TEMPERATURE AND
           AQUATIC  LIFE
           LABORATORY
          INVESTIGATIONS
              SERIES
TECHNICAL ADVISORY AND INVESTIGATIONS BRANCH

FEDERAL WATER POLLUTION CONTROL ADMINISTRATION

UNITED STATES DEPARTMENT OF THE INTERIOR

5555 Ridge Avenue

Cincinnati, Ohio  45213

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                       FOREWORD








        The Laboratory Investigations series was initiated




by the Technical Advisory and Investigations Branch in




1963.  The series was planned to describe laboratory




methods and techniques and to disseminate information that




may be of interest and use to other activities of FWPCA.



        The current addition to the series is a literature



review of the effects of thermal pollution on the aquatic




ecosystem.  Thermal pollution is a rapidly increasing




problem and this review will aid. in evaluating existing




problems and in the prevention of future problems.








                                   December 196?

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         TEMPERATURE AND AQUATIC LIFE
     Laboratory Investigations - Number
 Technical Advisory and Investigations Branch
          Technical Services Program
Federal Water Pollution Control Administration
   United States Department of the Interior
               Cincinnati, Ohio

                 December  1967

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CHEMICAL REACTIONS	JAMES L. HOLD/WAY
                                                 Chemist

BACTERIA	LOUIS A. RESI
                                                  Microbiologist

FRESHWATER FISHES	NELSON A. THOMAS
                                                  Aquatic Biologist

MARINE, ESTUARINE AND ANADROMOUS FISHES  .  .  . LOYS P. PARRISH
                                                  Aquatic Biologist

AQUATIC PLANTS AND BENTHOS 	 R. KEITH STEWART
                                                  Aquatic Biologist

EDITOR 	 KENNETH M. MACKENTHUN
                                                  Supervisory Aquatic
                                                  Biologist

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                              SUBJECT INDEX
CHEMICAL REACTIONS	    1



     Ammonia .....  	   12




     Biochemical oxygen demand  	    8




     Carbon dioxide	   10



     Dissolved Minerals	   ik




     Henry's law	    1




     Hydrogen	    8




     Hydrogen sulfide	    9




     Methane 	    7




     Nitrogen.  .	    7



     Oxygen	    5




     Sulfur dioxide	   13




     Summary	   l6




     References	   17



BACTERIA	   19



     Bacterial indicators	   26




     Bacterial survival	   25




     Microbiotic cycles	   21




     Self-purification  	   23



     Slime gro-wrths	«	   26




     Summary	   27




     References	   29

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                                                            Page

FRESHWATER FISHES 	    30

     Acclimation	    32

     .Benefits of thermal pollution	    ^6

     Effects on toxicities	    ^5

     General temperature effects	    30

     Maximum temperatures 	    3^

     Preferred temperatures 	    38

     Sudden temperature changes 	    31

     Summary	    k"f

     References	    i»9

MARINE ESTUARINE AND ANADROMOUS FISHES	    52

     Acclimation	    56

     Anadroraous fishes
        Eggs	    6?
        Young	    TO
        Adults	    T3

     Development.	    5k

     Distribution 	    55

     Ecology	    55

     General temperature effects	    52

     Marine and estuarine fishes
        Eggs	    57
        Young	    58
        Adults	    63

     Metabolism	    53

     Physiology	    53

     Reproduction 	    5^
                                   ii

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                                                            Page



     Tolerance	    57




     Summary	    75




     References	    77




AQUATIC FLAM'S AND BENTHOS	    83




     Bottom organisms	    87




     Fresh-water algae and other aquatic plants 	    85




     General	    83




     Summary	    93



     References	    9^




SELECTED BIBLIOGRAPHY 	    97




INTRODUCTION	    iv




SUMMARY	     v
                                   iii

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                                INTRODUCTION








       Temperature, a catalyst, a depressant,  an activator, a restrictor,




a stimulator, a controller, a killer, is one of the most important and




most influential water quality characteristics to life in water.  Temper-




ature determines those species that may be present; it activates the hatching




of young, regulates their activity and stimulates or suppresses their growth




and development; it attracts, and kills when the water becomes too hot or




becomes chilled too suddenly.  Colder water generally suppresses develop-




ment; warmer water generally accelerates activity and may be a primary cause




of aquatic plant nuisances when other environmental factors are suitable.




       Because of the importance of this single environmental facet to




aquatic ecology, this report was developed to consider some of the features




of temperature and its interrelationships.  It is divided into five segments;



these are:




         I.  Chemical Reactions




        II.  Bacteria




       III.  Freshwater Fishes




        IV.  Marine, Estuarine and Anadromous Fishes




         V.  Aquatic Plants and Benthos




An extensive temperature bibliography is appended.
                                     iv

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                                 SUMMARY








1.  Chemical reaction rates vary with temperature, generally increasing



    as the temperature is increased.




2.  The solubility of gases in water varies with temperature.  Dissolved




    oxygen is decreased by the decay or decomposition of dissolved organic




    substances; the decay rate increases as the temperature of the water




    increases reaching a maximum at about 30 C (86 F).




3.  The temperature of stream water, even during the summer, is below the




    optimum for pollution-associated bacteria.  Increasing the water tem-




    perature increases the bacterial multiplication rate when the environ-




    ment is favorable, and the food supply is abundant.  Increasing the




    water temperature within the growth range of the bacteria causes a more



    rapid die-off when the food supply is limiting.




k.  Warm water fish can survive temporarily in waters heated artificially




    "to 33-9 C (93 F); some fish populations, such as roach, perch,  and



    carp, are reduced at these high temperatures.  In cold weather,  stream




    temperatures should be substantially below 33-9 C (93 F) "to prevent




    mortalities when fish move through excessive temperature gradients.




    Cold water non-anadromous fish populations such as trout should not be




    subjected to temperatures exceeding 1^.5 C (58 F).  In cold weather



    stream temperatures should be below lU.5 C (58 F) to prevent mortalities




    of cold water fishes.

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 5.  Sudden changes in temperature can be more harmful to some species of




     fish than continued exposure to a higher temperature.




 6.  Fish can adapt to higher temperatures faster than to lower temperatures.




 7.  The maximum temperature for a given species of fish varies with the




     fish's rate of heating, size, and physiological condition.




 8.  Fish may starve at elevated temperatures because of their inability to



     capture food.




 9.  Fish seek out a preferred temperature at which they can best survive,




     which is several degrees below their lethal temperature.




10.  The toxic effects to fish of certain material increase with temperature.




11.  Temperature changes are most important to fish in enclosed areas in the




     marine environment such as estuaries and bays as opposed to open areas




     although tolerance to temperature fluctuations is greater in fresh-water




     and estuarine forms than in open water marine species.



12.  There are restricted ranges of temperature within which fish can reproduce




     successfully; larval development especially requires narrow ranges of




     temperature.  A fish population may exist in a heated area only by con-



     tinued immigration from the outside.  Fish may be absent from such areas




     during warm summer months and present in cold winter months.




13.  Increased temperatures may block the migrations of anadromous fish.




lU.  Cold is as important to fish populations as heat because of the inability



     of fish to acclimate quickly to rapid decreases in temperature.  Thus,



     in some areas fish populations may be limited by decreases as well as

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     increases in temperature.   The growth rate of fishes is reduced in waters




     colder than the optimum temperature range for the species.




15.  When water temperatures increase, the predominate algal species change



     from diatom to green algae and. finally at high -temperatures to blue-green




     algae.



16.  The number and distribution of bottom organisms decrease as water temper-




     atures increase above 90 F, which is close to the tolerance limit for a




     "balanced" population.  The adult stage of many species is  able to tole-




     rate higher temperatures than the eggs or young.



17.  A benefit of heated effluents is the defouling of intake pipes accomplished




     by reversing the flow of water through the pipes.




18.  Certain benefits, including open water winter fishing in otherwise ice




     covered areas, and a cold water fisheries downstream from deep reservoirs,




     can be derived from artificially induced temperature changes.  The




     benefits of fish being attracted to heated water in the winter months




     may be negligible compared to fish mortalities that may result when the




     fish return to the cooler water; lethal temperatures may result from




     heated discharges in the summer months.
                                     vil

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                          I.  CHEMICAL REACTIONS








Introduction




       All the impurities contained in a water result from intimate




contact of the water with such impurities during which a portion are




dissolved or suspended.  The process of solution is a chemical re-




action which proceeds as long as the water is in contact, with a solu-




ble substance or until equilibrium is reached (Camp, 19^3)•   In general



the solubility of solids and liquids may be considered a function of




temperature, unless extreme pressure conditions are involved.




       The solubility of nonreactive gases (gases that do not react




with water to an appreciable extent) at equilibrium with the atmos-




phere is proportional to the partial pressure of the gas in the atmos-




phere and follows Henry's law, Cs=KsP, where Cs is the saturation




concentration of the gas in the water, P is the partial pressure of




the gas phase and Ks is the proportionality constant called the co-




efficient of absorption.  Water is saturated with a gas when the pro-




portionality implied in Henry's law is fully established.  Rising




temperatures decrease the saturation value as do the salts of hard and




brackish waters (Fair and Geyer, 195*0.  The solubility of reactive




gases in water is modified because they ionize in and/or react with the




water.

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       The establishment of equilibrium in a given chemical reaction

implies that the reaction is reversible and that a point has been

reached where a balance between the reactants exists.  Mathematically,

this relation is expressed by the mass action law
where capital letters refer to types of molecules or ions taking part

in the reaction and lower case letters to the number of them.  On the

basis of thermodynamic principles the relation

       (C)P(D)q
       (A)m(B)n=k

is universally true, the parentheses indicating activities of the en-

closed substances.  The equilibrium constant k has a characteristic

value for each reaction that is dependent only on temperature (Fair

and Geyer, 1951!-).

       The effects of temperature on equilibria are given by the Van't
Hoff equation  d(log^0_ _
                  dT    ~ RT2

where K is the equilibrium constant, T is the Kelvin temperature, R is

the gas constant, AH? is the enthalpy change per gram-molecular weight

for the reaction from left to right.  Integration of the above equation

between the limits Tj_ and T2 gives
                    RT2

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for constant AH? and conversion to common logarithms
              " 2 . 303IT T2 " TI   1<-.576V TLT2


(Glasstone and Lewis, 1960).
       The rates of most chemical reactions increase as the tempera-

ture is raised.  A frequently used very approximate rule, enunciated

by Van't Hoff, is that the rate doubles for each rise in temperature

of 10 °C (18°F).

       Mathematically, the change in specific rate constant with tem-

perature for any simple chemical reaction is given by the Arrhenius


          d(logek)    E
             at
where k is the specific reaction rate constant, T is the Kelvin tem-

perature, R is the gas constant (1.99 cal/degree C), E is a constant

characteristic of the reaction and termed the activation energy.

Integration of the above equation between the limits T^ and T2 Sives
                    RT2 ~

for constant E and conversion to common logarthms


          ko     -E  ( 1  1 % _
       L0\^ 2.303R(kO?2"^I;~

(Glasstone and Lewis,  1960).

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       Two other methods for expressing the temperature dependence of


a reaction rate are often encountered.  They are:



       k£_   T2-T1

       ki  ®



whence


       9 = ^2 for T2-Ti  and  Q10 = ^ for To
           Kl                      Kl



here T2 and T]_ are measured in degrees centigrade (Fair and Geyer,


195^).
Dissolved Gases




1.  Non-Reactive Gases


       The gases that do not react to an appreciable extent vrith water


but which occur in sufficient quantities to be determined by chemical


analyses are oxygen, nitrogen, hydrogen and methane.

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                             Oxygen



       Since all living organisms are dependent on oxygen in one



form or another to maintain the metabolic processes that produce



energy for growth and reproduction, it is of great significance in



the aquatic environment.  The solubility of atmospheric oxygen (see



Figure l) in fresh waters ranges from lk.6 mg/1 at 0°C (32°F) to



6.6 mg/1 at 1
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a\
                    FIGURE  I     SOLUBILITY OF OXYGEN IN WATER
                 EXPOSED TO WATER-SATURATED AIR AT 760 mm Hg.
     o
8  10  12  14   16  18  20  22  24
              TEMPERATURE  °C
26  28  3O 32  34  36

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is also produced by chlorophyll-bearing algae and submerged aquatic



plants through photosynthesis.



       In natural waters deficient in dissolved oxygen, accelerated



atmospheric aeration (reaeration) is evident.  The rate of reaeration



in each unit of time is proportional to the remaining degree of un-



saturation of dissolved oxygen in the waters (Streeter, 1958).  Temp-



erature influences the rate of solution and affects the rate of dif-



fusion of oxygen.  As temperature increases the saturation capacity



declines and the rate of diffusion increases (Velz and Gannon, 1960).






                            Nitrogen



       The solubility of atmospheric nitrogen in water is about one-



half that of oxygen.  While the principal source of nitrogen is the



air, denitrifying bacteria will release nitrogen to water (Allee



et al., 19^9).  In a lake the waters of the hypolimnion become and



remain supersaturated with nitrogen as they get warmer; rapid warming



may cause nitrogen to escape as bubbles (Nordell, 1961).








                             Methane



       Methane is not a permanent constituent of the earth's atmos-



phere (Camp, 1963).  The primary source of methane in natural waters



is anaerobic decomposition.  Solubility of methane varies from 39•6



mg/1 at 0°C (32°F) to 15.9 mg/1 at kO°C (l(A°P) and 760 mm of mercury

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                                  8
(Nordell, 19^l).  Some ground waters may contain sufficient methane to



constitute a fire and explosion hazard.  Methane may persist even after



aeration.



                              Hydrogen



       The solubility of hydrogen varies from 1.93 mg/1 at 0°C (32°F)



to 1.48 mg/1 at to°C (104°F) in pure water in contact with the pure



gas at 760 mm of mercury (Nordell, 1961).  Hydrogen comprises less



than 0.001$ of the earth's atmosphere (Camp, 1963).  It is produced



in the water primarily from anaerobic decomposition of organic matter.








Pi ss olvedOrganic Sub stance s



       Organic substances in surface waters decay or decompose chiefly



by bacterial action and exert a demand on the dissolved oxygen of such



waters.  This biochemical reaction is similar to an unimolecular chemi-



cal reaction, that is, the rate is approximately proportional to the



remaining concentration of unoxidized organic matter.  Thus the bio-



chemical oxygen demand of a surface water is a measure of the concen-



tration of decomposable organic matter (Camp, 1963).  The reaction



rate ordinarily expressed as k rises as the temperature of the water



increases reaching a maximum at about 30°C (86°F), (Hoak, 1961).  This



increased rate results in a greater demand on the'dissolved oxygen in



the surface water.  Theriault (1927) has shown that at 20°C (68°F) the



biological oxidizability of polluted water increases by about 2$ for



each degree centigrade increase.

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2.  Reactive Gases

       Gases that react with water are hydrogen  sulfide, carbon

dioxide, sulfur dioxide and ammonia.  These either ionize in water

or react with the water to produce ions.


                        Hydrogen Sulfide

       The principal sources of hydrogen sulfide in natural waters

are anaerobic decomposition of organic matter and the discharge of

industrial wastes from oil refineries, leather tanneries, chemical

plants and paper mills (Camp, 1963); the solubility of hydrogen sul-

•fide ranges from 7,070 mg/1 at 0°C (32°F) to 2, 360 mg/1 at ifO°C

(10^°F) and 760 mm of mercury (Nordell, 1961) .  It ionizes in water

in two steps --
       2.  HS"^— > H++S=

An equilibrium relation is established for each step as follows


       i.  [H+UHS-]
       2.   [H+][S=]
              [HS-]

                                     <-r
vith equilibrium constants of l.lxlO~' at 25°C (T7°F) for step 1 and

1x10     at 25°C (77°F) for step 2. Equilibrium constants vary with
                                                                 o
temperature and corresponding values at l8°C (64.4°F) are 5.7x10"

and 1.2x10"    (Lange, 1961).  As indicated by the comparative magni-

tude of the equilibrium constants for hydrogen sulfide at 25°C (77°F)

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                                10





the equilibrium in step 1 lies to the side of the product, i.e. H*




and HS~, while the equilibrium in step 2 lies to the side of the




reactant, i.e. HS~.  Because the equilibrium constants for steps 1




and 2 at l8°C (64.4°F) are less than the corresponding values at 25°C




(77°F) it follows that reactions 1 and 2 decrease with a decrease in




temperature (Prutton, 1951).




       Sludge deposits in streams and estuaries produce hydrogen




sulfide as they undergo anaerobic decomposition in which sulfates




are reduced.  Wheatland (195^) has indicated that the rate of forma-




tion of sulfide increases with temperature, doubling approximately




for each 10°C (l8°F) rise, and that reduction of sulfate to sulfide




will occur at temperatures as low as 5°C (ifl°F); but even at 25°C




(77°F) is inhibited by traces of dissolved oxygen.  Hydrogen sulfide




is oxidized in the presence of dissolved oxygen to water and free




sulfur or to sulfate.





                         Carbon Dioxide




       Free carbon dioxide is found in most surface waters and may




range from 0 to 5 mg/1 in rivers.  Lake waters may contain from 0 to



2 mg/1 at the surface with significant increases as the depth increases




because of the processes of decay at or near the bottom (Allee, et al.,




19^95•  Since the oxidation of organic matter furnishes carbon dioxide,




much higher concentrations may be found in surface waters receiving




organic wastes.  Surface waters receiving acid mine drainage may show




a high content of carbon dioxide; ground waters contain appreciable

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                               11

amounts of free carbon dioxide, ranging from 1 mg/1 to several hundred

(Nordell, 1961).

       The solubility of pure carbon dioxide in water ranges from

3,350 mg/1 at 0°C (32°F) to 970 mg/1 at 1*0 °C (lO^°F) and 760 mm of
mercury (Nordell, 19&1) •  The average carbon dioxide content of the
air varies from 0.035 percent in the country to 0.06 percent in the
cities and the solubility of atmospheric carbon dioxide in water
ranges from 1.0 mg/1 at OdC (32°F) and 0.03 percent to 2.0 mg/1 at
0°C (32°F) and 0.06 percent.  At kO°C (10^°F) the solubility is 0.3
mg/1 and 0.6 mg/1 respectively at 760 mm of mercury (Kordell, 1961).
These values indicate that the carbon dioxide contributed to natural
surface waters and ground waters from the atmosphere is negligible

compared with that from decaying organic matter.
       Carbon dioxide reacts with water to form a weakly dissociated

acid-carbonic acid,   C02+H^0 _±zz>H2CO~

Carbonic acid is a dibasic acid ionizing in two steps:
       2.  HC05 <— ^ ffN-CO?
              ^    jp.      ^

An equilibrium relation is established for each step as follows:

       1.
       2.
            [HCO§]

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                               12





Equilibrium constant values for steps 1 and 2 respectively, are




4.31^10~7  and 5.6xlO~i:L at 25°C (77°F) (Lange, 196l).  At this




point carbon dioxide establishes an equilibrium relation with the




mineral content of the water.





                             Ammonia




       Ammonia is an intermediate product in the bacterial decomposi-



tion of nitrogenous organic matter and may be discharged to natural




waters as a waste product of industry.  The presence of free ammonia




in natural waters is indicative of recent organic pollution, since




the atmosphere is substantially free of this substance (Camp, 19&3) •




       The gas reacts with water to produce ammonium hydroxide (often




termed aqueous ammonia); the ammonium hydroxide in turn ionizes to




produce ammonium and hydroxal ions.
An equilibrium relation is established as follows:
With ionization constant values of 1.8xlO"5 at 25°C (TT°F) and



2. 0x10 ~5 at lK>°C  (104°F) (Lange, 1961).  The equilibrium is shifted



toward the product side, i.e-. N%; OH~ at higher temperatures.  Because



the hydroxyl ion  is a product of the ionization of ammonium hydroxide,



the degree of ionization can be related to the hydrogen ion concentra-



tion or pH.  Camp (1963) indicates that 99.99$ of the ammonia in

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                               13

dilute solutions at 25 °C (77°F) is in the form of ammonixn ion at a

pH of 5 while at pH 11 only 1.78$ is in this form.

