COLUM HERMAL EFFECTS STUD
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ENVIRONMENTAL PROTECTION AGENCY
WATER QUALITY OFFICE
WASHINGTON, D.C. 20242
MEMORANDUM JUN 29 1971
TO: See Below
FROM: Chief, Technical Studies Branch
Division of Technical Support
Office of Water. Programs
SUBJECT: Technical Study Reports
This memorandum transmits to you copies of two technical study
reports as outlined in the Technical Study Reporting System of
September 18, 1970. The two reports are as follows:
1. Columbia River Thermal Effects Study,
Volume I Biological Effects Studies
2. Columbia River Thermal Effects Study,
Volume II Temperature Prediction
The reports originated from EPA Region X and are primarily a product
of work conducted at the Pacific Northwest Water Laboratory. We
trust that the information contained in these reports will be of value
to your office.
Lowell E. Keup
Enclosures
Addressees:
Office of the Administrator
Office of Media Programs
Office of Categorical Programs
Office of Standards and Enforcement and General Counsel
Office of Research and Monitoring
R & D Project Reports System
EPA Public Information Office
Office of Water Quality Library
EPA Library
Interim Regional Coordinators
National Water Laboratory Libraries
WRSIC
Commerce Clearing House
Library of Congress
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COLUMBIA RIVER THERMAL EFFECTS STUDY
VOLUME I: BIOLOGICAL EFFECTS STUDIES
Environmental Protection Agency
in cooperation with
Atomic Energy Commission
and the
National Marine Fisheries Service,
U.S. Department of Commerce
January 1971
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FOREWORD
The Columbia River Thermal Effects Study was undertaken in pur-
suit of the policies and objectives of the Federal Water Pollution
Control Act, particularly as amended by the Federal Water Quality
Act of 1965, which required the establishment of water quality stan-
dards for the protection and enhancement of water quality throughout
the United States. In the process of establishing standards during
the period 1965-1968, the State and Federal water pollution control
agencies recognized water temperatures as an important factor affect-
ing water uses, both directly, as in the case of aquatic life, and
indirectly, as in the synergistic effects of temperature with other
parameters such as dissolved oxygen. In attempting to define temperature
requirements in the standards, however, pollution control authorities
encountered insufficient scientific knowledge and agreement on the
precise limits needed to protect water uses.
The Columbia River Thermal Effects Study was initiated in
July, 1968, in response to the specific problem of two inconsistent
temperature standards adopted for the Columbia River by the
States of Oregon and Washington, which share it as a border. Before
attempting to resolve these inconsistencies, the State and Federal
pollution control agencies could benefit from improved knowledge on
the temperature requirements and tolerances of the Columbia's Pacific
salmon and improved techniques for evaluation and prediction of the
temperature in the Columbia system. The report of the study con-
sists of two volumes. The first concerns the biological effects of
water temperature on Pacific anadromous fish in the Columbia River
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system. The second describes the application of mathematical models
to the Columbia River for prediction of water temperatures.
In the Pacific Northwest, standards were generally required to
protect the economically important Pacific salmon, a cold-water
anadromous species. The upriver runs of Columbia River fish resources
have been reduced and endangered by the physical alteration and blockage
of migration routes by the Nation's largest system of dams and
reservoirs. The quality of the aquatic environment has also been
modified by the discharge of pollutants and impoundment of the river's
flow in a series of reservoir lakes reaching into Canada. Particularly
regarding temperature quality, the Columbia River temperatures have
been both spatially and temporally altered by man's activities and
use of the water resources of the Region.
At about the time standards were established, public and
private electric power interests in the Northwest announced forecasts
of vastly increased power demands. The hydroelectric power potential
of the Northwest is nearly exhausted, and thermal power sources are
planned to meet future needs. This presented further potential for
modification of the thermal regime of the Columbia River system.
Initially, power producers assumed the possibility of using Columbia
River system waters for once-through cooling at thermal power plants.
The prospect, however, of numerous discharges of large quantities of
heated effluents to inland waters has since prompted the Region's
water pollution control agencies to issue policy statements which
require complete offstream cooling for thermal power plants located
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on inland waterways in the Basin. Power planners have accepted this
policy of offstream heat controls throughout the Basin.
Among the remaining environmental problems associated with in-
creased thermal power production is the projected use of the
existing hydroelectric system for power-peaking, with thermal units
providing the baseload, or firm power. The potential water quality
effects of exaggerated flow modification caused by these peaking
operations emphasizes the need to understand the existing thermal
regime of the Columbia River system. The prospects of industrializa-
tion, upon which the power demands are based, hold further potential
for environmental impacts which would require sound standards and
controls.
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CONTENTS
Chapter Page
I. INTRODUCTION 1
Purpose 1
Scope 1
Authority and Participants 5
Report Organization 7
Fish Species Studied 8
The Physical Environment 10
II. SUMMARY OF CONCLUSIONS 17
General 17
Adult Anadromous Fish 17
Juvenile Anadromous Fish 19
Non-Salmonid Fish 21
Secondary Production Organisms 21
III. GENERAL CONSIDERATIONS 23
IV. ADULT ANADROMOUS FISH 31
Adult Migration 31
Adult Migration Timing ' 31
Thermal Block 31
Location in Stream Cross-Section 34
Infectious Diseases 35
Gas Bubble Disease 38
Adult Thermal Resistance 40
Spawning 42
Effects of Temperature on Spawning 42
Ability to Spawn Volitionally 44
Egg Incubation 45
Importance of Natural Temperature
Fluctuation 45
V. JUVENILE ANADROMOUS FISH 49
Rearing 49
Growth Rate of Juvenile Salmonids 49
Attraction of Juveniles to Warmed Water 53
Juvenile Migration 53
Juvenile Migration Timing 53
Distribution of Juvenile Migrants in
Stream Cross-Section 58
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CONTENTS (Cont.)
Chapter Page
Passage of Juvenile Migrants Through
Thermal Plume 59
Thermal Resistance of Migrating Juveniles .... 61
Predation 68
Infectious Diseases 69
Gas Bubble Disease 70
Toxicity 72
Beneficial Effects 73
VI. THERMAL EFFECTS ON NON-SALMONID FISH 75
VII. SECONDARY PRODUCTION ORGANISMS 77
Specific Foods 77
Plankton 78
Thermal Effects on Secondary Organisms 78
BIBLIOGRAPHY 81
APPENDIX A. Manuscripts 89
APPENDIX B. Membership of the Technical Advisory
Committee for Biological Effects, Columbia
River Thermal Effects Study 97
APPENDIX C. State Water Temperature Standards 99
APPENDIX D. Scientific Names of Fish and Other
Aquatic Organisms 101
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FIGURES
Number Title Page
1 Columbia Basin Map 2
2 General Timing of Salmonid Activities,
Columbia River 11
3 Temperature-Flow Profiles of Columbia
River, 1967 13
4 Schematic Representation of Temperature
Requirements for Life Processes of the
Pacific Salmon 26
5 Thermal Response of a Hypothetical Cold-Water
Fish 27
6 Timing and Relative Abundance of Adult Anadromous
Fish Migrations Past Prescott, Oregon 32
7 Relative Abundance of Young Anadromous Fishes and
Sturgeon in the Lower Columbia River, in Relation
to Season and Water Temperatures 57
8 Thermal Resistance of Juvenile Salmonids 64
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I - INTRODUCTION
Purpose
The purpose of this report is to present the available knowledge
on temperature requirements and tolerances of anadromous fish in the
Columbia River. The material is intended for use by State and Federal
water quality agencies in their ongoing programs for prevention,
control and abatement of pollution, particularly in connection with
water quality standards.
Scope
The information and data presented in this report are limited to
conditions and species found in the main stem Columbia River in those
areas remaining accessible to anadromous fish. Figure 1 presents a
map of the area. The material includes information from the literature
and from the files of Northwest fisheries agencies, with particular
emphasis on new knowledge developed by the research studies conducted
as part of the Columbia River Thermal Effects Study (CRTES). Although
related scientific work is reported in the literature on thermal
effects on salmonids and other fish, much of it is not directly
applicable to Columbia River conditions or fish. Therefore, references
shown in the bibliography are limited to studies having direct
application to the Columbia River species and habitat.
The research studies conducted as a part of the CRTES, and listed
in Appendix A, were designed to develop immediate answers to the needs
of water quality agencies of the Northwest in considering the adequacy of
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FIGURE I
Columbia River Basin
Shading indicates
present spawning areas
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water quality criteria limits and goals for water temperatures
in the Columbia River. In all the temperature criteria adopted
in the Northwest, two conditions were set: maximum upper limits
for protection of aquatic life, and incremental increases within
those upper limits. Therefore, the studies were oriented to more
specifically define both the maximal temperatures for salmonids
and the tolerances of those fish to changes in temperatures within
that range. The studies were further oriented to the particular
temperature-modifying influences found or anticipated in the Columbia
Basin: impact of impoundment and reservoir releases and discharges
from thermal power plants. Although the latter thermal source has
since been obviated by water quality agency preventive policies,
much of the study emphasis related to questions of large thermal dis-
charges, and this material is included for academic value.
The field studies were limited to the most critical aspects of
the lethal and sublethal thermal effects at various life stages of
the anadromous fish. It was recognized that two years is not
adequate time to conclude research of a subject of this complexity.
However, by concentrating efforts on those areas of immediate concern,
much could be learned upon which to base the needed decisions. By
their nature, many of the projects conducted as a part of the study
are site-specific to conditions in the lower Columbia River near
Prescott, Oregon and in the Hanford reach of the Columbia River.
Prescott is the proposed site of the first privately-developed nuclear
power plant in the Northwest; the Hanford reach is the last
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5
unimpounded reach of the river above the estuary and the site of the
only existing nuclear plants in the Region, operated by the Atomic
Energy Commission.
Authority and Participants
The CRIES was authorized by the Secretary of the Interior in
February, 1968.' In his approval of the water quality standards
for the State of Washington, the Secretary recognized that the temp-
erature criteria set by the State for the Columbia River were in-
consistent with those set by the State of Oregon for the same waters.
Rather than disapprove the temperature criteria portion of Washington's
standards, or reconsider approval of Oregon's standards, the Secretary
directed that the Thermal Effects Study be completed to provide
further knowledge with which to reconsider the adequacy of temperature
criteria on the Columbia River.
The Northwest Regional Office of the Environmental Protection
Agency was directed to provide leadership in those studies. To take
_!_/ Two Presidential Orders for reorganization took effect in the
interim between study initiation and report publication. The first
was effective October 3, 1970, and created the National Marine Fisher-
ies Service in a new National Oceanic and Atmospheric Administration
in the Department of Commerce to replace the former Bureau of Commercial
Fisheries in the Department of the Interior. The second was effective
December 3, 1970, and created an independent Environmental Protection
Agency to incorporate many Federal programs concerning the environment
and abolishing the Federal Water Quality Administration (formerly the
Federal Water Pollution Control Administration) in the Department of
the Interior. The water pollution control responsibilities and author-
ities of the Secretary of the Interior were thus transferred to the
Administrator of the Environmental Protection Agency.
