

z
xxxxx

x
x
x

zuznzuzz
><
>4
><><><><
><

X

xxxx
x

X

x
xxx

X X X
7 4.3 4.7 14.2 13.8
X X
X X

U)

xxx°

X
X
X

Ismael
x x x x

1.7 2.0

E
E

X

X
X X
X

xxxxx
x

S, m Meister

S, parasitiga_ (Wm. Smith) Hust.

S, ulna_ (Nitz.) Ehr.

Tabellaria.fenestrata. (Lyngb.)
Kutz. 10.7 10.4 9.4 3.0 1.3

Centric sp. #1 X X

xxxxx

X
X
X X

x

 

Chlorophyceae

 

Actinastrgp hantzschii Lagerheim X X X
A, hantzschii Lagerheim var.
fluviatile Schroeder X X X X X

Means. (Cords)
Ralfs X X X X X

 

 

29
Table 2 (con't.)

Chlorophyceae

 

A, convolutus Corda

Binuclearia eriensis Tiffany

Botryococcus braunii Kuetzing.

.P. sudeticus Lemm.

Chlorella ellipsoidea Gerneck

Chlorella vulgaris Beyerinck

C. sp. #1

Chlorococcum infusionum (Schrank)
Meneghini

Coelastrum micrgporum Naegeli

g, sphaericum Naegeli

Cosmarium bipunctatum Boergesen

E, formulosum Hoffman

E, subcrenatum Hantzsch

Crucigenia apiculata (Lemm.)

Schmidle

irregglaris Wille

lauterbornei Schmidle

quadrata Morren

rectapgularis (A. Br.) Gay

tetrapedia (Kirch.) W. and

G. S. West

Dictyosphaerium ehrenbergianum
Naegeli

Gloeoactinium limneticug, G. M. Smith

Gloeocystis planktonica (W. and G. S.
West) Lemm.

Golenkina radiata (Chod. ) Wille

Haematococcgs lacustris (Girod. )
Wittrock

Kirchneriella contogta (Schmidle)
Bohlin

We Janene Hutch.)
Moebius

S, Obegg (W. West) Schmidle

Lagerheigia genevgnsis Chodat var.
subglohoga (Lemm) Chodat

L, gagggigggg, (Lemm.) G. M. Smith

 

 

 

 

 

 

 

 

 

 

 

 

unusuauauu

 

S, pusillum Fresenius

Mougeotia elegantula Wittrock

Oocystis borgei Snow
9, pugilla Hansgirg

O. submarine Lagerheim
_P_________andorina mo___r___um (Muell. ) Bory

W m (Turp ) Meneghini
2. finale; Haven

xx

xxxx
xxxx

xxx
xxxx

1.1

xxxxx

x

X X X X
12.8 20.2 11.8 12.7 1

xxxx xxx

x

xx

X

1.2
3.0

30
Table 2 (con't.)

 

 

 

 

 

 

 

 

 

 

 

Chlorophyceae l 3 6 8 9
P; implex (Meyen) Lemm. 3.2 2.0 6.0 1.4 5.5
P, tetras (Ehr.) Ralfs X X X X X
Planktosphggzlfi.gelaiinflaa. G.M.

Smith X X X X X
Polyedgiopgis gggggifiping_ G. M.

Smith X X
2, pinulosa Schmidle X X
Qggdrigula closterioidgs (Bohlin)

Printz. X X X X X
Scenedesmus abundans (Kirch.)

Chodat X X X X X
S, acuminatus (Lager.) Chodat X X X
S, anomalus (G. M. Smith) Ahlstrom

and Tiffany x x x x
S, bernardii G. M. Smith X X X
S, bijuga (Turp.) Lagerheim X X X X X
S, carinagug (Lemm.) Chodat. X
S, gengigglaggs_ Lagerheim X X X X
S, dimgrphg§_ (Turp.) Kuetzing X X X X X
S, Qpliensia P. Richter X X
S, gnggzigguga_ (Turp.) de Breb. X X X X X
S, quadricauda (Turp.) de Breb. var.

alternans G. M. Smith X X X X X
Schroederia setigera (Schroed.)

Lemm. X X X X X
Sphaerocystis schroeteri Chodat X X X X X
Spongylomorum quaterggriup_ Ehr. X X
Staurastrum chaetocerg§_ (Schroeder)

G. M. Smith X X X X X
Tetraedron arthrodesmiforme (G. S.

West) WOloszynska X X X
I, caudatum (Corda) Hansgirg X X
I, hastatum (Reinsch) Hansgirg X X
g, incus (Teiling) G. M. Smith X X X X
I} limneticum Borge X X
I} minimum (A. Br.) Hansgirg X X X X
I, pentraedricum W. and B. S. West X X X X X
T} regulare Kuetzing X
I; trigonum (Naeg.) Hansgirg X X X X X
Tetrastrum glabrum (Roll) Ahlstrom

and Tiffany X
I, heteracanthum (Nordst) Chodat X X X X X
g, staurogeniaeforme (Schroeder)

Lemm. X X X X X
Westella botgyoides (W. West)

de Wildermann X X 1.7 1.6 2.2
Green sp. #1 X X

 

31
Table 2 (con't.)

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Cryptophyceae 1 3 6 8 9
Cryptomonas ovata Ehr. 1.1 X 1.1 X X
Cyanophyceae
Agmenellum quadridpplicatum Breb. X X X X X
Anabaena circinalis Rabenhorst X X X X X
Anabaena sp. #1 X X X X X
Anacystis cyanae Drouet and Daily 1.6 1.6 1.2 2.2 1.3
A, dispersus Meneghini X X X X 1.5
S, incerta Drouet and Daily 10.3 4.1 8.7 9.0 8.5
A, marina Drouet and Daily X X X X
A. minutus Meneghini X X
A, montana Drouet and Daily 5.7 5.6 X 3.2 4.0
A, pulchra Meneghini X X X X
A, thermalis Drouet and Daily 1.9 3.1 2.1 4.8 6.5
Aphanizomenon flos-aquae (L.) Ralfs 7.1 7.0 9.6 6.6 6.2
Coccochloris peniocystis Drouet and
Daily 1.0 X
9, stagpina Sprengel X X
Gomphosphaeria lacustris Chodat X X X
Marssoniella elegans Lemm. X X X X X
Microcoleus vaginatus (Vauch.) Com. X X X X X
DinOphyceae
Ceratium hirundinella (O. F. Muell.)
Dujardin 2.7 2.9 3.2 X X
Peridinium quadridens Stein X X X X X
Euglenophyceae
Euglena acus Ehr. X X
S, ggacilis Klebs X X X X X
E} minuta Prescott X
S, polymorpha Dangeard X X X X
Trachelomonas pulchella Drezepolski 1.5 X X X X

 

 

32

The mean number of observed species among the four station replicates
and the distinctly different species accumulated among those samples indi-
cates the relative commonness of species (Table 3). In all cases, the
most abundant species were common to all replicates at a station. The
maximum mean number of species recorded for any set of samples amounted
to 51 species on 17 September in the discharge canal when the maximum
number of distinctly different algal species occurred. There was no
statistically significant difference (p < .05) between the number of
species at the discharge canal and river on that date (Appendix A28).

The fewest species samples (19) were collected on 16 April at lake station
3.

Within the total list of species, only twenty three contributed

1% or more of the mean volume at any one of the stations (Table 2).
Twenty six occurred only at one station and comprised less than 1% of
the total volume collected in 1971. These most abundant and rarest
species differed only slightly from rare and abundant species in 1970

collections (Marcus, 1972).

Annual Differences in Species Abundance: The discharge canal had

 

the greatest number of species of any station on 87% of the 1970 dates
compared to 75% of the 1971 dates. Apparently, the discharge canal

retained its study area dominance in 1971 with respect to species abundance
even though major physical changes occurred in the water masses when pump-
ing started in 1971. However, a reduction in the mean number of species
identified at the discharge site occurred in 1971 compared to 1970 (Marcus,
1972). The river station consistently was next highest in number of species
during 1971 but averaged fewer species per sample in 1971 relative to

1970. Although differences exist in the mean number of species per sample,

the species tabulation list of Marcus (1972) numerically approximates

33

Table 3. Number of distinctly different species accumulated temporarily
and spatially with the mean number of species at a station in

 

 

parentheses.
Station

Date 1 3 6 8 9
4-16-71 25 (23) 21 (19) 23 (23) 4o (37) 41 (39)
5-20 42 (34) 41(35) 28 (24) 54 (47) 44 (38)
6-17 39 (31) 44 (32) 41 (30) 54 (42) 48 (36)
7-14 41 (36) 4o (30) 26 (20) 54 (38) 51 (37)
8-16 34 (23) 29 (22) 28 (21) 56 (43) 45 (36)
9-17 58 (36) 58 (42) 41 (28) 73 (51) 61 (47)
10-15 56 (40) 52 (33) 48 (32) 65 (44) 60 (42)

11-12 53 (38) S4 (39) 43 (29) 41 (40) 49 (41)

 

34
(203 sp.) the 1971 list (184 sp.) using an equivalent number of replicates
(160). Therefore, it appears that there was slightly more temporal
segregation of species in 1971, meaning the average persistence time of
each species was shorter than during 1970. Species common to both lists
equaled 140. Some of the difference in species lists is undoubtedly
due to imperfect identification but much is also caused by annual shifts

in the abundance of rare species.

