Ecological Research Series
SEASONAL EFFECTS ON TEMPERATURE
PREFERENCE IN YELLOW PERCH,
PERCA FLAVESCENS
Environmental Research Laboratory
Office of Research and Development
U.S. Environmental Protection Agency
Duluth, Minnesota 55804
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EPA-600/3-77-088
August 1977
SEASONAL EFFECTS ON TEMPERATURE PREFERENCE
IN YELLOW PERCH, PERCA FLAVESCEHS
by
Robert W. McCauley
Department of Biology
Wilfrid Laurier University
Waterloo, Ontario, N2L 3C5
Grant No. R802U67
Project Officer
Kenneth E.F. Hokanson
Environmental Research Laboratory - Duluth
Monticello Ecological Research Station
Monticello, Minnesota 55362
ENVIRONMENTAL RESEARCH LABORATORY - DULUTH
OFFICE OF RESEARCH AND DEVELOPMENT
U.S. ENVIRONMENTAL PROTECTION AGENCY
DULUTH, MINNESOTA 5580^
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DISCLAIMER
This report has been reviewed by the Environmental Research Laboratory -
Duluth, and approved for publication. Approval does not signify that the
contents necessarily reflect the views and policies of the U.S. Environmental
Protection Agency, nor does mention of trade names or commercial products
constitute endorsement or recommendation for use.
ii
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FOREWORD
Our nation's freshwaters are vital for all animals and plants, yet our
diverse uses of water for recreation, food, energy, transportation, and
industry physically and chemically alter lakes, rivers, and streams. Such
alterations threaten terrestrial organisms, as well as those living in water.
The Environmental Research Laboratory in Duluth, Minnesota develops methods,
conducts laboratory and field studies, and extrapolates research findings
to determine how physical and chemical pollution affects aquatic life
to assess the effects of ecosystems on pollutants
to predict effects of pollutants on large lakes through the use of
models
to measure bioaccumulation of pollutants in aquatic organisms
that are consumed by other animals, including man.
This report describes the seasonal temperature preference of adult
yellow perch. This report complements other reports describing the
temperature requirements of this species and provides a basis for evaluation
of laboratory gradient tanks as a short-cut method to measure thermal effects.
Donald I. Mount, Ph.D.
Director
Environmental Research Laboratory
Duluth, Minnesota
iii
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ABSTRACT
Seasonal variations in temperature preferences of adult yellow perch
(Perca flavescens) were sought by acclimating fish captured in the fall to
5, 10, 15, and 20C in the laboratory and determining their preferred tempera-
tures in a horizontal temperature gradient trough. Temperatures selected in
winter for fish acclimated to 5C ranged from 12 to 14C, considerably above
the temperature region (6C and below) previously established for optimum
gametogenesis. Final preferenda as determined from preferred temperature-
acclimation temperature curves were 24 , 20 and 17C during winter, spring
and summer respectively. Those in late summer were below published values
(23.5C) for this species. It is concluded that there are no demonstrable
effects, at least in winter and early spring which might reflect changing
physiological needs and that temperature gradients in nature serve rather
to attract perch to warm temperatures suitable for spawning in spring and
conducive to-growth in summer.
iv
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CONTENTS
Foreword iii
Abstract iv
Figures vi
Acknowledgments vii
1. Introduction 1
Statement of problem 1
Factors influencing preferred temperature 3
2. Conclusions 7
3. Recommendations 8
k. Materials and Methods 9
Structure of gradient trough 9
Experimental fish. 9
Procedures of testing 10
Acclimation 10
Determination of final preferendum 11
5. Results 12
Behavior of fish in thermal gradients 12
Preferred temperature 12
Condition of the gonads of the experimental fish 13
6. Discussion 15
Review of preferred temperature studies on
yellow perch 15
The role of temperature preference in the
life of the yellow perch 19
References 22
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FIGURES
Number Page
1 Examples of preferred temperature - acclimation 4
temperature curves.
2 Preferred temperatures of a rainbow trout (Salmo 5
gairdneri) over a ten day period as recorded in
a behavioral chamber.
3 Seasonal variations in water temperature of Lake 10
St. Clair, 1974.
4 The relation between preferred and acclimation 14
temperature in late winter and early spring.
5 Summary of the preferred temperature relations 16
of adult yellow perch.
6 Observed vertical distributions of yellow perch 17
in relation to season, depth and temperature in
Lake Opeongo.