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full advantage of the expertise, programs, and facilities already
devoted to research on the effect of heat and anadromous fish in the
Columbia, the National Marine Fisheries Service (NMFS) and the
Atomic Energy Commission were included as participants in the research
studies. The Atomic Energy Commission's (AEC) research was conducted by
Battelle Northwest (BNW) which operates the AEC's Pacific Northwest
Laboratory (PNL) at Richland, Washington. The research program was
developed by the three agencies, with many ongoing studies and research
projects used or adapted as part of the CRTES. At the conclusion of
each project, the research report was furnished to the Environmental
Protection Agency (EPA) for incorporation into the biological effects
report. These studies are referenced as appropriate to the subject
and scope of this report and are not summarized in total. EPA has not
assumed responsibility for publication of the individual research reports,
The author agency should be contacted for copies or further information
on any of the contributing manuscripts, listed in Appendix A.
To advise in the conduct of the research program, the ad hoc
Technical Advisory Committee on Biological Effects was organized.
Advisory Committee membership (Appendix B) includes State and Federal
fisheries and water quality agencies, power company representatives,
and Federal power and water management agencies. The Advisory Committee
reviewed research study proposals, which improved the studies and
avoided duplication of effort.
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Report Organization
Chapter II presents a summary of the major conclusions
bearing on the water quality requirements of the Columbia River
fish resources. Chapter III, entitled "General Considerations,"
develops a framework of concepts and definitions which are
applicable to the biological information presented in this report
and which transcend the various life history stages which are
discussed in subsequent chapters. The remainder of the report
presents the compilation of available knowledge pertinent to the
temperature requirements of Columbia River fish. It is organized
into four chapters: adult anadromous fish, juvenile anadromous
fish, non-salmonid fishes, and secondary production organisms.
Discussion of thermal tolerance and temperature effects is presented
in each chapter as appropriate.
The discussion in Chapters III and IV roughly corresponds
with the succession of activities of an anadromous fish: e.g.,
juvenile migration timing, rate of movement, distribution, etc.
The relevance of some of the subjects to thermal effects may not
be obvious to the reader. If one were considering the effects on
a migrating salmon of a single point source of heat or a general
warming of a river, one would ask a variety of questions regarding
the fish's activities. This report presents information to answer
many of these questions.
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8
Fish Species Studied^/
To meet the immediate study needs, the Biological Effects Report
and studies center upon the fish of commercial and sports value in
the Columbia system, particularly the Pacific salmon. The Columbia
River is inhabited by four species of Pacific salmon: sockeye salmon,
chum salmon, coho salmon, and chinook salmon. The chinook salmon
has distinct spring, summer, and fall races, while the other species
do not. Another significant anadromous member of the salmonid family
is the steelhead trout, which has both summer and winter races. Other
anadromous fish of the Columbia Basin include several other species
of trout, smelt, and American shad.
As anadromous species, these fish seek out Columbia system waters
far inland from the Pacific Ocean, historically reaching as far as
Canada. They may presently be found in nearly all accessible tribu-
taries where water quality conditions allow. Figure 1 shows the areas
which remain accessible to anadromous fish since construction of the
system of dams and reservoirs on the river. The area of concern for
these studies was limited to the main stem, which is important princi-
pally as the migration route to all available spawning ground in the
Basin and for seaward migration to the Pacific Ocean. Spawning also
occurs in the only remaining unimpounded waters of the main-stem
Columbia, the fifty-mile reach below Priest Rapids dam, which produces
large numbers of chinook salmon.
2j Scientific names of fish and other organisms mentioned in
this report are listed in Appendix C.
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9
The size of the Columbia River fish resource is most easily de-
scribed in terms of the fish count over Bonneville Dam. In addition,
it should be remembered that the Willamette River, Cowlitz River and
smaller tributaries produce significant additional numbers of fish,
and that the commercial and sports catch of these fish in the River
and Pacific Ocean from Alaska to California should be added to the
Bonneville count. The thirty-one year record of fish passing over
Bonneville Dam gives an annual average count of 705,875 salmon and
steelhead of all species. The populations of the various species
can be summarized as shown in the following table. The total numbers
of fish entering the Columbia River in recent years is considered
by fisheries agencies to be as high as in the 1930's. However, the
size of the fish resource is now largely dependent upon artificial
production. Also, the distribution of species and races making up
the total has been changed by man's alteration of the river system.
AVERAGE ANNUAL FISH COUNTS AT BONNEVILLE DAM BY SPECIES
1938-1968
Chinook salmon 385,252
Sockeye salmon 100,777
Coho salmon (Three-year average 1965-1968) 77,289
Chum salmon 1,214
Steelhead trout 141,343
Source: Annual Fish Passage Report, Columbia River Project, U. S.
Army, Corps of Engineers, North Pacific Division, Portland,
Oregon.
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10
All of the salmonids have similar life histories or cycles, but
each species and race matures at different rates, presenting differ-
ences in the duration and time of occurrence for the various stages of
activity: spawning and incubation, rearing, juvenile migration,
growth at sea, and adult migration. For example, salmon fry spend
from a few weeks to over a year in fresh water before beginning sea-
ward migration, depending upon the species and race. Similarly, the
amount of time spent at sea also varies from less than one year for
o /
jack salmon^' to over five years, depending upon the species and race
of fish. Figure 2 illustrates the time of occurrence during the year
of the various stages of activity for the chinook, coho, and sockeye
salmon, and for steelhead trout. As can be seen, migrating adults or
juveniles can be found in the Columbia River throughout the entire
year. (See also Figure 6.)
The Physical Environment
The Columbia River remains accessible to anadromous fish as
far as Chief Joseph Dam, river mile (RM) 545. Throughout this
length, two reaches are unimpounded: the estuary to Bonneville Dam
(RM 145) and the Hanford reach between the head of McNary Pool (RM
347) and Priest Rapids Dam (RM 397). The remainder of the river
is impounded behind nine run-of-the-river dams, with shallow reservoirs
ranging from 54 to 165 feet in depth (Figure 1).
3/ Jack salmon are sexually precocious males.
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ADULT MIGRATION
SPAWNING * INCUBATIOM
REARING
JUVENILE MIGRATION
ADULT MIGRATION
SPAWNING fc INCUBATION
REARING
JUVENILE MIGRATION
ADULT MIGRATION
SPAWNING * INCUBATION
REARING
JUVENILE MIGRATION
ADULT MIGRATION
SPAWNING * INCUBATION
REARING
JUVENILE MIGRATION
CHINOOK £23 = period of actvity
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11! \ 1 :S ? 1 I ( j P i : i. £ ; &; = £ -Si: £ i ; ;i ::fe;Hil ;:;;ii;x:i;i:l^|||
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STEELHEAD
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.-.... . .".'. .'.- .' -,'. "- . <* '- ' '-' '' '-' ' ' '' ' ' X* ' ' "'." «' ' '' -"' " " -' '"'I"" ' * - "' " ' XM~. ^ '.' *. '-" ' ' f '-' ' '."'.*-" ." ' ' "' '-" ' f- ' '-'' ' ' '' '-'\
111 111 xiilf!!i il|| ill iii i ; 1 i I- ; ;l|! 11 iill Jli^liilliilill^llsliillii;^^^
JAN FEB MAR APR IWXY JUN JUL AUG SEP OCT NOV DEC
Figure 2. General Timing of Salmonid Activities,
Columbia River
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Flow reversals in the tidal reach of the Columbia River are com-
mon at least as far upstream as Prescott (RM 72.8) and occur most
frequently during the minimum flow period, August to October (Battelle
Northwest, anon 1969; Snyder and McConnell, ms 21)-^/. The effect of
a flow reversal is to accumulate effluents which may be adverse to
fish.
The surface runoff in the Columbia Basin is characterized by a
typical snowmelt regime, with low flows during the late summer and
winter and high flows during the spring and early summer. The volume
of runoff in the Columbia ranks it as the fourth largest in the North
American Continent; the average annual runoff at The Dalles, Oregon
is 140 million acre-feet. Mean annual flow is 195,400 cubic feet per
second (cfs); mean monthly discharges range from 95,700 cfs in
January to 494,700 cfs in June.
With minor exceptions, the water quality of the Columbia River
other than temperature and dissolved nitrogenis not considered
detrimental to the fish resources. Water temperatures in the main
stem Columbia River to Priest Rapids Dam range from maximums in the
70's during warm summer months to lows in the 40's and occasionally
in the 30's during winter. Figure 3 shows water temperature and dis-
charge profiles for various points on the Columbia River to
illustrate general temperature and flow conditions.
_4/ To enable the reader to determine whether a reference in the
text pertains to a study done in the CRTES or to other work, manuscripts
prepared as part of the CRTES are referred to by author's name and a
manuscript number which is listed in Appendix A.
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13
JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC
600
-MAX. MONTHLY MEAN TEMP
JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC
V) 80
UJ
UJ
CC 7O
LU
Q 6O
UJ 50
or
ID
CC
UJ
£L 3O
BONNEVILLE DAM
MAX. MONTHLY MEAN TEMP
600
500
400
300
200
100
UJ
t9
o:
<
I
u
<
LU
UJ
I- 80
70
60
50-
40'
30
JAN FEB MAR APR MAY JUN JUL AUG StP OCT NOV DEC
BEAVER ARMY
TERMINAL
MAX. MONTHLY MEAN TEMR
JAN FEB' MAR APR MAY JUN JUL AUG SEP OCT NOV DEC
600
500
400
300
200
100
0
K
Z
O
Figure 3. Temperature-Flow Profiles of
Columbia River, 1967
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14
Changes in the temperature regime of the Columbia River in terms
of time of occurrence of seasonal temperature shifts and location of
temperature peaks have been noted. Analysis by the U. S. Geological
Survey of temperature data at Bonneville Dam in the period 1938 to 1966
indicates that a man-caused increase in Columbia River temperatures
has occurred beginning in the mid-1950's (Moore, 1968). Other studies
by Jaske and Goebel (1967) and Davidson (1969) have reported changes
in the natural temperature regime in the Columbia River main stem,
with the most significant changes being above the confluence of the
Columbia and Snake Rivers. The principal effect has been the shifting
of water temperature maximums so that they occur later in the year.
The shift in seasonal temperature pattern has been attributed to the
construction of numerous dams in the Basin. The Hanford works is
one of the largest identifiable sources of advective heat on the
Columbia River.