                         Sulfur Dioxide

       Sulfur dioxide is formed in one of three ways: (l) as an inter-

mediate product in the oxidation of hydrogen sulfide under aerobic

conditions, (2) in the reduction of sulfate under anaerobic conditions

and (3) in the combination of elemental sulfur and oxygen.  This is

part of the sulfur cycle taking place in many natural waters (Sawyer,

1960).  Combustion fumes from industrial operations may contribute

to the sulfur dioxide content of natural waters.
       Sulfur dioxide reacts with water to form sulfurous acid; the

acid then ionizing in two steps --
       2.  HS03 < — >

and equilibrium relation  is established as follows:

       1.  [ff*"][ HSOJ]
             [^863]

       2.  [H*][SO^]
             [HSO^]

With ionization constant  values of 1.72xlO~2 and 6.24x10"^ at 25°C

(Y7°P) for steps  1 and 2, respectively (Lange, 1961).  In the

presence of  dissolved  oxygen,  sulfurous acid and the hydrogen sul-

fite ion are readily oxidized.  The  solubility of  sulfur dioxide

markedly decreases with increasing temperatures (Camp, 1963).

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Dissolved Minerals




       About 99$ of the dissolved mineral matter found in natural




waters embrace only 10 elements, namely; hydrogen, oxygen, sodium,




potassium magnesium, calcium, silicon, sulfur, carbon and chlorine.




These occur as ions, radicals or molecules.  At ordinary temperatures




of natural waters some complexing of major dissolved species may occur




but is limited to the formation of ion pairs [e.g., Na(COo)|, NagCOg]




(Garrels and Christ, 1965).  Helgeson (1964) in a study of the effects




of elevated temperatures on the dissociation of complex ions in solu-




tion indicated that little or no changes occur unless there is an




appreciable change in the density of the solution; with increasing




temperature and decreasing density complexing is expected to increase.




       At present only enough chemica-1 information is available to




permit calculation of the inter-actions that take place among the major




dissolved species at earth surface temperatures in media as concentrated




as sea water.  The results of such calculations indicate that more than




30$ of the sulfate and bicarbonate are tied up as ion pairs with cations,




whereas $0% of the total carbonate is complexed.  One-hundred percent




of the chloride is present in the ionic form.  Changes in temperature,




pressure and composition of the water will modify this distribution;




however, variations in temperature and pressure to which ocean waters




are subjected will produce little change in distribution.

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                                 15
       The effects of small -variations in temperature [5°C(iH°F) to




lj-0°C(l04°F)] on equilibria and reaction rates involving minor constit-




uents in natural waters can not be determined at this time because of




a lack of information on equilibrium constants, enthalpy changes and




activation energies (Garrels and Christ, 1965).

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                                   16
Summary



1.  Chemical reaction rates vary with temperature,  generally increasing as



the temperature is increased.  The change in the specific rate constant is



given by the Arrhenius equation.



2.  The solubility of gases in water varies with temperature.  Dissolved



oxygen content of a surface water is decreased by the decay or decomposition



of dissolved organic substances; the decay rate increases as the temperature



of the water increases reaching a maximum at about  30 C (86 F).

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                                    17


                             REFERENCES CITED
Allee, W. C., et al., 19^9-  Principles of animal ecology.  W. B. Saunders
Co., Philadelphia, 83? pp.

Anon., 1965.  Standard methods for the examination of water and wastewater.
Twelfth Edition, American Publ. Health Assn., Inc., New York, 769 pp.

Camp, T. R., 1963.  Water and its impurities.  Reinhold Publishing Corp.,
New York, 355 pp.

Dysart, B. C., and P. A. Krenkel, 1965.  The effects of heat on water
quality.  Proc. 20th Ind. Waste Conf., Purdue Univ., pp. 18-39.

Fair, G. M. and J. C. Geyer, 195^.  Water supply and wastewater disposal.
John Wiley & Sons, Inc., New York, 973 pp.

Garrels, R. M. and C. L. Christ, 1965.  Solutions, minerals, and equilibria.
Harper & Row, New York, if50 pp.

Glasstone, S. and D. Lewis, 1960.  Elements of physical chemistry.  D. Van
Nostrand Co., New York, 758 pp.

Helgeson, H. C., 19&f.  Complexing and hydrothermal ore deposition.  Pergamon,
New York, 128 pp.

Hoak, R. D,, 1961.  The thermal pollution problem.  Jour. Water Poll. Control
Fed., 33(12):  1267-1276.

Hutchinson, G. .E., 1957.  A treatise on limnology.  John Wiley & Sons, Inc.
New York, I:  1015 pp.

Lange, N. A., 1961.  Handbook of chemistry, tenth edition.  McGraw-Hill Book Co.,
New York, 1969 pp.

Nordell, E., 1961.  Water treatment for industrial and other uses.  Reinhold
Publishing Corp., New York, 598 pp.

Prutton, C. F. and S. H. Marion, 1951.  Fundamental principles of physical
chemistry.  The MacMillan Co., New York, 803 pp.

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                                    18

Sawyer, C. N., I960.  Chemistry for sanitary engineers.  McGraw-Hill Book
Co., New York, 36? pp.

Streeter, H. W., 1958.  The oxygen sag and dissolved oxygen relationships
in streams.  Oxygen Relationships in Streams, U. S. Dept. of Health, Educa-
tion, and Welfare, Robert A. Taft Sanitary Engineering Center, Cincinnati,
25-30.

Theriault, E. J., 1927.  The oxygen demand of polluted water.  U. S. Public
Health Service Bull. 173.

Velz, C. J. and J. J. Gannon, I960.  Forecasting heat loss in ponds and
streams.  Jour. Water Poll. Control Fed., 32(U):  392-^17.

Wheatland, A. B., 195^.  Factors affecting the formation and oxidation of
sulfides in a polluted estuary.  Jour, of Hygiene, Cambridge Univ. Press,
19U-210.

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                               II.  BACTERIA



Introduction




       Temperature changes in the aquatic environment affect ecological




relationships among the biota, processes of natural purification, and




growth and survival of microorganisms.  There are a wide variety of




microorganisms found in the aquatic environment.  The numbers and species




in the population vary depending on whether they are in ground water,




lakes, or streams.  Unpolluted water bodies have low concentrations of




microorganisms.  The microbial content of natural waters is approximately




proportional to the amoxmt of organic matter present.  Unpolluted waters




usually have a greater number of species in proportion to their total




population; conversely polluted waters usually have a greater- total



population in proportion to the number of bacterial species in their




environment.




       Many stream bacteria come from the air and soil.  Bacteria in the




air are aersols or suspended on dust particles that settle or fall with




precipitation.  Soil flora in water are due to precipitation and seeping




ground water that becomes surface run-off when entering or forming a




stream.  Many of the microorganisms that are native to natural waters




are especially adapted to the stream environment, and some are difficult




or now impossible to grow on culture media.




       Interest in the microbiology of water centers on the transmission




of disease via the water route.  Polluted waters have high concentrations




of microorganisms from municipal or industrial waste waters.  When a




stream is polluted with the excreta of warm blooded animals, it is most




likely to contain enteric pathogens.  Indicator organisms (coliforms) are





                                    19

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                                   20



used to measure bacterial concentrations and indicate the potential




presence of pathogens from sewage.




       Industrial wastes add organic and .inorganic materials to water and




in some cases large numbers of bacteria, but, in general, they do not




contain pathogenic organisms.  Industrial wastes may "be growth stimulating




or toxic to bacteria (Heukelekian, 1953).



       Water, an essential for all forms of life, can serve as a medium




for growth and reproduction for many microorganisms.  Environmental factors




affecting the growth of microorganisms are chemical, physical, and




nutritional.  Although these factors are interdependent,  the physical



factor of temperature is one of the most important.




       The relations of temperature to the growth of microorganisms are




complex.  Some hardy bacteria grow in a wide temperature range;  other



fragile bacteria grow in a narrow temperature range.  For each organism




there is a minimum, the lowest temperature at which growth can occur;  an




optimum, the temperature most favorable for growth; and a^maximum,  the




highest temperature at which growth and multiplication can occur.




       Bacteria may be classified according to their temperature require-




ments for growth.  Organisms having optimum growth temperatures under




80°C (68 F) are grouped as psychrophilejs or cold-loving;  these occur in




the soil and cold waters of the north.   Thermophiles,  heat-loving




organisms having optimum temperatures of 55° to 65° c (131 to lU9°F) are




found in soil, decaying organic matter, hot spring water and near the



discharge points of hot water effluents.  They are of little importance




in stream ecology.  The majority of bacteria are called mespphiles; they



are the intermediate group, having optimum temperatures in the range




between the extremes.  Many of the organisms found in natural waters and

-------
                                    21




soils are saprophytes (organisms that live on decaying organic matter);




they have optimum temperatures of 22° to 28° C (70 to 82°F) and "belong



to the mesophilic group.




       The parasitic bacteria have optimum temperatures of 37°C (98.6°F)



and include those microorganisms pathogenic to man.  Temperature changes



greatly affect the rate of activity of these organisms.



       The effect of temperature on the species of organisms cannot




always be considered separately from the other environmental factors.



Some species can be more abundant in the winter, others in the summer,




when their environmental conditions are varied (Burrows, 1959)•



Microbiotic^Cycles




       The interrelationships among aquatic biota are of primary importance



to the aquatic environment (ingrara, Mackenthun.  and Bartsch. 1966).  In




unpolluted waters, autotrophic algae and chlorophyll bearing bacteria




initiate microbioti-c cycles (Silvey and Roach, 196*0-  The metabolites



and decomposition products of the organisms provide nutrients for use




by gram-negative heterotrophic bacteria.  The principal gram-negative




organisms are Alcaligenes, Aerobacter, and Pseudomonas.  In waters




polluted with sewage, the Escherichia also increase, as do associated




types.  The new algal growth following the gram-negative bacilli may




be either diatoms or blue-green algae.  In cool waters the diatoms will




generally prevail and be followed in turn by the gram-positive heterotophic




bacilli.  In warmer waters, the gram-negative bacilli remain and grow




in coexistence with blue-green algae; then the actinomycetes grow and




develop an antibiotic effect,  reducing the gram-negative population.  The




gram-positive spore-forming bacilli follow the actinomycetes (Figure 2).

-------
                                       22
10
9
8
7
6
5
4
3
2
1
0
I
r-2XlO»
-2X10'
-2x10'
-2X10*
-2x10*
-2x10'
-2x10*
-2X101
-2X10°
-2X10-1
-2XJO"1
i
            -250
            -225
            -200
            - 175
            - 150
            - 125
            - ICO
            -  75
            - 50
            h- 25
            I-  0
                  Jan.   Feb.  Mir.   Apr.  May  Jun.   Jul.   Aug.  Sep.  Oct   Nov.   Dec
Tig. 2.  Annual Cycles of Gram-Positive  and 'Gram-Negative Heterotrophic Bacilli and
         Their Relationship to Blue-Green Algae and Aquatic Actinomycctes

The dotted curve w for blue-green algae, indicated i:t Wffs of areal standard units  on
Scale 1 at the left; tite solid curve is for gram-positive heterotrophs; the dashed curve,
gram-negative heterotrophs; and  the  dot-and-dash  curve,  actinomycetes.  Colony
counts of heterotrophs are indicated in IjOOO's by Scale 1; absolute numbers of bacilli
by Scale 2; a;\d actinojiiyceie plate counts in 1,000" s  by Scale 3.  Actinomycetes were
isolated on M^Bt agar.   Cram-positive hcterolropl'S were grown on vitamin-enriched
  Emerson's agar, and gram-negative heictqlroplis  on. epsin mrthylene blue agar.
     (From  Silvey  and Roach.  19o'K  p.  o4;

-------
                                   23



These various cycles may impinge upon each other when the stream or



reservoir is highly polluted.




Self-Purification




       When organic wastes are discharged into a receiving water a complex




chain of physical, chemical, biochemical and biological activities are



started which result in decomposition and degradation of the wastes.   This




complex process is self-purification, the details of which are still  unknown




(Heukelekian, 1953).



       The most significant function in natural purification is the




decomposition of organic matter by the microbial flora.  The saprophytic




organisms are the most active in this biochemical process; they have




optimum temperatures that are near the ambient temperature of many streams




during the summer months.  In a stream polluted with sewage, the pathogenic




and indicator organisms are also present and perform a minor role in  the



self-purification process.  The temperature of the stream water even  during




the summer is below the optimum for pollution associated bacteria.  Increas-




ing the water temperature increases the bacterial multiplication rate when



the environment is favorable and the food supply is abundant.  Increasing




the water temperature within the growth range of the bacteria causes  a more



rapid die-off when the food supply is limited.  The decrease of bacterial




numbers is higher during the summer than during the winter (Figure 3)•




       Unpolluted streams contain dissolved oxygen near saturation levels.




When organic wastes are discharged into the stream, the biochemical pro-




cess is aerobic - reducing the dissolved oxygen by oxidation and dilution.




When the reaeration rate of the stream is low, the oxygen may be




depleted by the bacterial metabolism of the increasing population.

-------
                       12
Figure 3. From Huekelekian,
       1953, p. 27

-------
                                    25





As the available oxygen diminishes, the aerobic organisms die-off




rapidly, sharply decreasing the natural purification process of




assimilating the organic waste load.  When this occurs there is a shift




in the flora in the stream to the facultative anaerobic organisms.  The




ultimate result of anaerobic decomposition may be the same as that of




the aerobic, but it is very slow and less -desirable (Fair, Geyer, and




Morris, 1958).




       As the temperature increases, the dissolved oxygen solubility




decreases.  When a warm water discharge is near a sewage treatment plant




outfall, self-purification can be very effective if the organic waste



load is not excessive.




       The impounding of water may improve  the water quality by




reducing sediments, color, bacteria and temperature.  Ingols (1957) found




also that the pattern of reservoir discharge permitted slime develop-




ment downstream during low flow and scouring of the slime during high




flows.  "Out of phase" dilution of the receiving stream could promote




or retard stream self-purification (Berger, 1961).  Berger as well as




Renn (1957) agree that the rate of stream reaeration increased at higher




temperatures.




Growth and Survival




       Chambers and Clarke (1966) state:  "Many bacteria reproduce in




water- among the genera that, will grow in water of unquestioned potable




quality are: (l) Pseudomonas,  (2) Xanthomonas, (3) Achromobacter.




(U) Escherichia, (5) Aerobacter,  (6) Streptococcus,  (7)  Desulfovibrio,




and (8) Crenothrix."



       Renn (1957) points out that elevating the stream temperature can




be favorable for those bacteria that can  multiply in water by inducing

-------
                                    26
the recurring cycles of life and death more rapidly.  However,  enteric




pathogens have highly selective requirements.  They cannot multiply or




survive well in natural water,  so they die-off more rapidly.




       Because higher temperatures in a stream polluted with sewage




generally result in increased bacterial numbers, low temperatures are




not conducive to rapid growth.   Stream temperatures of 1° to 8°C (33-8-




k6.k F) may surpress growth and multiplication, and act as a preservative




as in the storage of samples for bacterial analyses.  Freezing of water



can result in reduced microbial populations by killing off a majority




of the microorganisms.  Streams that have high organic waste loads and




low temperatures tend to develop slime organisms,  generally Sphaerotilus.




       A study of the Columbia River in Oregon showed Sphaerotilus




growths to be maximum at water temperatures of 10° to 15 C (50 -59 F).




Growth ceased when temperatures dropped below k°C (39.2°F) and resumed




when temperatures increased above U°c (39.2°F).  Infestations of



Sphaerotilus  may occur at temperatures below 10°C (50°F) if the growing




period is sufficiently long (Amberg and Cormack, 1960).  Beds of




Sphaerotilus slime may extend farther downstream from a waste outfall




in the winter than in the summer when warmer temperatures seem to inhibit




the efficiency of food conversion by the organism (Dondero, 1961).




       Clark et al. (19&0 assessed the value of bacterial indicators




of pollutions as indicators of viral pollution by studying the relative




survival of the organisms in water-  They observed that the lower the




temperature the longer the survival for both bacteria and viruses.  The




enteric bacteria had survival times in proportion to the degree of




pollution, the greater the pollution the longer the survival time.  The

-------
                                    27



increased quantity of nutrients present in the more polluted water may




account for the longer survival time of the bacteria.   The viruses




studied survived longer in the "clean" Little Miami River water and in




the grossly polluted raw sewage than in the moderately polluted Ohio




River water.  They also point out the difficulty in generalizing on




comparative survival times because the different genera of organisms




may have different survival times in the different stream environments




(Table l).




Summary




       The temperature of stream water, even during the summer, is




below the optimum for pollution-associated bacteria.  Increasing the




water temperature increases the bacterial multiplication rate when the




environment is favorable and the food supply is abundant.  Increasing the




water temperature within the growth range of the bacteria causes a more




rapid die-off when the food supply is limited.

-------
                                        ?8
                                   TABLE 1

          AVERAGE TIME IN DAYS FOR 99-9 PERCENT REDUCTION IN ORIGINAL
            TITER OF INDICATED MICROORGANISMS AT THREE TEMPERATURES


              (From Clarke, Berg, Kabler and Chang, p. 526, 196U)
                    Little Miami River       Ohio R-foer      	    	.Sewage
   Microorganisms   - 28<>c  2Qoc   ^c     ^^  2Q4C   ^    g8«c  ZQ9(,~  ^OG


Poliovirus I          1?    20     2?      11    13     19     1?    23    110
ECHO 7                12    16     26       $     7     15     28    1*1    130
ECHO 12                5    12     33       3     5     19     20    32     60
Coxsackie Aj?         < 8   < 8     10       5     8     20      6  No Data  12
A. aerogenes           6     8     15      15    18     W*     10    21     56
E. coli                67     10       551112    20     lf8
S. fecalis             6     8     17       9    18     57     ^    26     U8

-------
Amberg, H. R. and J. F. Cormack, 1960.  Factors affecting  slime  growth
in the lower Columbia River and evaluation of  some possible  control
measures.  Pulp Paper Mag. Can., 6l:T-70 to T-80.

Berger, B. B., 1961.  Does production of power pollute  our rivers?
Power Engineering, March, p 60-6l.

Burrows, W., 1959-  Textbook of microbiology.  W. B.  Saundefs  Co.,
Philadelphia, Pa.

Chambers, C. and N. Clarke, 1966.  Control of  bacteria  in  nondomestic
water supplies.  Advances in Applied Microbiology.  Academic Press Inc.,
N. Y., 8:105-1^3.

Clarke, N., G. Berg, P. Kabler, and S.  Chang,  196U.   Human enteric
viruses in water: sour.ce, survival and  removability.  Reprint  from
International Conference on Water Pollution Research.   London, 1962.
Pergamon Press - Oxford, London, N. Y.  and Paris, pp. 523-536.

Bonders, N. C., 1961.  Sphaerotilus, its nature and significance.
In: Advances in Applied Microbiology, £• 77 -10?.

Fair, G. M. , J. C. Geyer and J. C. Morris, 1958.  Water supply and
waste-water disposal.  John Wiley & Sons, Inc., New York.

Heukelekian, H., 1953-  Stream pollution and self -purification.  In:
industrial wastes their disposal and treatment.  (W.  Rudolfs,  Ed.)
Reinhold Pub. Corp., New York, pp 8-30.

Ingols, R. S., 195T-  Pollutional effects of hydraulic  power generation
stream pollution.  Sewage and Industrial Wastes, 29(3): 292-29?.

Ingram, W. M., K. M. Mackenthun. and A. F. Bartsch, 1966.  Biological
field investigative data for water pollution surveys.   U.  S. Department
of the Interior, Federal Water Pollution Control Administration, WP-13,
139 PP-

Renn, C. E. , 1957-  Warm-water effects  on municipal supplies.  Jour.
American Water Works Assoc., U
Silvey, J. K. G., A. W. Reach, 1964.  Studies on microbiotic cycles in
surface waters.  Jour. American Water Works Assoc., 56(l):60-?2.

-------
                          III.  FRESHWATER PISHES








Introduction




       Changes in fish populations can result from the many types of arti-



ficial cooling and heating of natural water*.  These changes result from



the discharge of condensed water from steam-electric generating plants,



distillery effluents, and irrigation waters.  Stream temperatures are



raised also by the removal of stream bank trees and other vegetation.



Water temperatures are often elevated in excess of the air temperature by



absorption of heat by the .stream bed.  Yet another type of thermal pollu-



tion results from the discharge of cold-water from stratified impoundments;



this water may provide an ideal habitat for trout and other cold-water fish



when sufficient dissolved oxygen is present.



General Effects



       The effects of temperature on fish are acute because fish do not



possess an efficient method to compensate their internal temperature against



a temperature change in the water in which they are immersed.  If a more



favorable temperature is available, fish do have the ability to seek it out.



       Chemical reactions are accelerated within the body cells with ele-



vated temperatures.  Prosser (1955) discusses four possible death mechanisms,



although he does not attach specific temperature values to various death



processes.  These are:  an enzyme inactivity caused by the acceleration of
                                     30

-------
                                    31
the enzyme reaction to such a state that it is no longer effective; coagula-




tion of cell proteins; melting of cell fats; reduction in the permeability




of cell membranes.  Cells may also be killed by toxic action of the products




of metabolism and, incomplete metabolism accumulating in the cells (Ellis,




19U7).