7 Seasonal depth distributions of yellow perch 18
(year 11+) in Lake Michigan.
vi
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ACKNOWLEDGMENTS
I wish to thank Dr. L.A.A. Read and Mr. J. Herringer of Wilfrid Laurier
University for assistance in carrying out experiments. Mr. Steve Nepszy,
Director of the Lake Erie Biological Research Station, Ontario Ministry of
the Environment, kindly supplied the specimens of yellow perch.
vii
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SECTION i
INTMIWIICTION
STATEMENT OF I'KOM.KM
Recant proliferation ol power general ing plants has intuited in «
corresponding Increase In the amount ol natural water used lor cooling. Watte
energy In the electric generating process Is Introduced to the water course
in Che form of a warm el fluent. The elleets on iiquatlc ticoityatems ol the warm
diluent known either as thermal enrichment or pollution, depending ""
circumstances, are now the themes ol an Increasing numbei ol Invent Igaiions
in Che new field ol thermal ecology. The perjomtive term "thermal pollution"
Implies that the addition ol calories (calnhtci U>n) has adverse effects on
other water uses. In some instances, however, calefacilon (a neutral
expression) may confer advantages to some spec Inn at l<'(>M found
that the range for the optimum development ol the embryos (I'-ioc)
Is somewhat below that for optimum growth ol juveniles (I7-1HO
(llokanson et al. 1977). There are numerous other example* ol changing thermal
requirements of a species with the Hie stage.
Perhaps the most detailed and thorough study of the thermal requirements
ol n species Is that carried out !>y Brett (1971) and his associates with the
sockeye salmon, Oncorhyneua nerki. They found from their studies on the
relation of performance, metabolism, circulation and growth that the
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physiological optimum for the species was centered around 15 C - also the
firmly established preferred temperature for the species (Brett, 1952). For
other fish a general agreement between the preferred or selected temperature
and the temperature for the optimum growth has been demonstrated.
The thermal requirements of various life stages of Perca flavescens
have also received attention, especially over the past several years (Hokanson,
1977). Gametogenesis in this species appears to require a prolonged period
of cool temperature (<10C) for several months (Jones e_t al. , 1976) but after
their successful maturation the range for spawning is shifted upwards to the
range 4-19 C encompassing an optimum of 8-11 C (Hokanson, 1977). Optimum
temperatures for growth of the postembryonic stages appear to be In the
region of 20 C (Hale and Carlson, 1972).
One of the urgent problems today is the setting of limits for thermal
increments to natural waters supporting desirable species of fish in the light
of the knowledge that thermal requirements of a species may change with life
history and with season. Since considerable time and expense are required
to establish thermal requirements of a species throughout its life it was
proposed in the present study that the preferred temperatures as determined
for various life stages and at various times of the year might provide
useful first approxlmationsof a species' changing thermal requirements.
Some fish species display marked seasonal changes in temperature
preference (i.e. changes independent of acclimation temperature) and that
these often occurred during that part of the year when much of gametogenesis
takes place. Sullivan and Fisher (1954) noted that brook trout (Salvelinus
fontinalis) underwent pronounced changes in temperature preference at various
seasons of the year. In March, temperatures selected increased abruptly by
some four degrees from the previous month even though the temperature of the
water in which they were being reared decreased slightly. In the fall of the
year there was a corresponding decrease of the same magnitude under similar
circumstances. Zahn (1963) found pronounced seasonal effects in the plaice
(Pleuronectes platessa) and in the bitterling (Rhodeus sericeus). The
temperatures selected by mature alewives (Alosa pseudoharengus) in Lake
Michigan increased five to six degrees during May and June, at the peak
of their onshore spawning migration (Otto, 1976). Investigators have usually
been aware of seasonal effects and nave usually taken them into account when
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carrying out preferred temperature studies. It was therefore felt that a
fruitful line of enquiry would be an examination of seasonal changes in
temperature selection in yellow perch. At the time the present research
project on temperature preference in yellow perch was drawn up the published
data were not conclusive and there was some evidence that seasonal effects
existed.
The relationship between preferred and acclimation temperature in this
species was reexamined at various times of the year to explore nonthermal
seasonal effects. It was anticipated that any abrupt change in this rela-
tionship might reflect changing thermal requirements and that the hypothesis
that temperature preference may be used as a first approximation of thermal
requirements of a species could be tested.
FACTORS INFLUENCING PREFERRED TEMPERATURE
Thermal History
Temperature preference for most of the species studied is influenced by
thermal history expressed either by thermal acclimation or acclimatization.
The relationship between preferred temperature and acclimation temperature
varies considerably in various species of fish. Typically preferred tempera-
ture rises with acclimation temperature although in some species there is
essentially no change (Brett, 1952; McCauley and Tait, 1971) or even, in
the instance of adult rainbow trout a decrease in preferred temperature
(Garside and Tait, 1958). A scheme of classification for the various types
of curves (Fig. 1) was compiled by Manfred Zahn (1962), from which he drew
Inferences about the general thermal requirements of various species.
Little work has been done on the role of acclimatization temperature
itself in determining temperature preference since investigators have
understandably favored the closely controlled laboratory experiments with
their reproducible results. As mentioned earlier there is evidence that
seasonal factors other than the annual march of temperature may influence
temperature preference in some species.
Pesticides and Anesthetics
Since the process of temperature selection is mediated by the nervous
system it would be expected that substances which effect nervous activity
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could also exert some influence on temperature selection. Ogilvie and
Anderson (1965) showed that temperature preference of salmonids which had
ingested DDT in their food underwent significant changes. Goddard, Lilley
and Tait (1974) noted that lake trout exposed to the anesthetic MS222 did
not display normal temperature selection until several days after treatment.