Potential for further modification of the temperature regime
of the river is offered by proposals for construction of a third
powerhouse at Grand Coulee Dam, additional storage in the Columbia
River treaty dams, proposals to construct Ben Franklin Dam to impound
the Hanford reach of the Columbia, effects of power peaking, and the
potential for numerous small thermal discharges accompanying the
economic growth predicted for the Region. Use of the Columbia system
of reservoir releases for temperature management has been demonstrated
by cooperative use of the Grand Coulee release program to lower
Hanford water intake temperatures (Jaske, 1966). The use of
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15
mathematical models for temperature prediction in the Columbia River
offers potential for developing limited temperature enhancement by
the water management system. The AEG simulation model COL HEAT is
reported by Jaske (op. cit.). The temperature prediction capability
developed by the EPA as a part of the Thermal Effects Study is
presented in Volume Two.
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II - SUMMARY OF CONCLUSIONS
This summary presents the generalized conclusions drawn from
studies described in the literature and those conducted as part of
the CRIES.
General
Many variables influence the thermal tolerance of Pacific salmon.
However, within broad ranges, the fisheries agencies of the Pacific
Northwest have recommended optimum temperatures which are conducive to
the production of fish resources in the Columbia Basin. These ranges
are:
Migration routes: 45 to 60 F (7.2 to 15.6 C)
Spawning areas: 45 to 55 F (7.2 to 12.8 C)
Rearing areas: 50 to 60 F (10 to 15.6 C)
Individual races of salmonids may have adapted to a wider range of
temperatures and other factors may ultimately affect the optimum range.
Nevertheless, these temperature ranges remain accepted as the most
desirable conditions which can be considered conducive to enhancement
of the fish resources of the Columbia Basin.
Adult Anadromous Fish
There is an increasing tendency for adult salmon and steelhead
trout to cease upstream migration at temperatures of 70 F (21.1 C)
and above. Existing evidence is not conclusive on whether migration
would be blocked or curtailed at temperatures below that level.
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18
Adult salmon and steelhead appear to have specific rather than
random migration routes and prefer to be close to the shoreline in
shallow water.
The frequency of infection from fish diseases endemic to the
Columbia River increases with increasing temperatures. Temperature is
but one of the stresses which may interact to instigate the effects of
disease organisms. Warm water infectious diseases are less likely to
cause mortalities at temperatures below 60 F (15.5 C).
Under conditions of nitrogen supersaturation, any temperature
rise increases the chance of producing gas bubble disease in salmon
and steelhead. Fish stressed by nitrogen supersaturation are more
susceptible to the effects of thermal stress and infectious diseases.
Laboratory experiments have shown that water temperatures at
69.8 F (21 C) and above are considered to be directly lethal to
more than half of adult salmon and steelhead exposed to that level;
that is, temperature alone would kill the fish at that level. However,
the effects of temperature in combination with other stresses (disease,
nitrogen supersaturation, etc.) can be considered more important
than direct temperature effects in terms of ultimate fish survival.
The discharge of heated effluents from the Hanford Atomic Works
has not been demonstrated to have a detrimental effect on salmon
spawning downstream or upon migrating adults.
Temperature stresses placed upon adult salmon and steelhead may
indirectly and adversely affect reproduction through excess energy
costs, disease, and increased toxicity
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The initial egg incubation period is critically sensitive to
temperature, and requires a range of 42 to 61 F (5.5 to 16.1 C)
although a lower upper limit is preferable for general application.
Diurnal temperature fluctuation at temperature levels within
the range 42 to 65 F (5.5 to 18.3 C) is of significant benefit
to the growth and survival of eggs and fryas opposed to constant
temperature levels.
Juvenile Anadromous Fish
The growth of juvenile salmonids is enhanced at temperatures
within the range of 41 to 62.6 F (5 to 17 C). Heat additions to
colder waters which would bring the water temperatures within this
range might be considered beneficial, while heat additions which
would result in temperatures above this range would be considered
detrimental to juvenile growth. The optimum temperature for growth
changes with seasons and depends on food availability.
The timing of juvenile downstream migration has been signifi-
cantly delayed due to impoundment by dams; the delay has many adverse
effects on the fish, including subjection to unfavorable temperatures.
Fall chinook produced in the Hanford reach apparently maintain their
historical migration pattern (March until mid-June) from that point,
although their fate downstream has not been determined. The period
of peak outmigration for all species in the lower Columbia River is
from March to June, with smaller numbers moving throughout the year.
The smaller juveniles migrate closer to shore, while some larger
fish and selected species are found more evenly distributed throughout
the stream or in mid-channel.
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20
Based on Hanford area research, the number of downstream juvenile
migrants affected by the Hanford thermal plumes is probably small.
The risk of a directly lethal dose is also small, due to the velocity
of the stream and rapid mixing.
In defining the thermal resistance of juvenile salmonids, the
principles of thermal dose and acclimation apply: the effect of
rapid temperature increase is directly proportional to duration of
exposure. Acclimation to higher temperatures within the tolerance
zone significantly increases thermal resistance. Fish stressed by
nitrogen supersaturation, toxicants, or disease have reduced
temperature resistance.
The upper allowable temperature for any species of juvenile
salmonid should be a minimum of 5.4 deg. F (3 deg. C)' below the
ultimate upper lethal temperature to avoid significant curtailment
of activity. Temperatures above 68 F (20 C) are considered to be
adverse for juvenile salmonids. Temperatures near 62.6 F (17 C) may
be considered to be at the upper end of the optimum temperature
range. Between 62.6 and 68 F (17 and 20 C), any increase in temper-
ature probably is of little benefit to juvenile salmonids and
increases the likelihood of disease infection and other indirect
effects .
5_/ To distinguish between temperature changes and actual degrees
of temperature, "15 deg. C" means a temperature change by that amount
from some base temperature; "15 C" refers to exact temperature on a
temperature scale.
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21
Under controlled laboratory conditions, predators were shown to
selectively prey upon fish whose behavior has been altered by temper-
ature stress as compared to unstressed control fish.
Nitrogen supersaturation in the river is a significant detrimental
factor weighing against successful downstream migration of juveniles,
and the degree of supersaturation is accentuated by temperature
increases. At nitrogen saturation levels over 115 percent, any
increase in temperature could be damaging due to temperature-gas
solubility relationships.
Non-Salmonid Fish
Columbia River smelt have a lower temperature preference than
salmonids. The adult females may be more susceptible to detrimental
effects of temperature than other fish.
Yellow perch and three-spine stickleback have a higher thermal
tolerance than salmonids; however, they require a chill period.
Secondary Production Organisms
Aquatic insects form a major part of the diet of chinook fry
in the Hanford reach of the Columbia River and of all salmonid
species migrating through the Lower Columbia in spring and fall
months. Zooplankton are the dominant food organisms from July
through October in the lower river.
The most abundant plankter (Daphnia pulex), which is an
important salmon food in the Lower Columbia River, is relatively
resistant to thermal shock (30 C for 15 minutes). Below the Hanford
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22
discharges, the maturity of an aquatic insect (Caddis fly) was
advanced two weeks due to a temperature increase of about 1 deg. C,
No changes in the growth pattern could be observed in another
bottom invertebrate, the Columbia River limpet.
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Ill - GENERAL CONSIDERATIONS
A discussion of the general effects of temperature on aquatic
life is presented in this chapter; that is, those concepts of thermal
effects which transcend the discussions of various life stages in the
following chapters. The material developed here will establish a
framework of definitions and concepts which will make succeeding
chapters more understandable.
The significance of water temperatures to aquatic life is better
understood if it is remembered that fish are cold-blooded; they
depend upon their environment to provide a livable temperature
range to allow their normal life functions without internal bodily
regulation. The extent of influence of temperatures upon the fish
has been described by Brett (1956).
Temperature sets lethal limits to life; it conditions the
animal through acclimation to meet levels of temperature
that would otherwise be intolerable; it governs the rate of
development; it sets the limits of metabolic rate within
which the animal is free to perform; and it acts as a
directive factor resulting in the congregation of fish
within given thermal ranges or movements to new environ-
mental conditions.
In the evolution of the species, the Pacific salmon and steelhead trout
have adapted their life activities to the naturally occurring conditions
in the Columbia Basin, so that the flow and water quality (including
temperature) characteristics of the Columbia River and its tributaries,
or even small creeks, have become the environmental requirements for
particular species and races of salmon and steelhead.
The anadromous fish exhibit a more complex pattern of life
activities than do resident fish that do not migrate. In their unique
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24
journey from the inland spawning ground downstream to the Pacific and
their return, years later, to spawn and die, the salmonids must adapt
to a variety of environmental changes and stresses. Their migratory
nature takes them from the clear and cold creeks with relatively
small volume into the mighty Columbia and finally into the saline
estuary and Pacific Ocean. The adaptation from fresh to salt water
alone is a physiological strain.
Thus, it should be remembered that, while temperature can be a
stress in itself, it may be of more importance to the survival of
the fish species as a synergistic agent, an influence upon the success-
ful adaptation to other water quality and physical changes which the
fish must face to fulfill their migratory instincts. Temperature
influences the levels of dissolved oxygen and nitrogen in the water,
the incidence of disease in the fish, their susceptibility to preda-
tion, as well as directly affecting their ability to migrate, spawn,
and develop. Thus, temperatures which may be within the thermal
tolerance zone, i.e., the temperatures "... at which the animal
will never die from the effects of temperature alone" (Fry, 1947),
can profoundly influence the impact of other environmental factors
upon the productivity of the fish resources.
The thermal requirements and tolerances of Pacific salmon and
steelhead cannot be precisely defined except in terms of several
variables: the species and race of fish, the stage of development,
previous or concurrent exposure to factors which interact synergistically
with temperature (e.g., disease, toxicity, nitrogen supersaturation),
previous acclimation temperatures, and duration as well as degree of
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25
exposure to a given temperature. The graphic generalization presented
in Figure 4 (Brett, 1960) depicts the concepts employed by biologists
in defining the thermal requirements of Pacific salmon. As can be
seen, the tolerance zone of temperatures which are conducive to repro-
duction is narrower than the range of temperatures necessary for growth
of the fish. Outside the tolerance zone, the resistance of the fish
to lethal temperatures (represented by the undulating line) is dependent
upon several factors mentioned above.
An important concept defining the resistance of fish to lethal
temperatures is acclimation. Fish acclimated to higher temperatures
within the tolerance zone are most resistant to temperatures above
that zone. In other words, acclimation of a fish to a warm tolerance
temperature will result in a higher lethal limit for that fish, and
vice versa. The influence of acclimation on thermal tolerance can be
seen in Figure 5. (For purposes of comparability among investigators,
the incipient lethal temperatures are generally defined as the levels
at which fifty percent of the test population would die.) As can be
seen from the figure, the upper level of the tolerance zone rises as
the temperature to which the fish has been acclimated rises. However,
in all cases there is an ultimate level at which the influence of
acclimation gives way and the temperatures become lethal regardless of
previous acclimation. This level is termed the ultimate incipient
lethal temperature. The diagonal line in Figure 5 is a reference line
to provide a ready indication of where the lethal temperature and the
acclimation temperature are the same.