         According to Brett (i960), temperature acts in a variety of different




ways; it can be lethal, cause a reduction of activity, and limit reproduction.



The slow rate of acclimation appears to result in greater mortalities from




cold despite the ability of fish to withstand lower temperatures.




Sudden Temperature Changes




         The effect of thermal shock on fish can be more harmful than continued




exposure to a higher temperature (Cairns, 1956).  In studies with rainbow




trout, Threinen (1958) found that death /would result from an instantaneous




or rapid increase (shock) of U.1°C (20°F) above an acclimation temperature




of 12.2°C (5^°F), however, a similar increase of 8.4°C (l5°F) could be




tolerated from a temperature of 10.6 C (51 F).  A rise of the acclimation




temperature from 12.2 to l8.lj°C (5U-65°F) during a 2U-hour period permitted




trout to withstand a temperature of 23.^ C (7^ F) with only minor distress




for short periods.



         Fish having the ability to adapt to higher temperatures faster and




over a larger gradient often are attracted to artificially heated water with-




out a resultant mortality.  However, mortality often results when these fish

-------
                                     32
return to cold water.  Agersborg (1930) found fish dying when they attempted



to return from heated water (26.1°C - 79°F) to the colder stream (0°C - 32°F);




death occurred even when fish moved into water that was 5.6°C (10°F) cooler.



Falkner and Houston (1966) found that the mean erythrocytic (red blood cell)




volume underwent a transient decrease while total blood iron (and presumably




haemoglobin and mean erythrocytic iron content) fell slightly after goldfish




which had been acclimated to 20°C (68°F) were subjected to an abrupt increase




of 10°C (l8°F).  Heinicke and Houston (1965) concluded that while thermal




shock induces initial deviations in iono- and osmoregulatory ability the




goldfish can compensate for these changes during the acclimation period




through respiratory activities, and restore its original ionic status.




       A rise in temperature from 10°C to 20°C (50-68°F) reduced resistance




to a decrease in oxygen in perch, roach, and mirrorcarp.  In rainbow trout




the resistance was lowered considerably between a rise in temperature from




10°C to 16°C (50-60.8°F) (Downing and Merkens, 1957).




Acclimation




       The importance of acclimation temperatures has long been known to



fish hatchery personnel and physiologists working on lethal temperatures.




Much of the work on lethal temperatures is of little value because holding




temperatures and durations are not given.  Springtime mortalities often re-



sult from fish being subjected to warmer water temperature after acclimation




to cold winter temperatures.

-------
                                     33
       Doudorof£ in Brown (l951\ discussing the work of Pry, et al.,



and Fry (19^7) concludes that fish could stand brief exposures to considerably



higher temperatures without showing distress when they had been acclimated to



the maximum possible temperature.  However, the fish suffered mortality when



they had been acclimated to low temperatures.  Similarly, Doudoroff in dis-



cussing the work of Hart (1952) noted considerable geographic, seasonal, and



other variations of the resistance to heat of some species of fish acclimated



to the same temperature.



       Doudoroff in discussing the rate of acclimation summarized the work



of many workers and concluded that the increased heat resistance (the ability



to withstand increased water temperatures) is acquired usually at a very fast



rate in the high temperature range from 26°C to 30°C (T8.8-86.0°F) although



there may be a latent period of one or longer days in which virtually no



change takes place in the upper lethal temperatures.  Most of the resulting



increase was achieved in a period of one to three days.  There was little



or no loss of resistance in the first three days.  Thus, if a fish has



acquired a higher heat resistance it will not be lost rapidly on subsequent



exposures to low temperatures.



       Jones (196^) in discussing the work of Sumner and Wells noted that the



tolerance to high temperatures once acquired may persist for considerable



periods after return of the fish to the acclimation temperature.   The time

-------
of acclimation need not be continuous.  An intermittent exposure to a



different temperature for sufficient hours per day can produce the same



acclimation temperature as a continuous exposure.



       The acclimation of fish is important in determining the maximum en-



vironmental temperature in which fish can survive.  Jones (1964) discusses



how the resistance time shortens with a progressive rise in temperature until



the fish succumbs to an ultimate lethal temperature.  As the acclimation



temperature rises the. thermal death point rises, but it rises at a slower



rate.  Accordingly, experiments on roach show that for every 3-degrees rise



in acclimation temperature the thermal death point rises only 1-degree C.(l.8 F).



Maximum Temperatures



       Maximum temperatures have been determined for numerous species of



fish (Table 2_).  Tnese temperatures are important in determining the absolute



temperature at which a fish can survive, but they are often higher than the



maximum temperature at which a population can survive.



       Alabaster (1962) found that heated effluents, by virtue of their



high temperatures only, may be lethal to caged trout and coarse fish



acclimated to normal river temperatures during the summer and may also



occasionally kill free-living fish which are near effluent outfalls when



temperatures increase rapidly.  Small free-living fish are principally



affected, large fish apparently are able to swim away to safety.  He con-



cludes further that where the water temperature of the whole river is above



normal because of mixing with continuous discharge of heated effluent,

-------
                                      35

                                 TABLE 2

                    TOLERANCE LIMITS FOR CERTAIN FISHES

   Values are LD5Q temperature tolerance limits, i.e., water temperatures
survived by 50 percent of the test animals. Counts were made by observing
or estimating the number killed during exposure, or within a reasonable time
thereafter in which it could be safely assumed that all deaths were attributable
to the temperature effects.

                 (This Table Taken in Part From Anon.,  1962)

                                                Lower  ijimTCUpper  limit
      FiSh                  O,-,        /O-ciN       Or>      f °-C<\     -rr    O
                                               C      (°F)     Hr    °C      (°F)	Hr
Bass , largemouth
(Micropjterus salmoides
f loridanus )
Bluegill (Lepomis
macrochirus macrochirus )
Bluegill (L. macrochirus
purpurescens)
Bullhead (Ameiurus n.
nebulosus, A. n.
narmoratus )
Catfish, channel
(ictalurus lacustris
20.0°C
30.0°C

10.0°C
30.08C
15.0°C
30.0°C
20.0°C
30.0°C

15.0°C
25.0°C
(68
(86

(50
(86
(59
(86
(68
(86

(59
(77
.0°F)
.0°F)

.O°F)
.0°F)
.0°F)
.0°F)
-0°F)
,0°F)

.0°F)
.0°F)
5.0°C
11.0°C



3.0°C
11.0°C
1.0°C
7.0°C

0.0°C
6.o°c
(41
(51



(37
(51
(33
(44

(32
(42
.0°F)
.8°F)



.4°F)
.8°F)
.8°F)
.6°F)

.0°?)
.6 .-')
24
24



24
24
24
24

24
24
32.0°C
24.0°C

29.0°C
36.0°C
?,1.0°C
34.o°c
32.0°C
35.o°c

30.o'c
34.o°c
(89
(93

(82
(96
(87
(93
(89
(95

(86
(93
.6°F)
.2°F)

.4°F)
.9°F)
.8°F)
.2°F)
.6°F)
.0°F)

.0°F)
.2°F)
72
72

24
24
60
60
96
96

24
24
lacustris, I.I punctatus)
Chub, creek (Semotilus
a. at romaculatus )
5
25
Dace, blacknose (Rhinich- 5
thys a. atratulus, R.a. 25
meleagris )
Goldfish (Carassius
auratus )
Greenfish (girella
nigricans)
Killifish (Fundulus
heteroclitusl
Minnow, fathead
(Pimephales promelas)
Minnow, blunt-nose
(Hyborhyncnus notatus)

2
24
37
12
18
14
20
20
30
15
25
.0°C
.0°C
.o°c
.o°c
.o°c
.o°c
.o°c
.o°c
.o°c
.o°c
.o°c
.o°c
.o°c
.o°c
,o°c
(41
(77
(41
(77
(35
(62
(75
98
(53
(64
(57
(68
(68
(96
(59
(77
.0°F)
.0°F)
.0°F)
.0°F)
.6°F)
.6°F)
.2°F)
.6°F)
.68F)
.4°F)
.2°F)
.0°F)
.0°F)
.0°F)
.0°F)
.0°F)

5.0°C
0.0°C
5.0°C
15.0°C
5.0°C
13.0°C
1.0°C
2.0°C
2.0°C
1.0°C
8.0°C

(41
(32
(41
59
(41
(55
(33
(35
(35
(51
(33
(46

.0°F)
.O°F)
.O°F)
.O°F)
.0°F)
.4°F)
.8°F)
.6°F)
.6°F)
.88F)
.8°F)
.4°F)

24
14
14
14
120
72
48
48
24
24
24
24
25.0°C
32.0°C
30.0°C
29.0°C
28.0°C
34.0°C
(77.0°F)
(89.6°F)
(8o.6°F)
(84.2°F)
(82.4°F)
(93.2°F)
36.0°c (96.8°F)
42.o0c(l07.68F)
30.o°c (86.0°F)
31.0°C (87.8°F)
32.0°C
34.o°c
32.0°C
33«0°C
31.0°C
33-0°C
(89.6°F)
(93-2°F)
(89.6°F)
(9l.4°F)
(87.8°F)
(9l.4°F)
96
96
340
340
14
14
14
14
120
120
133
133
133
133

-------
TABLE 2, Continued
                                      36
Fish
Acclimated to
°r" ( °TP\
v \ £ /
Lower Limit
O/i / O-ri \
u \ -T /
Hr
Upper Limit
°C (°F)
Hr
Mosquito fish            15.0°C
(Grambusia affinis       35.0°C
affinis, G.a. holbroki)

Perch (Perca flavescens)  5«0°C
Winter                   25.0°C
Summer

Shad, gizzard
(Dprosoma cepedianum)

Shiner, common
(Uptropis cornutus
frontalis)

Shiner, common
(Notropic .cornutus
chrysocephalus)

Shiner, lake
(N. atherinoides)
                         25.0°C

                         25.0°C
                         35.0°C

                          5-0°C
                         25.0°C
                         30.0°C

                         25.0°C
                         30.0°C
                          5.0°C
                         15.0°c
                         25.0°C
Shiner, golden           20.0°C
(Notemigonus c.          30.0°C
crysoleucas, N.c, auratus)

Sucker, common           15-0°C
(Catostomus commersoni)  25.0°C
Sunfish
(Lepomis gibbosus)
                         10.0°C
                         30.o°c
                                  (59.
                                  (95.
    0°F)   2.0°C (35.6°F)  2lf  35.0°C (95.0°F)  -66
    0°F)  I5.o°c (59.0°F)  2k  37.0°c (98.6°F)  ,66
(Ul.O°F)                       21.0°C (69.8°F)   96
(77.0°F)   k.O°C (39.2°F)   2k  30.0°C (86.0°F)   96
(77.0'F)   9.0°c (U8.2°F)   2k  32.o°c (89.6°F)   96
(77.0°F)  11.0°C (51-8°F)
(95.0°F)  20.0°c (68.0°F)  2k  37.o°c (98.6°F)   kQ
                           2k  3k.O°C (93.2°F)
(Ul.O°F)
(77.0°F)
(86.0°F)   8.0°c (U6.

(77.0°F)
(86.o°F)
                               27-0°C (80.6°F) 133
           k.o°c (39-2°F)   2k  31.0°C (87.8°F) 133
                       !•)   2k  3l.O°C (87.8°F) 133
                               32.0°C (89.6°F)  133
                               3U.O°C (93.2°F)  133
(ifl.O°F)
  ).0°P)
                               23.0°C (73.U°F)  133
(59.0°F)   2.0°C (35.6°F)   2k  29.0°C (84.2°F)  133
                                  (77.0°F)   8.0°c (U6J
Trout, brook              3.0°C   (37.J
(Salvelinus fontinalis)  20.0°C   (68.0°F)
                         25.0°C   (77.0°F)
                           2k  3i.o°c (87.8°F)  133
(68.0°F)   8.o°c (k6.k*F]   2k  32.o°c (89.6°F)   66
(86.0°F)  11.0°C (51.8°F)   2k  35.0°C (95.0°F)   66


(59-0°F)
(77.0°F)   5.o°c fUl.O°p)   2k  29.o°c (84.2°F)  133

(50.0°F)                       28.0°C (82.U°F)   2k
(86.0°F)                       2k.O°C (75.2°F)   2k
                                                                 23.0°C (73.U°F) 133
                                                                 25.0°C (77»0°F) 133
                                                                 25.0°C f77.0°F) 133

-------
                                     37
coarse fish populations may "be reduced locally when the mean daily tem-

perature reaches 30°C (86°F) and increased when the water is not warmed

to more than 26°C (T8.8°F).

       Wells (191^) concluded that the resistance of fish to temperature

varies with species and size of fish.  There is no definite maximum tem-

perature for a given species of fish; it varies with the fish's rate of

heating, size, and physiological condition.

       A temperature need not kill the fish directly for it to be lethal.

Brook trout were found to be comparatively slow in catching minnows at

1?.2°C (63°F) and virtually incapable of catching minnows at 21°C (TO°F).

This resulted in the trout virtually starving to death (Anon., 1962).

       As a maximum temperature for cold water fishes the Pennsylvania

Department of Health recommends*:  that no wastes or waters shall be added

from any source having temperatures in excess of that of the receiving

waters except that during the period October through May, when stream

temperatures are below 1^.5°C (58°F) the temperature of wastes discharged

to the streams shall not exceed lk.5°C (58°F).  To allow for the normal

production of aquatic life in warm water lakes and streams it is recommended

that water temperatures resulting from thermal discharge shall not exceed
*  Anon., 1962.  Heated discharges . . . their effect on streams.  Rep. by
   the Advisory Committee for the control of stream temperatures to the
   Pa. Sanitary Water Board.  Pa. Dept. Health, Harrisburg, Publ. No. 3,
   108 pp.

-------
                                     38
30 C (93 F) exclusive of the required mixing zone and in no case shall this



peak temperature prevail for more than eight hours in any 2U-hour period.



Preferred Temperature



         It is generally acknowledged that fish can live for short periods



of time in higher than normal temperatures,  but at these temperatures fish



cannot perpetuate their populations.  Pish are extremely sensitive to tem-



perature and seek out the temperature that is best for their survival.



The temperature that fish seek out is termed "preferred temperature;" these



are listed for several species of fish in Tables 3 and h and Figures U and 5.



         Windermere char eggs hatched in U5 days at 10°C (50°F), and in 95



days at k°C (39.2°F) (Swift, 1965).  Mortality to some extent occurred at



8°C (U6.U°F), with total mortality occurring at 12°C (53.6°F).



         Ferguson (1958) concluded that the level of thermal acclimation in-



fluences the range of temperature preferred.  In general, the preferred



temperature is considerably higher than the acclimation temperature at



lower thermal acclimations, but this difference decreases up to the final



preferred temperature where both coincide.  A final preferred temperature



and the relation between acclimation and preferred temperature  is charac-



teristic for the species.



         Tarzwell (1957) concluded that while temperatures higher than the



optimum, and high temperatures of short duration, 23.9 to 2?.8°C (75 to 82°F),

-------
                                         39
                                    TABLE 3
            THE FINAL TEMPF.RATUKE PKEFERENDA FOR VARIOUS SPECIES OF FISH
                      AS DETERMINED BY LABORATORY EXPERIMENTS

           Young of the Year or Yearling Fish Were Used, Except as Noted.
                   (This Table Taken in Part From Ferguson, 1958)
        Species
       Final
   Preferendum
        Authority
Bluegill
(Lepomis macrochirus)

Bass, Largemouth
(Micropterus salmoides)

Carp
(Cyprinus carpio)

Pumpkinseed
(Lepomis gibbosus)

Goldfish
(Carasjius auratus)

Bass, Smallmouth
(Micropterus dolomieu)

Grass Pickeral
(ESOX vermiculatus)

Yellow Perch
(Perca flavescens)

Muskellange
(ESOX masquinongy)

Burbot
(Lota lota lacustris)

Yellow Perch
(Perca flavescens)

Brown Trout
(Salmo trutta)

Brook Trout
(Salvelinus fontinalis)

Rainbow Trout
(Salmo gairdnerii)

Lake Whitefish
(Coregonus clupeaformis)
      32.3°C
     (90.1°F)

 30.0-32.0°C
  (86-89. 6°F)

      32.0°C
     (89.6°F)

      31.5°C
     (88.7°F)

      28.l°c
     (78.8°F)

      28.o°c
     (82.1f°F)

      26.6°c
     (78.8°F)
     (75.6°F)
     (75.2°F)

      21.2°C
     (70.2°F)

      21.0°C
     (69.8°F)
(57.2~60.8°F)

      13.6°C
     (56.5°F)
Fry and Pearson
 (MS, 1952)

Fry
 (MS, 1950)

Pitt, Garside and Hepburn
 (1956)

Anderson
 (MS, 1951)

Fry
Fry
(MS, 1950)

Berst and Lapworth
(MS, 1950)

Ferguson
(1958)

Jackson and Price
(MS,
Grossman, Ireyawa and Pecicock
(MS, 1953)

McCracken and Starkma
(MS,
                         Tait
                         (MS,  1958)

                         Graham
      12.7°C
Garside and Tait
(MS, 1958)

Tompkins and Fraser
(MS, 1950)
Lake Trout
(Salvelinus namaycuch)
      12.0°C
     (53-6°F)
McCauley and Tait
(MS, 1956)

-------
                                   TABLE k

                FIELD OBSERVATIONS ON VARIOUS SPECIES OF FISH
                         AND ASSOCIATED TEMPERATURES

  Some temperatures are estimates derived from Ferguson's tabled or figured
data.  August distributions and temperatures were used wherever possible.   These
figures represent the temperature of the w ater strata in which the fish seemed
to concentrate.  It is judged these represent preferred natural temperature.