Species Diversity and Equitability

Inshore stations most often were among the stations with the highest
phytoplanktonic diversity while the lake stations had consistently lower
diversity (Figure 7, Appendix A29). The species equitability, or evenness
of species relative abundance, did not regularly vary with diversity,
except on one notable occasion during the algal bloom on 16 August when
both population parameters were low (Appendix A30). At that time, the
lake stations had relatively few total species compared to the remainder
of dates but relatively high numbers/liter among a few species. The
inshore stations had slightly more species but exhibited the same low
equitability on that date. Usually, diversity differences among sites
were due more to the equitability component than the number of species
but both directly affected variations in diversity.

Monthly diversity and equitability varied erratically at the discharge
station as well as at the other stations. The discharge canal had the
highest or next to the highest diversity on 87% of the dates in 1971
versus only 38% of the comparisons in 1970. Absolute diversity levels at
the canal station in 1971 were higher than 1970 levels 75% of the time.
That station also exhibited higher equitability in 1971 on 75% of the

dates which coincided with the higher diversity dates. However, the mean

35

.Amufiafinmufisvollmmxon umoao mxuwmuo>wvllmoxon monoumnv

 

pump mafiaaamm some so coquum >9 huHHHnmufisvo woo hufimpm>fiv vascuxcmaaoumna com: .5 muawwm
«. .32 $.60 5.58 o. .233 2 :2. t 2:... ou 2.: 8 52
2.5a: >h..=m<h50m
mm. 3. 5. sf} . . . _ . «0.. . 4 pm. 4 .no. 8. no. 8. no.
a a H _ .

 

 

 

 

\\\

 

 

 

 

 

 

 

 

 

 

 

 

 

 

0d.

nun

 

O;

 

 

 

W\\

\4

 

 

 

 

Hli

 

e W

_\\\

 

 

 

 

 

 

 

_\\

 

 

 

R\

 

 

 

 

[5

 

 

 

 

 

 

Fbrbprb P

 

awn O.

— nun .0;

 

TA

fi\\

 

 

m

 

 

 

 

 

\

 

 

 

fl,

 

 

m
(WIN

m\\

\

 

 

 

.Hn 0a nun

.25: >h.m¢m>_o

NOILVLS

36

number of species in discharge samples was higher during 1970 than

1971.

Seasonal and Yearly Statistical Comparisons

 

The following parameters were compared through factorial analysis:
(1) number of species, (2) numerical diversity, (3) numerical equitability,
(4) total numbers/liter, (5) blue-green algal volume, (6) green algal
volume and (7) diatom volume using year and month station factors. Results
indicated a statistically significant two-factor interaction term for
all variables except numerical equitability, total counts/liter, and
green algal volume (Appendix A31). But a three—factor interaction term
was also significant for all factors except green algal volume. There-
fore, a year test averaging all months at each station produced incon-
sistent and non-independent means. This is primarily due to the wide
variation in replicates incorporated in the mean and the non—additivity
of the model which violates an assumption for analysis of variance. A
separation of 1970 and 1971 data by the three growing seasons sampled
was performed to ascertain whether real differences existed between years
for a given season. Seasonal testing has more biological basis because
phytoplankton flourish seasonally in lake systems. It is evident that
statistical differences existed in the number of species in the spring
and fall, in species diversity during the fall season, and for equit-
ability measured during the fall as well as diatom volume in the summer
(Table 4). The reduction in the number of species affected the diversity
and equitability comparisons made for the fall season but no effect
occurred in the spring comparison. Diatom volume increased in the summer
of 1971 independently from non-significant changes in other algal class

volumes.

37

.commmm aamm onu pom o3u vow umaasm msu pom mounu .comfiumaaoo magnum osu pom pom: whoa mmumv mounHN

.mumw% cowsumn commmm om>aw m pom mfimxamcm mnu cw vmownaoo ouo3 m was m .o .m .H mcoaumumH

 

A¢ aowumum
unmoxm Han ma umnwfin
mum3 mcofiumum Hamv

 

 

A.e.evee.o n e eemooo.o u e A.e.evma.o n e eoUfiH\ma5Ho> acumen Awe
A.m.cvea.o u a A.m.cvmm.o I a A.m.ovmw.o I a umu«H\ma:Ho> Human compo Amy
A.m.cvnom.o I a A.m.cvmm.o I a A.m.cvon.o I a umuHH\ma:Ho> Human amouwloaam on
“.m.evaa.o . a A.m.evam.o u e A.m.ev-H.o u e nouea\maeao> Hams Hence Ame
A.m.evem.o u e A.m.evoe.o a e i.e.eveae.o a e emuea\6ue=ou Hence Ase
AON¢H cw nonwan
mums mcoaumum HHmv
«qo.o I a A.m.cvmam.o I a A.m.cvwn.o I m muwawnmuwsvm Hmowuoaaz AMV
Am “emcee case as Haema;
mum: mcowumum Hamv
«moo.o u e A.m.evon.o u e A.m.evmmo.o - e suamum>eo Hmeeuoaaz Ame
house as “memes Acnmfi an “meme:
mum3 mcoquum Hamv puma mcowumum Hamv
eeaoo.o u e A.m.eveme.o n e 45~o.o . e meeueem mo emeasz Ase
Hana umaaom magnum umumamumm
commm
~.H m

 

.Am x we
mumsvm coma cofiuomuouca ecu co momma uaumwuoumum ecu mo muwaanmnoua mocmofiwaawwm a nu“: mucuomm Amy
cofiumum use va you» osu mp muoumamuma msofluo> wcwumou mocmfium> mo mammamcm mm3loau o no muaammm .q manna

38

An appraisal of statistical comparisons indicates that no striking
dissimilarities in the phytoplankton population parameters occurred tempor—
ally or annually in the study area. Examining the standing crap at monthly
intervals without doubt lends itself to wide fluctuations in populations.

A look at the physiological responses of the phytoplankton community may

illustrate any significant alterations due to plant operation.

Community Metabolism
Effects in Depth

Average depth profiles of gross primary productivity (GPP) indicate
a significant reduction in carbon assimilation at approximately the 1.5
m depth for all stations both in 1971 and 1970 (Figure 8). Among the
three stations, the river was least productive at all depths. The
average GPP for the river surface was almost as great as values for the
lake surface during 1971 but not nearly as high as at the surface in the
discharge canal.

A higher GPP occurred at the 0.5 m depth relative to the average
surface rate in the lake during 1971. This contrasts with the sharp
decline from the surface to 0.5 m exhibited in 1970 at the lake station.
No apparent yearly change occurred at the canal station in average sur-
face GPP but higher productivity was indicated at the 0.5 and 1.5 m depths
in 1971 compared to 1970. Below the surface, average GPP at the discharge
station simulated average lake GPP at each measured depth. Average
river gross primary productivity was slightly higher during 1971 at the
surface and 0.5 m depths but was comparatively reduced below 0.5 m.

A respiratory profile with depth.may have indicated slight decreases
in carbon release at stations from the surface to 2.5 mbut variation

was so great that no statistically significant change could be identified.

 

0.55

LO'i

LS‘

Depth (meters)

2.04

2.59

04

0.5-

1.04

15“

Depth (metere)

2.0'

0.5‘

1.01

LS"

Depth (meters)

2.0d

 

39

Lake

 

 

  
  

Discharge

 

River

  

 

 

2.5

Figure 8.

 

 

 

U I U j I 1 1 ‘U' j

.02 .04 .06 .08 .10 .I2 .14 .lb )8 .20 .22 .24 .26

mgC/l/ht.

Depth profile of average gross primary productivity measured
during the midday interval for the years 1970 ( ) and
1971 ( ----- ) at the three stations in the study area

(1: x 1 8.13:.)

 

Dark bot
than did
experien
three st

the 0.5

consumpt

Spatial

w
or disc?
the high
were mac‘
lore ox}
the disc
that ex}
the rim
1972 me;
river d1

fly
and bio.
““8113 1.
fungi, 4
Each C01
tion. 1
on two ‘
"at“ d.

pit.atim

40

Dark bottle respiration exhibited more temporal deviation from the mean
than did light bottle changes, causing wider standard errors. The river
experienced a greater average respiratory rate at the surface among the
three stations but the average for the discharge canal was greatest at
the 0.5 and 1.5 m depths. A significant decrease in average oxygen

consumption occurred at 2.5 m in the discharge canal.

Spatial and Temporal Effects

 

Midday Productivity: Gross primary productivity, as measured by

 

in §i£g_light and dark bottles, was lower in the river than in the lake

or discharge on all but one occasion (Table 5). The discharge canal had
the highest midday GPP on six of the study dates when complete sets

were made. On the remaining five dates the lake environment evolved

more oxygen. Maximum GPP of 498 i 9 mg C/m2/hr occurred on 16 July in

the discharge canal. Heating of intake water reached a A t of 8 C in

that exposure. Highest GPP for the lake was recorded on 18 June and for
the river on 2 October. Mean midday rates for 1971 were similar to spring
1972 mean rates (Table 5) at the lake but higher in the discharge and
river during 1971.