Diel Rhythms
There is not much evidence for the existence of diel rhythms in selected
temperature although with the recent development of the behavioral thermo-
regulatory devices such effects even though subtle, may now be easily inves-
tigated. Figure 2 shows consecutive mean preferred temperatures for adult
rainbow trout held for two weeks in a behavioral chamber constructed in our
laboratory. No marked diurnal variations in temperature preference is
apparent from thermograph tracing. Of ecological significance is the daily
vertical migration of sockeye salmon described by Brett (1971) in which fish
may feed in the epilimnion at night at 18 C but spend the day in deeper
water at 6 C where low temperatures promote efficient utilization of food.
CO
3
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o
<5
ex
CD
»*
CO
ol
rainbow trout - adults
(Snlmo gairdneri)
lake trout (Salvelinus namaycush)
yellow perch (Perca flavescens)
Figure 1
Acclimation temperature
Examples of preferred temperature - acclimation temperature curves
(Zahn, 1962).
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Figure 2. Preferred temperatures of a rainbow trout (Salmo fiairdneri)
over a ten day period as recorded in a behavioral chamber.
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Age
Ferguson (1958) in his review of the distribution of fish in laboratory
and field temperature gradients pointed out that discrepancies between data
obtained by both methods may be attributed to the differences in age of
laboratory subjects (conveniently small and therefore juvenile) and adult
fish observed in the field. There is a general belief among investigators
that the juvenile of a species are found in the warmer part of the natural
temperature gradient than those frequented by adults. Comparisons from the
literature, however, are made difficult because of variations in the type of
apparatus and techniques followed (Zahn, 1962). McCauley and Read (1973)
showed that in yellow perch the final preferendum of the young when tested
under similar experimental conditions, exceeded that of the adults by several
degrees Celsius. These results are not only in keeping with observations on
the distribution of this species in the field but also with the data of other
investigators who worked separately on either the adult or the juveniles
(McCracken and Starkman, 1948; Ferguson, 1958; Barans and Tubb, 1973).
Other Factors
At the time of writing, there are some investigations being carried out
in other laboratories on the effects of other environmental identities and
factors on temperature preference. Social interaction (prey-predator and
territoriality) health and light, under certain conditions, can effect the
distribution of fish in a natural gradient or disrupt an otherwise perfectly
well-planned laboratory experiment. Needless to say, the art of conducting
experiments in temperature preference includes the suppression or control
of these factors so that the directive effects of temperatures alone can be
described. The reader is referred to Richards, Reynolds and McCauley (1977)
for a detailed discussion of techniques of determining temperature preference
in fish.
6
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SECTION 2
CONCLUSIONS
Seasonal, non-thermal variations in preferred temperatures of yellow
perch were not found using standard laboratory methods. It has been
documented that this species requires temperatures below IOC for several
months during the winter for successful gametogenesis but this change in
thermal requirements was not reflected in the temperature preference of
perch during this season. It is concluded that preferred temperatures,
at least in laboratory acclimated fish cannot be utilized as first
approximations of the thermal requirements of all life stages. It is
suggested that temperature preference of perch in late winter functions
with other environmental stimuli to direct fish during seasonal migrations
to warm nearshore waters.
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SECTION 3
RECOMMENDATIONS
The traditional laboratory approach followed in the present investi-
gation failed to reveal any seasonal effects in preferred temperature in
adult yellow perch independent of acclimation temperature. This was contrary
to expectations since thermal requirements of the species do change during
winter. Exposure of fish to constant temperatures for extended periods of
time (i.e. acclimation) may alter normal responses to temperature gradients
and thus result in experimental artifacts. The ambiguous findings of the
present study re-emphasize the need to examine laboratory techniques and
to continuously validate laboratory results by field observations. Field
acclimatized fish, unlike laboratory acclimated fish, are exposed to many
environmental stimuli which may modify distribution in nature. It is
recommended that, when possible, acclimatized subjects be tested throughout
the year especially when seasonal variations reflecting changing thermal
requirements or annual migrations are sought.
8
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SECTION 4
MATERIALS AND METHODS
STRUCTURE OF GRADIENT TROUGH
Temperature gradients were produced in a trough three meters long, 30
cm deep and 20 cm wide. Preheated water maintained at a constant tempera-
ture by a thermostatically controlled heater entered at one end, was pro-
gressively cooled by a heat exchanger consisting of a refrigerated coil
along one side of the tank. Vertical thermal stratification, a disadvantage
of horizontal gradient troughs, was greatly reduced by currents of compressed
air introduced through air breakers spaced at regular intervals in the water
along the side of the tank. By adjusting the rate of flow of water through
the tank and by altering the temperatures of the fore-chamber along with
that of the coolant, it was possible to produce gradients of about 8 C which
were usually stable throughout the course of an experiment. A serious dis-
advantage of this gradient trough, however, was the inordinate amount of
adjustment required by the operator to produce a gradient and this disturbed
the scheduling of the experiments.