               (This Table Taken in Part from Ferguson, 1958)
Species Temperature
Bass, Largemouth 26. 6-27. 7 °C
(Micropterus .s^Jnoides) (80.0-8l.9°F)
Bass, Spotted 23.5-2l*.l*°C
(Microjpterus punctulatus) (7l*.l-75.9°F)
Walleye 20.6°C
(Stizostedion v. vitreum) (69.1°F)
Walleye 22. 7-23. 2 °C
(Stizostedion v. vitreum) (72.9-73-8°?)
Gizzard Shad 22.5-23.0°C
(Dorosoma cepedianum) (72,5-73.1*°F)
Freshwater Drum 21. 6-22. 2 °C
fAplodinotus grunniens ) (70 . 9-72 . 0 °F )
Rock Bass ll*.7-21.3°C
(Ambloplites rupestris) ( 58 . 5-70 . 3 °F )
Rock Bass 20.7°C
(Ambloplites rupestris) (69.3°F)
Yellow Perch 21.2°C
(Perca flavescens) (70.2°F)
Yellow Perch 2l.O°C
(Perca flavescens) (69.8°F)
Yellow Perch 12.2°C
(Perca flavescens, small) (5l*.0°F)
Yellow Perch 20.2°C
(Perca flavescens, larger) (68.1*°F)
Water
Norris
Reservoir
Norris
Reservoir
Trout
Lake
Norris
Reservoir
Norris
Reservoir
Norris
Reservoir
Lakes
Streams
Lake
Opeongo
Costello
Lake
Muskellunge
Lake
Muskellunge
Lake
Location
Tennessee
Tennessee
Wi scons in
Tennessee
Tennessee
Tennessee
Wisconsin
S. Ontario
Ontario
Ontario
Wisconsin
Wisconsin
Author
Dendy
19^8
Dendy
Hile and
Juday, 1<
Dendy
191*8
Dendy
Dendy
Hile and
Juday, 1<
Hallara
1958
Present
Work
Present
Work
Hile and
Juday, IS
Hile and
Juday, 1£

-------
TABLE  k, continued
Species Temperature
V7ater
Location
Author

Yellow Perch 20. 2° C
(Perca flavescens, larger) (68;^°F)
21.0°C
(69.8°F)
20.8°C
(69.5°F)
19.7 °c
(67.5°F)
Bass, Smallmouth 20.3-21.3°C
(Micropterus dolomieui) (68.5-70.3°F)
21. VC
(70.5°F)
Sauger l8.6-19.2°C
(Stizostedion canadense ) (65.1-66.6°F)
Brook Trout llf.2-20.3°C
(Salvelinus fontinalis) (57.6-68.5°F)
15.7°C
(60.3°F)
" " 12.0-20.0°C
(53.6-68.0°F)
Mottled Sculpin l6.5°C
(Cottus bairdii) (6l.7°F)
Brook Trout x Lake Trout 13.1°C
(Salvelinus hybrid) (55.6°F)
White Sucker 11.8-20.6°C
(Catostomus commer sonnii ) (53.3-69.1°F)
White Sucker
(Catostomus c. lU.l-l8.3°C
commersonnii) (57.1f_64.9°F)
Round Whitefish 13. 9- 17. 5 °C
(Prosopium cylindraceum) (57.0-63.5°F)
Alewife h.h- 8.8°C
(Pomolobus pseudoharengus) (39. 9-1*7 .8°F)
Silver
Lake
Nebish
Lake
Trout
Lake
Lake
Nipissing
Nebish
Lake
Streams
Norris
Reservoir
Moosehead
Lake
Streams
Redrock
Lake
Streams
Wisconsin
Wisconsin
Wisconsin
Ontario
Wisconsin
R. Ontario
Tennessee
Maine
S. Ontario
Ontario
S . Ontario
Jack L. Ontario
Sproule Lake
Musk, Trout
Silver
Moosehead
Lake
Moosehead
Lake
Cayuga
Lake
Wisconsin
Maine
Maine
New York
Hile and
Juday, 19^1
Hile and
Juday, 19^1
Hile and
Juday, 19^1
Present
Work
Hile and
Juday, 19^1
Hallam
1958
Bendy
19U8
Cooper and
Fuller
Hallam
1958
Baldwin
19*48
Hallam
1958
Martin and
Baldwin, 1958
Hile and
Juday, 19Ul
Cooper and
Fuller, 19^5
Cooper and
Fuller, 19^5
Galligan
1951

-------
TABLE U  , continued
                                       1*2
Species
Lake Trout
(Salvelinus naraaycush)
tt ii
it ti
it it
American Smelt
(Osmerus mordax)
it it
Temperature
10. 0-15. 5 °C
(50.0-59.0°F)
lk.O°C
(57.2°F)
n.o-ii.5°c
(51.8-52 .?°F)
8.0-10.0°C
(46.1f-50.0°F)
12.8°C
(55.0°F)
6.6- 8.3°C
(^3.9-47.9^)
Lake Whitefish 11.4-11.9°C
(Coregonus clupeaf ormis ) (52. 5-53. 5 °P)
Longnose Sucker
(Catostoimis catostomus)
Burbot
(Lota lota maculqsa)
Coregonys
(Leucichthys artedi)
ii ti
(Cisco or Lake Herring)
ll.o-ll.6°c
(51.8-53-0°F)
io.8-ii.U°c
(51.4-52. 5°F)
8.0-10.0°C
(46.4-50.0°F)
5.5- 7.2°C
(4l.9-44.8°F)
Water
Cayuga
Lake
White
Lake
Moosehead
Lake
Louisa
Redrock
Lake
Charaplain
Cayuga
Lake
Moosehead
Lake
Moosehead
Lake
Moosehead
Lake
Lake
Nipissing
Cayuga
Lake
Location
New York
Ontario
Maine
Ontario
New York
New York
Maine
Maine
Maine
Ontario
New York
Author
Galligan
1951
Kennedy
19^1
Cooper and
Fuller, 19^5
Martin
1952
Greene
1930
Galligan
1951
Cooper and
Fuller, 19^5
Cooper and
Fuller, 19^5
Cooper and
Fuller, 19^5
Fry
1937
Galligan
1951

-------
 U_
 o
    82.4
    78.8
    75.2
    71.6
    68.0
 iu  64.4
 cc

 5  60'8
 QL
 S  57.2
 S
 LU
 h-  53.6
    50.0
    46.4
    42.8
FIGURE  4
o
o

 I
LU
ID
H
Cd
LU
a.
2
LU
28

26

24

22

20

18

16

14

12
   8
          APLODINOTUS GRUNN1ENS
           (FRESHWATER DRUM)
            tMOTTLED SCULPIN)
             COTTUS  BAIRDIJ
          •(WHITE LAKE ONTARIO)-
           SALVELINUS NAMAYCUSH
             (LAKE  TROUT)
            (CAYUGA  LAKE)
                        (MOOSEHEAD  LAKE)
                             LOUISA
                          REDROCK LAKES
                                                MICROPTERUS  SALMOIDES
                                                             BASS)
                                   MICROPTERUS PUNCTULATUS
                                     (SPOTTED BASS)	j
   (SMALLMOUTH BASS)
-C  M. DOLOMIEU   ~
                                  -C  AMBLOPTTTETS  SP.
                                       (ROCK BASS)
PROSOPIUM  CYLINDRACEUM
     QUADRILATER1.E
    ROUND WHITEFISH
  SALVELINUS HYBRID
                                   CORE60NUS CLUPEAFORMIS
                                     (LAKE WHITEFISH)
                                    COREGONUS (WHITEFISH)
                                    (LAKE  NIPISSING)
                                   COREGONUS (WHITEFISH)
                                      (CAYUGA  LAKE)
FIELD  OBSERVATIONS  OF  FISH  AND  ASSOCIATED  TEMPERATURES
DURING  MIDSUMMER  (AUGUST  MOSTLY).  THE  DEPTH   OF  EACH
RECTANGLE  CORRESPONDS  TO   THE TEMPERATURE  RANGE.  POINTERS
ON  LATERALLY  ROUNDED  FIGURES  REPRESENT A DERIVED  AVERAGE.
VERTICAL  RELATIONS  ONLY ARE   IMPORTANT  (FERGUSON, 1958),

-------
 lAJ
82.4 28
78.8 26
75 2 24



16 22

fiR O PO
\J O . \y . . c. w
fid. 4 i IP
O t .T- | 1 0
6O.8 £ 16
-^
_/
C -» /> ^f t jt
57.2 
-------
                                    1*5
may not kill trout they produce environmental conditions favorable for the




production of coarse fish.




       One unusual set of data collected on preferred temperatures of rainbow




trout by Garside and Tait (1958) showed that the preferred temperatures were




inversely related to the acclimation temperature.  They state, "Fish cannot




lose heat because they must pass considerable quantities of water through




their respiratory system in order to compensate for the low quantities of



dissolved oxygen contained in waters possessing a higher temperature.  With




the animal passing higher quantities of heated water across their gills the




body temperature of the animal must rise."




Effect ofTemperature on Toxicity




       Effects of artificially induced temperature changes can result in




fish mortalities; as temperature increases the toxiclty of certain




materials increases.




       The Prevention Subcommittee of the Central Water Committee, Ministry




of Health, England, (Anon., 19^9) states, "...increase in temperature also




increases the lethal effect of toxic substances to fish."  For example a




rise in temperature from 8°C (1*-6°F) to l8°C (6V"F) approximately doubled the




toxicity of a low concentration of potassium cyanide.




       The toxicity of chloride concentrations has been shown to be dependent




on temperature.  The temperature has a significant effect on the time of




both initial and final mortalities, the rate of mortality, and the duration

-------
of the mortality for rainbow trout.  It has been postulated that the metabolic




rate of the fish, which affects the rate to which fluoride is toxic to rainbow




trout, is affected by the increased temperature of the fish, Angelovic (1961).




Benefits




       Trembley (i960) concluded that most fish species are attracted to and




invade heated water areas from late September until early June.   Attraction



to heated water has been observed in England, and has been reported frequently




in America.  This adds to the recreational value of localized areas, because




angling can be continued throughout the winter when there may be little or no




fishing in other areas.  Trembley found that one of the disadvantages to




providing winter fishing is that fish leave the heated-water zone in the



hot summer months.




       Another benefit of artificially induced temperature changes is the




production of trout and other cold water fish in the reach downstream from



reservoirs.  Low level penstock discharges from stratified reservoirs often




lowers the temperature in the receiving stream to 12.8°C (55°F)  and it may




not exceed 20°C (68°F) even in summer (Mackenthun et al.,

-------
Summary




1.  Warm water fish can survive temporarily in waters heated artificially




    to 33-9°C (93°F)j even at. 30°C (86°F) coarse fish populations,  such




    as roach, perch, gudgeon, tench and carp,  are reduced.  In cold weather,




    stream temperature should be substantially below  33-9°C (93 F) to




    prevent mortalities when fish move through excessive stream gradients.




2.  Streams supporting cold water non-anadromous fish populations should




    not receive heated effluents that will raise receiving stream tempera-




    tures above 1^.5 C (58 F).  In cold weather, stream temperature should




    be below l4.5°C (58°F) to prevent mortalities.




3.  Sudden changes in temperature can be more harmful to some species o^




    fish than continued  exposure to a higher temperature.




k.  Fish can adapt to higher temperatures faster than to lower temperatures.



5.  The maximum temperature for a given species of fish varies with the fish's




    rate of heating, size, and physiological condition.




6.  Fish may starve at elevated temperatures because of their inability to




    capture food.




T.  Fish seek out a preferred temperature at which they can best survive




    which is several degrees below their lethal temperature.




8.  The toxic effects to fish of certain materials increase with tempera-




    ture.



9.  Certain benefits, including open water winter fishing in otherwise




    ice covered areas and a cold water fisheries downstream from reservoirs,




    can be derived from artificially induced temperature changes.  The

-------
benefits of fish being attracted to heated water in the winter




months may be negligible compared to fish mortalities that may




result when the fish return to the cooler water; lethal tempera-




tures may result from heated discharges in"the summer months.

-------
                            REFERENCES CITED
Agersborg, H. P. K. , 1930.  The influence of temperature  on  fish.   Ecology,
11(1) :  136-1M4-.

Alabaster, J. S., 1962.  The effect of heated effluents on fish.
Internat. Conf. on Water Poll. Resrch., London, 1962.  Air and Water
Poll. 7(6/7):  5^1-563, 1963.  Bio. Abstr. , 14.5(13), Abstr. No. 5^305,
Angelovic, J. W. , W. F. Sigler, and J. M. Neuhold.  Temperature and  fluorosis
in rainbow trout.  Jour. Water Poll. Control Fed., 33:   371-381, Apr.
Anon., 19^9.  Discharge of heated liquids into  streams.  Rep.  of Rivers Poll.
Prevention Sub -Commit tee of the Central Water Committee, Minist. of Health,
England, pp. 69-76.

Anon, 1962.  Heated discharges . .  . their effect on  streams.  Rep. by the
Advisory Committee for the control  of stream temperatures to the Pa. Sanitary
Water Board.  Pa. Dept. Health, Harrisburg, Publ. No.  3, 108 pp.

Brett, J. R. , I960.  Thermal requirements of fish - three decades  of study,
191*0-1970.  Bio. Problems in Water  Poll. , Trans. 1959 Seminar, Robert A.
Taft Sanitary Engng. Center Tech. Rep. W60-3, Cincinnati, pp.  110-117.

Brown, M. E. , 1957.  The physiology of fishes.  Vol.  I.  Metabolism.  Vol. II.
Behavior.  Academic Press.

Cairns, 3., Jr., 1956.  Effect of heat on fish.  Indus. Wastes, 1(5):
180-183.

Downing, K. M. , and C. J. Merkens,  1957.  The influence of temperatures on
the survival of several species of  fish in low  tensions of dissolved oxygen.
Ann. Appl. Bio., ^5(2):  261-267.

Ellis, M. M. , 1914-7.  Temperature and fishes.  U. S. Fish and Wildl. Serv, ,
Fish Leaflet 221.

Falkner, N. W. , and A. H. Houston,  1966.  Some  haematological  responses to
sublethal thermal shock in the goldfish, Carassius auritus L.  Jour. Fish.
Resch. Bd. Can., 23(8):  1109-1120.

-------
                                   50
Ferguson, R. G. , 1958.  The preferred temperature of fish and their mid-
summer distribution in temperate lakes and streams.  Jour. Fish Resch.
Bd. Can., 15:  607-62k.  Spo. Fish. Abstr., Ml), Abstr. No. 2275, 1959-

Fry, F. E. J. , 19^7.  Effects of the environment on animal activity.
Univ. of Toronto Study Bio. Ser. 55:  Publ. Ont. Fish. Resch. Lab.,
68:  1-62.

Fry, F. E. J. , J. S. Hart, and K. F» Walker, 19^6.  Lethal temperature
relations for a sample of young speckled trout, Salvelinus fontinalis,
Univ. Toronto Stud. Bio. Ser. 5^, Publ. Ont. .Fish Resch. Lab., 66:  1-35.

Garside, E. T., and J. S. Tait, 1958.  Preferred temperature of rainbow
trout (Sjalmo gairdrieri Richardson) and its unusual relationship to acclima-
tion temperature.  Can. Jour. Zool., 36:  563.  Spo. Fish. Abstr., 3(3) }
Abstr. No. 1676, 1958.

Heinicke, E. A., and A. H. Houston, 1965.  Effect of thermal acclimation
and sublethal heat shock upon ionic regulation in the goldfish Carassius
auratus L. , Jour. Fish Resch. Bd. Can., 22(6):  1^55-1476.

Jones, J. R. E., 1964.  Thermal pollution:  the effect of heated effluents.
Fish and River Pollution, Chap. 13:  153-168,  Butterworth and Co., Ltd.,
Washington, D. C.

Mackenthun, K. M. , W. M. Ingraza, and R. Porges, 196U.  Limnological aspects
of recreational lakes.  U. S. Dept. of Health, Educ. and Welfare, GPO
176 pp.

Prosser, C. L., 1955*  Physiological variations in animals.  Bio. Rev.,
30(3)i  229-262.

Swift, D. R., 1965.  Effect of temperature on mortality and rate of
development of the eggs of the Windermere Char (Salvelinus alpinus ) .
J. Fish. Res. Bd. Canada,
Tarzwell, C. M. , 1957.  Water quality criteria for aquatic life.  In:
Biological Problems in Water Pollution.  Robert A. Taft Sanit. Engng.
Center, Cincinnati, Ohio, pp. 2U6-272.

Threinen, C. W. , 1958.  Cause of mortality of a midsummer plant of rainbow
trout in a southern Wisconsin lake, with notes on acclimation and lethal
temperatures.  Prog. Fish. Cult., 20:  27.  Spo. Fish. Abstr., 3(2), Abstr.
No. 15^3, 1958.

-------
                                   5L
Trembley, F. J,, I960.  Research project on effects of condenser dis-
charge water on aquatic life, progress report, 1956 to 1959.  Inst.
of Resch., Lehigh Univ., 15^ pp.  Water Poll. Abstr., 3^(ll), Abstr.
No. 2157, 1961.

Wells, M. M., 191k.  Resistance and reactions of fishes to temperature,
Trans. Illinois Acad. Sci., J:  ^8-59.

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               IV.  MARINE, ESTUARIHE AND ANADROMOUS FISHES








Introduction




       Because a growing demand for electricity supplied by steam or nuclear



generators has increased the need for ample water, both for cooling and




thermal waste assimilation, larger areas of fresh and marine waters will




receive significant temperature elevations (Mihursky and Kennedy, 1967).



Future emphasis will be away from the use of inland waters, toward the uti-




lization of estuarine and marine resources (Naylor, 1965).  The protection




of fishes in the estuarine and marine areas, as well as anadromous fishes that



must move through areas of heated water is becoming an increasing problem.




General Temperature Effects




       Researchers,, studying the effects of fluctuating temperatures on




fishes, have taken two approaches:  one method of study is to observe the



reactions of fishes in their natural habitat; the other method is to remove




representative samples of fish from the natural habitat to the laboratory




and observe them under selected test conditions.  Both methods of study have




been used equally well.




Physiology



       The physiology of fishes is directly affected by temperature.  Fishes




are classed as poikilothermus or animals whose body temperature follows

-------
                                    53
changes in environmental temperatures rapidly and precisely.  In such animals



the factors favoring heat loss tend to equal the factors producing body heat



and the "body temperature approaches environmental temperature (Prosser, et al.,



1950; Kinne, 1963).  In a majority of fishes the body temperature differs from



that of the surrounding water by only 0.5-1.0°C (0.9-1.8°F) (Nikolsky, 1963).



Therefore, one of the fundamental requirements of fishes is that the external



temperature must "be best suited to internal tissues (Brett, 1956).  Cells



exposed to heat undergo an increase in the viscosity of protoplasm.  This



increase is reversible to a point beyond which heat death occurs (Gunter, 1957).



Various functions of an organism, such as reproduction, locomotion and growth



may have different temperature ranges and these ranges should be known in



order to evaluate the effects of temperature on that organism (Kinne, 1963).



For example, according to Kinne (1963) and Naylor (1965), marine and brackish



water organisms may increase or decrease osmotic regulation as a function of



temperature.



       Temperature fluctuations act on an organism in a variety of ways:



a) metabolic rates are changed, b) reproduction is affected, c) distribution



may be increased or decreased and d) tolerance limits are widened or narrowed.



Metabolism



       Rates of metabolism and activity increase with increasing temperatures



over most of the tolerated temperature range and then often cease suddenly



near the upper lethal temperature.  Such rates vary with different species,

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processes and levels or ranges of temperature and may be modified "by salinity




and oxygen factors (Kinne, 1963).  Changes in metabolic rates, because of




temperature fluctuations, may be signaling factors for spawning or migra-



tion (Nikolsky, 1963).




Reproduction




       The effects of temperature on reproduction in many animals are con-



fined to narrower ranges than the majority of functions (Kinne, 1963 and




Gunter, 1957).  Most marine animals have restricted temperatures for breeding.




Rising temperatures in the spring induce development of the gonads and actual



spawning takes place when a certain temperature level is reached, which varies




for different species.  Some fish spawn on a drop in temperature while others



respond to a rise in temperature (Gunter, 1957).  Because of narrow breeding




requirements the survival of a species in heated waters does not preclude




the possibility that the species may be prevented from breeding and may exist



in an area by continued recruitment from outside (Naylor, 1965).




Development




       Temperature changes affect fish development in several ways.  Meristic



characters and shape may be changed as well as embryonic development.  Low




temperatures slow down development and in some cases many marine and brackish




water animals attain a larger final size because of their slow, long continued



growth rather than rapid growth (Kinne, 1963).

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                                    55
Distribution and Ecology




       Since temperature is the most important single factor governing the




occurrence and behavior of life, it not only affects the distribution of a




single species, it may also modify the species composition of a community




or an ecosystem (Gunter, 1957 and KLnne, 1963).  Tropical and subtropical




fishes are more stenothermal (tolerate a narrow range of temperatures) than




those of boreal and higher latitudes and marine forms are more stenothermal




than fresh water ones (Nikolsky, 1963).  In his publication on temperature




effects on marine organisms Naylor (1965) noted that estuarine foims were




more tolerant of heated effluents than marine forms and littoral species,




and concluded that some coldwater stenothermal forms may be eliminated by




heated discharges and some eurythermal (tolerate a wide range of temperatures)




species may be increased.  He also noted that temperature effects seem to




be more pronounced in enclosed areas of estuaries and bays, while heat




effects in open estuaries are least striking.  In tropical areas, species




live close to their thermal limits and effects of heated effluents are more




pronounced, while in northern (Arctic) areas species may be 13 to 16 C




(23.^ to 28.8 F) below their death temperatures and may not be as severely




affected.




       By testing species in the laboratory Brett (1956) noted that a slow




rate of decrease in environmental temperature is of greater importance for

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                                    56
maintaining life than a slow rate of increase.  Lethal cold can be more




important than lethal heat as a factor limiting the distribution of marine




fish and as a hazard to some in their native habitats (Doudoroff, 1957).



Acclimation




       The capacity to acclimate depends on the genetic background, en-




vironmental history and present physiological condition and age of the




organism involved (Kinne, 1963).  For example, the resistance of animals




to cold is much more variable than resistance to heat and resistance to cold




varies with size, smaller fish resisting best (Gunter, 1957).



       Acclimation to different temperatures may involve changes in orienta-




tion, migration, and other behavioral aspects such as territorialism as




well as biological rhythms (Kinne, 1963).  In his experiments with marine




fishes, Doudoroff (1957) noted that acclimatization to heat may be acquired




very rapidly, the speed varying with heat.  Also, brief or intermittent




exposure to high temperatures can result in markedly increased resistance




to heat which is not readily lost on subsequent exposure to low temperatures.




However, it is the rapidity of the onset of low temperatures that probably




causes death, outstripping the ability of fish to acclimate and resulting




in greater mortalities that are due to cold in nature (Brett, I960).  Deaths



resulting from the inability of fish to rapidly acclimate to lowering temp-




eratures have been reported by Gunter and Hildebrand (1951) and Galloway (1951).

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                                    57
According to Kinne (1963) acclimation to low temperature usually tends to



shift the lower thermal limits downward and acclimation to high temperatures



tends to shift the upper limits upward.  As a result the ability to acclimate



affects the temperature ranges that a fish can tolerate.