Midday Respriation: Community respiration includes total chemical

 

and biological oxygen demand but the chemical demand is assumed to be
negligible. The enclosed biological community contains heterotrophic
fungi, bacteria and zOOplankton as well as autotrOphic phytoplankton.
Each component contributes an unknown amount to the total oxygen consump-
tion. Respiration was consistently lowest in the lake (Table 5), except
on two extraordinary occasions which may have been influenced by river
water due to westerly winds on those dates. Most of the time lake res-

piration averaged about one-half that in the river and discharge canal.

y (GPP) and community respiration rates (R) estimated during the

hr (X i l S.E.).

tivit
El

Mean gross primary produ
midday interval in mgC/m

Table 5.

 

Stations
Discharge

River

Lake

 

GPP

GPP

GPP

A t1

Date

 

272 i 3 232 t 21 122 i 2 480 i 99
206

132 i 22
368

348 i 12

4
1
8
7

5-21—71
6-3

78 i 18

‘H

24

26

H

137 i 12

58

+

263 i 48

6

536 i

66 t 16
98
108

398 i 31

4
5
9
7
18

266 i
236
473 i 25

448 i 33 140 i 42

184 i

6-18
6-30

'H

51
176

+l

373 1 21
299

+|

4
4

147 i 10
178

+

114
466

2

4

+l

6

+

15
22

+

'H

498
324
362

+

280 i 16
252

8
9
8
4
5
4

7-16
7-29
9-15
10-2

0.0

H

210

1

95 i
104
245

+

378

+

1
0

+

41

538 i 10

0.5

2
2
21

+

604 i 38

+

.—

3
l
10

+|

+1

375

6

109 t

+l

4
14

-H

193
284
272

26 i
22

354 i 13

6
4

287 i

t

120
147

+

1
8

+4

272
242

"H

103 i 14
181 i

10-16
10-30

2 191 i

8

+

—

8

+|

42

4

+l

41

62 i 11

11-13

 

152.3 308.2 314.4 112.9 265.6

259.1

Yearly Mean

 

22 1
173 1

4
2
7

+l

-11

38 1

2-29-72
5-12
6-1

57 i 42

208 i 10
354 i

6

'H

101

8

136 i 12

H

88 i
102

3 70 1
5

7

117

8
4

0

+1

21

+|

232

4

+

150

59 i

428 i

5

\O

 

64.5 113.3 180.3 50.5 281.0

272.5

Spring Mean

 

 

138.8 263.2 283.5 102.5 268.2

261.2

Grand Mean

 

The A t corresponds to the difference between intake and discharge temperatures.

1

42
Respiratory rates were similar in magnitude for both years at the lake
station, i.e.: 1971 yearly averages and 1972 spring averages were nearly
alike. Hourly respiration rates in the discharge canal during 1971
relative to 1970 averaged (by season): (1) spring, 23% lower; (2) summer,
166% higher; and (3) fall, 134% higher.

Diurnal Gross Primary Productivity: The diurnal change in GPP,
measured on one occasion each in the summer of 1971 and late spring 1972,
indicates peak photosynthesis in the early afternoon (Figure 9), close to
the time that midday productivity was normally estimated.

Midday GPP peaks at the lake station were not much greater than
mid-morning rates. Lowest secchi transparencies during the study period
were recorded at the lake on those dates. The magnitude of diurnal
productivity appeared to vary as much spatially as it did temporally.
Real mean differences in respiratory rates for 26 August were evident:
(l) the discharge averaged 54 mg C m3/hr, (2) the lake averaged 18 mg
C/mslhr, and (3) the river averaged 6 mg C/m3/hr. Temperature differ—
ences were minor for that date (1C). Respiratory estimates on 1 June
1972 were similar for all stations with a A t = 4C recorded on that date.

Daily and Seasonal Gross Primary Productivity: To obtain daily
estimates of GPP, the two sets of diurnal data were combined with the
midday productivity measurements by proportioning the midday estimates
to the average diurnal change in GPP (based on the two diurnal studies)
measured over the rest of the day (Table 6). This procedure is of course
subject to error since a constant proportion of total daily GPP is
assumed to exist outside the midday ig_§i£g_period. Another technique
suggested by Schindler (1971) revealed similar daily and seasonal GPP
to that obtained by the proportional method in Table 6. Schindler's

technique incorporates the ratio of total daily solar radiation (It)

‘4

2: some: :. .zan\OoE

I‘

up: some: :- .E\nE\OuE

.r .n.51..fltrnn.

rigure 9

\

43

3604
32m
280- , \
24m /

I204 I

001 ff /f “\‘K \\
40+ 1 / \.\\

mgC/m’lhr In upper In3

 

 

 

 

2 4 6 8 7512141818 02224
TIIIE (8-28-71)

320 (I
2804

240'1

1601

I201

mgC/m’lhr In upper m3

4m

 

 

 

K“

was (45-1-71)

Figure 9. Average gross primary productivity estimated during the
diurnal period on two study dates (lake . ; discharge
----- ; river ).

44

Table 6. Mean daily and seasonal gross primary productivity during 1971
and 1972 (gC/m2/day and season) compared to 1970 means (Marcus,

 

 

 

 

 

 

 

1972).
Station
Marcus Marcus Marcus
Date Lake (1972) Discharge (1972) River (1972)
5-21—71 3 8 2.8 1.3
6-3 3.0 1.4 0.3
6-18 5.1 2.8 0.7
5 daily 4.0 0.9 2.3 2.0 0.8 0.8
x seasonal 361.0 78.3 212.3 185.8 68.7 71.0
6-30 2 1 2.6 1.1
7-16 3.2 5.2 1.2
7-29 2.7 3.2 1.0
9-15 3.5 3.2 0.9
5 daily 2 9 1.7 3.6 2.7 1.0 1.5
x seasonal 261.6 159.2 323.1 250.7 94.4 141.2
10-2 3.1 3.9 2.1
10-16 0.9 2.1 0.1
10-30 1.4 1.8 1.1
11-13 0.5 --- ---
g daily 1.5 0.7 2.6 2.2 1.4 0.7
x seasonal 135.7 62.9 239.2 204.6 124.7 63.7
Total gC/m2 for
three seasons 758.3 300.4 774.6 641.0 287.8 275.9
2—22772 0.3 -0.1 --—
§ daily 0.3 -0.1 ---
x seasonal 30.0 -6.8 ---
5-12 -—- 1.0 ---
6-1 1.3 1.1 1.1
6-14 4.9 1.6 0.0
35 daily 3.1 1.2 0.6
X seasonal 285.2 113.5 50.2

 

 

.
.i—x-r-‘ov - -

to part
(See AP
synthes
into ac
that th
Se
of year
Novembe
had the
dischar
1971.
more th
the grc
discha:
a reduc
spring.
limitec
S4
conSis1
spring
AVErag.
lower

Steatl

IO

13 Con
inatan

callal’

45
to partial daily radiation during the bottle incubation interval (Ip)
(See Appendix A4). This method assumes a linear response of photo-
synthesis to light energy incident at the water's surface but it takes
into account definite changes in the percentage of total daily GPP
that the midday value comprises, regardless of the study day.

Seasonal summaries of GPP indicate no obvious relation between time
of year and GPP except that GPP was lowest during the late fall (13
November) and winter (29 February) sampling (Table 6). The lake usually
had the highest productivity during the spring of 1971 and 1972 while the
discharge productivity was usually greatest in the summer and fall of
1971. Based on 1971 seasonal estimates, the river GPP was at most slightly
more than one-half the GPP recorded at the other sites. During 1971,
the growing season GPP for the lake station was slightly lower than the
discharge GPP during the same period. Spring data from 1972 suggests
a reduced seasonal GPP at the discharge and lake sites from the previous
spring. Winter GPP in the study area appears relatively low based on
limited sampling.

Seasonal average hourly GPP at the discharge canal in 1971 was
consistently higher than GPP that Marcus (1972) reported, i.e.: a 15%
spring increase, a 33% summer increase and an 18% fall increase (Table 6).
Average hourly GPP in the discharge canal during the spring of 1972 was
lower than the two previous spring GPP's. Lake hourly GPP in 1971
greatly exceeded 1970 rates but differences at the river station were minor.

GPP/R Ratios: Ratios of diurnal GPP and daily respiration (GPP/R)
reveal whether or not the community is autotrophic (producing more than
is consumed) or heterotrophic (producing less than is consumed). In no
instance during the study period was GPP/R > 1 in the river or discharge

canal, indicating heterotrOphy (Table 7). This contrasted the lake site

 

Table

46

Table 7. Ratios of gross primary productivity/community respiration
at the sampling stations (based on gC/m2/day) with seasonal
and yearly averages for the years 1970 and 1971.