EXPERIMENTAL FISH
Two lots of adult yellow perch were obtained in the fall of two
successive years from Lake St. Clair. Fish were accumulated over several
weeks at the Lake Erie Fisheries Research Station and held in troughs
supplied by running water before being transferred to the Waterloo laboratory.
At Waterloo they were divided randomly into four experimental groups and
reared at 5, 10, 15, and 20 C, respectively, in running water from the
municipal water supply. They were held under a photoregime supplied by
normal laboratory fluorescent lighting controlled by a time switch set each
week in accordance with the natural photoperiod prevailing in Waterloo,
Ontario. Figure 3 shows the natural yearly march of temperature of Lake St.
Clair. The fish were trained to accept commercial trout food which was
supplemented by meat and live minnows. The condition factors of the
experimental fish after a year of holding was similar to that of fish in
the local lakes.
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30-
25-
o
£ 20H
0)
Q.
15H
10H
Maximum temperature
Minimum temperature
Month
Figure 3. Seasonal variations in water temperature of Lake St. Clair,
1974.
PROCEDURES OF TESTING
Acclimation
Temperatures selected in a laboratory gradient by a sample of fish depend,
10
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for the most part, on thermal history expressed as thermal acclimation. For
this reason, it is necessary to standardize the response of fish to a tempera-
ture gradient by holding them at a temperature level sufficiently long enough
to erase the effects of previous thermal history. While information is
available on the rate of gain and loss of thermal acclimation as measured by
thermal resistance, there are very little data for this process with respect
to temperature preference. It has been an article of faith with investigators
that acclimation rates are approximately the same for changes in preferred
temperature. The only published reference is that of Javaid (1972) who
found that young-of-the-year salmonids could acclimate from 10 to 20 C within
two days. Our own ancillary study on behavioral thermoregulation in rainbow
trout illustrates how fish in a temporal temperature gradient increased their
preferred temperature during the course of an experiment (Figure 2). As in
most other studies on temperature preference, we took the conservative
approach in regard to acclimation period, maintaining our samples of fish at
the acclimation temperature for at least one month before determining
preferred temperatures.
Determination of Final Preferendum
The final preferendum is the single value which characterizes the thermal
preference of a species since it represents its eventual stable choice in an
infinite range of temperatures. The most direct way of determining this
value consists in leaving a sample of fish in the gradient apparatus until
the region of temperatures favored by them does not change with .time. Up to
the present, this method has not been followed at least with spatial gradients
because of the physical difficulties of maintaining temperature gradients for
so long a time in an apparatus and holding fish for long periods in the
confined space of a tank. In practice, workers have systematically determined
preferred temperatures for fish acclimated to a series of acclimation tempera-
tures, plotted these values against acclimation temperature, and by interpola-
tion, graphically estimated that point for which preferred and acclimation
temperatures coincide. When this method is adopted, care must be taken that
the fish are kept in a thermal gradient for periods less than a day because
of a gradual change in acclimation level resulting from exposure to higher
or lower temperatures.
11
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SECTION 5
RESULTS
BEHAVIOR OF FISH IN THERMAL GRADIENTS
In the routine determination of temperature preference in the labora-
tory, attempts are made to minimize or at least take into consideration
gradients of other environmental identities; i.e. light, gravity, water
velocity, etc. Time was allowed the experimental subjects to habituate to
the environment of the test tank and for this reason the perch were intro-
duced 24 hours into the gradient tank maintained initially at the acclimation
temperature. In many instances the fish tended to congregate at the two ends
of the trough, perhaps responding to a slight light gradient in which the
ends of the tank were the darkest regions of the tank or displaying behavior
known as a "fright huddle". It was found that the "end effect" bias could
be over-ridden by directing a small, bright light on both end walls of the
tank. By this means, distributions in the tank were largely influenced by
temperature rather than by other factors. In spite of patience and resource-
fulness of the operators in creating conditions in the gradient trough
conducive to behavioral thermoregulation, the experimental subjects often
did not respond to the thermal gradient. Much of the data of the present
study obtained could not be used since it was apparent that the fish were
not responding to temperature gradients. This imprecision in selecting
temperature in this species had also been noted by other investigators and
is reflected in the wide standard deviations quoted in their data (Barans
and Tubb, 1973).
There was no noticeable trend to increased preferred temperature with
sojourns of up to 24 hours in the gradient trough. The time for the perch to
become habituated to the apparatus was considered to be three hours, and
distributions of fish in the tank were not used in calculation of temperature
preference before this period had elapsed.
PREFERRED TEMPERATURE
Modal temperatures of distributions of fish held at the four acclima-
tion levels and tested at three seasons of the year for two years are
12
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summarized in Table 1. The data for winter and early spring are plotted in
Figure 4 along with those of McCracken and Starkman (1948) who also studied
the effect of acclimation temperature on preferred temperature at this time
of the year. In view of possible non-thermal seasonal influences our late
winter data may be compared with those of the previous study. The curve
connecting the points of the 1948 study intersects the 45 degree diagonal
in the vicinity of 20 C indicating a final preferendum of this value. Up
to the 20 C acclimation level the data of the present study are in good
agreement with those of the earlier work although they indicate a higher
final preferendum.