-------
80
70
60-
SO-
40
32
25
20-
10
LETHAL ZONE
TOLERANCE ZONE
RESISTANCE ZOh/E
LETHAL ZONE
2
N>
RESISTANCE ZONE
o J. _
P F f
o > £
3 o a
o ~
o:
a.
ui
v.
I
AGE-YRS
EGG-HATCH-EMERGE-MIGRflrE YEARUNG GRILSE MATURING ADULT-SPAWNING STAGE
FALL WN. SR SU. FA. WN. SP SU. FA. WN. SP SU. FA. WN. SR SU. FA. WN. SEASON
Figure 4. Schematic representation of temperature requirements for life
processes of the Pacific salmon (adapted from Brett, 1960).
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27
30 -
25
o
o
2O
15
o
c
IO ~
Temperature
of "instantaneous" Death
UPPER ZONE OF THERMAL RESlSTANC
\\N\\\\\\\\\\\\\\\\\\\\
Ultimate Incipient Let no I Temperature
so%
Mortality
X:XZONE OF THERMAL TOLERANCE
\ LOWER ZONE
OF THERMAL
RESISTANCE \
5 10 15
Acclimation Temperature °C
Figure 5. Thermal response of a hypothetical
cold water fish (adapted from Brett,
I960, by Coutant, 1968).
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28
The levels of temperature which will be lethal to a fish are
further influenced by the time of exposure. The resistance to an
abrupt increase in temperature is a function of both the degree of
temperature rise and the time of exposure to that temperaturereferred
to as the thermal dose. Exposure to a lethal temperature for a sublethal
time will not result in fish death.
The fisheries agencies of the Pacific Northwest recognize the
influences of these many variables upon the thermal tolerance of
Pacific salmon. Within broad ranges, however, a committee of these
agencies has recommended optimum temperatures which are conducive to
the production of fish resources in the Columbia Basin (Snyder et al.,
1966).
Migration routes: 45 to 60 F (7.2 to 15.6 C)
Spawning areas: 45 to 55 F (7.2 to 12.8 C)
Rearing areas: 50 to 60 F (10 to 15.6 C)
The committee developed their recommendations after considering natural
temperatures for fish activities throughout the basin, as well as at
fish hatcheries where temperature problems have developed.
The technical committee's recommendations reinforced the work of
Burrows (1963) in which he similarly defined the optimum temperatures
for Pacific salmon, with one added stage. During egg and fry incubation,
after the 128-cell stage of development is reached, Burrows concluded
that temperature ranges may vary but should remain within 32 and 55 F
(0 and 12.8 C). The effect of fry size and time of migration on sur-
vival in different areas makes it impossible to define the optimum
temperature range during this stage more precisely.
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29
Although individual races of salmonids may have adapted to a
wider range of temperatures and other factors may ultimately affect
the optimum range, these recommended temperatures remain accepted by
the fishery agencies as the most desirable conditions which can
be considered conducive to enhancement of the fish resources of
the Columbia Basin. Even though temperatures which occur in the Columbia
River now exceed these optimums for periods during the year, those
conditions must be considered detrimental to the fish.
A number of adverse effects of existing river conditions have
been noted, as will be seen in succeeding chapters. The remainder
of this report concentrates primarily upon studies of the effects
of temperatures outside these optimum ranges and of abrupt changes
in temperature within these ranges. The optimum temperature levels
and concept should be borne in mind as these studies are reviewed.
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IV - ADULT ANADROMOUS FISH
The material presented in this chapter concerns the effects of
temperatures on adult anadromous fish from the time they enter the
Columbia River from the Pacific Ocean to the completion of spawning
in the upper reaches of the Columbia River and its tributaries.
The discussion centers first upon the migration period, then turns
to spawning conditions.
Adult Migration
Adult Migration Timing
Few studies have related temperature to the migration of adult
anadromous fish, although a number of biologists believe there is
evidence that temperature is an important factor. The timing of
migration and relative abundance of adult anadromous fish at
Prescott, Oregon, was developed by Coutant and Becker (1968) and is
shown in figure 6. Fish counts at the dams on the Columbia River
and major tributaries also show timing of migration.
Thermal Block
The question of whether increased water temperatures could creat<
a block to inland migration of adult fish has been raised. A number
of observations seemed to indicate a relationship between temper-
atures and migration.
In an extreme temperature year (1941), sockeye and chinook
salmon and steelhead trout were observed congregating in small,
hitherto unused, cold creeks near Bonneville and Rock Island Dams
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water Tempt., Bonneville Dam 1965
JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC
Figure 6. Timing and relative abundance of adult anadromous fish migrations past
Prescott, Oregon (Adapted from Coutant and Becker 1968).
a/
~~ Broken portion of temperature line represents levels above established criteria.
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33
when temperatures in the Columbia rose to 71 to 75 F (21.7 to 23.9 C)
(Fish and Hanavan, 1948). Similarly, high water temperature has
been attributed as a major cause of delayed migration of sockeye
salmon into the Okanogan River. When the Okanogan was 70 F (21.1 C)
or rising above that level, the fish would not leave the cooler
Columbia River to enter their natal Okanogan. Migration was resumed
when the temperature began to fall. Below 70 F (21.1 C) migration
was not blocked by rising or stable temperatures (Major and Mighell,
1967) .
To test the influence of temperatures on the selection or
avoidance of migration routes, the National Marine Fisheries Ser-
vice conducted tests at the Fisheries Engineering Research Laboratory
at Bonneville Dam. Although the results were not conclusive in
defining the existence of a temperature block, the adult salmon
and steelhead did show a preference for ambient or cooler temperatures
over migration channels whose waters were above 70 F (21.1 C). No
preference or avoidance of either cooler or warmer waters was shown
as long as the ambient water temperatures remained between 50
and 70 F (10 to 21.1 C) (Anon, 1965-1968).
It has been speculated that large thermal discharges to the
Columbia might cause a thermal block similar to that noted above
for natural situations. Tracking sonic tagged steelhead trout
and summer chinook salmon at Hanford has shown no such block,
however, at that location (Coutant, MS 35). Fish apparently
avoided the warmest areas of the river and travelled through
the section essentially unimpeded. There were no statistically
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34
significant differences in migration rates between downstream reaches
heated 3.6 to 5.4 deg. F (2 to 3 deg. C) above normal and the cooler
upstream reaches at times when summer water temperatures remained
generally below 68 F (20 C). Some fish changed course of migration
when they entered shoreline seepage areas of warmed water.
From available evidence it is concluded that there is an in-
creasing tendency for salmon and steelhead trout to cease upstream
migration at temperatures of 70 F (21.1 C) and above. Existing
evidence is not conclusive on whether migration would be blocked
or curtailed at temperatures below that level.
Location in Stream Cross-Section
To provide a sounder basis for water quality management
decisions, the question was raised whether salmon and steelhead
trout exhibit a preferred location in the stream cross-section,
either in distance from shore or in depth. Little data are avail-
able to relate fish location to the temperature stratification
which occurs in reservoirs and the mixing and dispersion patterns
of heated discharges.
Sonically-tagged adult steelhead traced upstream through
Ice Harbor Reservoir during the fall of 1969 usually travelled
relatively close to the reservoir shore in waters forty-feet
deep or less. The fish followed a fairly specific route through
the reservoir with distinct travelling, milling and resting areas.
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35
The fish favored the bottom area during September and October
but were surface oriented during November and December. The
slower rate of travel through the reservoir in November and
December is presumed to be due to lower water temperature (Monan,
Liscom and Smith, ms 2). Other studies of migration of sonic-tagged
fish in the unimpounded Hanford reach of the Columbia indicated that
both steelhead trout and chinook salmon migrated principally along
shorelines, in water less than 5 meters deep. Throughout the dis-
tance between Richland and Priest Rapids Dam, fish selected the
north (or east) shoreline. This selection was most pronounced in
the area of operating reactors which were located on the opposite
shoreline. The pattern was essentially the same in two years,
1967 and 1968 (Coutant 1969). The reason for the shoreline specificity
has not been determined.
Although the knowledge of migration routes in the Columbia
is sparse, the available evidence indicates that adult salmon and
steelhead appear to have specific rather than random migration
routes and that they prefer to be close to shoreline in shallow
water.
Infectious Diseases
Three diseases endemic to the Columbia River are recognized to
be the most serious to adult salmon and steelhead trout: columnaris,
furunculosis and ceratomxya. The first two are bacterial diseases
while the latter is due to a protozoan. High mortalities of the
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36
adult fish prior to spawning, particularly where they are confined,
e.g., a spawning channel, have been attributed to one or more of
these diseases. Pathogenic agents appear to be more of a problem
to adults than to juveniles during migration (Fujihara, 1968). Columnaris
has been subjected to the greatest investigation, especially with
regard to adult salmon. The relationship of disease incidence and
severity to water temperature conditions has been demonstrated by
several investigations (Ordal and Pacha, 1963).
Natural outbreaks of columnaris disease in adult salmon have
been linked to high water temperatures in the Fraser River, British
Columbia. Adult sockeye salmon were exposed to columnaris as they
entered the river, with resident fish probably providing the source
of infection. The pathological effects of the disease became
evident when water temperatures along the migration route, and in
spawning areas, exceeded 60 F (15.5 C). Prespawning mortality
reached 90 percent in some tributaries. Columnaris in the infected
sockeye spawners was controlled when temperatures fell below 57-58 F
(13.9-14.4 C) and mortalities were reduced (Colgrove and Wood, 1966).
In laboratory experiments using Columbia River water, adult sockeye
and coho salmon were examined for columnaris infection after being
held at temperature levels of 50, 62, 68, and 73 F (10, 16.7, 20, and
22.2 C). At 68 and 72 F (20, 22.2 C) dead fish frequently showed
the lesions typical of columnaris infection which are sufficient to
_7/ Pathological effects is the identification of disease
organisms by the technique of streaking from suspected areas of
infection onto nutrient plates for incubation and examination.
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37
account for death. Columnaris and other diseases were believed to
be responsible for the death of many of these fish, although 72 F
(22.2 C) was considered directly lethal to most of the adult salmon over
a period of a week. Adult sockeye and coho were thoroughly exposed
to columnaris disease at all temperature levels but infections failed
to develop at the 50 F (10 C) level, and were not lethal at 62 F
(16.7 C) (Bouck, et al., ms 1).
Other studies of the occurrence of columnaris in the Columbia
River fish recognize the influence of higher temperatures on in-
fection, but also emphasize that many factors interact to cause
infection. Battelle Northwest studies in the Hanford area draw attention
to the resident infection carrier:
"The presence of immune carrier fish capable of releasing large
numbers of C^ columnaris organisms to the water is probably
the primary cause of salmonid infection in the Columbia River.