 

 

 

 

 

 

 

Marcus Marcus Marcus

Date Lake (1972) Discharge (1972) River (1972)
5-21-71 1.20 0.53 0.11
6-3 0.34 0.28 0.16
6-18 1.52 0.29 0.05

Spring Mean 1.02 0.38 0.32 0.24 0.11 0.12
6-30 0.77 0.29 0.91
7-16 0.29 0.72 0.28
7-29 0.76 0.35 0.20
9-15 0.82 0.22 0.07

Summer Mean 0.66 0.66 0.40 0.56 0.36 0.32
10-2 4.84 0.84 0.80
10-16 1.73 0.31 0.01
10-30 1.37 0.28 0.24
11-13 0.51 -—-- ---—

Fall Mean 2.11 1.23 0.48 1.64 0.35 0.13
2-29-72 0.22 0.19 0.19

Winter Mean 0.22 ---- 0.19 ---- 0.19 -—--

Yearly Mean 1.0 0.76 0.36 0.81 0.25 0.19
5-12-72 ---- 0.24 ----
6-1 0.78 0.34 0.22
6-14 3.45 0.29 ----
Spring Mean 2.12 0.29 0.22

 

 

which e
annual
within
lake DE
removir
the yea
the in:
incorpc
oxygen
8 littl
over-e:
relati‘
Y1
iCant c
Water 1
and fa
ratioS
Of 197
were c
higher
but a
1aCked

ratiOS

Coolin‘

sourCe,

47

which exhibited autotrophy (GPP/R > 1) on several occasions. Estimated
annual averages, although far from precise because of great variability
within seasons and inadequate cold weather sampling, indicate that the
lake most often is near an autotrOphic condition during the year. But
removing the two relatively high ratios for the lake station would reduce
the yearly mean ratio below 1.0, i.e.: heterotrophy would occur. However,
the inshore sites are markedly heterotrophic. Daily respiratory rates
incorporated in the ratios are based on the assumption that metabolic
oxygen loss remains constant over the 24 hour period when R is probably
a little higher during the day than it is at night. This could create
over-estimates in respiration which would reduce the GPP/R ratios but
relative comparisons are not likely to be as greatly affected.

Yearly variations in average seasonal GPP/R ratios suggests signif-
icant differences in the productive and consumptive capacity of the sampled
water masses (Table 7). The lake exhibited a higher ratio in the spring
and fall of 1971 as well as during the spring of 1972 compared to 1970
ratios. Lower ratios at the discharge canal during the summer and fall
of 1971 contrasts the changes at the lake station in 1971. River ratios
were characteristically low and similar during all three years. Slightly
higher yearly ratios were obtained for the lake and river site in 1971,
but a much reduced ratio was recorded at the canal site. Marcus (1972)
lacked any winter sampling which likely would have reduced his yearly

ratios even more than the tabulated values.

Sources of Power Plant Effects
On 18 August 1971 (D-1) and 14 October 1971 (D-Z) experimental
cooling system studies were completed to test the effect of various

sources of potential alteration to in situ community metabolism as water

 

All-l}i

is used
on ambi
tempera
(4) me«
of the
system
times

was 11

this a

aCCOm'
in GP

metab

the d
cont!
GPP 1
char;
intiu
ture,
ulat‘

tempe

t 1V3
influ

the d

48

is used for cooling (Figure 10, Table 8). Potential sources of effects

on ambient metabolism included: (1) mixing of river and lake water, (2)
temperature alteration, (3) length of exposure to increased temperature,
(4) mechanical damage and temperature alteration combined. An estimate

of the ability of the community to recover after exposure to the cooling
system was also included. Discharge samples were obtained twice during
times when chlorine effects should have been minimal and once when chlorine
was likely present (0825) according to plant Operational schedules but

this was not validated by direct measurement.

The effects of the cooling system were in some ways very consistent
on both dates and in other respects much less consistent. On D-1 and
D-2, all water returned to ambient temperatures of the lake showed
relatively lower GPP and respiration rates. The temperature alteration
accompanying those shifts was about a 10 C decrease. Uniform recovery
in GPP to that of the lake suggests only temporary changes in community
metabolism.

In August, water from the lake (22 C) and intake (25 C) incubated in
the discharge canal (33 C) either maintained or exhibited lower GPP than
control samples at those sites. Discharge water had a slightly lower
GPP than intake water on that date but intake water transferred to dis—
charge conditions was inhibited (+ 8 C change). In October, the GPP of
intake water was apparently stimulated by exposure to discharge tempera-
tures (+ 9 C). Plume origin water (lower discharge canal) was also stim-
ulated by exposure at the discharge temperatures (+ 2 C change). Ambient
temperature appears to govern the different responses between dates.

Mechanical effects appeared to be inhibitory in October but stimula-
tive in August. In addition, length of exposure to heated water has little
influence on the maintenance of GPP which was as high in the plume as in

the discharge on both dates.

 

mmh.m >O=hm

49

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.w manna

H8. .88 m 633

51

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52

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53

Respiration stays the same or is increased whenever a significant
temperature elevation occurs whereas respiration significantly decreases
when incubation takes place at cooler temperatures. Mixing of lake and
river water at the intake decreased respiration over that of the river

water by dilution with lake water that has a relatively low oxygen demand.

DISCUSSION

Changes in Community Structure

 

Algal community composition did not exhibit any significant changes
after plant Operation. Comparing mean 1970 class volumes (Marcus, 1972)
with 1971 class volumes suggests maximal variation was as much as 10
times different between 1970 and 1971 for each class except when blue-
green algae were not present in 16 April 1971 samples. This variation
could easily lie within experimental chance. Neither did any of the
dominant phytoplankton species of pre-operational (1970) studies show
pronounced responses to cooling water use during 1971 studies. The num—
ber of species was lower in 1971 but this was consistent over the whole
study area, nor did species diversity significantly change at the lake and
inshore sites between years. Slightly higher than predicated (Table 10),
algal biomass (volume) occurred in the discharge canal during 1971 but
mean cell size in the discharge canal showed similar fluctuations between

the two years.

Annual Variation in Cross Primary Productivity

The most useful tool for discovering short-term integrated physio-
logical shifts in phytoplankton metabolism seems to be their ability to
actively fix energy. It was expected that a shift in average daily and
seasonal photosynthetic rates at the discharge station would accompany
plant operation. Marcus' (1972) measurements of mean daily GPP in the
river during 1970, when cooling water was not being pumped, closely

approximated measurements made at the river station in 1971 (Table 9).

54

55

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Av m v
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56

But the 1971 mean daily GPP at the lake station increased 155% over the
previous year while the daily rate at the discharge site increased only
22% from the 1970 mean GPP.

A list of possible factors which influence the phytoplankton com-
munity aid in explaining yearly differences in GPP ( Table 9). Average
concentrations of important macronutrients were essentially similar in
1970 and 1971 and therefore were probably not limiting algal growth. It
is impossible that observed changes in lake GPp were caused by any change
in nutrient concentrations. Temperature differences from 1970 to 1971
were negligible so that temperature was not responsible for higher GPP
in the lake during 1971. ZOOpIankton abundances (grazers and non-grazers)
may have slightly increased in the lake over 1970, probably as a response
to higher GPP in 1971 rather than decreased predation because fish
abundances in the lake were similar in 1970 and 1971 (Cole, 1973). Wind
velocities on study dates were similar but overall slightly calmer
weather occurred in 1971. This may have been partly responsible for
reduced suspended solids in the lake during 1971. Suspended solids in
western Lake Erie have been demonstrated to affect the variation in
light penetration with other factors being equal (Verduin, 1954). Thus,
the significant yearly differences in daily GPP at the lake were caused
by differences in light penetration. Average depth profiles of GPP in
the lake indicated a real light inhibition at the surface during 1971 in
contrast with 1970 when surface values were the greatest. Gross primary
productivity in the lake was higher in 1971 only at subsurface depths.
The fact that other parameters influencing GPP are uniformly distributed
with depth except wind disturbances implies that light indeed does control
GPP in the lake. Cody (1972) also reported a surface inhibition of

primary productivity in western Lake Erie (Bass Island region) with a

57

maximum reached between 1 and 3 meters followed by a rapid decline in
02 production and carbon assimilation below 3 meters. Similar sharp
decreases in GPP with depth were eXperienced in the study area. The
higher GPP/R ratios also reflect the significant increase in yearly GPP
at the lake in 1971 because respiration only slightly increased during
1971 (Table 9). But average ratios in 1971 at the discharge site have

remained about the same as 1970 values.

Influence of Power Plant Operation on Intake Communitngetabolism

 

Further yearly comparison of the discharge canal is constrained by
the fact that in 1971, it was actually a flow-through environment unlike
the standing situation of 1970. The effects of strong currents and river
mixing were negligible in 1970 as opposed to 1971. Therefore, an analysis
of observed vs. expected changes at the site for 1971 seems applicable.
Based on the calculated average mixing ratio of lake water to river water
(6.7:1), expected values of parameters are compared to observed values in
the discharge canal (Table 10). The difference between expected and
observed probably indicates the integrated effects of temperature change,
river and lake mixing, and mechanical effects. But it should also be
remembered that lake values were determined over one kilometer from the
intake and considerable differences could occur between the lake water
at station 3 and lake water at the intake before it reaches the cooling
system. The lake station GPP and respiration must be assumed to represent
the lake contribution at the plant intake because GPP measurements were
not made near the mixing zone of the intake.

Passage through the cooling system apparently stimulated hourly
and daily GPP and respiration in the discharge canal over that expected

from the simple mixing prOportions. Mechanical and river effects may

..