Table 1. MEAN PREFERRED TEMPERATURES OF SAMPLES OF YELLOW PERCH HELD AT
FOUR CONSTANT ACCLIMATION TEMPERATURES THROUGHOUT THE YEAR (C,
standard deviations are shown in brackets)
Year Season AccHimation temperature, (C) Final
preferendum,
5 10 15 20 (C)
1974
1975
Winter
Spring
Summer
Winter
Spring
Summer
13.0
12.3
13.8
12.0
13.1
13.5
(1
(2
(1
(1
(2
(1
.9)
.5)
.6)
.5)
1)
.7)
15.5
18.8
13.5
14.2
18.1
14.1
(2.3)
(2.0)
(1.8)
(2.1)
(1.9)
(1.6)
18.4
20.2
17.6
18.0
21.0
18.1
(2.2)
(1.8)
(2.6)
(1.9)
(1.6)
(2.3)
24.2
20.2
16.1
23.2
19.5
17.0
(3.0)
(2.4)
(2.1)
(2.4)
(2.5)
(2.2)
25
21
17
30
21.1
18
CONDITION OF GONADS OF THE EXPERIMENTAL FISH
At the end of the series of preferred temperature determinations, experi-
mental fish were sacrificed, the gonads weighed and histological sections
prepared. The groups held at 5 and 10 C in 1974 showed evidence of sexual
maturation when three females at 5 C and two at 10 C spawned in May. The
phenomenon was noted the following year when two females spawned at the same
two temperatures and one at 15 C. No tests were made, however, to determine
egg fertility. Logarithms of ovary weight were plotted against logarithms of
body weight. The scattering of points indicated that there were no real
differences among gonadosomatic indices of fish reared at various temperatures.
13
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Since the ovaries of the test fish are likely a heterogenous mixture of eggs
in various stages of resorption and new development, no conclusions may be
drawn. Interpretation of the apparent differences of the ovaries awaits
the judgment of an experienced reproductive fish physiologist.*
o>
-a
2?
a.
60
25-
20-
15 -
10-
5 -
,
BJO "/
CO /"
o / o
! V
I /
° /
y^ late winter
/ late spring
/ D late spring
S D late winter
/ O McCracken
/ Starkman
0_
0 5 10 15 20
/ °
/
0
1974
1974
1975
1975
and
1948
I
25 3(
Acclimation Temperature C
Figure 4. The relation between preferred and acclimation temperature in
late winter and early spring.
* The slides are at present in possession of the writer and are available
for inspection by any interested investigator.
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SECTION 6
DISCUSSION
REVIEW OF PREFERRED TEMPERATURE STUDIES ON YELLOW PERCH
In spite of a variety of techniques and apparatus there appears to be
good general agreement among the published preferred temperature data of
perch. When the data of various authors are plotted on the same graph the
relation between preferred temperature and thermal history expressed as
acclimation or acclimatization temperature may be described by an envelope
of positive slope enclosing the experimentally determined points (Fig. 5).
The similarity between curves for both acclimated and acclimatized fish is
evidence that there is no strong influence for seasonal effects apart from
water temperature. The data of McCauley and Read (1973) seem to be anomalous
since they lie several degrees below the envelope. No reason is apparent
for their depressed values. Since they were tested in the early fall in a
vertical gradient it is suggested that they may have displayed a type of
"predictive thermoregulation" as defined by Neill (1976). Neill proposed
that some fish species may anticipate seasonal changes in spatial thermal
gradients and move towards the future region of preferred temperature. Since
yellow perch in the lake move to deep water with the passing of summer, fish
in the vertical gradient of McCauley and Read may have responded seasonally
to a hydrostatic gradient and swam lower in the tank. Some support for this
speculation is provided by field data depicting graphically the depth and
thermal distribution of adult perch in Lake Opeongo, Algonquin Park, from
June to September (Fig. 6). During July and August the mean of the thermal
distributions lies between 20 and 21 C but drops precipitously about the
beginning of September - a time marking the beginning of spermatogenesis
and secondary growth of ova (Turner, 1919).
Figure 7 is an interpretation of La Rue Well's 1972 study of the seasonal
distribution of fish species in Lake Michigan as determined by bottom trawls.
He recorded numbers of fish captured in relation to depth and temperature at
increasing distances from shore. Adults were usually found in the warm,
inshore waters displaying a final preferendura slightly under 20C in July.
These field observations are in essential agreement with laboratory data
15
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although any non-thermal seasonal effects are obscured because of decreasing
temperature gradients when water bodies cool from August onwards.
30 H
25 H
o 20
£
a.
15 -
10 -
5 "
2
/
/
* D
"1 /
/
*
*
V summer 1975
T winter 1975
V spring 1975
winter 1974
O spring 1974
summer 1974
D Barans and Tubb 1973
A Otto unpubl. 1973
* McCauley and Read 1973
5
10
15
l
20
25
i
30
Acclimation 1 Temperature ( C)
Acclimatization J
Figure 5. Summary of the preferred temperature relations of adult yellow
perch.