Tolerance



       The temperature tolerance of fish varies with their development,



area of distribution and physiology.  As noted earlier, estuarine forms are



more tolerant of heated effluents than marine forms and littoral species



(Naylor, 1965).  However, Kinne (1963) reports that in general, the total



range of temperature tolerated in the state of active life is smallest in



marine forms and largest in brackish and fresh-water forms.  Gradual changes



are tolerated much better than sharp changes.  Some species can stand a



gradual change up to 30 or 35°C (86 or 95°F), but at the upper extreme,



many organisms are killed by temperatures not far above those to which they



are accustomed.



                               MARINE  FISHES*



       Marine fishes that inhabit the shore line areas, estuaries or bays



are most often affected by temperature changes.  The problem of thermal



shock to pelagic (living or occurring in the open ocean) life histories



is extremely critical in marine environments (Mihursky and Kennedy, 196?).
       The effects of salinity and temperature on the eggs of Pacific Cod



 (Gadus macrocephalus) were studied by Forrester (196*0 and, Forrester and








 *  In this  report the term marine includes  estuarine  species.

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Alderdice (1966).  In the study by Forrester, eggs were held to completion



of hatching in various combinations of salinity and temperature.  Maximum



hatching success was in the vicinity of 19 parts per thousand (ppt) salinity



and 5 C (Ul.O F).  Forrester and Alderdice observed that the relationship



between rate of development of cod eggs and water temperature was linear at



temperatures of 5-ll°C (^1-51.8°F).  Time to 50 percent hatching ranged from



8.5 days at 11°C (51.8°F) to 1? days at 5°C (Ul°F).  Most successful hatching



occurred at the lower temperature.  One of the most noted variables in the



study on eggs of the American smelt (Osmerus mordax) in Maine was the large



increases in mortality during extreme fluctuations in daily water temperature



of as much as 7°C (12.6°F) as observed by Rothschild (1961).  Striped bass



e£&s (ROCCUS saxatilis) were found to survive in constant fluctuations of



water temperatures ranging from 12.8-23.9°C (55-75°F) Albrecht (196^).  The



tolerance of eggs of four marine fishes was studied by Hubbs (1965).



California killifish (Fundulus parvipinnis), topsmelt (Atherinops affinis),



California grunion (Leuresthes tenuis) and mussel blenny (Hjvpsoblennius sp.)



were incubated at a variety of temperatures.  Larvae successfully hatched



at temperatures between l6.6°C and 28.5°C (6l.9-83.1°F), 12.8 (-)°C and 26.8°C



(55-80.1°F), 1U.8° C and 26.8°C (58.6-80.1°F) and 12.0 (-) and 26.8 (-f)°C



(53.6(-)-80.l(+)°F) respectively (Table _5_).



Young



       Larvae of some marine fish are pelagic in the early part of their life



history, and temperature of the surrounding water determines the rate of

-------
                        59

Table 5 -  Temperature ranges reported for the hatching of
          eggs from various species of marine and anadromous
          fish.
Species
Roccus
saxatilis
Fundulus
paryipinnis
Atherinops
af finis
Leuresthes
tenuis
Hypsoblennius
sp.
Petromyzon
marinus
n
Oncorhynchus
nerka
0. tshawytscha
ti ti
n n
All Salmon
Lower
Temp, (c)
12.8
16.6

12.8
1^.8
12.0(-)

15.0
15.6

U.lf-5.8
5.8
9.H
5.6
5.8
Upper
Temp. (C)
23-9
28.5

26.8
26.8
26.8(+)

25.0
21.1

12.8-11*. 2
lU.2
llf.U
Ik.k
12.8
Remarks Source
Survived Constant Albrecht, ~L
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                                    60
development from a pelagic form to an actively swimming form.  The rate of



development of the lemon sole (parophrys vetulus) determines the number of



young that reach the nursery grounds in Hecate Strait, British Columbia



(Ketchen, 1956).  Small annual differences in the temperature of sea water



produce marked differences in the duration of the pelagic stage.  Below



average temperatures result in the larvae being carried by the currents



for a longer period of time and more larvae are deposited on the nursery



ground.  Thus, temperature may govern the strength of a year class.  A



temperature of 6.2°C (43.1°F) seems to produce the best deposition of



larvae.  Increases in temperature increase the rate and shorten the time



of development of herring (Clupea pallasi).  A temperature increase from



^.4-10.7°C (ifO-51.1°F) shortened development time from Uo to 11 days



(Blaxter, 1963).



       In the estuarine environment, fishes are more susceptible to heat



changes.  However, as noted in the general discussion, fish in estuarine



waters seem to tolerate a wider range of temperatures.  Striped bass



fingerlings (Roccus saxatilis) were able to tolerate 35°C (95°F) in lab-



oratory tests (Talbot, 1966).  According to Talbot (1966), Merriman (191*!)



studied the striped bass of the Atlantic coast and found the maximum



temperature in the New England area to be 25°C to 27°C (77.0 to 80.6°F)



with fish kills occurring at these temperatures.  Juvenile striped bass



have survived transfer between salt and fresh-water at temperatures in the

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                                    61
range of 12.8-21.1°C (55-70°F) "but they are not tolerant to changes from



fresh-water to saltwater at ?.2°C (U5°P) (Tagatz, 196l).  Striped "bass



acclimated at h.k°C (UO°F) and tested for eight hours with increases in



temperature of 2.3°C (3.5°F) at one hour intervals, had a median lethal



dose (LD  ) of 23.9°C (75°F), (Trembley, I960).



       The larvae of Atlantic Menhaden (Brevoortia tyrannus) were able to



survive longer when acclimated at cooler temperatures, than when acclimated



to warmer temperatures.  Acclimation temperature was more important to larval



survival at test temperatures below 5.0°C (ifl°F) than at 5.0°C (Ul°F) and



above.  Larvae acclimated at 7 and 10°C (hk.6 and 50°F) survived over twice



as long at ^.5°C (1*0.1°F) as those acclimated at 12.5 or 15°C (5^.5 or 59°F)»



(Lewis, 1965).  The effects of salinity on temperature tolerances were checked



by Lewis (1966), who found a temperature of 6°C (42.8°F) and below with zero



ppt salinity lowered larval survival time to only a few hours.  At a salinity



of 5-30 ppt and a temperature of U°C (39.2°F) larval survival was good.  Lower



and upper limits of salinity tolerance were increased with increasing tem-



perature .



       The relation of menhaden (jB. tyrannus) to estuaries was studied by



Reintjes and Pacheco (1966).  A water temperature of 3°C (37.^OF) may be



critical to larval survival.  Larval menhaden can suffer mass mortalities



when water temperatures fall below 3°C (37.^°F) for several days or chill



rapidly.  The matter of chill seems to be very important to estuarine fishes.

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                                    62
Doudoroff (19^2) studied the resistance and acclimation of marine fishes



to temperature changes.  Young greenfish (Girella nigricans) were used in



his tests.  Heat resistance was gained rapidly and lost slowly.  Resistance



to chilling was lost slowly on warming and acquired no more rapidly on



cooling.  The changes of resistance to heat and to cold were found to be



more or less independent and distinct phenomena.  Acclimation to cold is



slow, to heat, fairly fast.  For these reasons injury by chilling is no



less important as a possible limiting factor in the distribution of marine



fishes than heat injury.



       More active fishes are able to avoid harmful temperatures and to exercise



selection in experimental gradient (Doudoroff, 1938).  In the case of pelagic



marine larvae, circumstances may dictate their survival.  A mortality of



marine fish larvae was noted in an area off Georges Bank.  Currents carried the



larvae from cold 7.8°C (U6°F) water into warm layers of 20°C (68°F) water and



large mortalities resulted (Colton, 1959).



       A study of the effect of extreme temperatures on herring larvae (C.



harengus) revealed an upper lethal temperature 22.0 to 2U.O°C (?1.6-75.2°F)



and a lower lethal temperature of -0.75 to -1.8°C (30.6-29.1°F).  Larvae



were 6-8 millimeters long and acclimated to temperatures between 7.5 and



15.5°C (58.2-72.9°F) (Blaxter, I960).  Young topsmelt (Atherinops affinis)



acclimated to temperatures of 20 C (68 j?) had an upper hQ hour median tolerance



limit (TLm) of 31.8°C (89.1°F) and a lower U8 hour TLm of 10.1°C (50.1°F),



(Doudoroff, 19^5).  Young greenfish (Girella nigricans) acclimated to

-------
                                    63
temperatures of 12-28 C (53.6-82.U F) exhibited a lower U8 hour TL  of U.I
                                                                  m
                                 ft



to 13°C (39.lt-55.lft) and a upper TL  of 28.7 to 31.5°C (8l.<9-88.8°F),
                                    m


(Doudoroff, 19^2).  Temperature ranges reported for young fish are listed



in Table 6 .



Adults



       Mult fish are usually able to select their preferred temperatures,



unless they are trapped in shallow waters or forced to migrate through



heated or chilled areas.  Fish kills have been reported in areas of shallow



water.  Atlantic round herring (Etrumeus sadina) and chum mackerel (Scomber



colias) were observed dead and dying after several days of cold weather had



dropped water temperatures in Pamlico Sound, North Carolina, to 5.2 C (Ul.U F),



(Wells, Wells and Gray, 196l).  The effects of winter water conditions were



also observed by Schwartz (196U).  He noted that most fish sank when killed



by lowered temperatures and would probably not be observed.  In the area of



high temperatures, alewives (Alosa pseudoharengus) died of heat shock after



being herded into water of 26.7 - 32.2°C (80-90°F).  The same species showed



no effects when they entered a lagoon with 22.8°C (73°F) water (Trembley, I960)



Herring, in nature, have been found in almost all temperature ranges permitted



by their resistance to temperature extremes.  Herring at the appropriate



season have an upper lethal temperature of 19.5 to 21.2°C (67.1 to 70.1 F)



depending upon size and can survive short exposure 'to temperatures below



-1.0°C (30.2°F), (Brawn, 1960).

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               Table 6 -  Temperature ranges reported for young
                          marine and anadromous fish.
Species
0. tshawytscha

0. kisutch

0. nerka

0. gorbuscha
0. keta

Clupea
harengus
Atherinops
af finis
Girella
Acclimation Lower Lethal Upper Lethal
Temp.(c) Temp.(C) Temp.(c) Remarks Source
23 7-k
20 25.1
23 6.k
20 25.0
23 6.7
20 2k.k
23-9
23 7.3
20 23.8

7.5-15.5 -0.75to-1.8 22.0-2*f.O

20 10.1 31.8
Approx.
Approx.
Approx.
Approx.
n
Approx.
Approx.
Approx.



k days Brett,
1952
7 days "
14- days "
7 "
k days "
7 days "
7 days
k days "
7 days "

Blaxter,
I960

U8 hr TL Doudorof
m 19U2b
nigricans
12-28
U.l-13
28.7-31.5
U8 hr TL
                                                                   m
Doudoroff,

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                                    65
       Adult white perch (Roccus americana) acclimated to k.k°C (UO°F) and



tested for 8 hours with increases of 2.0°C (3.5°F) every hour exhibited a



median lethal dose (LD  ) of 27.8°C (82.0°F), (Trembley, I960).  In a flow-



ing water test with heat increases of 1.1°C (2°F) per hour saltwater killifish



(Fundulus heteroclitus) acclimated at 7«2°C (U5°F) had a LD Q of 37°C (99°?)



(Trembley, 196l).  Adult California killifish (P. parvipinnis) were tested



by Doudoroff (19^5).  At acclimation temperatures of lU-28°C (57.2-82.^°F)



the upper TL fs, for k8 hour tests, were 32.3 to 36.5°C (90.1-97.7°F).  The



lower U8 hour TL  was 30°C (86°F) for fish acclimated at 20°C (68°F).  Striped
                m


bass occur in wide ranges of temperatures in the estuary (Talbot, 1966).  They



will spawn between lU.U°C (58°F) and 21.1°C (70°F).  Ranges of temperature



tolerated are 6.0-7.5°C (U2.8-U5.5°F) to 25-27°C (77-80°F).  Temperature



tolerances of three- marine fishes have been determined by Hoff and Westman



(1966).  The common silverside (Menidia menidia) acclimated at temperatures



ranging from 7-28°C (M*.6-82.U°F) had an upper kQ hour TL  range of 22.5 to 32.5°C



(72.3-90.3°F) and a lower range of 1.5 to 8.7°C (3U.8-U7.8°F).  Winter flounder



(Pseudo pleuronectes) acclimated at temperatures of 21-28 C (69.8-82.U- F)



had a lower U8 hour TL  range of 1.0-5.^°C (33.8-^1.6°F).   Flounder acclimated
                      m


at temperatures from 7-28°C (kk.6-d2.k°F) had an upper range of 22-29°C



(71.6-8U.2°F).  Northern swellfish (Spheroides maculatus)  were acclimated at



temperatures of 1^-28°C (57.2-82.^°F) and had a lower 1*8 hour TL  of 8.^-13°C
                                                                m


(J±7.1-55.1|0F).  Fish acclimated at temperatures of 10-28°C (50-82.k°F) had



TL 's of 28.2-33.0°C (82.9-90.k°Y).  Temperature ranges for adults are listed



in Table 7 .

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Table 7 - Temperature ranges reported for adult
          marine and anadromous fish.
Species
Clupea harengus
Pundulus
parvlpinnis
it
Roccus saxatilis

Menidia menidia

Pseudo pleuronectes

Spheroides
maculatus

Mult Salmon
Acclimation
Temp.(C)

1U-28
20

7-28
21-28
7-28

1H-28
10-28

Lower Lethal
Temp.(C)
- 1.0

30
6.0-7.5
1.5-8.7
1.0-5.k

8.U-13
0.0
Upper Lethal
Temp.(c)
19.5-21.2
32.3-36.5

25-27
22.5-32.5
22-29

28.2-33.0
26.7
Remarks

U8 hr TL
m
ii it it
Tolerated in
Estuary
U8 hr TL
m
1*8 hr TL
m
it it n

it it ii
it ii ii
Survival Temp.
Source
Brawn, I960
Doudoroff, 19U2b
it ti
Talbot, 1966
Hoff & Westman,
1966
it
it

it
it
Anon. , 1966
                                                                              CTN

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                                    67
       In a statement about the observed presence or absence of fish in



heated areas, Trembley (1960) reported that fish may be eliminated from



heated zones during warm months, but may congregate in heated areas during



winter months.



                             ANADROMOUS FISHES



       Anadromous fish are unique in their life histories.  Eggs incubate



in fresh water and the resulting young spend a period of a few months to



several years in fresh water, then migrate to saltwater, where they grow



into adults.  As adults the fish mature in salt water and return to fresh



water to spawn.  During their life cycle, anadromous fish are subjected to



various stresses such as salinity (osmotic) change, physical change, predators,



and temperature (Brett, 1957)*



Eggs



       Investigations have shown that thermal requirements in the very early



stages are more exacting than in the adult (Brett, 1956).  Eggs of the sea



lamprey (petromyzon marimis) require the most exacting thermal levels



(McCauley, 1963).  The range of constant temperatures necessary for successful



hatching is narrow, being 15-25°C (59-77°F).  The range could be extended to



12-26°C (53.6-78.8°F) if the eggs were able to develop to the head stage



before they were subjected to increased temperatures.  Similar results were



noted by Piavis (1961) who reared sea lamprey eggs at low constant temperatures and



was unable to grow viable burrowing larvae at any temperature below 15.6 C



(60r) or above 21.1 C (70°F).  Constant temperatures were used in the

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                                     68
incubation of chinook and sockeye salmon eggs (Oncorhynchus tshawytscha and



0. nerka) (Combs, 1965).  Chinook salmon eggs which had developed to the 128



cell stage could tolerate 1.7°C (35°F) water for the remainder of the incuba-



tion period.  Sockeye salmon eggs were less resistant to high temperatures



and more resistant to cold temperatures.  Their lower threshold temperatures



for normal development were k.k-5.8°C (kQ-k2.5°F) and upper threshold tem-



peratures were 12.8-lU.2°C (55-57.5°F).  A lower threshold for chinook eggs



was established at 5,8°C (U2.5°F) and an upper threshold at 1*1.2°C (57.5°F),



(Combs and Burrows, 1957).  Mortalities occurred when eggs were incubated



above or below these temperatures.



       Hayes, in 19^9j subjected salmonid eggs to extreme temperatures and



noted that certain tissues will exhibit cell multiplication without differentia-



tion.  Salmon embryos were incubated by Hayes, Pelluet and Gorhan (1953) in



temperatures within the limits for survival.  Hatching of the embryos tended



to appear precociously at low temperatures.



       According to Johnson and.Brice (1953) reservoir water could be used for



incubation when the daily mean temperatures were below 12.2°C (5U°F).  Chinook



eggs incubated over 15.6 C (60 F) suffered excessive mortality.  Results from



laboratory tests conducted by Olson and Foster (1957) and Nakatani and Foster



(1966) were slightly higher.  They reported that chinook eggs, especially in



cold water, could begin incubation at temperatures as high as 16.1 C (6l F),



without significant loss.  Seymour (1956) reported that young chinook eggs

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                                     69
should "be reared at temperatures ranging from 9.^°C to 1U.U°C  (1*9 to 58°F)




for best results.  Abnormal fry and the hatching period were increased by




higher and lower temperatures.  A slightly wider range of temperatures 5.6-




ik.k C (U2-58°F) was suggested by Leitritz (1962).  As a general range for




all Pacific salmon eggs LT  's were reported at 2.5°C (36.5°F) and l6.0°C




(60.8 F) but less than normal survival was noted below 5.8°C (U2.5 F) and



above 12.8°C (55°F), (Anon, 1966).




       In the natural environment, McNeil (1966) studied the effect of low



temperatures in the spawning beds of pink (£. gorbuscha) and chum (0. keta)




salmon and determined that freezing was important only when the maximum day-




time temperatures remained below 0°C (32°F) for at least two days.  Eggs and




larvae of pinks and chum are able to survive at low temperatures and high




salinities (Rockwell, 1956).




       In a summary of the significance of temperatures on salmon egg in-




cubation (Anon, 1966) the following points were emphasized; a) the effects




of temperature vary with many things including species and race, b) mortality




attributable to temperature is also a function of duration of exposure,



c) temperature during the initial incubation period is critical and, d) if




the initial incubation temperatures are below 5.6°C (U2°F) or above 12.8°C




(55 F) less than normal survival can be expected.  Temperature ranges re-




ported for the hatching of eggs are tabulated in Table _5__.

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                                     70
Young



       Young of anadromous fish, especially salmon, spend much of their life



in fresh-water and most research on them pertains to this environment.  De-



termination of tolerance limits for different species of fish may be very



difficult and time consuming.  Brett (i960) has reported the ultimate upper



lethal levels can differ between species by as much as 17°C (31°F).



       Five species of Pacific Coast salmon were tested by Brett (1952) for



their temperature tolerances.  The five species of salmon tested were spring



or Chinook (Oncorhynchus tschawytscha), silver (0. kisutch), pink (0.



gorbuscha), sockeye (0. nerka) and chum (0. keta).  Fish used in the tests



were less than one year of age with an average length of U05 centimeters



and an average weight of 1 gram.  The maximum acclimation temperature was



2k C (75.2°F).  Springs were reported to be very active and good feeders at



2k C (75.2 F) but growth was poor.  Pinks, sockeyes, chum and cohos were all



intolerant to 2^°C (75.2 F) water.  Of the species tested, springs and cohos



were most tolerant to prolonged high temperatures, sockeyes intermediate,



and pink and chum least tolerant.  The upper lethal temperatures were as



follows:  spring - 25-l°C (77.2°F); coho - 25.0°C (77.0°F); sockeye - 2k.k°C



(76.0°F); pink - 23.9°C (75.0°F); and chum - 23.8°C (jk.B0?).   Acclimation



temperatures for all species were 10-20°C (50-68°F).  The lower lethal tem-



peratures for the highest acclimation of 23°C (73.U°F) were spring, 7.U°C



     °F); coho, 6.kQC (U3.5°F); sockeye, 6.7°C (^.0°F); and chum,  7.3°C



   .1°F).  For all species the region of greatest preference was 12-l4°C

-------
                                    71
(53-6-57.2 F), (Table 6 ).  In a report on the Columbia River salmon (Anon.,



1956) lethal tolerances for 50 percent of the juveniles tested were listed



as 0.0°C (32.0°F) and 25.1°C (77.2°F).  However, poor growth was reported



for temperatures below k.k°C (kO°F) and above l8.3°C (65°F).  In contrast



with the above results Kerr (1953) tested young chinook salmon and reported



a maximum temperature of 26.7°C (80°F) tolerated by them.   He also reported



them able to tolerate a rise in temperature of 9°C (l6°F)  in the cooling



water from a steam generated electric plant.