 

58

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59
interact with temperature elevation to produce higher GPP. As indicated

by the two entrainment studies, mixing at the intake of lake and river
water appeared to stimulate GPP over that which the lake experienced
before the in_§i£g community was exposed to a temperature rise. Therefore,
the A t is probably not the only stimulant. The average decrease in GPP/R
ratios suggests more energy is being used to maintain the community than
expected. Slightly higher cell carbon (biomass) occurred at the discharge
site over expected concentrations which hints of a higher standing crop

of phytoplankton. An increase over expected was also experienced in the
particulate organic carbon measurements. This may seem contrary to the
lower GPP/R ratios but it indicates that the stimulation of respiration

is primarily due to increased decomposition of detritus from the river-
lake mixture while a relatively high proportion of the GPP was retained

in cell biomass. But the GPP/B ratio also was higher than expected which
suggests that GPP has increased more than biomass in the discharge canal.
Margalef (1968) believes pOpulation stability is inversely correlated
with the GPP/B ratio and directly correlated with species diversity. A
relatively high GPP/B ratio usually is indicative of an unbalanced,
productive and eutrophic system where export of organic production
predominates. But the discharge water had a comcomitant increase in
respiratory consumption that increased at a faster rate than the algal
biomass. The source of this community respiration appears to be hetero-
trophic and undoubtedly must be concentrated on detritus breakdown
because the observed algal biomass also was higher in the discharge

canal. This sounds completely contradictory to the description of a
"eutrOphic system" but Marcus (1972) also expressed the importance of
organic import (detritus) to the study area. This is particularly

emphasized in the low GPP/R ratios common to the study area.

60
The higher species diversity than expected is usually not character-

istic of a system that exhibits high GPP/B ratios but Marcus (1972) also
reported a relatively high species diversity in the discharge canal in
conjunction with high GPP. The high diversity is probably due to the
constant import of rare species to the canal from surrounding marsh
habitats.

Any physiological alterations produced in the phytoplankton community
from cooling system effects appeared to be only temporary because GPP
recovered almost completely when bottles from the discharge canal were
returned to ambient lake conditions. This also implies that exposure to
river water and mechanical effects had no persistent after-effects on
lake phytOplankton metabolism. Temperature seemed to be the major if not
the only regulatory factor involved.

The initial ambient water temperature before passage could be involved
in determining photosynthetic inhibition or stimulation. At initial
temperatures of 26 C, a depression in GPP resulted but at 16 C, an
increase in GPP was exhibited. Similar water temperature regimes reported
by Morgan and Stross (1969) and Warinner and Brehmer (1966) produced
equivalent results in carbon fixation rates of enclosed brackish water
and river communities respectively. Heating in cooler seasons to stimulate
GPP may be desirable and advantageous.

Chlorine effects probably did not occur on days primary productivity
was measured because average hourly GPP rates were stimulated rather than
inhibited in the discharge canal during 1971. Recent studies have shown
inhibition of photosynthetic capabilities or destruction of algal cells by
chlorine (Brook and Baker, 1972; Hirayama and Hirano, 1970).

In general, I suspect that plant activities in 1971 stimulated gross

primary productivity of intake water. A localized effect probably occurred

61

from this stimulation because discharge canal GPP recovered to that
measured in the lake. The lowered GPP/R ratios appear to be caused by
detritus import into the discharge canal followed by heterotrOphic
respiration rather than algal respiration. Algal cells will eventually
become part of this detritus system unless primary consumers ingest

the cells to satisfy their energy needs. The characteristically con-
sumptive conditions of the study area would tend towards a more balanced
(GPP/R - 1) system if foreign organic import was reduced but may be
offset by accumulations of algal biomass in the vicinity of the discharge
canal due to stimulated GPP.

Because the Monroe Power Plant produced not more than 25% of its
maximum potential electrical output during post-operational studies of
1971 and early 1972, more studies are needed to predict long-term effects
of the cooling water discharge on the near-shore environment of western
Lake Erie. In particular, an examination of the ability of photosynthetic
organisms to recover from cooling system passage might be sought by
in_§i£2_laboratory culture of post-condenser communities. By exposing
these communities to various temperatures over ambient, a time factor
for recovery may be obtained as measured by such parameters as oxygen

production, carbon fixation or ATP activity in those cultures.

LITERATURE CITED

American Public Health Association. 1971. Standard methods for the
examination of water and wastewater, 13th ed. Washington. 874 p.

Brook, A. J. and A. L. Baker. 1972. Chlorination at power plants:
impact on phytOplankton productivity. Science 176: 1414-1415.

Cody, T. E. 1972. Primary productivity in the western basin of Lake
Erie. Ph.D. thesis, Ohio State Univ., Columbus. 113 p.

Cole, R. A. 1972. Physical and chemical limnology along the western
shore of Lake Erie. Tech. Rpt. 13, Inst. Water Res., Mich. State
Univ., E. Lansing, Mich. 120 p.

. 1973. An ecological evaluation of a thermal discharge:
summary of early post-Operational studies. Tech. Rpt. 32.0, Inst.
Water Res., Mich. State Univ., East Lansing, Mich. 43 p.

 

Davis, C. C. 1964. Evidence for the eutrophication of Lake Erie from
phytoplankton records. Limnol. Oceanogr. 9(3): 275-283.

Drouet, F. 1968. Revision of the classification of the Oscillatoriaceae.
Acad. Natur. Sci. Philadelphia, Monogr. 15. 370 p.

and W. A. Daily. 1956. Revision of the coccoid myxophyceae.
Butler Univ. Bot. Stud. 12: 1-218.

 

Federal Water Pollution Control Administration. 1968. Lake Erie Report:

A plan for water pollution control. U.S. Department of the Interior,
104 p.

Gebelein, N. 1971. Analysis of plankton and periphyton, p. 31-59. 13:
Williams, C. C., g£_§l, Studies on the effects of a steam-electric
generation plant on the marine environment at Northport, New York.
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Stony Brook, N. Y. 119 p.

Giese, A. C. 1972. Cell physiology. 2nd edition. Saunders, Philadelphia
592 p.

Hirayama, K. and R. Hirano. 1970. Influences of high temperature and
residual chlorine on marine phytOplankton. Mar. Bio. 7: 205—213.

Hoar, W. S. 1966. General and comparative physiology. Prentice—Hall,
Inc., New Jersey. 815 p.

62

63

Hustedt, F. 1930. Bacillariophyta (Diatomeae). Die susswasser-flora
mitteleuropas, Part 10. Jena. 466 p.

MacArthur, R. H. 1965. Patterns of species diversity. Biol. Rev.
40: 510-533.

Marcus, M. D. 1972. The distribution of phytoplankton and primary
productivity near the western shore of Lake Erie. M.S. thesis.
Mich. State Univ., E. Lansing, Mich. 96 p.

Margalef, R. 1968. Perspectives in ecological theory. Univ. of Chicago
Press, Chicago. 111 p.

McNabb, C. D. 1960. Enumeration of freshwater phytoplankton concentrated
on the membrane filter. Limnol. Oceanogr. 5(1): 57-61.

Michigan Department of Natural Resources. 1970. Great Lakes Algae
Monitoring Program--1969. Bureau of Water Management. Lansing,
Mi. 16 p.

Morgan, R. P. II and R. G. Stross. 1969. Destruction of phytoplankton in
the cooling water supply of a steam electric station. Chesapeake
Sci. 10(3 and 4): 165-171.

Nalepa, T. F. 1972. The distribution of 200p1ankton along the western
shore of Lake Erie. M.S. thesis. Mich. State Univ., E. Lansing,
Mi. 112 p.

Parkhurst, B. R. 1971. The distribution and growth of the fish populations
along the western shore of Lake Erie at Monroe, Michigan, during
1970. M.S. thesis, Mich. State Univ., E. Lansing, Mich. 71 p.

Patrick, R. 1969. Some aspects of temperature on freshwater algae, p.
161-165. In; P. A. Krenkel and F. C. Parker (eds.) Biological
aspects of thermal pollution. Vanderbilt University Pres.

and C. W. Reimer. 1966. The diatoms of the United States,
Vol. I. Acad. Nat. Sci. Phila. Monogr. 13. 688 p.

 

, B. Crum, and J. Coles. 1969. Temperature and manganese
as determining factors in the presence of diatom or blue-green
algal floras in streams. Proc. Nat. Acad. Sci. USA, 64: 472-478.

 

Pielou, E. C. 1969. An introduction to mathematical ecology. Wiley,
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Prescott, G. W. 1962. Algae of the western Great Lakes area. W. C.
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Schindler, D. W. 1971. Production of phytOplankton and 200plankton in
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64

Steele, R. G. D. and J. H. Torrie. 1960. Principles and procedures of
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plankton from cell volume or plasma volume. Limnol. Oceanogr.
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Strickland, G. D. H. 1960. Measuring the production of marine phyto—
plankton. Bull. Fish. Res. Bd. Can. 122. 172 p.

Suchanek, T. H., Jr. 1971. Analysis of primary productivity, p. 21-29.
12; Williams, C. C., g£_§1, Studies on the effects of a steam-
electrical generating plant on the marine environment at Northport,
N. Y. Tech. Rpt. 9, Marine Sciences Res. Ctr., State Univ. of New
York, Stony Brook, N. Y. 119 p.