16
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MAY
JUNE
JULY
AUGUST
SEPT.
14 -
Gill Nets
1938
Horizontal scale
Number of fish captured
MAY JUNE
Minnow Traps
1938
~ 200. 9 .290
Horizontal scale
Number of fish captured
Figure 6. Observed vertical distributions of yellow perch in relation to
season, depth and temperature in Lake Opeongo. Frequency poly-
gons depict numbers of fish captured at various depths. Modified
from Ferguson (1958).
17
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THE ROLE OF TEMPERATURE PREFERENCE IN THE LIFE OF THE YELLOW PERCH
One of the shortcomings of laboratory investigations on temperature
preference of fish is the unnatural compression of the temperature gradients
in experimental tanks. The data from which preferred temperatures of yellow
perch were calculated by Ferguson (1958) and McCracken (1948), for instance,
were obtained using a vertical tank having a depth of about one metre. The
field information with which the laboratory data was compared, on the other
hand, was obtained by observing fish in natural thermal gradients of greater
dimensions. In some field studies on the movements of fish in and around
thermal effluents, the "natural" thermal gradients are compressed and a
phenomenon resembling behavioral thermoregulation has been recorded (Neill
£t aK 1972; Spigarelli e_t al_. 1972). It seems difficult to conceive,
however, that fish spread out some fifty metres in the vertical temperature
gradient existing in a lake in the temperate region during the period of
summer stratification are continually moving up and down the water column
many times a day as they do in a laboratory gradient tank, in an effort to
regulate their body temperature. It is clear that, as usual, laboratory
findings must be applied with reservation to the field situation since other
gradients (light, gravity, water velocity, food availability, etc.) and
other non thermal inputs affecting temperature selection are acting
simultaneously.
From perusal of the literature and from contemplation of the results of
the present study the significance of temperature preference in the life of
the yellow perch is considered to be summarized below. The selected tempera-
ture response in perch serves (along with other gradients) to direct the
movements of perch to thermally favourable habitats. During the winter
season the environment of the species is cooler than ten degrees C for a
period long enough to ensure development of the gonads (Jones ejt al. 1976).
The results of the present investigation along with those of Barans and Tubb
(1973)indicate that adult perch in mid-winter would choose temperatures
above 12 degrees C if these were present in the environment. Since tempera-
tures of this range do not occur naturally until early spring, the opportunity
to move to these warmer waters is not normally present. The proximity of
warm effluents, of course, could lure fish into unfavorably high temperatures.
19
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Temperature preference during the growing season likely serves to direct
fish into waters which have the optimum temperature range. The upward
ascension of fish up the temperature gradient early in spring ensures that
lake dwelling populations move shorewards to warmer waters where spawning
and incubation of embryos take place. Our work and that of others indicate
that, unlike some species (see introduction) there are no obvious seasonal
effects apart from temperature as a function of season on the temperature
selection in yellow perch. Barans and Tubb (1973), argue however, that in the
perch of their study which were collected seasonally from Lake Erie and
kept in a long, horizontal temperature gradient for periods up to a week,
there were differences in the final preferendum over the year. Even granting
this, the apparent final preferendum for adult perch sampled in midwinter
was 15 C, some 10 degrees C above the optimum for gametogenesis.
There is a hint of seasonal differences in the data of the present
investigation although there is a suggestion (see Fig. 5) that perch in
lakes move to cooler waters in September, these may be ascribed to artifacts
in the particular experimental method followed herein. Exposure of fish to
constant temperatures may have interfered with the normal process of matura-
tion and consequently affected the response to temperature gradients,
especially in the summer. Samples held at 15 and 20 C, for instance, did
not spawn indicating that they were egg bound and were resorbing eggs. This
unusual physiological state may have affected their responses to temperature
gradients and given rise to artificial temperature preferences. Further
research is necessary to separate the effects of the experimental method on
preferred temperature value from real seasonal influences. The technique
of maintaining fishes at constant temperatures for an unnaturally long duration
(i.e. spanning several seasons) should be checked by determining the final
preferendum of acclimatized fish. In principle this would involve allowing
fish to remain in a laboratory temperature gradient until the selected
temperature becomes stable. In practice this has been difficult because of
problems of maintaining stable thermal gradients in tanks and holding fish
in confined space.
Recent development of temporal gradients in the last five years have
allowed this approach to be taken at least with some species (Neill et al.
1972; Reynolds, 1973; Beitinger, 1974). Neill e_t a_l found that juvenile
20
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yellow perch regulated temperatures in the shuttlebox so that they fell in
the preferred temperature range although they pointed out that data for perch
were more variable than those for the other five species studied.
The results of the present study emphasize the need for the re-examina-
tion of techniques used in the determination of temperature preference in
fish. In response to this need,the writer and four other workers in the
field held a symposium (Richards, Reynolds, and McCauley, 1977) in which
temperature preference methodology was reviewed and evaluated. The data
of the yellow perch study also underline the necessity of reviewing the
ecological significance of temperature preferenda of species of fish in
general.