Fish ladders with high population density of coarse fishes
during migration of salmonids are the most likely sites of
exposure and infection (Fujihara, 1967). Since non-exposed
fish appear to be very susceptible to infection, the release
of heavy concentrations of the pathogen with the correspond-
ingly low water flow in ladders would create ideal conditions
for infection." (Fujihara and Nakatani, ms 43).
Data collected on antibody levels in Columbia River fish "...suggest
peak yearly effective infection of at least 70 percent to 80
percent of most adult river fish species" (Fujihara and Hungate,
ms 42). Occurrence of the disease was generally associated with
temperatures above 55 F (12.8 C); the authors further suggest that
the incidence of columnaris may be increased more by extended periods
of warm temperatures than by peak summer temperatures. Incidence of
columnaris was generally most severe in the Snake River and in the
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38
Columbia River for some distance below its confluence with the Snake
River. This may be due to Snake River temperatures being 1.8 to
5.4 deg. F (1-3 deg. C) higher than the Columbia River (Fujihara
and Hungate ms 42).
The above studies demonstrate that certain diseases endemic to the
Columbia River become more infectious with increasing temperatures.
However, the disease cause and effect relationship is understood only
in general terms of stress factors, one of which is temperature.
Other factors including the general condition of the fish, nutritional
state, size, presence of toxicants, level of antibody protection,
exposure to nitrogen supersaturation, level of dissolved oxygen,
and perhaps other factors interrelate in the infection of fish
by diseases. However, the diseases discussed here are of less
importance at temperatures below 60 F (15.5 C) ; that is, in most
instances mortalities due to columnaris are minimized or eliminated
below that level.
Gas Bubble Disease
Gas bubble disease occurs when the river water is supersatur-
ated with dissolved nitrogen gas. The disease is characterized by
the appearance of macroscopic gas bubbles under the skin or in
the fins and in the blood. Early observations of this phenomenon
were related to hatchery water supply systems (Rucker and Hodgeboom,
1953). The histopathology of gas bubble disease has been described
as interruption of the blood supply to an organ when gas bubbles
(embolism) form in the blood vessels. An embolism reaching the
heart can cause death (Pauley and Nakatani, 1967; Bouck, et al., 1970).
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39
Nitrogen supersaturation has been recognized as a critical
problem in the Columbia River and, during severe periods, essentially
all impounded areas and certain free flowing sections of the Columbia
and Snake Rivers which are accessible to salmon and steelhead are
significantly supersaturated. The condition occurs when the release
of water over the spillways at dams forces the normal nitrogen levels
(100 percent equilibrium with air) to rise to supersaturation.
Some reduction in dissolved nitrogen levels would result from
passage of more water through turbines and less over the spillway.
Investigations into possible methods of reducing spill at the
dams on the Columbia and Snake Rivers and consideration of
other methods for controlling nitrogen leave little hope for
a significant reduction of the problem in the near future.
During the fall and winter, water is generally not spilled
over the dams and nitrogen saturation levels are normal; during
the spring and summer, spilling from the dams causes nitrogen
supersaturation of 120 to 146 percent (Ebel, 1969; Ebel, 1970,
A and B).
The harmful effects of high water temperatures on adult sal-
mon are worsened by prior or simultaneous exposure to nitrogen super-
saturation. Survival time of jack chinook acclimated to 62.6 F
(17 C) which were subjected to 71.6 water (22 C) was drastically
shortened when the warmer water contained dissolved nitrogen gas
levels in excess of 115 percent of saturation (Coutant and Genoway,
1968). Acclimation of the fish to water supersaturated with nitro-
gen reduced subsequent survival time at 71.6 F (22 C), regardless
of the nitrogen levels in the warmer water.
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40
During a period of high nitrogen supersaturation in the
Columbia River, adult sockeye salmon were removed from the river
and subjected to air equilibrated water at temperatures varying
between 50 and 72 F (10-22.2 C). Loss of eyes or blindness,
which was attributed to gas bubble disease, occurred in a third
of the fish at the 50 and 62 F levels (10-16.7 C), but increased
to one-half to two-thirds of the fish when temperatures were 68
and 72 F, respectively (20, 22.2 C) (Bouck, et al., ms 1).
The scope and exact nature of this nitrogen-temperature interaction
in fish is not well understood; however, sufficient evidence exists
to warrant the conclusion that any temperature increase intensifies
harmful effects of gas bubble disease, primarily due to temperature-
gas solubility relationships. Further, fish stressed by nitrogen
supersaturation are more susceptible to the effects of thermal
stress and infectious diseases (Bouck, et al., op cit.). Although the
full impact of nitrogen supersaturation upon fish is not completely
understood, evidence indicates that the condition results in signi-
ficant mortality of both adult and juvenile salmon and steelhead.
In addition, the disease can cause sub-lethal damage, such as blind-
ness, which may be essentially equivalent to death in terms of
damage to natural reproductive potential (Westgard, 1964; Bouck,
et al., ms 1).
Adult Thermal Resistance
For the purposes of this report, the concept of thermal
resistance refers to the resistance of fish to temperature levels
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41
outside their preferred or optimum range, and to abrupt changes
in temperature within that range.
The precise temperature level which would cause direct thermal
death to adult Pacific salmon has not been defined, due in part to
the difficulties in testing such large animals. The physiological
causes of direct thermal death to fish are still unknown, but are
presumed to involve a complex disruption of the cellular processes
by purely thermal (physical) means. One hypothesis suggests that
oxygen delivery to the nervous tissue lags behind metabolic usage
at the lethal temperature. In one study, 69.8 F (21 C) was esti-
mated to be near the ultimate incipient lethal temperature for
adult coho (Coutant, ms 38). (The incipient lethal level of water
temperatures in these studies was defined as the level at which
fifty percent of the animals died but the remainder survived for
an indefinite period of time.) For steelhead, the incipient lethal
level was estimated to be near 69.8 F (21 C), and for chinook,
near 69.8-71.6 F (21-22 C) (Coutant, op. cit.). These studies
"...suggest that elevation of general environmental temperatures
above 21 C would be directly lethal to adult salmon and steelhead"
(Coutant, op. cit.).
In other studies designed to determine the effects of temper-
ature during simulated migration of adult sockeye salmon, (Bouck,
et al., ms 1) the fish were subjected to four temperature levels:
a. 50 F (10 C), a theoretical optimum temperature;
b. 62 F (16.5C), Columbia River temperature on July 1, 1969;
c. 68 F (20 C), Columbia River legal temperature limit;
d. 72 F (22 C), an estimated directly lethal temperature.
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42
In these studies, adult sockeye salmon survived an average of 3.2
days at 72 F (22.2 C) and 11.7 days at 68 F (20 C). Because of the
sudden onset and rate of mortality, 72 F (22.2 C) was considered
to be directly and acutely lethal to adult sockeye salmon. Survival
of a few adult sockeye salmon for about one month indicated that
68 F (20 C) is not directly lethal.
However, as will be emphasized throughout this report, the
effects of temperatures in conjunction with other stresses is
equally or more important than direct lethal effects in terms of
ultimate fish survival. For example, in the studies above, 68 F
(20 C) was not considered an ultimately safe level for adult sockeye
salmon because the effects of other stresses such as gas bubble
disease and infectious diseases were aggravated at that temperature.
The added stress of holding the fish was no doubt an additional factor.
Spawning
Effects of Temperature on Spawning
The effects of water temperatures upon salmon spawning have
not been well defined in the literature, although the preferred
range is considered to be similar to that for egg incubation, 42-55 F
(5.5-12.8 C) . The range reported by the Columbia River Fishery
Technical Committee for spawning of all species in the Columbia River
tributaries is 34-67 F (1.1-19.4 C) (Snyder, et al., 1966).
Since the Hanford reach has been subjected to the possible
detrimental effects of heated effluents of the atomic reactors
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43
and is the only Columbia River main stem section remaining suitable
for salmon spawning, more studies have been directed to the con-
ditions in that area than elsewhere. Nakatani (1969) states that
most of the local fall chinook spawn in water temperatures about
50-59 F (10-15 C) but the late spawners in a cold year may deposit
their eggs in temperatures of about 41 F (5 C), well below the op-
timum .
Some of the conclusions reported by Watson (ms 39) on studies
of fall chinook salmon spawning in the Hanford reach of the Col-
umbia River over the period 1947-69 are: (1) Throughout this
period, no apparent relationship existed between numbers of fall
chinook salmon spawning in the Hanford reach and river temperature,
flow, or elevation during the spawning season. (2) Closure of
reactors immediately upstream from major spawning areas did not
alter the general distribution of the spawning fish. (3) The
assessment of any subtle biological effects of the reactors on
local salmon populations was not possible in the study. (4) Other
changes in the ecology of the river, such as produced by dams,
appear to be of greater influence on the numbers of locally spawn-
ing salmon than the reactor operation.
The major spawning areas between RM 363 and RM 376 (km 585-
605) lie downstream from heated effluent outfalls and have been sub-
jected to incompletely mixed effluents. In several years, salmon
spawning was observed within a hundred meters downstream from effluent
outfalls, but the exact nature of the temperature profile in this
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44
region of incompletely mixed thermal plume was not known.
A study in an area of incompletely mixed effluent during
the winter of 1954-55 showed the temperature difference between
water above and below the gravel surface was usually less than
1 deg. C. The distribution of spawning salmon in the river down-
stream from the effluent of four reactors showed little change after
the reactors were no longer in operation. The location of spawning
areas appears to be primarily influenced by gravel size, current
velocity, and water depth (Nakatani, 1969).
Overall, these studies indicate that there is no evident
relationship between the operation of the Hanford reactors and
numbers of fish spawning in the Hanford reach of the river.
Ability to Spawn Volitionally
The effect, if any, of water temperatures upon the ability
of the salmon to spawn has been subjected to only limited study.
Information gained by Bouck et al. (ms 1) in adult salmon studies
at Bonneville showed no apparent adverse effects to eggs in
utero of coho salmon from prolonged exposure of the adult fish
to 62 F (16.7 C). At 68 F (20 C), prespawning mortalities were
sufficiently high to preclude judgment of temperature effects on
eggs in utero. Aside from the more direct effects of disease on
maturing salmon, it was observed that the increased metabolic rate
at higher temperatures manifests itself in abnormal body weight
loss and a corresponding decrease in weight of sex products.
These changes cannot be viewed as beneficial to migration or survival
of the fish to spawn.
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45
The additional stresses placed on a fish by higher temper-
atures may indirectly and adversely affect reproduction in terms
of excess energy costs, disease, and increased tpxicity. In other
words, although the exposure of the adult female to adverse tempera-
tures may have secondary effects which cause prespawning mor-
talities, the detrimental effects on the eggs in utero have not
been demonstrated.