       Burrows (1963) suggests that a range in temperature for maximum pro-



ducitivity in fingerlings should be between 10-15.6 C (50-60°F).  His research



indicated that to attain maximum productivity the water temperature must not



only remain within the tolerance level of the fingerling but in species with



more than a minimum stay in fresh-water, the temperature must reach that



necessary for optimum growth level as well.



       Tests designed to reveal the effects of temperature on the physiology



of young salmon showed internal temperatures of smolts reached equilibrium



with the external environment in 3-5 minutes (Harvey, 196U).  Effects of



temperature on fin ray and vertebrate counts were checked by Seymour (1959)



who noted that the average number of vertebrae per lot of fish was less for



lots reared at temperatures in the middle portion of the 3«9-16.7 C (39 F-



62°F) range than for lots reared at either extreme of the  range.

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                                    72
       Several researchers have checked the effects of temperature on



swimming speed and metabolism.  Optimum cruising speeds occurred at 15 C



(59 F) for young sockeye and 20°C (68°F) for young coho (Brett, Hollands



and Alderdice, 1958).  When young sockeye salmon swimming at a speed of 1



foot per second were subjected to a temperature change of 10-15°C (l8°P



to 27 F) their metabolic rate increased by more than 50 percent (Anon.,



1962).



       Migration of young fish from fresh-water to saltwater may subject



them to wide ranges of temperature.  Effects of temperature on pink salmon



(0. gorbuscha) were studied by Sheridan (I960 and 1961).  He reported an



interaction between air temperature and snow that may cause fry to migrate



to sea at unfavorable times.  There is presumably one "best" time for fry



to enter saltwater and the normal time of seaward migration may be best for



food supplies and saltwater temperatures or other unknown factors.  In addi-



tion to the temperature and stream flow in the river, the  temperature and



salinity of the marine environment during early life are also very important



(Vernon, 1958).



       During their stay in fresh-water young fish are subjected to diseases



that may or may not be influenced by temperature.  Both the literature and



research show the effects of some diseases are increased by temperature in-



creases.  There is one disease, however, that may be reduced by increased



temperature (Ordal ani Pacha, 1963).  The myxobacterium Cytophaga psychrophila



is a disease of salmon fingerlings in low temperature water.  Losses due to

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                                     73
this disease can "be reduced by increasing temperatures above 6.1 C (^3 F)



in the spring.  Water temperature remaining relatively stable either above



or below the optimum range for extended periods is conducive to disease



development which may result in reduction of fingerlings produced (Burrows,




1963).



Adults



       Mult anadromous fishes spend most of their life cycle in the marine



area and enter fresh-water only to migrate up a stream to spawn.  During



their migration salmon do not take in food after reaching the estuary and



heading upstream.  High water temperatures increase their metabolic rate



and may result in fuel depletion before the fish can spawn (Anon., 1962).



Temperature was listed as one of the factors affecting timing of spawning



runs of pink salmon (0. gorbuscha) studied by Sheridan (1960).  For pinks,



a stream temperature near 10°C (50°F) seemed to be best.  Adult salmon



have been reported to survive 0.0°C (32.0°F) to 26.7°C (80°F) but spawning



effectiveness may be reduced below 7.2°C (^5°F) and above 15.6°C (60°F)



(Anon., 1966), (Table _7_).



       The migration of fish has been hampered by unfavorable temperatures.



Brett (1957) noticed the curtailment of the migration of sockeye salmon



through lakes in the spring and Major and Mighell (1966) concluded that



rising or stable temperatures above 21.1°C (70°F) tended to block the entry



of migrating fish from the Columbia River to the Okanogan River.  A study,



by Massmann (1957), of the relationship of water temperature and shad catches

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in the York River for a three year period showed that greatest catches-per-unit



effort were made at water temperatures of 7.2°C (U5°F) to 15°C (59°F).  Below



a water temperature of k-.h C (ho jf) the fish stopped migrating and no catches



were made.  The death of migrating Atlantic salmon (Salmo salar) in Nova Scotia,



due to low water and high temperatures was recorded "by Huntsman (19^2).  Fresh



run grilse died at about 29.5°C (85.1°F) and acclimated grilse at about 30.5°C



(86.8°F).



       As noted earlier with juvenile salmon, temperature increases usually



result in an increase in disease which lowers the surviving numbers of spawn-



ing fish.

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                                    75
Summary



       Temperature fluctuations affect the metabolism, reproduction, dis-



tribution, ecology and tolerance of fishes.  The effect of a fluctuation



depends on the species of fish, the stage in the life history of the fish,



the rate of decrease or increase in temperature and the amount of thermal



fluctuation.  In the marine environment temperature changes are most important



in enclosed areas such as the estuaries and bays as opposed to open areas.



Tolerance to temperature fluctuations is least in marine forms and greatest



in estuarine and fresh-water forms.



       Pelagic forms are most susceptible to temperature fluctuations since



they are dependent upon water currents for much of their movement.  Adult



fish are usually able to select their preferred temperature gradient unless



trapped in shallow or enclosed areas or forced to migrate through heated or



chilled areas.  Most fish have restricted ranges of temperature within which



they can reproduce successfully.  Larval development also requires narrow



ranges of temperature.  For these reasons a fish population may exist in a



heated area only by continued recruitment from the outside.  In such areas



fish may be absent during warm summer months and present in cold winter



months.  In some areas populations of widely tolerant species may replace



stenothermal species.



       Increasing temperatures may block the migrations of anadromous fish



and increase the effects of diseases on those fish.  However, there are reports

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                                   76
of temperature increases reducing losses of young salmon from disease and




increasing survival of eggs.




       Cold is as important to fish populations as heat because of the




inability of fish to acclimate quickly to rapid decreases in temperature.




Thus, in some areas fish populations may be limited by decreases as well as




increases in temperature.

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                                    77

                             REFERENCES CITED
Albrecht, A. B., 196^.  Some observations on factors associated with survival
of striped bass eggs and larvae.  California Fish & Game Jour., 50:  2,
100-113.

Anon., 1962.  Annual report of the Fisheries Research Board of Canada,
1961-62, for the Fiscal year ended March 31, 1962, 206 pp., Water Poll.
Abstr. (Brit.), 36, 889, 1963.

Anon., 1966.  Columbia River water temperature conditions and research
requirements.  Water Supply and Water Poll. Control Subcommittee, Task Force
on Water Temperature Evaluation, Columbia Basin Interagency Committee,
v-i to v-3.

Blaxter, J. H. S., I960.  The effect of extremes of temperature on herring
larvae.  Jour. Mar. Bio. Assoc. Unit. King., 39, 605-60&.

Blaxter, J. H. S., 1963«  The behavior and physiology of herring and other
clupeids.  In:  Advances in Marine Biology, F. S. Russel (ed.), Academic Press,
New York, 261-393-

Brawn, V. M., I960.  Temperature tolerance of unacclimated herring (Clupea
harengus L.), Jour. Fish. Res. Bd. Canada, 17(5):  721-723.

Brett, J. R., 1952.  Temperature tolerance in young Pacific salmon, genus
Oncorhynchus.  Jar. Fish. Resrch. Bd. Can., 9(6):  265-323.

Brett, J. R., 1956.  Some principles in the thermal requirements of fishes.
Quarterly Rev. of Bio., 31(2):  75-87.

Brett, J. R., 1957.  Salmon research and hydroelectric power development.
Jour. Fish. Resrch. Bd. Can., Bull. No. 114, 26 pp.

Brett, J. R., I960.  Thermal requirements of fish - three decades of study,
19i4O-1970.  Bio. Problems in Water Poll., Trans. 1959 Seminar, Robert A.
Taft Sanitary Engng. Center Tech. Rep. W60-3, Cincinnati, pp. 110-117.

Brett, J. R., Hollands, M., and Alderdice, D. F.,  1958.  The effect of
temperature on the cruising speed of young sockeye and coho salmon.  Jour.
Fish. Resrch. Bd. Can., 1500:  587-605.

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                                    78
Burrows, R. E., 1963*  Water Temperature requirements for maximum productivity
of salmon.  In:  Water temperature influences, effects and control.  Proc.
of the Twelfth Pacific Northwest Symposium on Water Poll. Resrch., Pacific
Northwest Water Laboratory, U. S. Public Health Service, Corvallis, Oregon,
pp. 29-38.

Colton, J. B., 1959»  A field observation of mortality of marine fish larvae
due to warming.  Limnol. and Oceanogr., ^(2):  219-222.

Combs, B. D., 1965.  Effect of temperature on development of salmon eggs.
Prog. Fish. Cult., 27(3):  13^-137.

Combs, B. D.  and R. E. Burrows, 1957.  Threshold temperatures for the normal
development of chinook salmon eggs.  Prog. Fish. Cult., 19(l):  3-6.

Doudoroff, P., 1938.  Reactions of marine fishes to temperature gradients.
Biol. Bull.,  75:  U9U-509.

Doudoroff, P., 19^2.  The resistance and acclimation of marine fishes to
temperature changes.  I.  Experiments with Girella nigricans (Ayres).
Bio. Bull., 83:  219-2kk.

Doudoroff, P., 19^5.  The resistance and acclimatization of marine fishes to
temperature changes.  II.  Experiments with Fundulus and Atherinops.  Bio. Bull.,
88(2):  19^-206.

Doudoroff, P., 1957.  Water quality requirements of fishes and effects of
toxic substances.  In:  The Physiology of Fishes (M. E.Brown, editor), 2:
403-U30, Academic Press, Inc., New York, 503 p.

Forrester, C. R., 196U.  Laboratory observations of embryonic development and
larvae of the pacific cod (Gadus macrocephalus Tilesius).  Jour. Fish. Resrch.
Bd. Can., 2l(l):  9-16.

Forrester, C. R., and D.F. Alderdice, 1966.  Effects of salinity and temperature
on embryonic  development of the pacific cod (Gadus macrocephalus).  Jour. Fish.
Resrch. Bd. Can., 23(3):  319-3^0.

Galloway, J.  C., 1951.  Lethal effects of the cold winter of 1939/^0 on marine
fishes at Key West, Florida.  Copeia, 2:  118-119.

Gunter, G., 1957.  Temperature.  Chapter 8.  In:  Treatise on Marine Ecology
and Palaeoecology, I.  (Ed. by J. W. Hedgepeth).  Geol. Soc. Amer. Mem. 67,

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                                    79
Gunter, G., and H. H. Hildebrand, 1951.  Destruction of fishes and other
organisms on the south Texas coast by the cold wave of January 28-February 3>
1951.  Ecol., 32(4):  731-735.

Harvey, H. H., 1964.  Dissolved nitrogen as a tracer of fish movements.  Verh.
int. Ver. Limnol., 1962, 15, 947-951.  Water Poll. Abstr. (Brit.), 38, 878,
1965.

Hayes, P. R., 1949.  The growth, general chemistry, and temperature relations
of salmonid eggs.  Quart. Rev. Bio., 24(4):  281-308.

Hayes, F. R., D. Pelluett, and E. Gorham, 1953.  Some effects of temperature
on the embryonic development of the salmon (Sa-lmr> salar).  Can. Jour. Zoology,
31(1):  42-51.  Water Poll.Abstr. (Brit.), 26, 7, 1953.

Hoff, J. G. , and J. R. Westman, 1966.  The temperature tolerances of three
species of marine fishes.  Jour. Marine Res., 24(2):  131-140.

Hubbs, C., 1965.  Developmental tempperature tolerance and rates of four
southern California fishes, Fundulus  parvipinnis, Atherinops affinis,
Leuresthes tenuis, and Hypsoblennius sp.  Calif. Fish and Game7 51(2):
113-122.

Huntsman, A. G., 1942.  Death of salmon and trout with high temperature.
Jour. Fish. Resch. Bd. Can., 5(5):  485-501.

Johnson, H. E., and R. F. Brice, 1953.  Effects of transportation of green
eggs, and of water temperature during incubation, on the mortality of chinook
salmon.  U. S. Fish and Wildlife Service, Prog. Fish. Cult., 15(3):  104-108.

Kerr, J. E., 1953.  The fish rescue project at the Pacific Gas and Electric
Company's  Contra Costa steam plant.  Proc. Am. Soc. Civ. Engrs., 79(264):
10 pp.  Water Poll. Abstr. (Brit.), 27, 8,1954.

Ketchen, K. S., 1956.  Factors influencing the survival of the lemon sole
(Parophrys netulus, Girard) in Hecate Strait, British Columbia.  Jour. Fish.
Resch. Bd. Can., 13(5):  647-694.

Kinne, 0., 1963.  The effects of temperature and salinity on marine and
brackish water animals.  I.  Temperature.  Oceano. Mar. Bio. Ann. Rev.,
1:  301-340.

Leitritz, E., 1962.  Trout and salmon culture.  State of California, Department
of Fish and Game, Fish Bull. No. 107, 169 pp.

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                                    80
Lewis, R. M. , 1965.  The effect of minimum temperature on the survival of
larval Atlantic menhaden, Brevoortia tyrannus.  Trans. Am. Fish. Soc.,
Lewis, R. M. , 1966.  Effects of salinity and temperature on survival and
development of larval Atlantic menhaden, Brevoortia tyrannus.  Trans. Am.
Fish. Soc., 9500:
Major, R. L., and J. L. Mighell, 1966.  Influence of Rockey Reach Dam and
the temperature of the Okanoogan River on the upstream migration of sockeye
salmon.  U. S. Fish & Wildlife Service, Fish. Bull., 66(l):  131-
Massmann, W. H. , and A. L. Pacheco, 1957.  Shad catches and water temperatures
in Virginia.  Jour. Wildlife Mgmt., 2l(3):  351-352.

McCauley, R. W. , 1963.  Lethal temperatures of the developmental stages of the
sea lamprey, Petromyzon marinas L.  Jour. Fish. Resch. Bd. Can. , 20,
McNeil, W. J. , 1966.  Effect of the spawning bed environment on reproduction
of sink & chum salmon.  U. S. Fish & Wildlife Service, Fish. Bull., 65(2):
^95-523.

Merriman, D. , 19^1.  Studies on the striped bass (jRoccus saxatilis) of the
Atlantic coast.  U. S. Fish & Wildlife Service, Fish. Bull. 50(35):  77 pp.

Mihursky, J. A., and V. S. Kennedy, 1967.  Water temperature criteria to
protect aquatic life.  Symposium on water quality criteria to protect
aquatic life.  Am. Fish. Soc., Special Publ. No. k, pp. 20-32.

Nakatani, R. E., and R. F. Foster, 1966.  Hanford temperature effects on
Columbia River fishes.  Mimeo., 15 p.

Naylor, E. , 1965.  Effects of heated effluents on marine and estuarine organisms.
In:  Advances in Marine Biology, Sir Frederick S. Russell, Editor, Academic
Press, 3:  63-103.

Nikolsky, G. V., 1963.  The ecology of fishes.  Academic Press, New York,
352 p.

Olson, P. A., and R. F. Foster, 1957.  Temperature tolerance of eggs and young
of Columbia River Chinook salmon.  Trans. Am. Fish. Soc., Eighty- fifth Annual
Meeting, 1955, p. 203-207, Spo. Fish. Abstr. , 3, 1387, 1958.

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                                     81
Ordal, E. J. , and R. E. Pacha, 1963.  The effects of temperature on disease
in fish.  In:  Water Temperature Influences, Effects and Control.  Proc. of
the Twelfth Pacific Northwest Symposium on Water Poll. Resch., Pacific
Northwest Water Laboratory, U. S. Public Health Service, Corvallis, Oregon,
PP. 39-56.

Piavis, G. W. , 1961.  Embryological stages in the sea lamprey and effects
of temperature on development.  U". S. Fish & Wildlife Service, Bu. of
Commercial Fisheries, Fish. Bull. 182, 6l:  111-1^3, Sport Fish. Abstr. .
6:  3, 1961.

Prosser, C. L. , F. A. Brown, D. W. Bishop, T. L. John, and V. J. Wulff, 1950.
Comparative animal physiology.  W. B.Saunders Co., Phila. , Pa., 888 pp.

Reintjes, J. W. , and A. L. Pacheco, 1966.  The relation of menhaden to
estuaries.  In:  A Symposium on Estuarine Fisheries, Am. Fish. Soc., Special
Publ. No. 3, PP. 50-58.

Rockwell, J., Jr., 1956.  Some effects of sea water and temperature on the
embryos of the Pacific salmon, Oncorhynchus gorbuscha (Walbaum) and Oncorhynchus
keta (Walbaum).  Dissertion Abstr., 16(5):  880; Spo. Fish. Abstr., 2, 1, 1956.

Rothschild, B. J. , 1961.  Production and survival of eggs of the American
smelt, Osmerus mordax (Mitchill), in Maine.  Trans. Am. Fish. Soc., 90(1)*
U3-kd.

Schwartz, F. J. , 196U.  Effects of winter water conditions on fifteen species
of captive marine fishes.  Am. Midi. Nat., ?l(2):
Seymour, A. H. , 1956.  Effects of temperature upon young chinook salmon.  Ph.D.
Thesis, Univ. Wash., Seattle, 127pp.  Diss. Abstr. l6(ll):  22^9.   Spo.
Fish. Abstr., 2:  953, 1957.

Seymour, A. , 1959*  Effects of temperature upon the formation of vertebrate
and fin rays in young chinook salmon.  Trans. Am. Fish. Soc., 88:  58-69.

Sheridan, W. L. , I960.  Relation of stream temperatures to timing of pink
salmon escapements in Southeast Alaska.  Symposium on Pink Salmon,  H. R. MacMillan
Lectures in Fisheries.  Univ. of British Columbia, Vancouver, Canada, pp. 87-102

Sheridan, W. L. , 196l.  Temperature relationships in a pink salmon stream in
Alaska.  Ecology, U2:  91-98.

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                                    82
TagatZj M. E., 1961.  Tolerance of striped bass and American shad to changes
in temperature and salinity.  U. S. Fish & Wildlife Service, Spec. Sci. Rept.-
Fish., 388, 8 p.

TaTbot, G. B., 1966.  Estuarine environmental requirements and limiting
factors for striped bass.  In:  A Symposium on Estuarine Fisheries, Am.
Fish. Soc. Special Publ. No. 3, pp. 37-U9.

Trembley, F. J., 1960.  Research project on effects of condenser discharge
water on aquatic life.  Institute of Resrch., Lehigh Univ. Progress Report
1956 to 1959, 151 P.

Trembley, F. J., 1961.  Research project on effects of condenser discharge
water on aquatic life.  The Institute of Resrch., Lehigh Univ. Progress Report.

Vernon, E. H., 1958.  An examination of factors affecting the abundance of
pink salmon in the Fraser river.  Progr. Rep. int. Pacif. Salm. Fish. Comm.,
52 p.

Wells, H. W., M. J. Wells, and I. E.Gray., 196l.  Winter fish mortality in
Pamlico sound, North Carolina.  Ecol., k2i  217-219.

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                     V.  AQUATIC PLANTS AND BENTHOS





General



       Temperature regulates molecular movement and thus largely determines



the rate of chemical reactions and consequently the rate of metabolism



and activity of all organisms, both those with a relative constant temper-



ature (homoeothermus) and those with a variable temperature that is identical



with, or close to, their environment (poikilotherraus).  Because of its



regulative capacity in determining the rate of metabolism, temperature



presumably is the most important single environmental entity concerning



life and life processes.



       Variations in temperature of streams, lakes, estuaries, and oceans



are normal events that result from climatic and geologic phenomena.  The



range of temperatures in waters that support some form of aquatic life



other than viruses or bacteria is from -3°C (26.6°F) in super-cooled sea



water to 85°C (l85°F) in fresh-water thermal springs; most aquatic poi-



kilotherms,  if not all, tolerate only those temperature changes that occur



within a much narrower range whether it be high, intermediate, or low on



this scale of temperatures.



       Within the same species the biological effects of a given temperature



or temperature pattern may be different in different populations, at different



ages, in different life cycle stages, or in the two sexes (Sprague, 1963),



and such effects may depend on the temperature history of the individual

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tested (Prosser, et al., 1952), as well as on present or past effects of



other environmental factors.  Many organisms experience temperature changes



in their natural habitat, and these changes can "be an important prerequisite



for their well "being and completion of their life cycles.  A constant tem-



perature of 20 C (68 P), temperatures fluctuating rapidly and irregularly



between 15° and 25°C (59 and 77°F) with an average value of 20°C (68°F),



and temperatures fluctuating gradually and periodically between 15° and 25°C



(59  and 77 F) with an average of 20°C (68°F) do not necessarily have the



same biological effects.  Thus, even if other prerequisites are satisfactory,



the absolute temperature values of a body of water are only one measure of



its suitability for a normal assemblage of aquatic life; consideration also



must be given to temperature patterns or dynamics.  For example, distinctions



must be made between constant and fluctuating temperatures, between gradients,



ranges, averages, frequency and intensity of changes, duration of a given



pattern, and total summation (Kinne, 1963).