21
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REFERENCES
Barans, C.A.,and R.W. Tubb. 1973. Temperatures Selected Seasonally by Four
Fishes from Western Lake Erie. J. Fish. Res. Bd. Canada 30;1697-1703.
Beitinger, T.L. 1974. Thermoregulatory Behavior and Diel Activity Patterns
in the Bluegill, Lepomis macrochirus, Following Thermal Shock. U.S.
Fish. Wildl. Serv., Fish Bull. 72:1087-1093.
Brett, J.R. 1952. Temperature Tolerance in Young Pacific Salmon Genus
Oncorhynchus. J. Fish. Res. Bd. Canada £:265-323.
Brett, J.R. 1971. Energetic Responses of Salmon to Temperature. A Study
of Some Thermal Relations in the Physiology and Freshwater Ecology of
Sockeye Salmon (Oncorhynchus nerka). Amer. Zool. 11:99-113.
Ferguson, R.G. 1958. Preferred Temperatures of Fish and Their Midsummer
Distribution in Temperate Lakes and Streams. J. Fish. Res. Bd. Canada
L5:607-624.
Fry, F.E.J. 1947. Effects of the Environment on Animal Activity. Univ.
Toronto Studies, Biol. Ser. No. 55. Publ. Ont. Fish. Res. Lab. No. 68,
62 pp.
Fry, F.E.J. 1958. Experimental Study of the Behavior of Fish. Proc. Inco-
Pacific Council 111:37-42.
Garside, E.T. 1966. Effects of Oxygen in Relation to Temperature on
Development of Embryo Brook Trout and Rainbow Trout. J. Fish. Res. Bd.
Canada 23:1121-1134.
Garside, E.T.,and J.S. Tait. 1958. Preferred Temperature of Rainbow Trout
(Salmo gairdneri Richardson) and Its Unusual Relationship to Acclimation.
Can. J. Zool. 36:563-567.
Goddard, G.I., J.W. Lilley, and J.S. Tait. 1974. Effects of MS222 Anaesthe-
tization on Temperature Selection in Lake Trout. J. Fish. Res. Bd.
Canada 31:100-103.
Hale, J.G., and A.R. Carlson. 1972. Culture of the Yellow Perch in the
Laboratory. Prog. Fish-Cult. 34:195-198.
Hokanson, K.E.F. 1977. Temperature Requirements of Some Percids and
Adaptations to the Seasonal Temperature Cycle. J. Fish. Res. Bd. Canada
(In Press).
Hokanson, K.E.F., C.F. Kleiner, and T.W. Thorslund. 1977. Effects of
Constant Temperatures and Diel Temperature Fluctuations on Specific
Growth and Mortality Rates and Yield of Juvenile Rainbow Trout, Salmo
gairdneri. J. Fish. Res. Bd. Canada 34:639-648.
22
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Javaid, M.Y., and J.M. Anderson. 1967. Thermal Acclimation and Temperature
Selection in Atlantic Salmon, Salmo salar, and Rainbow Trout, Salmo
gairdneri. J. Fish. Res. Bd. Canada 24:1507-1514.
Javaid, M.Y. 1972. The Course of Selected Temperature During Thermal
Acclimation of Some Salmonids. Nucleus £:103-105.
Jones, B.R., K.E.F. Hokanson, and J.H. McCormick. 1976. Temperature
Requirements for Maturation and Spawning of Yellow Perch, Perca
flavescens (Mitchill). Unpublished Manuscript, U.S. Environmental
Protection Agency, Duluth, Minnesota.
McCauley, R.W., and W.L. Pond. 1971. Temperature Selection of Rainbow Trout
(Salmo gairdneri) Fingerlings in Vertical and Horizontal Gradients.
J. Fish. Res. Bd. Canada 28:1801-1804.
McCauley, R.W., and L.A.A. Read. 1973. Temperature Selection by Juvenile
and Adult Yellow Perch (Perca flavescens) Acclimated to 24C. J. Fish.
Res. Bd. Canada 30:1253-1255.
McCauley, R.W., and J.S.Tait. 1971. Preferred Temperatures of Yearling
Lake Trout Salvelinus namaycush. J. Fish. Res. Bd. Canada 27:1729-1733.
McCracken, F.D., and S.H. Starkman. 1948. Preliminary Observations on the
Preferred Temperatures of the Perch. Unpublished Manuscript, Ont. Fish.
Res. Lab. Libr., Toronto, Ontario. 4 pp.
Neill, W.H. 1976. Mechanisms of Behavioral Thermoregulation in Fishes.
Texas Agricultural Station, College Station, Texas. Contri. No. TA-
12354. 5 p.
Neill, W.H., J.J. Magnuson, and G.C. Chipman. 1972. Behavioral Thermoregu-
lation by Fishes: A New Experimental Approach. Science 176:1433-1445.
Ogilvie, D.M., and J.M. Anderson. 1965. Effect of DDT on Temperature
Selection by Young Atlantic Salmon, Salmo salar. J. Fish. Res. Bd.