Egg Incubation
Biologists concerned with hatchery production in particular
have been concerned with the effects of water temperatures on
salmon eggs at various stages of development. Several investigators
have explored the effects of high or low temperatures on salmon egg
incubation (Combs and Burrows, 1957; Combs, 1965; Olson and Foster,
1955) .
In studies of embryonic development, the order of appearance
of anatomical features in the embryo and certain meristic character-
istics of fish were shown to be dependent on temperature of incubation
(Hayes, et al., 1953; Hubbs, 1922). Premature hatching at low
temperatures increases the proportion of abnormal fry produced. The
number of vertebrae also depends on temperature.
Burrows (1953) found that egg and fry incubation temperatures,
after the 128-cell stage of development is reached, may vary but should
remain within the range of 32-55 F (0-12.8 C). A number of researchers
have reached the same general conclusion: to avoid excessive mortali-
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46
ties of eggs and resulting fry the initial egg incubation should not
be at a temperature higher than 59-61 F (15-16.1 C). The Columbia
Basin Fishery Technical Committee reached the following conclusions
after review of the results of several investigators (Snyder, et al.,
1966):
1. Differences in results of the several investigations on
temperature limits for incubation of salmon eggs are not too impor-
tant from a practical standpoint.
2. Temperature during initial incubation is critical.
3. If initial incubation temperature is below 42.5 F (5.8 C)
or above 55 F (12.8 C) less than normal survival can be expected.
4. Mortality attributable to temperature is also a function
of duration of exposure.
Experiments at Hanford in 1965 (Olson and Nakatani, 1968)
showed higher mortalities in eggs incubated in cold river water than
in lots incubated in water warmed by addition of reactor effluent.
Nakatani (1969) believes that warming the colder waters of the
Columbia River in early winter might actually be beneficial for
egg incubation.
From the many studies of egg incubation and temperatures, it
is generally agreed that the initial egg incubation temperature is
critical and should be within the range of 42-55 F (5.5-12.8 C) and
should remain below 58 F (14.4 C) at all times but may go as low as
32 F (0 C) after the 128-cell stage.
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47
Importance of Natural Temperature Fluctuation
Many of the laboratory studies to define the necessary tempera-
ture range for egg incubation were conducted at controlled constant
temperature levels. Since natural temperatures in the stream
fluctuate daily under the influence of solar radiation, the signifi-
cance of fluctuating temperatures should also be noted, although
studies of this aspect of egg incubation have been limited.
Olson and Nakatani (ms 45) investigated the effect of fluctuating
temperatures on incubating chinook salmon eggs and fry at increments
ranging 1, 2, and 3 deg. F above and below the basic river temperature.
No difference in survival was found as compared to nonoscillating
temperatures.
The Oregon Fish Commission and National Marine Fisheries Service
in 1969 cooperated on experimental incubation of fall chinook eggs
(Fulton, unpublished manuscript). On a simulated natural diurnal
temperature regime, peak temperatures were from 60 to 65 F C15.5-18.3 C)
with a 4 deg. F fluctuation. The results in both constant and
fluctuated temperature groups generally agreed with other investi-
gators; that is, above the optimum range the warmer the water, the
lower the egg survival. However, there was a significantly greater
survival in eggs incubated at fluctuating temperatures with peaks
above 63 F (17.2 C) and a significantly better survival for fry at all
temperatures (with one exception) in the fluctuated temperature group
when compared with constant temperature groups. This indicates that
there may be significant benefit to eggs and fry from a diurnal
temperature fluctuation at all levels within the zone of tolerance
42 to 65 F (5.5 to 18.3 C).
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V - JUVENILE ANADROMOUS FISH
The temperature requirements and tolerances of juvenile
anadromous fish, from the fry stage through seaward migration to the
Pacific Ocean, is the subject of this chapter. The discussion
centers first on the rearing stage in fresh water, a period which
may vary from one week to over a year depending upon the fish species
and race. Juvenile migration to the sea is then discussed in terms
of temperature effects.
Rearing
Growth Rate of Juvenile Salmonids
Two aspects of the relationship between juvenile growth and
water temperatures require attention. First, the effect of incre-
mental increases in water temperature over natural conditions should
be known; and second, the desirable range of temperatures for
enhancement of juvenile growth should be defined.
The effect of incremental increases in temperature over the
Columbia River seasonal pattern has been observed on young salmon and
trout (Olson and Foster, 1955; Olson and Nakatani, 1968). Increased
water temperatures clearly accelerated fish growth in all lots.
Fish in the warmest lots were as much as eight times heavier than
those in the coldest lots at the conclusion of the tests. Excessive
mortalities resulted from temperature increases in excess of 4 deg. F
over the river temperature for the lots spawned on October 30; but
an incremental increase of 12.5 deg. F over the base temperature for
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50
lots spawned on December 8 produced a mortality of only 12.4 percent.
The significance appears to be that 4 deg. F over the October 30
temperature results in the water being over 60 F, while 12.5 deg. F
added to the December 8 temperature produces 60 F or below. The
researchers believe their studies indicate that young fall chinooks
can safely withstand greater thermal additions during the winter and
suggest, further, that elevated temperatures will favor the survival
of the eggs spawned in the colder late months of the year over those
spawned earlier.
In other studies the effect of incremental increases of 2, 4,
and 4.7 deg. F (1.7, 2.2, and 2.6 deg. C) above Columbia River
temperatures was tested on the growth of juvenile steelhead (Olson
and Templeton, ms 46). The warmest of these heat increments tended
to slow the rate of growth when added to summer maximum temperature
waters (average of 70 F, 21.1 C) but favored the growth during the
colder seasons of the year.
Brett- et al. (1969) studied growth of fingerling sockeye salmon
in relation to water temperature and the metabolic rate of food
utilization. The most favorable range for growth was found to be
between 41 and 62.6 F (5-17 C), with a physiological optimum in the
vicinity of 59 F (15 C). The amount of food necessary to maintain
the fishes weight increased rapidly as temperatures rose above
53.6 F (12 C), with no growth occurring at approximately 73.4 F
(23 C), despite the presence of excess food.
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51
In field and lab studies with juvenile coho at Oregon State
University, Averett (1969) found that the range of temperatures at
which maximum efficiency of food utilization for growth appears,
changes with season and depends upon the range of consumption rates
being considered within a season. His findings show the most
efficient growth, within consumption ranges believed to occur in
nature, is at the temperature of 41 to 57.2 F (5 to 14 C) in early
spring, 51.8 to 57.2 F (11 to 14 C) in early summer, 57.2 to 62.6 F
(14 to 17 C) in late summer, 51.8 to 62.6 F (11 to 17 C) in fall,
and 41 to 46.4 F (5 to 8 C) in late winter.
Additional laboratory studies, also at Oregon State University,
compared growth rates of juvenile coho salmon kept at different
fluctuating temperatures (L. B. Everson, MS Thesis). Temperatures
were incrementally increased approximately 6.3 and 12.6 deg. F
(3.5 and 7.0 deg C) from control temperatures that followed the daily
and seasonal temperature changes of a natural stream. Relationships
were established between rates of food consumption and growth during
short-term (30 day) and long-term (8 month) studies. At comparable
ration sizes the growth rates of fish kept at elevated temperatures
were generally lower than those kept at control temperatures. The
response to elevated temperature was markedly influenced by ration
size; the greater the ration, the more nearly were growth rates
comparable at all temperatures studied. Comparison of results of
short and long-term studies indicated that the fish did not benefit
from long-term acclimation to elevated temperatures. Seasonal
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52
differences in food consumption, and consequently growth, that are
not directly related to temperature are indicated from comparison of
results obtained during different seasons but over similar ranges
of temperature.
Studies with juvenile coho salmon at control and elevated
temperatures 9 deg. F (5 deg. C) increment in two large outdoor
model streams at Oregon State University have shown that heating
enhanced growth only during the winter of the first year of operation.
Growth subsequently has been greater in the control than in the
treatment stream. The level of abundance of benthic invertebrates
appears to be closely related to the differences in growth of the
coho (R. A. Iverson, personal communication).
Becker (ms 40) studied the food, feeding, and growth of juvenile
chinook salmon at stations above and below nuclear reactor effluent
discharges in the Columbia River. He found a lack of detectable
thermal effects which he attributed to the fact that the thermal dis-
charge plumes are in mid-river and the effluents are well mixed before
reaching inshore feeding areas. The transient nature of the fish
populations, and the availability of food organisms in the river
drift complicated efforts to evaluate any subtle thermal effects.
To summarize, the growth of juvenile salmonids is enhanced at
temperatures within the range 41-62.6 F (5-17 C) depending on season
and availability of food. Heat additions to colder waters which
would bring the water temperatures within this range might be
considered beneficial, while heat additions which would result in
temperatures above this range would be considered detrimental to
juvenile growth.
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53
Attraction of Juveniles to Warmed Water
During cold-water periods in the Hanford reach of the Columbia,
seining in the inter-gravel seepage area near the B-C reactor showed
that small numbers of chinook fry were present in the area which
was warmed by the seepage to 50 to 53.6 F (10-12 C). The warming of
the areas is the result of seepage from an effluent disposal ditch.
No fry were available in the other areas of the river where the river
temperatures during March remained at 35.6 to 39.2 F (2 to 4 C).
Becker (ms 40) speculates that the young chinook select the warmer
areas when the river temperature is below a lower thermal preferendum
53.6 to 57.2 F; 12 to 14 C). He postulates that, because the river
temperatures in early spring are well below the preferred level for
juvenile salmon, the addition of heat by the Hanford discharges during
early spring is not harmful and may actually be beneficial to the
emergent fry (Becker, op. cit.).
Juvenile Migration
Juvenile Migration Timing
Although the annual timing of seaward migration of juvenile
salmonids varies according to species and race of fish, each, does
exhibit distinct timing patterns which are preferred and beneficial
to successful downstream migration. Migration has historically been
timed to coincide with favorable temperature and flow conditions
during the year. The construction of dams and reservoirs in the
Columbia system has altered the historical migration patterns.
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54
On the Columbia and Snake Rivers, studies have been made of the
time of juvenile salmonid migration under both pre- and post-
impoundment conditions. Mains and Smith (1964) found that before
impoundment the migration of zero-age chinook from the upper Columbia
River occurred during March through July. After construction of four
dams, Park (1969) found that the migration period had shifted to
April through August at Priest Rapids Dam, with peak numbers in
August during 1965 and 1966. However, since juvenile chinook were
found by Becker (ms 41) to be in the Hanford reach of the Columbia
from March until mid-June, he concluded that the progeny of fall
chinook spawning in the Hanford reach retain the historical migration
pattern detected by Mains and Smith (1964) above.
The migration rates of yearling chinook salmon in the Columbia
and Snake Rivers were measured by Raymond (1968, 1969). Marked fish
took 32 days to travel from the Salmon River to Bonneville Dam in
1966, a distance of 415.6 miles (669 km). He predicted that, with
additional impoundment of the Snake River, an additional 36 days
(total of 68 days) would be required for salmon to migrate from the
Salmon River and Grand Ronde River to Bonneville Dam.