Taft, C. E. 1945. The desmids of the west end of Lake Erie. Ohio J.
Sci. 45: 180-205.

and C. W. Taft. 1971. The algae of western Lake Erie. Bull.
Ohio Bio. Survey 4(1): 1-189. Ohio State Univ., Columbus. 189 p.

 

Tiffany, L. H. 1934. The plankton algae of the west end of Lake Erie.
Contrib. F. T. Stone Lab., Ohio State Univ., No. 6: 1-112.

Verduin, J. 1954. Phytoplankton and turbidity in western Lake Erie.
Ecology 35(4): 550-561.

Warinner, J. E. and M. L. Brehmer. 1966. The effects of thermal effluents
on marine organisms. Air and Water Poll. Int. J. 10: 277—289.

Weber, C. I. 1971. A guide to the common diatoms at water pollution
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Cincinnati, Ohio. 101 p.

Westlake, D. F. 1969. Interpretation of results: units and compar-
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methods for measuring primary production in aquatic environments.
Blackwell Sci. Pub., Oxford and Edinburgh, England.

APPDENICES

65

Appendix Al. Water temperature elevation in the cooling system at the
one meter depth, Raisin River flow, and pumping rate at the
Monroe Power Plant site recorded at the time of plankton
collections and primary productivity studies.

 

 

River flow Pumping Rate

Date m3/sec1 m /sec A t (over intake)
4-16 15.4 0 0
5-20 6.3 23 5
5-21 5.9 23 4
6-3 5.5 23 1
6-17 5.6 23 6
6-18 4.9 23 8
6-30 2.6 23 7
7-14 1.9 23 8
7-16 2.0 23 8
7-29 1.9 23 9
8-16 1.6 23 2
8-18 1.2 23 10
8-26 1.1 23 1
9-15 1.2 23 8
9-17 1.4 23

10-2 3.0 23

10-14 2.7 23 10
10-15 2.6 23 8
10-16 2.3 23 5
10-30 3.7 23 4
11-12 2.8 23 9
11-13 2.7 23 7
2-29-72 5. 7 462 1
5-12 25.2 23 8
6-1 9.2 46 4
6-14 5.2 46 5

 

1City of Monroe contributes not more than a 10% additional discharge to
river flow before it reaches the plant intake.

2Doubled because pumping units doubled.

66

Appendix A2. Periodic total residual chlorine concentrations (ppm) con—
densed from plant Operational reports of 1971 and 1972.

 

 

Date ppm1 Date ppm Date ppm
6-24-71 0.52 10-10 0.05 2-27-72 0.35
6-29 0.42 10-12 0.05 2-28 0.
7-1 0.52 10-15 0.05 4-11 0.5
7-6 0.42 10-20 0.2 4-20 0.
7—8 0.72 10-22 0.05 5-2 0.07
7-12 0.52 10-27 0.1 5-4 0.05
7-15 0.152 11-1 0.05 5-9 0.06
7-21 0.152 11-5 0.1 5-11 0.04
7-23 0.152 11-11 0.1 5-16 0.07
7-27 0.22 11-12 0.1 5-18 0.12
8-10 0.22 11-16 0.0 5-25 0.0
8-17 0.152 11-19 --- 5-27 0.1
8-20 0.202 11-22 0.05
9-8 0.152 11-27 0.10
9-10 0.752 12-7 0.2
9-11 0.752 12-14 0.25
9-15 0.13 12-16 0.25
9—17 0.13 2-1-72 0.35
9-22 0.13 2-3 0.35
9-25 0.053 2—8 0.25
9—28 0.053 2-10 0.25
9-30 0.053 2-15 0.20
10—5 0.05 2-17 0.20
10-8 0.05 2-21 0.30

 

1All values are average total residual unless indicated otherwise

2Maximum total residual chlorine

3Free chlorine

67

mhmuma m CH wawHHHm

coomumumm I m

wchuos I B
“swamp Acov ououmummaoa

 

 

H
n.0NIm.ON N.O o.HNIo.HN H.O o.mHIo.HN H.O HHIo
NNINN H.O o.mHION 0.0 o.wHIm.wH m.o HIo
o.mHIo.mH 0.0 o.HHIm.HH III O.OHIo.0H N.H NHIm
m.HIo.n o.H IIIIII III o.HIo.H O.N NNImNIN
o.HHIo.HH N.O n.0Io.N H.O O.NIo.N o.H MHIHH
mHINN H.O O.NHIm.wH H.O m.OHI.NH m.o OmIOH
o.mHIm.mH m.o o.mHIm.mH H.O o.HHIm.¢H o.H OHIOH
o.mNIo.mN m.o O.OHIm.OH m.o o.mHIo.mH 0.0 HHIOH
mNION H.O o.HNImN m.o ONIm.HN N.H NIOH
m.mNIm.mN 0.0 o.MNIm.mN m.o O.NNIo.NN m.o mHIm
m.MNIo.HN Amvm.o Havo.o o.qNIo.qN Amvm.o Aavm.o m.NNIo.mN Amvn.o navo.c ONIw
o.NmIm.mm 0.0 o.HNIo.ON m.o NNIo.MN m.o wHIm
o.omIm.Hm 0.0 MNImN 0.0 NNINN H.H mNIN
m.mNI.Nm m.o m.mNIo.mN 0.0 mNIMN o.H OHIN
m.NmIo.Hm m.o ONIm.NN H.o mNIn.ON m.o omIo
O.OmIm.Nm III HNImN III O.NNIm.mN III wHIo
mHINN III m.mHION III o.wHINN III nIo
HNINN III o.oNIm.oN III m.NHIm.NH III HNIHNIn
manumuOQEOH Haooom ousumuomamh Hauomm munumuomama Aav Hsoomm Omen
MazdmomHo mm>Hm H mxda

ZOHH¢Hm

 

.mmHooum zuH>Huo=ooum mumaHum mo maHu may um omouoomu mumumamumm HOOHmhsm .m< xHomumm<

68

Appendix A4. Measured daily solar radiation in conjunction with primary
productivity studies of 1971 and 1972.

 

 

Langleys/discharge It/
Date Langleys/day in situ period Ip
4-15-71 517 239 _-_.
5-21 565 245 2.33
6-3 604 284 2.13
6-18 626 277 2.27
6-30 534 239 2.22
7-16 581 237 2.44
7-29 582 249 2.33
8-18 510 --- ---
8-26 263 --- ---
9-15 342 128 2.63
10-2 294 137 2.17
10-14 212 --- --_
10-16 231 104 2.22
10-30 198 112 1.79
11-13 200 104 2.22
2-29-72 190 140 1.35
5-12 698 309 2.27
6-1 423 275 1.54

6-14 632 296 2.13

 

69

 

NH Oz H.HH ON HO OHIOH
OH HO 0.0 OO OO OHIOH
OH :O O.OH ON OO HHIOH
NN OO O.N ON NN NIOH
OH Oz O.NH ON OO NHIO
ON 32 O.HH ON NO OHIO
ON 3O O.HH ON HO ONIO
HN OO N.O 0.0 OOH OHIO
ON O N.HH OO NO OHIO
OH 3 N.O OO ON ONIN
HN 3 O.NH ON OO OHIN
HN :z 0.0H ON NO HHIN
NN Oz O.HH OH OO OOIO
HN OO N.OH OH OO OHIO
HN mm 0.0 0.0 OOH NHIO
OH 2 O.HH OO HO OIO
OH 32 0.0H OH HO HNIO
OH : 0.0H ON ON ONIO
OH HO N.N OH OO OHIH
Hoov ouaumumm aoHuouan A.ua\axv woman um>oo hen How mmHnm Ouma
Isms HHH OOH: OOH: mOmHo>< New N Iesm HHOHOOOO N
wcHHHm>Oum

 

.AanHNmHv mmHosum OuH>Huo=ooum one scuxumHm Om mechuuom GOHOOEMONOH HOOHwOHOHOOOOZ .m< Nvaumm<

70

 

OH
«H
«H

OH
NH

3m
3m

Mm
mz
3m
mm
m2

N.mH
N.MH
O.cH
O.¢H
m.qH
m.OH
0.0

m.HH

OO
OO
ON
OOH
OO
OOH
Om
ON

Nn
NN
OOH
mm
mm

mm
«O

OHIO
HIO
NHIm
NNIONIN
MHIHH
NHIHH
OmIOH
OHIOH

N.O.OOOV n< menumad

71

Appendix A6 . Mean total algal numbers (1971) (X 103/liter).

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Date Station

4-16-71 1 3 6 8 9
1791 1844 3546 3755 4004

5-20—71 6 9 8 1 3
2808 4462 4618 4926 5348

6-17-71 6 3 8 9 l
1977 6582 10014 10995 12857

7-14-71 6 3 1 9 8
889 1628 1827 5265 8537

8-16-71 3 6 l 9 8
3640 4811 11948 18948 23289

9—17-71 1 6 3 9 8
2910 3157 3283 4122 5022

10-15-71 6 3 9 l 8
6238 8536 9312 11998 13636

11-12-71 1 3 6 8 9
1732 1781 1908 2987 4004

 

 

 

72

Appendix A7. Mean total algal volume (1971) (X 10-4 ml/liter).