Canada 22:503-512.
Otto, R.G., M.A. Kitchel, and J.O. Rice. 1976. Lethal and Preferred Tempera-
tures of the Alewife (Alosa pseudoharengus) in Lake Michigan. Trans.
Am. Fish. Soc. 105:96-106.
Reynolds, W.W. 1973. Orientation Responses of Laboratory-Reared Larval and
Juvenile Gulf Grunion (Leuresthes sardina) to Artificial Gradients.
Ph.D. thesis, Univ. Arizona, Tucson. 96 pp.
Richards, F.P., W.W. Reynolds, and R.W. McCauley. 1977. Temperature Pre-
ference Studies in Environmental Impact Assessments: An Overview with
Procedural Recommendations. J. Fish. Res. Bd. Canada 34:728-761.
23
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Spigarelli, S.A. 1972. Fish Tagging Studies at the Point Beach Nuclear
Power Plant. II. The Movements of Tagged Fish. In Radiological and
Environmental Research Division Annual Report. U.S. Atomic Energy
Commission Report ANL-7960, Part III. Argonne, Illinois, p. 83-88.
Sullivan, C.M., and K.C. Fisher. 1954. Seasonal Fluctuations in the
Selected Temperatures in Speckled Trout Salvelinus fontinalis (Mitchill)
Biol. Bull. 107:278-288.
Turner, C.L. 1919. The Seasonal Cycle in the Spermary of the Perch.
J. Morph. 32:681-711.
Wells, L. 1968. Seasonal Depth Distribution of Fish in Southeastern Lake
Michigan. Fishery Bulletin 6_2(1):1-15.
Zahn, M. 1962. Die Vorzugstemperaturen zweier Cypriniden und eines
Cyprinodonten und die Adaptationstypen der Vorzugstemperatur bei
Fischen. Zool. Beitr. _7:15-25.
Zahn, M. 1963. Seasonal Variations in the Preferred Temperatures of
Plaice and Bitterlings. Verhandlungen der Deutschen Zoologischen
Gesellschaft in Munchen. 17 p. (Transl. from German by Fish. Mar.
Serv. of Canada. Transl. Ser. No. 3501, 1975).
24
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TECHNICAL REPORT DATA
(Please read Instructions on the reverse before completing)
1. REPORT NO.
EPA-600/3-77-088
2.
3. RECIPIENT'S ACCESSION-NO.
4. TITLE AND SUBTITLE
Seasonal Effects on Temperature Preference in
Yellow Perch, Perca Flavescens
5. REPORT DATE
August 1977 issuing date
6. PERFORMING ORGANIZATION CODE
7. AUTHOR(S)
Robert W. McCauley
8. PERFORMING ORGANIZATION REPORT NO.
9. PERFORMING ORGANIZATION NAME AND ADDRESS
Wilfrid Laurier University
Department of Biology
Waterloo, Ontario, N2L 3C5
10. PROGRAM ELEMENT NO.
1BA608
11. CONTRACT/GRANT NO.
Grant No. R802467
12. SPONSORING AGENCY NAME AND ADDRESS
Environmental Research Laboratory - Dul., MN
Office of Research and Development
U.S. Environmental Protection Agency
Duluth, Minnesota 5580U
13. TYPE OF REPORT AND PERIOD COVERED
Final Report; May 1973-May
14. SPONSORING AGENCY CODE
975
EPA/600/03
15. SUPPLEMENTARY NOTES
16. ABSTRACT
Seasonal variations in temperature preferences of adult yellow perch (Perca
flavescens) were sought by acclimating fish captured in the fall to 5, 10, 15, and
20C in the laboratory and determining their preferred temperatures in a horizontal
temperature gradient trough. Temperatures selected in winter for fish acclimated to
5C ranged from 12 to 14C, considerably above the temperature region (6C and below)
previously established for optimum gametogenesis. Final preferenda as determined from
preferred temperature-acclimation temperature curves were 24, 20, and 17C during
winter, spring and summer respectively. Those in late summer were below published
values (23.5C) for this species. It is concluded that there are no demonstrable
effects, at least in winter and early spring which might reflect changing physiological
needs and that temperature gradients in nature serve rather to attract perch to warm
temperatures suitable for spawning in spring and conducive to growth in summer.
17.
KEY WORDS AND DOCUMENT ANALYSIS
DESCRIPTORS
b.lDENTIFIERS/OPEN ENDED TERMS C. COSATI Field/Group
Temperature gradients
Spatial distribution
Freshwater Biology
Animal behavior
Animal migrations
Water temperature
Preferred temperature
Final preferendum
Seasonal
Perch adults
06/F
18. DISTRIBUTION STATEMENT
RELEASE TO PUBLIC
19. SECURITY CLASS (This Report)
UNCLASSIFIED
21. NO. OF PAGES
33u
20. SECURITY CLASS (This page)
UNCLASSIFIED
22. PRICE
EPA Form 2220-1 (9-73)
25
r U. S. GOVERNMENT PRINTING OFFICE 1977-757-056/6505 Region No. 5-11
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