Park (1969) observed that the peak movement of juvenile chinook
in 1966 at Bonneville Dam occurred two months later (June versus
April) than in the period 1946-1953. He also indicated that the peak
movement at Bonneville was influenced by the late release of many of
these fish from upstream hatcheries.
From these studies, it appears the time of downstream migration
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55
of juvenile salmonids through the upper and mid Columbia River has
been significantly delayed due to impoundment of the river by dams.
Migration of fall chinook produced in the Hanford reach of the
Columbia appears to commence in the same period as the historical,
pre-impoundment patternalthough whether delays in migration of
these fish occur in reservoirs downstream has not been considered.
The effect of delays in juvenile migration is to subject the
fish to increased stresses and adverse conditions. The detrimental
aspects of delay in migration and of the alteration of the physical
condition of the migration route into a series of reservoir environ-
ments have been described by a number of studies (Park, 1969;
Davidson, 1965; and Raymond, 1970, B). Juvenile salmon and steelhead
that are delayed in downstream migration may be forced to migrate
during an unfavorable time period in terms of seasonal temperature
rises in the Columbia River. Downstream juvenile migrants have
historically migrated during periods of high flows and moderate
temperature; delays in this pattern bring them into peak summer
temperatures and curtailed flow conditions.
Delayed entry into salt water also is considered detrimental.
Baggerman (1959) has suggested that the disposition for seaward
migration of juvenile Pacific salmon is directly related to an
observed salt-water preference. "Juvenile salmon exhibit a preference
for salt water prior to, during, and after the normal period of down-
stream migration." Therefore, it has been suggested that delays in
reservoirs could adversely affect production, since either premature
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56
or delayed entry into salt water appears to reduce the survival rate
(Andrew and Geen, 1960).
When delayed in the reservoir, the migrants encounter increased
temperature conditions caused by seasonal warming characteristics.
Several related studies support this theory. Apparently, salmonids
stop their migration when they encounter a temperature gradient
upon entering a reservoir from a stream (Durkin, et al., in press).
Other observations at storage reservoirs are unpublished. Juvenile
salmon and steelhead that are delayed may lose their desire, or even
ability, to migrate and become residual in the reservoir where they
are subjected to unfavorable temperature, nitrogen supersaturation,
increased predation, and disease.
The relative abundance of young anadromous fish in the lower
Columbia River in relation to season and temperature is discussed by
Coutant and Becker (1968) and is shown in Figure 7.
The generalized conclusions relative to juvenile migration
timing, are that the downstream migration from the upper and mid-
Columbia has been significantly delayed due to impoundment by dams;
the delay has many adverse effects on the fish, including subjection
to unfavorable temperatures. Fall chinook produced in the Hanford
reach apparently maintain the historical migration pattern of
departure from that point. Although inconsistent with the above
information, the period of peak outmigration in the lower Columbia is
from March to June, with smaller numbers moving throughout the year
(McConnell and Snyder, ms 12). Water temperature during this period
is within the optimum range for migration.
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I Chinook Salmon
Coho Salmon
SocKeye Salmon
American 'Shod
Ste«lhead Trout
20
19
o
o
"10
n
I
ft 5
Vertical Dimensions Indicate
RelativeAbundance Only
WATER TEMP BONNEVILLE DAM. 1965
JAN I FEB I MAR I APR I MAY I JUN I JUL I AUG I SEP I OCT I NOV I DEC
32
Figure 7. Relative abundance of young anadromous fishes and sturgeon in the
lower Columbia River, in relation to season and water temperatures
(Coutant and Becker, 1968)
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58
Distribution of Juvenile Migrants in Stream Cross-Section
The distribution of seaward migrant Chinook salmon in the stream
cross-section was studied by Mains and Smith (1964) near the con-
fluence of the Columbia and Snake Rivers. The horizontal distribution
was similar on both rivers, with some preference for the shore at
both locations. Vertical distribution favored the surface on the
Columbia, but there was more general distribution in the Snake River.
From several years of sampling in the Columbia River estuary,
the National Marine Fisheries Service has generally established
the migration patterns of juvenile salmon migrants from the mouth
of the river to Bonneville Dam. Although each species is different,
generally the smaller fish which migrate in their first year (fall
chinook, chum) are found in shallow water near the beaches. The
yearling migrants (spring chinook, coho, sockeye, and steelhead) are
most abundant in deeper water and more evenly distributed or, in
the case of sockeye, concentrated in mid-channel. A proportion of
the fall chinook has been found to hold for varying but extended
periods of time throughout the year in the vicinity of Puget
Island (RM 44).
The cross-section studies generally indicate that smaller
downstream migrants move closer to shore, while some larger fish
and selected species are found more evenly distributed or in
mid-channel.
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59
Passage of Juvenile Migrants Through Thermal Plume
When the current studies were being planned, the question per-
haps uppermost in the minds of fishery biologists considering thermal
effects was the fate of downstream migrating salmonids who might pass
through a thermal plume created by a thermal power plant discharge.
Since the Hanford Atomic Works had been discharging heated effluents
to the Columbia for many years and the AEC had contracted for studies
of the biological effects of the discharges, attention centered upon
those studies. Few others had seen the plumes and none had access to
the classified information concerning the temperature of the effluent
or its mixing characteristics. The AEC researchers subsequently
submitted papers for publication describing studies designed to answer
some of these questions (Becker and Coutant, ms 36; Becker, Coutant
and Prentice, ms 37).
Theoretically, an unknown portion of the downstream migrating
salmonids would pass through the mixing zones below the heated dis-
charges. To test the effects of this theoretical occurrence, a float-
ing trap was "fished" in the effluent plume (K reactor) for four days
in May 1968. Water temperatures in the trap ranged from 52.7 to 59.9
F (11.5 to 15.5 C) during this period. The catch consisted of 174
small chinook; eight mortalities occurred in this group which, were
attributed to mechanical injury. In a similar effort in October and
November of the same year, no fish were trapped.
Additional tests of thermal effects in 1968 and 1969 involved
drifting of caged juvenile salmon through the center of a plume.
Most drifts resulted in no loss of fish; however, one drift through
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60
the main K-reactor plume in mid-river resulted in total mortality.
Most drifts through the K-reactor plume resulted in low thermal
"doses" because the exposure duration was too short to cause excessive
mortality even though 77 F (25 C) was exceeded. The researchers
concluded from the two years of study that, under certain environ-
mental conditions (low river flow-high river temperature), the
potential clearly exists for lethal exposure to migrating juvenile fish.
The proportion of the juvenile salmonid migrants which would
enter the mixing zone in the mid-stream plume is not known. However,
it is considered to be small due to the disproportionately low ratio
of effluent volume to total water mass. The velocity of river flow
through the plume is 0.16 to 3.3 feet per second CO.5 to 1 meters per
second), and the lethal temperatures within the plume exist for only
a relatively short distance in the mixing turbulence before dispersal
to sublethal levels even at low flows. Approximately 80 percent of
the difference between plume temperature and background river tempera-
ture is lost within five seconds of discharge as measured at the
surface (Becker, Coutant, and Prentice, ms 37).
The area of perhaps greater potential danger is the inshore K-
reactor shoreline leakage area where total mortalities occurred on two
drifts of salmon through the area. The lower velocities and reduced
mixing rate make this area and any similar situations of potentially
greater concern.
General conclusions regarding the effects of thermal plumes in
the Hanford area upon downstream juvenile migrants, are that the
number of fish affected is probably small, as is the risk of a
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61
directly lethal dose, due to the velocity of the stream and rapid
mixing. The sublethal, or indirectly lethal, effects of plume
exposure are discussed under predation and gas bubble disease.
Thermal Resistance of Migrating Juveniles
As in the case of adults, the thermal resistance of juvenile
salmonids to temperatures outside their tolerance range, or to drastic
changes in temperature within their tolerance range, is a factor of
both the degree of temperature difference and the length of exposure.
In fact, thermal dose (time and temperature) is more significant in
the case of juveniles since their smaller mass allows the fish tissues
to heat up quicker. The preferred temperature range for juvenile
salmonids is 41 to 62.6 F (5 to 17 C) (Brett, et al., 1969; Averett,
1969).
The maximum swimming performance of juvenile coho salmon after
exposure to abrupt thermal changes was tested by Groves (ms 14);
temperature changes were within the range from 41.4 to 73 F (5 to 23 C)
He concluded that, if temperature alone is considered, a thermal
change does not reduce the ability of the fish to make maximal swim-
ming efforts. "Migrant juvenile coho salmon probably could routinely
experience sudden prolonged thermal changes spaced anywhere within
their acceptable thermal range and still be able to make the maximal
swimming efforts demanded by such life emergencies as rapid evasion
and defense."
In studies to further define the lethal effects of thermal shock,
four species of juvenile salmon plus steelhead were used in thermal
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62
shock studies by the NMFS (Snyder and Blahm, ms 8). Fish were ex-
posed to shock temperatures of 78.8 and 84.2 F (26 and 29 C) from
acclimation temperatures ranging from 39.2 to 68 F (4 to 20 C) .
One hundred percent mortality occurred in less than three hours at
78.8 (26 C) and in twenty-five minutes at 84.2 F (_29 C) . Fifty
percent mortality was recorded in less than two hours at 78.8 F (26 C)
and within eight minutes at 84.2 F (29 C). The death process was
found to be reversible at 78.8 F (26 C) but not at 84.2 F (29 C) if
the fish were returned to the control water temperature as soon as
loss of equilibrium was noticed.
A program of research on thermal resistance using Columbia River
water for experimentation has been conducted by the National Marine
Fisheries Service. Water pumped from the river was thermally adjusted
to simulate temperature conditions (Snyder, Blahm, and McConnell, ms 29).
All fish were acclimated in the river water at the experimental facility
prior to the test. Acclimation temperatures in the tests ranged be-
tween 50 and 68 F (10 and 20 C). In some cases, little or no mortality
occurred at a temperature below 73.4 F (23 C), while in a few cases
significant mortality occurred at temperatures as low as 62.6 F (1.7 C)-
The investigators concluded that:
1. Water quality other than temperature played a critical role
in these tolerance tests.
2. High nitrogen (No) gas saturation probably produced the most
important indirect effect with imposed temperature increases.
3. Resistance levels were reduced in most cases 5.4 to 9 deg. F
(3 to 5 deg. C) from perviously published laboratory findings and time
to death was shortened.
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63
4. Mortalities occurred at test temperatures which were similar
to naturally occurring river water temperatures.