       The temperature range tolerated by many species of organisms is



narrow during very early development, then increases somewhat and finally



decreases again in the "old adult."  Similarly, it often is more restricted



during the sexual phase than during other phases.  Upper lethal temperatures



may be lower for animals from cold water than for closely related species



from warm water (Prosser and Brown, 1961).  A similar aspect presumably could



be found among closely related algae as well as other plants.  Many mobile

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organisms such as fish, some zooplankton, certain algae, and bottom-




associated animals can avoid critical temperatures by vertical and




horizontal migration into more suitable conditions.  Other such organisms



may be attracted to areas with critical temperatures and succumb when




these are attained.




Fresh-Water Algae and Other Aquatic Plants




       Except for fresh waters in tropical areas, relatively broad tem-




perature ranges naturally occur seasonally and diurnally, but even in




tropical waters there may be some narrow seasonal changes or major changes




resulting from unusual climatic events.  Although tropical waters usually




experience only minor temperature changes, significant qualitative and




quantitative modifications occur in their associated flora.  Similar changes




have been found during summer in temperate and north-temperate fresh water




apparently with no attendant temperature variation, and such phenomena




have led to the hypothesis that temperature, per se, bears no relationship




to floristic changes in these and other waters (Blum, 1953; Pearsall, 1923;




and Butcher, 1924).  Nevertheless, more recent studies employing laboratory




procedures wherein variables other than temperature were static have shown




that temperature, per se, does have a profound influence on aquatic algae




and other plants (Phinney and Mclntyre, 19&5; Owens and Maris, 196U), and



many others provide nonlaboratory data that strongly implicate temperature




as one of the causes of both qualitative and quantitative changes in aquatic




flora (for example:  Cairns, 1956; Wallace, 1955; Trembley, I960; Palmer,

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                                   86
          Patrick, 19^).  Extracellular algal products, nutrients, or other



factors can cause algal population changes (Mackenthun, et al., 196U;



Mackenthun, 1965; Hartman, I960; and Fogg, 1962), and these may have "been



responsible for those population changes mentioned above where there were no



attendant temperature variations.



       Algae and other plants, like poikilothermous animals, lack physiological



mechanisms to maintain constant internal temperatures, and have tissue tem-



peratures identical with or very close to that of their environment.  Ter-



restrial plants are subjected to much wider temperature ranges than those



living in aquatic environments, but a few species of aquatic algae tolerate



temperatures higher than any terrestrial plants.  For example, several



authors have reported that some species of algae can tolerate water tem-



peratures as high as 85°C (l?5°F) as found in thermal springs (Mann and



Schlicting, 1967; and Kinne, 1963), but optimal temperatures for the same



or similar species may range from 51° to 56°C (123.8° to 132°F) according



to Brock and Brock, 1966.  Other algae, notably certain diatoms, can tolerate



low temperatures near 0 C (32 F), and some may remain viable after freezing.



Ulothrlx zonata, a filamentous green algae, grows best below 15 C (59 F)J



and can produce reproductive bodies, zoospores, at temperatures near 0 C



(32°F) in ice water (Oltmanns, 1922-23, as cited by Blum, 1953).



       Such tolerances to high or low temperature extremes are not universal



among algae and other aquatic plants; similarly, an individual plant may

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                                    87
not thrive at all temperatures between those extremes mentioned above.



Rather, there appears to be particular temperature ranges that are



tolerated by each species and by closely related species or groups of



species.  A similar concept applies to optimum temperatures for aquatic



algae.  Thus, Cairns (1956) indicated that in an unpolluted stream diatoms



grow best at 18° to 20°C (6k.k to 6.8°F); green algae at 30°C (86° to 95°F);



and blue-green algae at 35  to ^4-0 C (95 to 10^ F).  If environmental tem-



peratures near 10 C (50°F) are increased either naturally or artificially



to about 38  (100.U F), the predominance of groups of species changes



correspondingly from diatoms to green algae and finally at the uppermost



temperatures to blue-green algae (Wallace, 1955).  A few of the more high-



temperature-tolerant species belonging to algal groups other than the blue-



green may persist with the predominant blue-green species in such cases,



and several less tolerant species of blue-greens may succumb with the diatoms



and green algae at these higher temperatures.




Bottom Organisms



       In a study by Strangenberg and Pawlaczyk (1961) on



the effect of warm-water discharges on

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a river they found that river-bottom plants and animals decreased in




number when the water temperature exceeded 30 C (86 F).  The macro-




invertebrate riffle fauna of the Delaware River was adversely affected




by heated water effluents.  The macro-invertebrate biomass was reduced




from I.Ok to 0.09 grams per square foot throughout the summer in the




area of maximumly heated water, as compared with a control station.  A




35 C (95 F) water temperature at the time of sampling was found to be




causing a detrimental effect on many organisms, especially the caddisfly,




Hydropjsyche sp., many of which were dead, while those alive were extremely




sluggish.  The data suggest  that there is a tolerance limit close to




32.2°C (90°F) for a variety of different kinds of animals in the popula-




tion structure of benthos with extensive losses in numbers and diversity




of organisms accompanying further temperature increase (Coutant, 1962).




       Another classic demonstration on the effects of increasing water




temperatures upon the change in the composition of a macro-invertebrate




population is presented by Walshe (19**8).  The thermal index (22-hour




LD_-) of seven species of midge larvae reflect the probable sequence




of preferred temperatures.  These seven species and their thermal in-




dices are as follows:  Tanytarsus brunnipes, 29 C (81*.2 F); Prodiamesa




olivacea, 30°C (86°F); Anatopynia nebulosa, 30.^°C (95°F);  C.  long!stylus,




35°C (95.9°F); and Anatopynia varia, 38°C

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                                    89
       In studies on the shift of the composition of macro-invertebrate




populations (Wurtz and Henn, 1965)5 it was shown that no immediate kills



resulted from thermal shock of lU C (25 F).  However, persistent exposure




to 35 C (95 F) over 2U hours brought about changes in the composition of



the macro-invertebrate population.




       Studies of particular species of macro-invertebrates have shown




that lethal temperatures vary considerably with the type of organism.




Noland and Reichel (19^3) in studying the fresh-water snail (tymnaea




stenalis) found that cultures died when the water temperature reached




30.5 C (89.6 F).  Fresh-water snails (Vivlparus malleatus) died when




held at a temperature of 37.5°C (99-5°F) (Hutchinson, 19^7).



       The highest 2^-hour median tolerance limit lethal temperatures




that could be obtai-ned by raising acclimation temperatures from 10 C




(50°F) to 20°C (68°F) were estimated to be 3^-6°C (9^.2°F) for the sowbugs




Asellus intermedius, Forks, and the scud, Gammarus fasciatus, Say, 33.2 C



(91.8°F) for the scud, Hyallella -azteca (Saussure), and 29.6°C (85.3°F)




for the scud, Gammarus pseudolimnaeus Bousfield (Sprague, 1963).




       The fresh-water snail (Physa gyrina) has been found to live and




reproduce in a waste water ditch between 28° and 35°C (82.h° to 95°F)




(Agersborg, 1932).



       Many marine bottom-associated organisms have stenothermal or narrow




temperature ranges.  In some cases, a particular species may be stenothermal

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                                    90
for one developmental stage, and eurythermal for another.  Breeding or



spawning requirements are generally stenothermal.  The time of spawning



for molluscs is highly dependent upon temperature.  Most molluscs with



specific temperature-breeding relationships are spring and summer spawners,



and many do not spawn until a certain temperature is reached (Allen, 1963).



Spawning of the American oyster (Crassostrea virginica) takes place between



15 and 32 to 3^°C (59 and 89.6 to 93.2°F), depending on condition of the oyster,



and the spawning process is usually triggered by a rise in temperature



(Galtsoff, 196U).



       A large number of species are able to tolerate higher temperatures



than those at which they can breed.  For example, Carcinus maenas thrives,



but does not breed in lU-28°C temperatures (57.2-82.U°F) (Naylor, 1965).



In the case of the European lobster, temperature controls a different part



of reproduction.  Larvae require a minimum temperature of 15 C (59 F) even



though the developing eggs, and adults, will tolerate lower temperatures



(Gunter, 1957).  For the above two cases, temperature limits the popula-



tions and recruitment of organisms must occur from outside the heated area.



       Physiology, metabolism, and development are all affected by



temperature.  At a temperature of 6-7°C (42.8-4U.6°F), C. virginica



ceases feeding.  Above 32 C (89.6°F) ciliary action, responsible for move-



ment of water, rapidly decreases; and almost all functions of the body



cease, or are reduced to a minimum at U2.0°C (107.6°F).  The European

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                                    91
oyster, Ostrea lurida, has a tendency to close it's shell in response to




falling temperatures.  At l4-6°C (39.2-U2.8°F) the shells of oysters remain




closed most of the time; at 6-8°C (U2.8-U6.4 F) the shells open for about




6 hours per day; and at 15 C (59 F) the shells remain open for 23 hours



per day (Galtsoff, 1964).  Very little is known about prolonged effects




of temperature above 32-3^ C (90-9^ F) on oyster populations.  Long, con-




tinued exposure to high temperatures may impede the normal rate of water




transport.  When either low or high temperatures cause a closing of shells




or a ceasing of ciliary action, oysters cease to feed and lose weight.




Thus, temperature may produce an effect similar to chronic toxicity.




       Acclimation and tolerance of bottom-associated organisms may be




affected by temperature changes.  The crab (Hemigrapsus nudus) can regain




tolerance to high temperatures, after a low temperature history, in less




than a week.  Shore crabs (Pachygrapsus crassipes) may require a half time




of six days in order to acclimate to a temperature change of 7.5 C (13'5 F)




(Kinne, 196?).  The giant scallop (Placopecten magellanicus)  acclimates




rapidly to a rise in temperature of 1.7 C (3.IF) per day, but may take




as long as three months to lose this acclimation to high temperatures




(Dickie, 1958)-  The opossum shrimp (Neomysis americana) is very intolerant




to temperature increases, and does not appear to survive at temperatures




above 31°C (8?.8°F) in the Chesapeake estuary (Mihursky and Kennedy,




1967).

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                                     92
       Distribution of benthic organisms may be controlled by temperature.



Reef-forming corals will not live where temperature falls below 18-19 C



(6U.U-66.2 F).  The American oyster (C. virginica) on the Gulf coast is



present in water that may vary from U-3U°C (39.2-93.2°F); while the European



oyster (0. edulis) is restricted to water with temperatures of 0-20 C



(32-68 F) (Gunter, 1957).  In a study on the York River, in Virginia, Warinner



and Brehmer, 1966, found that the community composition and abundance of



marine benthic invertebrates in the river were affected by thermal discharge



over a distance of 300-UOO meters from the discharge outfall.  They con-



cluded that during the months of high normal river temperatures there was



clear evidence of biological stress.



       Cold is as important as heat in its effects on marine organisms.



Cold water may kill directly, or in some cases indirectly when organisms



are "numbed" or rendered inactive and unable to protect themselves from



predators (Gunter, 1957).



       One of the benefits derived from heated water is the defouling of



intake pipes.  Experiments have shown that fouling by Etytilus edulis and



M. californianus could be controlled by tri-weekly reversals of heated



discharge water either for periods of one hour at 38-Ul°C (100.U-105.8°F)



or for seven hours at 3^.5°C (91.U°F) (Naylor, 1965).

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                                   93
Summary



1.  Temperature is one of the most important single environmental entities



concerning life and life processes.  The various functions of an or-



ganism may have somewhat different temperature ranges, and if these are



not provided in the habitat the organism will die.



2.  When water temperatures are increased, the predominance of groups of



algal species changes correspondingly from diatom to green algae and



finally at higher temperatures to blue-green algae.



3.  The number and distribution of bottom organisms decrease as water



temperatures increase with a tolerance limit close to 90°F for a "balanced



population structure.  Studies of particular species of macro-invertebrates



have shown that lethal temperatures vary considerably with



the type of organism.  In some cases a particular species may be steno-



thermal for one developmental stage, and eurythermal for another.  Thus,



a large number of species are able to tolerate higher temperatures than



those at which they can breed.



k.  Cold is as important to aquatic plants and benthos as is heat.



5.  One of the beneficial uses of heated effluents is the defouling of



intake pipes; accomplished by reversing the flow of water through the



pipes for a specified period of time.

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                                  94

                             REFERENCES CITED
Agersborg, H. P. K. , 1932.  The relation of temperature to continuous
reproduction in the pulmonate snail.  Nautilus, 45(4): 121-123.

Allen, J. A., 1963.  Ecology and functional morphology of molluscs.  Oceano.
Mar. Bio. Ann. Rev., 1,' Harold Barnes, Ed., 253-288.

Blum, J. L. , 1953.  The ecology of algae growing in the Saline River, Michigan,
with special reference to water pollution.  Doc. Thesis, Univ. Mich., ix +
176 pp.

Blum, J. L. , 1956.  The ecology of river algae.  Bot. Rev., 22:  291-341.

Brock, T. D. , and M. L. Brock, 1966.  Temperature optima for algal develop-
ment in Yellowstone and Iceland hot springs.  Nature, 209, No. 5024, 733-34.

Butcher, R. W. , 192*4-.  The plankton of the River Wharf e (Yorkshire).  Naturalist,
pp. 175-180, 211-21*1.

Cairns, J. , Jr., 1956.  Effects of increased temperatures on aquatic or-
ganisms.  Ind. Wastes, 1(4):  150-152.
Coutant, C. C. , 1962.  The effect of a heated water effluent upon the macro-
invertebrate riffle fauna of the Delaware River.  Perm. Acad. Sci., 37:
58-71.

Dickie, L. M. , 1958.  Effects of high temperatures on survival of the giant
scallop.  Jour. Fish Resch. Bd. Can., 15(6):  1189-1211.

Fogg, G. E., 1962.  The importance of extra- cellular products of algae in
the aquatic environment.  Bio. Prob. Wat. Poll., 3rd Seminar, PHS Publ. No.
999-WP-25, C. M. Tarzwell, Ed., 424 pp.

Galtsoff, P. S., 1964.  The American oyster Crassostrea virginica Gmelin.
U. S. Fish & Wildlife Serv. , Spec. Sci. Reps., Fish. No. 64^ 480 p.

Gunter, G., 1957.  Temperature.  Treatise on marine ecology and palaeoecology I.
(Hedgepeth, J. W. , Ed.), Chap. 8, Geol. Soc. Am. Mem., 67, 159-184.

Hartman, R. T., I960.  Algae and metabolites of natural waters.  In:  The
Ecology of Algae, Spec. Publ, No. 2, Pymatuning Lab. of Field Biology, Univ.
Pittsburgh, Pittsburgh, Pennsylvania, pp. 38-55.

Hutchinson, L. , 194-7.  Analysis of the activity of the fresh water snail,
Viviparus malleatus (Reeve).  Ecol., 28(4):  335-345.

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                                    95
Kinne, 0., 1963.  The effect of temperature and salinity on marine and
brackish water animals.  Temperature.  In:  Oceano. and Mar. Bio., Ann.
Rev., 1 (Harold Barnes, Ed.), 301-3^0.

Kinne, 0., 19&7.  Physiology of estuarine organisms with special reference
to salinity and temperature:  general aspects.  In:  Estuaries, George H.
Lauff, Ed., Amer. Assoc. Adv. Sci., Publ. No. 83, 525-5*4-0.

Maekenthun, K.  M. , W. M.  Ingram,  and  R.  Forges.   1961*.   Limnological Aspect-s of
Recreational Lakes.  U.  S.  Public Health Service, Publ.  No.  116?:   176 pp.

Mackenthun, K. M.
1965.  Nitrogen and phosphorus in water, an annotated selected bibliography
of their biological effects.  U. S. Public Health Service, Publ. No. 1305:
in pp.

Mann, J. E. and H. E. Schlichting, Jr., 1967.  Benthic algae of selected
thermal springs in Yellowstone National Park.  Trans. Amer. Microscopical
Soc., 86(1):  2-9.

Mihursky, J. A., and V. S. Kennedy, 1967.  Water temperature criteria to pro-
tect aquatic life.  Symposium on Water Quality Criteria to Protect Aquatic
Life, Am. Fish. Soc., Spec. Publ. No. k, 20-32.

Naylor, E., 1965.  Biological effects of a heated effluent in docks at
Swansea, S. Wales.  Proc. Zool. Soc., London, Ihk,:  253-268; Water Poll.
Abstr. , 39(6):  Abstr. No. 997.

Naylor, E., 1965 •  Effects of heated effluents upon marine and estuarine
organisms.  In:  Advances in Marine Biology, 3> Academic Press, New York,
63-103.
Noland, L. E. , and E. Reichel, 19^3*  I^e cycle of Lymnaea stagnalis
completed at room temperature without access to air.  Nautilus , 57 (1 ) :  8-13.

Oltmanns, F. , 1922-23.  Morphologic und biologic der algen.  2 auff . , 3 vols.

Owens, M. , and P. J. Maris, 196^-.  Some factors affecting the respiration of
some aquatic plants.  Hydrobiologia, 23:  533-5^3-

Palmer, C. M. , 1965.  Phytoplankton periodicity in a, Newfoundland pond.
Phycologia, 1:  39^0.

Patrick, R. , 19^8.  Factors affecting the distribution of diatoms.  Bot.
Rev., 1U(8):  U78-52U.

Pearsall, W. H. , 1923.  A theory of diatom periodicity.  Jour. Ecol., 11:
165-183.

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                                    96
Phinney, H. K. , and C. D. Mclntire, 1965.  Effect of temperature on
metabolism of periphyton communities developed in laboratory streams
Limnol. Oceanog. , 10(3 ):
Prosser, C. L., et al., 1952.  Comparative animal physiology.  W. B. Saunders
Co., Philadelphia, Pa., 888 pp.

Prosser, C. L. , and F.A. Brown, Jr., 1961.  Comparative animal physiology.
Second Edition, W. B. Saunders Co. , Philadelphia, Pa. , 688 pp.

Sprague, J. B. , 1963.  Resistance  of four freshwater crustaceans to lethal
high temperature and low oxygen.   Jour. Fish. Resch. Bd. Can., 20(2} :
387-^15.

Trembley, F. J. , I960.  Research project on effects of condenser discharge
water on aquatic life, progress report, 1956 to 1959«  Institute of Resch.
Lehigh Univ.,  15*4- pp.; Water Poll. Abstr. , 3^(ll), Abstr. No. 2157.
Wallace, N. W. , 1955.  The effect of temperature on the growth of some fresh-
water diatoms.  Notulae naturae of the Acad. Nat. Sci, of Philadelphia,
280:  1-11.

Walshe, B. M. , 19^.  Oxygen requirements and thermal resistance of chironomid
larvae from flowing and still water.  Jour. Exp. Bio., 25:  35.

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

Wurtz, C. B.,  and C.E. Renn, 1965.  Water temperatures and aquatic life.
Prepared for  Edison Electric Institute Research Project No. ^9, 99 PP«

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                         TEMPERATURE and AQUATIC LIFE

                           A Selected Bibliography


Abbott, B. J., I960. A note on the oxygen and temperature tolerance  of
the triclads, Phagocata gracilis  (Haldeman) and Dugesia tigrina  (Girard) .
Virginia Jour. Sci., 2(1): 1-8.

Adlung, K. G. , 1957-  The toxicity of insecticides to fish and its
dependence on temperature.  Nat. Wiss., UU: 622-623.  Water Poll. Abstr.,
31(12), Abstr., No. 2Mj£, 1958.  Spo. Fish. Abstr., Ml), Abstr. No.
2166, 1959-

Agersborg, H. P. K. , 1930.  The influence of temperature on fish.  Ecology,
11(1): 136 -3M.

Agersborg, H. P. K. , 1932.  The relation of temperature to continuous
reproduction in the pulmonate snail.  Nautilus, U5(4); 121-123.

Alabaster, J. S., 1962.  The effect of heated effluents on fish,
Internat. Conf. on Water Poll. Resrch. , London, 1962.  Air and Water
Poll. 7(6/7): 5^1-563, 1963.  Bio. Abstr., ^5(13), Abstr. No. 5^305, 196^.