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Date Station

4-16-71 1 3 9 8 6
488 576 586 645 1528

5-20-71 6 9 8 3 l
712 942 1171 2730 2992

6-17-71 6 8 9 l 3
985 2901 4780 4787 5633

7-14-71 6 l 3 9 8
505 859 1342 3412 4505

8-16—71 3 6 l 9 8
1851 2059 5922 8877 10222

9-17-71 9 3 8 6 l
1605 1689 2014 2062 2226

10—15-71 9 6 8 3 1
2321 2713 3405 3628 5940

11-12-71 9 l 6 3 8
611 695 733 768 1704

 

 

Appendix A8.

73

Mean individual phytoplanktonic volume (1971) in u3.

 

 

 

 

 

 

 

 

 

 

 

 

 

Date Station
4-16-71 9 8 l 3 6
1463 1714 2733 3152 4277
5-20—71 9 6 8 3 l
2330 2515 2586 5114 5890
6-17-71 8 l 9 6 3
2952 3928 4308 5069 8538
7-14-71 1 8 6 9 3
5034 5533 5553 6441 8072
8-16-71 8 6 9 1 3
4396 4415 4755 5026 5088
9—17-71 9 8 3 6 l
3926 3946 5072 6207 7524
10-15-71 9 8 6 3 1
1692 3674 4247 4332 4956
11-12-71 9 6 l 3 8
1531 3869 4004 4249 5718

 

 

 

74

Appendix A9. Mean volume of blue-green algae (1971) (X 10..6 ml/liter).

 

 

 

 

 

 

 

 

 

 

 

 

 

Date Station
4—16-71 1 3 6 8 9
0 0 0 O 0
5-20—71 6 8 9 l 3
40 2687 3461 7464 9170
6-17—71 6 3 8 l 9
1228 4714 6586 8156 20236
7-14-71 6 1 3 9 8
541 554 1524 13894 22090
8-16-71 3 6 l 9 8
9848 12805 49445 74124 90836
9-17-71 9 3 8 l 6
1536 4257 6072 6807 8876
10-15-71 9 8 6 3 l
2667 13384 14229 14788 29718
11-12-71 9 6 3 l 8
326 689 1198 1448 1684

 

 

Appendix A10.

75

Volumetric dynamics of Aphanizomenon flos-aguae (X 10-

 

 

 

 

ml/liter).
Station

Date 1 3 6 8 9
4-16-71 (1)1 0 0 o 0 0
5-20 (3) 7.5 44.8 4.0 8.8 32.3
6-17 (5) 6.9 3.2 15.5 10.6 0
7-14 (7) 44.8 38.6 19.5 76.3 83.3
8-16 (9) 2112.6 614.7 906.0 3744.4 2867.7
9-17 (11) 53.8 17.7 197.2 30.6 41.7
10-15 (13) 503.4 469.5 305.4 281.0 165.9
11-12 (15) 28.4 25.9 38.3 57.2 13.5
1Designates a coded date.

76
5

 

 

 

Appendix.All. Volumetric dynamics of Anacystis incerta (X 10- ml/liter).
Station

Date 1 3 6 8 9
(1) 0 0 0 0 0
(3) 0 0 0 O O
(5) 75.99 75.99 0 0 1114.5
(7) 0 21.2 0 1101.9 227.98
(9) 1773.1 141.2 211.7 3580.0 3039.7
(11) 338.8 183.5 561.2 63.5 63.5
(13) 1891.3 405.3 861.2 296.4 0
(15) 25.4 0 0 0 0

 

Appendix A12. Volumetric dynamics of Anacystis thermalis (X 10.5 ml/liter).

 

 

 

Station
Date 1 3 6 8 9
(1) 0 0 0 0 0
(3) 0 0 0 0 10.6
(5) 59.3 29.7 5.3 389.8 779.6
(7) 0 63.8 34.5 635.4 779.6
(9) 144.1 14.1 30.1 508.4 632.7
(11) 106.3 178.8 40.2 124.0 10.6
(13) 259.9 192.1 118.6 0 0

(15) 14.2 24.1 10.6 0 15.9

 

77
5

 

 

 

Appendix A13. Volumetric dynamics of Anacystis gyanae (X 10- m1/1iter).
Station

Date 1 3 6 8 9

(1) 0 0 0 0 0

(3) O 0 0 0 0

(5) 0 0 18.6 0 0

(7) 0 28.9 0 12.4 44.5

(9) 593.6 190.2 105.4 949.7 791.4
(11) 24.8 0 61.1 310.1 0

(13) 118.7 0 0 612.2 0

(15) O 0 0 O 0

78

Appendix A14. Mean volume of green-algae (1971) (X 10..6 m1/liter).

 

 

 

 

 

 

 

 

 

 

 

 

 

Date Station
4—16-71 1 6 3 9 8
42 49 62 315 1013
5-20-71 8 6 9 1 3
1707 1905 2026 2356 2481
6-17-71 6 8 1 9 3
4944 12809 20852 21546 35056
7-14-71 6 1 3 8 9
2678 3199 9598 17498 18254
8-16-71 1 6 8 3 9
896 2124 3136 4396 10505
9-17-71 3 6 8 1 9
2623 3776 7087 8962 9229
10-15—71 9 6 8 3 1
6363 6829 9922 14010 18234
11-12-71 9 8 3 1 6
220 440 2218 2251 2683

 

 

 

79
5

 

 

 

Appendix A15. Volumetric dynamics of Houseotia elegantula (X 10- ml/
liter).
Station
Date 1 3 6 8 9
(1) O 0 0 0 0
(3) 0 0 0 0 0
(5) o o o o 34.0
(7) 0 0 0 0 0
(9) 0 0 0 0 45.4
(11) 240.5 85.4 0 237.3 170.9
(13) 635.9 620.8 238.5 270.5 61.7

(15) 60.8 29.6 33.2 23.7 14.2

 

8O

Appendiqu16. Volumetric dynamics of Pediastrum duplex (X 10-5 m1/1iter).

 

 

 

Station
Date 1 3 6 8 9
(1) 0 0 0 0 0
(3) 52.0 103.9 0 0 0
(5) 1885.5 3294.5 415.7 1118.9 1740.5
(7) 215.2 652.4 164.5 1391.2 1409.0
(9) 0 358.0 0 0 165.8
(11) 150.1 34.6 10.0 173.2 277.1
(13) 718.3 331.5 124.3 536.9 155.9
(15) 110.8 159.3 112.6 0 0

 

Appendix A17. Volumetric dyanmics of Pediastrum simplex (X 10-5 m1/liter).

 

 

 

Station
Date 1 3 6 8 9
(1) 0 0 0 0 0
(3) 0 0 0 0 0
(5) 0 0 0 0 o
(7) 0 58.8 .22.0 0 0
(9) 0 0 146.9 0 656.1
(11) 470.1 58.8 350.4 176.3 381.9
(13) 281.2 281.2 210.9 88.1 293.8

(15) 0 O 88.1 0 O

 

Appendix A18 .

Mean diatom volume (1971) (X 10-

81

6

m1/1iter).

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Date Station

4-16-71 1 8 9 3 6
4833 5438 5544 5690 15232

5-20-71 9 6 8 3 1
3805 4973 7023 15392 20103

6-17-71 6 9 8 1 3
2999 5111 7789 12481 13879

7-14-71 6 3 9 1 8
1665 1679 1722 3719 5179

8-16-71 3 6 1 9 8
895 1021 2068 3859 5374

9-17-71 9 8 1 6 3
3755 5080 5356 7042 9814

10-15—71 6 3 8 1 9
5860 6980 10380 10920 13927

11-12-71 1 6 3 9 8
3188 3898 - 3992 5548 14912

 

 

 

 

82

 

 

 

Appendix .Al9. Volumetric dynamics of Coscinodiscus radiatus (X 10-5

m1/1iter).

Station

Date 1 3 6 8 9
4—16-71 73.6 68.1 22.7 61.5 109.6
5-20 253.8 77.9 83.3 66.9 10.6
6-17 22.9 470.4 265.9 47.1 28.4
7-14 182.9 89.7 104.1 257.3 82.97
8-16 185.5 82.3 93.4 323.6 271.98
9-17 410.6 524.8 602.6 403.6 269.7
10-15 404.9 174.3 158.3 196.9 196.4
11-12 83.0 113.1 124.5 61.9 77.9

Appendix A20. Volumetric dynamics of Melosira granulata (X 10-5 m1/1iter).