Alabaster, J. S. and A. L. Downing,  I960.  The behavior of roach (Rutilis
rutilis L. ) in temperature gradients in large outdoor tanks.  Proc.
Indo-Pacific Fish. Coun. Hl(U9).

Alabaster, J. S. and A. L. Downing,  1966.  A field and laboratory
investigation of the effect of heated effluents on fish.  Fish. Invest.
(Minist. of Agr., Fish, and Food of Unit. King.), Ser. I, 6(k) : k2.

Alabaster, J. S. , D. W. M. Herbert, and J. Hemens ,  1957.  The survival
of ranibow trout (Salmp gairdneri Richardson) and perch (Perca fluviatilis
L. ) at various concentrations of dissolved oxygen and carbon dioxide.
Ann. Appl. Bio. If?: 177-188.

Alabaster, J. S., and K. G. Robertson,  1961.  Effect of diurnal changes
in temperature, dissolved oxygen, and illumination on the behavior cf
roach (Rubilus rutilus L. ), bream (Abramis brama L. ) and perch ( Perca.
f luviatilus L. ) .  Animal Behavior, 9(3-4): l8j.

Alabaster, J. S., and 'A. Swain,  1963-  Heated water and fish.  Ann.
Rep. Challenger Soc., 3(15) : 39-

Alabaster, J. S. , and R. L. Welcomrne,  1962.  Effect of concentration
of dissolved oxygen on survival of trout and roach in lethal temperatures.
Nature, London, 19^: 107.  Water Poll. Abstr., 36: 672, 1963.  Spo. Fish.
Abstr., 9(2), Abstr. Ho. 658?,
                                  97

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                                 98

Alderdice, D. F.,  1963.   Some  effects of simultaneous variation in
salinity, temperature and dissolved oxygen on the resistance of young
coho  salmon to a toxic  substance.   Jour.  Fish.  Res.  Bd.  Can.,  20:  525-550.
Spo.  Fish. Abstr.  8(3), Abstr. No.  5905,  1963.   Water Poll.  Abstr.
37(1), Abstr. No.  178,  1964.

Allanson, B. R., and R. G.  Noble,   1964.   The tolerance  of Tilapia
mossambica (Peters) to  high temperature.   Trans.  Am.  Fish.  Soc.  93(4):
323-332.  Spo. Fish. Abstr. 10(l),  Abstr.  No.  7192,  1965.

Allbaugh, C. A., and J. V.  Manz.,  1964.   Preliminary study of the
effects of temperature  fluctuations on developing walleye  eggs and
fry.  Prog. Fish-Cult.  26(4):  175-180.

Allee, W. C., A. E. Emerson, 0.  Park,  T.  Park,  and K.  P. Schmidt,  1949.
Principles of animal ecology.  W.  B.  Saunders   Co.,  Philadelphia,  Pa.

Allen, K. R., 1940.  Studies on  the biology of the early stages  of the
salmon, (Sa'lmo salar)'.  I.  Growth  in the  River Eden.   Jour.  Animal
Ecol. 9(1): 1-23.

Ames, A. M., and W. W,  Smith,  1944.   The  temperature  coefficient of the
bactericidal action of  chlorine.   Jour, of Bact.,  47:  445.   Water  Poll.
Abstr., 17, Jul. 1944.

Anaichev, A. V., 1959.  Digestive  enzymes of fish and seasonal changes
in their activity.  Biokhimiia (Transl.),  24(6):  952-958.  Spo.  Fish.
Abstr., 6(1), I960.

Anderson, R. C., 1951.  Preferred  temperature  of  a sample of Lepomis
gibbosus, the pumkinseed.   Manu. in the Ontario Fish.  Resrch.  Lab.
Library, Toronto.

Andrews, C. W., 1946.  Effect  of heat on  the light behavior  of fish.
Trans. Roy. Soc. of Can., Ser. 3,  40,  Soc.  5:  27-31.

Angelovic, J. W., W. F. Sigler,  and J. M. Neuhold.  Temperature  and
flourosis in rainbow trout.  Jour.  Water Poll.  Control Fed., 33: 371-381,
Apr.  1961.

Anon., 1949.  Discharge of  heated  liquids  into  streams.  Rep.  of Rivers
Poll. Prevention Sub-Committee of the  Central Water Committee, Minist. of
Health,. England, pp; 69-76.

Anon., 1954.  Changes in the dissolved-oxygen content  of river water
used  for direct cooling.  Rep. No.  10, Generating  Station Operation-
Resrch. Liaison Committee.  Brit. Electr. Author.

Anon., 1955.  First progress report.   Aquatic Life Advisory  Committee
of ORSANCO, Sewage and Industrial Wastes,  27(3):  321-331-

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                                99
Anon., 1956.  Feed heavy on a rise in temperature.  Prog. Pish Cult.,
18(1): kk.

Anon., 1956.  Proceedings of the fifth annual water symposium, February
1956.  Louisiana State Univ., Engng. Exp. Sta. Bull. 55.

Anon., 1956.  Aquatic life water quality criteria  (Second progress
report of the Aquatic life Advisory Committee of the Ohio River Valley
Water Sanitation Commission).  Sewage and Indus. Wastes, 28(5):
678-690.

Anon., 1958.  Oxygen relationships in streams, proc. of seminar at Robert
A. Taft Sanitary Engineering Center, Cincinnati, Ohio, Oct. 30 - Nov. 1,
1957-  Publ. Health Serv. Tech. Rep. No. W58-2.

Anon., 1960.  Is heat a pollutant?  Pensylvania thinks so.  Chem. Engng.
Prog. Staff, Chem. Engng. Prog. 56: 33.

Anon., 196l.  Effect of water temperature on stream re-aeration. Proc.
Am. Soc. Civ. Engrs. 87, SA6: 59-71.

Anon., 1962.  Heated discharges . . . their effect on streams.  Rep. by
the Advisory Committee for the control of stream temperatures to the
Pa. Sanitary Water Board.  Pa. Dept. Health, Harrisburg, Publ. No. 3,
108 pp.

Anon., 1963.  Water temperature influences, effects, and control.  Proc.
of the 12th Pacific Northwest Symposium on Water Poll. Resrch., Pacific
Northwest Water Lab., Publ. Health Serv., U. S. Dept. of Health, Education
and Welfare,  160 pp.

Anon., 1963.  Sediment transportation mechanics:  density currents.
Prog. Rep., Task Committee on Preparation of Sedimentation Manual,
Committee on Sedimentation, Jour, of the Hydraulics Div., ASCE,
89(HY5h Proc. Paper 3639, pp. 77-87.

Anon., 1965.  Symposium on streamflow regulation for quality control.
U. S. Publ. Health Serv. Publ. No. 999-WP-3C-, k2O pp.

Anon., 1965.  Minutes, third conference on Patuxent estuary studies,
Nov. 13-1^, 196^.  Chesapeake Bio. Lab. Ref. No. 65-23.

Anon., 1966.  Nuclear power plant proposal raises pollution issues.
Water Control News, l(l), May 16.

Anon., 1967.  Temperatures for hatching walleye eggs.  Prog. Fish Cult.,
29(1): 20 (Jan.).

Ansell, A. D., 1963.  Venus mercenaria L.in Southampton water.  Ecology
     : 396-397. '

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                                 100
Ansell, A. D., 1963-  The biology of Venus  mercenaira in British waters,
and in relation to generating  station  effluents.   Ann.  Rep.  Challenger
Soc. 3(15): 38.

Ansell, A. D. , and F. A. Loosemore,  1963.   Preliminary observations on
the relation between growth,, spawning  and condition in experimental
colonies of Venus mercenaria L.   Jour, du Conseil,  28:  285-29**-.

Ansell, A. D. , K. F. Lander. J. Coughlan, and F. A.  Loosemore.  196*4-.
Studies on the hard-shell clam, Venus  mercenaria,  in British waters.
I Growth and  reproduction in natural and experimental colonies.   Jour.
Appl.  Ecol., l: 63-82.

Ansell, A. D. , F. A. Loosemore, and K. F. Lauder,  196*4-.   Studies on
the hard-shell clam, Venus mercenaria, in British  waters.   II  Seasonal
cycle in bio-chemical composition.  Jour. Appl. Ecol.,  1:  83-95-

Anthony, E. H. , 196l.  The oxygen capacity  of goldfish blood (Carassius
auratus) in relation to thermal environment.  Jour.  Exp.  Bio.  38:  93-107.

Armitage, K. B. , 1962.  Temperature and Op  consumption of Ore homonella
chilensis (Heller) (Amphipoda: Gammeroida) .  Bio.  Bull.  123(2):  225- 232.

Arnold, G. E. , 1962.  Thermal pollution of  surface  supplies.   Jour.  Am.
Water Works As so. , 5^(11): 1332-13*4-6.

Bailey, R. M. , 1955-  Differential mortality from  high temperature in
a mixed population of fishes in southern Michigan.   Ecol.,  36(3):  526-528.
Spo. Fish. Abstr.' l(U), Abstr. No. 519, 1956.
Bakastov, S. S., I960.   Some data  on bottom temperatures  in the Rybinsk
Reservoir when the  surface  is  frozen.  Bull. -Inst.  Bio. Vodohranilishch
Akad.  Nauk SSSR 8/9: 62-66.   Spo. Fish. Abstr.  7(4),  1962.

Bakshtanskii, E. L. , 1961.  The role of  feeding  and of warming of the
water in the artifical rearing of  salmon above the  Arctic circle.
Rybnoe Khoz. 10: 15-18.  Referat.  Zhur., Bio. 1962,  No. 9178.   Spo.
Fish. Abstr. 8(2),  1963.

Baldwin, E. , 19*4-8.  An introduction to comparative  biochemistry.
Cambridge Univ. Press, 16*4-  pp.

Baldwin, N. S., 1957- ..Food consumption  and growth  of  brook trout at
different temperatures.  Trans. Am. Fish. Soc. 86: 323-328.   Spo.  Fish.
Abstr. 3(1), Abstr. No.  1362,  1958.

Banta, A. M. , and T. R.  Wood,  1928.  A thermal race of cladocera
originated by mutation.  Zeitschr. Induct.  Abstain - u. Vererbugsl.
Supplement b, 1: 397-398.

Baranov, I. V., 1961.  Thermic and hydrochemical condition in  the
Gorikovskii Reservoir.   Trudy  Inst. Bio. Vodokhranilishch Akad. Nauk
SSSR *4-(7): 29*4-320.  Referat.  Zhur., Bio.,  1962, No. HZh332.  Spo. Fish.
Abstr., 8(2), 1963.

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                                 101


Bardach, J. E., 1955-  Certain biological effects  of thermocline
shifts.  Hydrobiologia, 7(4): 309-324.

Bardaeh, J. E., and R. G. Bjorklund, 1957.  The temperature  sensitivity
of some American freshwater fishes.  Am. Nat. 91(859): 233-251.   Spo.
Fish. Abstr. 3(1), Abstr. No. 1322, 1958.

Barges, H. M., 1950.,  Pish distribution  studies, Niaugua Arm of the
Lake of the Ozarks, Missouri.  Jour. Wildl. Mgnt.  19(l): 16-33.

Barrington, E. J. W., and A. J. Matty, 1954.  Seasonal variation  in the
thyroid gland of the minnow, Phoxinus phoxinus L.  with some  observations
on the effect of temperature.  Proc. Zool. Soc. of London, 124: 547-564.

Basu, S. P., 1959.  Active respiration of fish in  relation to ambient
concentrations of oxygen and carbon dioxide.  Jour. Fish. Resrch.
Bd. Can. 16(2): 175-212.

Bata, G. L., 1957-  Recirculation of cooling waters in rivers and
canals.  Jour, of the Hydraulics Div. ASCE, 83(HY3), Proc. Paper  1265.

Battle, H. I., 1926.  Effects of extreme temperature on muscle and
nerve tissue in marine fishes.  Trans. Proc. and Roy. Soc. Can.,
5(20): 127-143.

Battle, H. I., 1929.  Effects of extreme temperatures and salinities
on the development of Snchelyopus cimbrius L.  Contr. Can. Bio. (N.S.)
5: 109-192.

Baudin, L., 1926.  Variation des echanges respiratoires des  poissons
en function des la pression atmospheriques  et de  la temperature.  Mem.
Soc. Vaudoise, Sci. Nat. 4(l): 1-40.

Beamish, F. W. H., 1964.  Respiration of fishes with special emphasis on
standard oxygen consumption.  II. Influence of weight and temperature
on respiration of several species.  Can. Jour. Zool. 42: 161-188.
Water Poll. Abstr. 37(10), Abstr. No. 156^, 1964.

Beaven, G. F., I960.  Temperature and salinity of  surface water at
Solomons, Maryland.  Chesapeake Sci., l(l): 2-11.

Becker, H. G., 1924.  Mechanism of absorption of moderately  soluble
gases in water, and its use in determining the rate of solution of
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                                  103
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Bisset, K. A., 19^6.  The effect of temperature on non-specific infections
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Bisset, K. A., 19^8.  Seasonal changes in the normal flora of fresh water
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Bisset, K. A., 19^8.  The effect of temperature upon antibody production
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Black, E. C., 1958.  Hyperactivity as a lethal factor in fish.  Jour. Fish.
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Blaxter, J. H. S., I960.  The effect  of  extremes  of temperature on
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Borodin, N. A., 193**.  Survival of fish  in freezing temperatures.
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Bowen, E. S., 1932.  Further studies in the aggregating behavior  of
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Brawn, V. M., I960.  Temperature tolerance of unacclimated herring
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Brett, J. R., 19^.  Some lethal temperature relations of Algonquin
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                                 105

Brett, J. R. , 1952.  Temperature tolerance in young pacific salmon,
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Brett, J. R., and D. F. Alderdice, 1958.  The resistance of cultured
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                                 106
Brown, L. A., 1929-  The  natural history of cladocerans in relation
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Brown, M. E. , 1946.  The  growth of brown trout  (Salmo trutta_ L.)  Ill  The
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Bullock, T.  H. , 1955-  Compensation for temperature in  the  metabolism
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Bullough, W. S., 1939.  A study of the reproductive cycle in  the minnow
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Burdick, G.  E., M. Lipschuetz, H. J. Dean, and  E. J. Harris,  1954.
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Burger, J. W. , 1939-  Some experiments on the relation  of the external
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Burnson, B., 1938.  Seasonal temperature  variations in  relation to
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Burrows, R.  E. , 1963.  Water temperature  requirements for maximum
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Burton, G. W. , and E. P. Odum, 1945.   The distribution  of stream fish
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                                107

Butler, P. A., 1965.  Biological problems in water pollution: reaction
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Butterfield, C. T., and E. Wattie, 19^6.  Influence of pH and temperature
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Cairns, J., Jr., and A. Scheier, 1958.  The effects of temperature and
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Cairns, J. , Jr., and A. Scheier, 196^.  The effects of sublethal levels
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Carlisle, D. B., 1957.  On the hormonal inhibition of moulting in
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Chadwick, W. L., F. S. Clark, and D. L. Fox, 1950.  Thermal control of
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                                 108
Chang, S. L. , M. Buckingham, and M. P. Taylor, 19^8.   Studies  on
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Chang, S. L. , and G. M. Fair, 19*4-1.  Viability and destruction of
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Chellis, R. D. and E. Ireland, 1959-  Site  studies for a steam power
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Chidester, F. E., 192^.  A critical examination of the evidence for
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Churchill, M. A., 19^7-  Effect of density  currents upon raw water
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Churchill, M. A., 19^5.  Discussion of "translatory waves.  "  Trans. ASCE
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Churchill, M. A., 195^-  Analysis of a stream's capacity of assimilating
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Clark, D. , 1959-  River inadequate for cooling needs of new power
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                                  109
Clemens, H. P. , and K. E. Sneed, 1958.  Effect of temperature and
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Cocking, A. W. , 195T.  Relation between the ultimate  upper lethal
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Cocking, A. W. , 1959-  The effects of high temperature on roach  (Rutilus
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Coker, R. E., 1934.  Reaction of some fresh water copepods to high
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Cole, W. H. , 1939-  The effect of temperature on the color change in
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Collins, G. B., 1952.  Factors influencing the orientation of migrating
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Colton, J. B., 1959-  A field observation of mortality of marine
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Combs, B. D. , 1965.  Effect of temperature on development of salmon
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Combs, B. D. , and R. E. Burrows, 1957.  Threshold temperatures for the
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                                  110

Cory, R. L., and H. F. Davis, 1965.  Automatic data system aids thermal
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Crisp, D. J. , 1957-  Effect of low temperature on breeding of marine
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                                  in
Damann, K. E. , 19^1.  Quantitative  study of  the phytoplankton of Lake
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Dendy, J. S., 19^.  Further studies of  depth distribution of fish,
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                                  112
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                                  115
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Fry, F. E. J., V. S. Black, and E. C. Black, 1947.   Influence  of
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                                  118
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                                  119
Gradzinski, Z., 1950.  The  susceptibility of the heart in the sea-trout
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                                  120
Hal-beck, G. E., Jr., 1953.  The use of reservoirs and lakes  for the
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                                  121
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                                  122-
Kerry, S., 1959-  Pollution of rivers by heated discharges.  Bull.
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                                  12k


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                                  125
Hutchinson, L., 1947-  Analysis of the activity of the  fresh water
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                                  126
James, M. C., 0. L. Meehean, and E.  J.  Douglass,  19^.   Fish stocking
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                                  127
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                                   128
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                                  129
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                                  130
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                                  131
Mariner, L. T. , and W. A. Hunsucker, 1959-   Ocean  cooling water systems
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                                  132
McEwen, G. P. , 1929-  A mathematical theory of the  vertical distribution
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                                  133
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Parson, J. W. , and E. Crittenden, 1959-  Growth of redeye bass  in
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                                  139


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Rao, K. P. , 1953-  Threshold concentrations  of oxygen in water for
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Shaw, P. A., 1946.  Oxygen consumption of trout and salmon.  Calif.
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Shatter, D. S. , and M, J. Whalls, 1955.  Effect of  impoundment  on water
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Shultz, L. P., 192T.  Temperature controlled variation in  the golden
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Spector, W. S.} 1956.  Handbook of biological data.  W. B. Saunders
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Strandberg, C. H. , 1962.  Dispension and diffusion  of heated  coolant
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Swift, D. R., 1959.  Seasonal variation  in the activity of the  thyroid
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Teter, H. E. , I960.  The "bottom fauna  of Lake Huron.   Trans.  Am.  Fish.
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Tsukuda, H. , I960.  Heat and cold tolerances in relation to body size
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Tsukuda, H. , and W. Ohsawa, 1959-  The heat and cold  coma temperatures
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Verduin, J. , I960.  Letter in science 131 p. 232, January 22, I960.

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Vliet, R. V., 1957.  Effect of heated condenser discharge water  upon
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                                   150
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Wickwire, G. C., L.  D. Sager, and W. E. Surge, 1929.  Comparative
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Williams, A. E., and R. K. Burris, 1952.  Nitrogen fixation "by blue-
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Wingfield, C. A., 19*tO.  The effect of certain environmental factors on
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                                  151
Wood, A. H., 1932.  The effect of temperature on the growth and
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Woynarovich, E., 196l,  Oxygen consumption of Dreissena polymorpha
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Wuhrmann, K. , and H. Woker, 1955.  Influence of temperature and oxygen
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Wurtz, C. B., 1962.  Zinc effects on freshwater mollusks.  Nautilus,
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Wurtz, C. B., 1962.  The effect of heated discharges on aquatic life
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Wurtz, C. B., and C. H. Bridges, I960.  A study of the effects of
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Wurtz, C. G. , and T. Dolan, I960.  A biological method used in the
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Yamamoto, T. , 1937.  Influence of temperature on the embryonic development
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Yamamoto, T., 1937-  Influence of temperature on the embryonic development
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                   LABORATORY INVESTIGATIONS SERIES








1.  Sargent Polarographic Oxygen Analyzer; Thermo-Fishometer



    Water Thermometer; Modified Methods for Turbidity; Modified



    Methods for Color; Suspended Solids Determination (June 2kt




    1963).*



2.  Recovery of Simple Cyanides by the Serfass Distillation



    Procedure as Compared with the Williams Cuprous Chloride



    Method (April 7, 19614-).



3.  Nitrate Determination in Saline and Estuarine Waters



    (October 13, 1964).



k.  Dissolved Oxygen Determinations by Oxygen Meter (January



    31, 1966).



5.  Picture-Key to the Genera of Aquatic Midges (November, 196?).



6.  Temperature and Aquatic Life (December, 196?).
   Out of Print

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LABORATORATORY  INVESTIGATIONS
SERIES -  NUMBER  SIX

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