 

 

 

Station
Date 1 3 6 8 9
(1) 92.1 222.2 222.4 120.4 53.7
(3) 141.98 50.9 0 87.4 41.6
(5) 134.7 49.3 5.4 122.9 0
(7) 4.1 4.8 0 35.98 13.5
(9) 4.2 0 4.2 13.2 14.0
(11) 21.7 192.6 12.6 30.98 31.1
(13) 137.9 85.3 51.7 110.1 96.2

(15) 27.9 34.97 43.0 45.4 50.9

 

83

 

 

 

 

 

 

 

Appendix A21. Volumetric dynamics of Stephanodiscus astraea (X 10-5
mllliter).
Station
Date 1 3 6 8 9
(1) 2.8 18.7 52.1 86.8 120.9
(3) 185.5 357.6 106.1 161.4 142.4
(5) 102.7 80.0 1.0 33.2 10.0
(7) 1.5 0.87 0.4 0 2.4
(9) 3.0 3.2 4.2 9.5 10.1
(11) 15.7 31.8 9.0 11.2 5.6
(13) 416.3 273.0 211.1 585.6 797.4
(15) 35.2 48.3 74.9 1146.1 243.0
Appendix A22 . Volumetric dynamics of Stephanodiscus tenuis (X 10.-5
m1/liter).
Station
Date 1 3 6 8 9
(1) 50.1 45.0 82.5 47.4 30.2
(3) 0 2.9 4.1 7.4 14.0
(5) 23.6 3.7 0.3 29.4 6.2
(7) 0.9 0.14 0.12 18.2 1.5
(9) 0.24 0.0 0.12 14.1 6.3
(11) 5.5 7.2 0.7 26.1 34.9
(13) 7.7 5.99 7.7 74.3 221.6
(15) 2.9 1.96 0.8 13.6 37.1

 

84
5

 

 

 

 

 

 

 

Appendix .A23. Volumetric dynamics of Fragilaria capucina (X 10- ml/liter).

Station
Date 1 3 6 8 9
(1) 1.6 0 43.2 0.0 0.0
(3) 29.0 47.6 19.2 15.3 0.0
(5) 38.4 8.2 0.0 0.0 0.0
(7) 15.81 10.4 0.0 0.0 0.0
(9) 0.0 0.0 0.0 0.0 0.0
(11) 0.0 14.6 20.9 6.7 3.0
(13) 39.9 71.6 102.4 34.3 22.1
(15) 113.9 132.4 108.0 86.9 62.8
Appendix A24. Volumetric dynamics of Fragilaria crotonensis (X 10.-5

m1/liter).

Station
Date 1 3 6 8 9
(1) 78.9 47.1 421.8 18.1 0.0
(3) 321.4 80.4 67.4 40.4 21.5
(5) 74.3 47.3 0 36.6 19.0
(7) 20.8 10.9 0.0 0.0 0.0
(9) 0.0 0.0 0.0 8.3 10.5
(11) 15.3 0.0 0 0.0 0.0
(13) 42.0 16.8 41.97 0.0 0.0

(15) 0.0 0.0 8.1 0.0 0.0

 

85

 

 

 

Appendix A25. Volumetric dynamics of Tabellaria fenestrata (X 10-5
m1/liter).
Station

Date 1 3 6 8 9
(1) 84.8 121.4 480.9 63.5 40.2
(3) 1004.2 800.4 205.3 227.0 75.7
(5) 823.5 705.4 0.0 343.2 134.0
(7) 122.1 42.7 61.4 46.0 0.0
(9) O O O 0 O
(11) 51.3 171.95 51.6 0 O
(13) 0 19.66 0 0 0
(15) 7.8 54.5 0 21.4 0

-5
Appendix A26. Volumetric dynamics of Cryptomonas ovata (X 10 m1/1iter).

 

 

Station

Date 1 3 6 8 9
(1) 0 0 0 0 0
(3) 0 0 0 0 0
(5) 121.9 156.3 56.4 64.5 34.1
(7) 23.0 5.2 1.1 1.2 16.7
(9) 14.3 16.3 1.6 55.8 27.9
(11) 30.3 0 77.1 75.4 43.96
(13) 51.0 33.5 10.2 6.99 10.7
(15) 6.8 4.7 6.2 0 1.2

 

86

 

 

 

Appendix A27. Volumetric dynamics of Ceratium hirundinella (X 10-5
m1/liter).
Station

Date 1 3 6 8 9
(1) 0 0 O 0
(3) O O 0 O
(5) 0 57.1 0 0
(7) 80.9 52.99 11.9 0 O
(9) 532.5 296.8 421.3 152.1 0
(11) 74.2 0 47.7 47.7 47.7
(13) 0 0 0 0

(15) 0 19.1 0 0

 

87

Appendix A28. Mean number of species at each station in 1971.

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Date Station
4-16-71 3 1 6 8 9
19 23 23 37 39
5-20-71 6 1 3 9 8
24 34 35 38 47
6-17-71 6 1 3 9 8
30 31 32 36 42
7—14-71 6 3 1 9 8
20 3O 36 37 38
8—16-71 6 3 1 9 8
21 22 23 36 43
9—17-71 6 1 3 9 8
28 36 42 47 51
10—15-71 6 3 l 9 8
32 33 40 42 44
11-12-71 6 1 3 8 9
29 38 39 40 41

 

 

Appendix A29. Mean phytoplanktonic diversity (1971).

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Date Station
4—16-71 1 3 6 8 9
2.73 2.83 3.07 4.04 4.26
5-20—71 6 3 1 9 8
3.59 3.72 3.79 4.02 4.21
6-17-71 1 9 6 3 8
2.68 3.15 3.25 3.58 3.93
7-14-71 6 1 3 8 9
3.38 3.64 3.72 4.17 4.30
8-16-71 6 1 3 8 9
1.02 1.35 1.63 2.02 2.19
9-17-71 6 3 l 9 8
2 69 4.05 4.08 4.19 4.36
10-15-71 9 8 3 6 1
3.16 3.36 3.76 3.88 4.00
11-12-71 9 6 3 8 1
3.58 3.79 4.16 4.26 4.36

 

 

 

 

Appendix A30. Mean phytoplanktonic equitability (1971).

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Date Station
4-16-71 1 6 3 8 9
.29 .36 .37 .45 .48
5-20—71 3 8 1 9 6
.38 .39 .41 .43 .49
6-17-71 1 9 6 8 3
.21 .24 .32 .36 .37
7-14—71 1 3 8 6 9
.34 .42 .47 .52 .53
8-16-71 6 8 1 9 3
.10 10 .11 .12 .14
9-17—71 6 3 9 8 1
24 .39 .39 .40 .46
10—15-71 9 8 1 3 6
21 .23 .40 .42 .46
11-12—71 9 3 8 6 1
.30 .46 .47 .47 .53

 

 

Appendiqu3l. Analysis of variance of factorial design comparing year,

station and month factors.

 

(1) Number of Species

 

 

 

Source df M.S. F Stat.
Year (A) 1 4890.63 635.75**
Station (B) 4 2310.22 300.31**
A x B 4 94.21 12.25**
Months (C) 7 644.82 83.82**
AC 7 361.30 46.97**
BC 28 98.54 12.81**
ABC 28 99.18 12.89**
Error 240 7.69
(2) Phytoplanktonic Numerical Diversity
Source df M.S. F. Stat.
A 1 0.26 3.72*
B 4 4.21 60.18**
AB 4 0.35 4.96**
C 7 10.34 147.96**
AC 7 4.94 70.70**
BC 28 1.61 23.06**
ABC 28 1.18 16.88**
Error 240 0.07
(3) Phytoplanktonic: Numerical Equitability
Source df M.S. F Stat.

A 1 0.41 164.82**
B 4 0.02 9.48**
AB 4 0.003 1.23N.S.
C 7 0.296 118.49**
AC 7 0.079 31.71**
BC 28 0.047 19.01**
ABC 28 0.044 17.81**
Error 240 0.002

Appendix A31(con't.)

(4) Total Phytoplanktonic Numbers (Units)

 

 

 

Source df M.S. F. Stat.
A 1 0.227 15.82**
B 4 2.112 147.28**
AB 4 0.011 0.76N.S.
C 7 1.722 120.101**
AC 7 0.764 53.32**
BC 28 0.293 20.46**
ABC 28 0.232 16.24**
Error 240 0.014

(5) Total PhytOplanktonic Cell Volume

Source df M.S F. Stat.
A 1 0.007 0.376N.S.
B 4 0.777 39.70**
AB 4 0.072 3.67*

C 7 2.722 139.12**

AC 7 1.231 62.90**

BC 28 0.363 18.53**
ABC 28 0.242 12.39**
Error 240 0.0196

(6) Blue-green Algal Volume

Source df M.S. F. Stat.
A 1 37.99 49.29**

B 4 1.36 1.77N.S.
A x B 4 2.56 3.32*

C 7 122.12 158.44**
AC 7 19.37 25.14**

BC 28 1.77 2.29**
ABC 28 1.84 2.38**
Within 240 0.77

Appendix A3l(con't.)

(7) Green Algal Volume

92

 

 

Source df M.S. F. Stat.

A 1 32.34 107.51**

B 4 1.76 5.84**

AB 4 0.16 0.52N.S.

C 7 13.75 45.71**

AC 7 4.43 14.74**

BC 28 1.09 3.63**

ABC 28 0.42 1.38N.S.
Within 240 0.30

(8) Diatom Volume

Source df M.S F. Stat.
A 1 0.18 9.88**
B 4 0.95 52.29**
AB 4 0.35 19.05**
C 7 5.50 301.17**
AC 7 1.55 84.7**
BC 28 0.35 19.06**
ABC 28 0.22 12.35**
Error 240 0.02

 

* - significant at the a =

** — significant at the a

.05 level.

.01 level.

710

“NW

2
3

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