The ultimate upper lethal temperature for juvenile salmon is
apparently between 73.4 and 77 F (23 and 25 C) if other stresses are
minimal. (Refer to Chapter III and Figure 5.) However, in the
presence of additional stresses (e.g., sublethal nitrogen supersatura-
tion), a temperature of 68 F (20 C) or lower may be lethal within one
week. Temperature in excess of 68 F (20 C) ". . . may be considered
suboptimum for survival of juvenile salmonids in river ecosystems"
(Becker, Coutant, and Prentice, ms 37).
In order to summarize the results of the many studies of thermal
resistance of juvenile salmonids, the pooled data was used to construct
the minimum mean thermal resistance curves in Figure 8. Data are in-
cluded from Brett (1952), Coutant and Dean (ms 33), Mighell (ms 17),
Blahm and McConnell (ms 3), Blahm and Parente (ms 4), Snyder and
Blahm (ms 5), Snyder and Craddock (ms 6), Blahm and McConnell (ms 7),
and Snyder and Blahm (ms 8) from tests on chinook, sockeye, coho, and
steelhead. Mean survival time (in minutes) is scaled on the ordinate
with temperature on the abscissa.
In thermal resistance tests it is standard procedure to measure
the median resistance time (time to 50% mortality) of test animals
exposed to a given temperature. Survival time increases at an ex-
ponential rate at progressively decreasing test temperatures. Connecting
most of the points on a graph of the logarithm of time plotted against
temperature produces a straight, sloping line (Figure 8). A point
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64
10,000-
5.00O-
I.OOO-
500
01
u
2 IOO--
u
P 50
cc.
V)
UJ
5--
Pooled data from several
studies (all species)
-National Marine Fisheries
service dofa from
Prescott Studies
-4-
' 0 22 24- 26 28
TEMPERATURE °C
30
Figure 8. Thermal Resistance of Juvenile Salmonids
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65
is reached at the upper end of a line (for a given acclimation
temperature of test animals) at which 50% mortality will never occur
regardless of time of exposure. This is termed the "incipient lethal
temperature."
Figure 8 was constructed by plotting the results on semi-log
graph paper of all comparable studies for a given acclimation temper-
ature and drawing a line (by inspection) through most of the points.
While this method gives an approximation, the error is not signifi-
cant from a practical standpoint and permits simplification of a
complex subject. Some results, particularly those of the National
Marine Fisheries Service at the Prescott Laboratory, were noticeably
different from the others and are shown by the dashed line. Reasons
for these differences are discussed on page 60.
For every degree of temperature increase it is noted that the
survival time is decreased from 50 to 75 percent regardless of the
acclimation temperature. It is also apparent that as the temperature
increases the difference in acclimation temperatures is less signifi-
cant (the resistance lines become closer together).
To test the element of time of exposure to juvenile thermal
resistance, fall chinook fry which were acclimated to 48.2 F (9 C)
were abruptly exposed to 86 F (30 C) waters over intervals ranging
from 5 to 35 seconds (Groves and Mighell, ms 15). The fish were
affected in direct proportion to the duration of exposure. At 5
seconds exposure, no loss of equilibrium or mortality was observed.
At a 10 second exposure, equilibrium loss occurred in 16 percent of
the fish and increased at a linear rate up to where 100 percent showed
equilibrium losses at 35 seconds exposure.
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66
Yearling sockeye, coho and steelhead were tested to determine
the effects of their previous temperature acclimation upon their
resistance to high lethal temperature (Mighell, ms 17). Fish
acclimated at 41, 50, 59, 68, and 73.4 F (5, 10, 15, 20, and 23 C)
were tested at exposure to 82.4, 86, and 89.6 F (28, 30 and 32 C) .
Results showed that, at all test temperatures, the trend was for
increased resistance to loss of equilibrium and death as acclimation
temperatures increased. The author notes the mean survival time in
relation to previous acclimation temperature increased markedly at
82.4 F (28 C), particularly for coho and steelhead, and suggests
that this may be a critical upper limit in order for acclimation
to significantly influence resistance to heat. Resistance was
greatest in steelhead, followed by coho and sockeye. Size of the
fish within the ranges tested did not affect resistance times to
death or loss of equilibrium.
Using a test flume and actual passage through a condenser tube,
Kerr (1953) tested thermal resistance of juvenile chinook salmon at
the Contra Costa stream plant on the Sacramento River. In flume tests,
fish ranging from 1.34 to 2.4 inches long were exposed for ten
minutes without acclimation to temperature rises in increments from
0 deg to 27 deg F (0 - 15 deg. C) and from a minimum water temperature
of 55-56 F (12.8-13.3 C) to a maximum water temperature of 83 F
(28.3 C). "It was found from these experiments that small yearling
salmon could withstand an instantaneous temperature rise of 25 deg. F
(13.9 deg. C) without loss of life and that their maximum temperature
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67
tolerance was approximately 83 F (28.3 C) " (Kerr, op. cit.). In
condenser tests, fish were exposed to 16 deg. F (8.8 deg. C) temper-
ature rises for 3.5 to 5 minutes with no mortalities. No mortalities
occurred in flume-tested fish held for 24 hours; a three percent
mortality occurred to condenser-tested fish held for ten days. (The
fish in these experiments did not receive a lethal dose).
Generalized conclusions regarding the thermal resistance of
juvenile salmonids indicate that the effects of a rapid temperature
increase is directly proportional to duration of exposure. Acclimation
to higher temperatures significantly increases temperature resistance.
Differences between species are small, and all investigators using
fish that were not prestressed achieved essentially the same results.
Fish stressed by nitrogen supersaturation, toxicants, or disease have
reduced temperature resistance; therefore, temperatures should desirably
be maintained well below the upper lethal limits to insure ultimate
survival. Brett (1960) recommended that the upper temperature limit
"required" for any species be 5.4 deg F (3 deg C) below the ultimate
upper lethal temperature to avoid significant curtailment of activity.
Temperatures above 68 F (20 C) are considered to be adverse for
juvenile salmonids, while temperatures near 62.6 F C17 C) may be
considered to be a maximum optimum temperature. Between 62.6 and
68 F (17 and 20 C), any increase in temperature is of no benefit to
juvenile salmonids while increasing requirements for food, the
possible synergistic action with toxic materials, and the liklihood
of disease infection.
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68
Predation
Increased susceptibility to predation of juvenile salmonids has
been assumed to occur when the fish have been subjected to the debili-
tation or shock of adverse water temperatures. Several studies to
define the relationship between temperature and predation have been
conducted.
Coutant (ms 31) speculated that, since loss of equilibrium was
an obvious behavioral response to a sublethal dose of a lethal temper-
ature, then smaller doses would also have a behavioral effect. A
series of laboratory experiments were designed to test the differential
predation rates at varying temperature levels. The tests sought to
determine if non-detectable behavioral changes would be reflected in
predator success when compared with untreated control fish. Thermally
shocked juvenile rainbow trout and chinook salmon were selectively
preyed upon by larger fish when shocked and unshocked fish were
subjected simultaneously to the predators (large rainbow trout).
Statistically significant increases in predation rates occurred at
thermal doses which were a fraction of the dose which, causes visible
equilibrium lossten percent in the case of chinook and 20 percent
in the case of rainbow.
Further experiments were conducted to determine if the predator
response paralleled thermal dose responses of visible equilibrium loss
and death (Coutant, ms 32); that is, is predation response a function
of temperature and exposure time? Rainbow trout fingerlings ac-
climated to 59 F (15 C) were subjected to three levels of lethal
temperature for various lengths of time. The fish were removed from
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69
the shock temperatures at intervals and immediately exposed to the
large predator rainbow trout along with control fish. A statistically
significant difference in predation on the two groups was evident
in 0.55 minutes at 30 C, at two minutes at 28 C, and 32 minutes at
20 C. The treated fish were more readily captured by the predators.
Similar results were found by Sylvester (ms 18) using coho for
predators and underyearling sockeye for prey, although no control
groups were used. In a series of experiments at acclimation temper-
atures of 44.5, 53.6, and 62.6 F (7, 12, and 17 C) he found that a
thermal dose of 18 deg. F (10 deg. C) for sixty seconds significantly
increases a prey's susceptibility to predation. The results were
similar for five second exposure to incremental temperatures ranging
from 23.4 to 41.4 deg. F (13 to 23 deg. C).
It should be remembered that the prey fish in these laboratory
experiments had no opportunity for excape as would be available in
natural conditions. However, the general conclusion that predators
select fish whose behavior has been altered by temperature stress
appears well substantiated.
Infectious Diseases
Infectious diseases of juvenile salmonids have been extensively
studied in connection with hatchery operations, but less work has
been done relating disease incidence with higher temperature than
has been done with adults.
Studies with hatchery-reared juvenile rainbow trout showed that
a number of factors are involved in the relationship between temperature
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70
and columnaris, including immunity, species, age, condition, other
disease, crowding and difference in columnaris strain virulence
(Fujihara, Olson, and Nakatani, ms 44). They pointed out that the
highest mortalities occurred during the initial temperature rise and
not during the peak temperatures of the summer.
Recent studies on the relationship between water temperature and
mortalities of rainbow trout caused by Ceratomyxa shasta, a myxospor-
idian, have been conducted at Oregon State University. Little
evidence was found for a trend between temperature and total percent
infectivity due to C^. shasta. A strong relationship, however, was
found between temperature and survival time of the host. One hundred
percent mortality of exposed fish occurred in 17 days at 69 F (20.6 C)
and 95 percent mortality in 56 days at 54 F (12.2 C) (Udey and Fryer,
unpublished). The full significance of these findings cannot be
appreciated without further study.
Gas Bubble Disease
Gas bubble disease is considered to be one of the most signi-
ficant water quality problems causing mortalities of downstream
migrating salmonids in the Columbia River. (For a description on the
disease and the nitrogen conditions in the Columbia River, see the
chapter on adults.) Although much of the evidence is as yet circum-
stantial, the physical extent of the problem has been defined by Ebel
(1970 C). Of the downstream migrants arriving at Ice Harbor Dam on
the Snake River, 25-45 percent of the chinook and 30-58 percent of
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71
the steelhead fingerlings exhibited visible gas bubble disease
symptoms. These observations support those of Raymond (1970) that
70 percent of the population of downstream migrant chinook from the
Salmon River was lost between Whitebird and Ice Harbor Dam. An
estimate for steelhead produced from Dworshak hatchery indicated a
25-30 percent loss to Ice Harbor Dam and about a 70 percent loss to
McNary Dam. Dissolved nitrogen supersaturation levels during the
period of downstream migration reached a peak of 146 percent satura-
tion in the forebay of Lower Monumental Dam in 1970.
Preliminary studies have been reported by Coutant (ms 34) in
which he explores the possible effects of nitrogen supersaturation
upon juvenile salmonids passing through the Hanford thermal plumes.
He attempted to define the levels of gas supersaturation and temper-
ature change required to produce identifiable synergistic effects
resulting in mortality of juvenile salmonids. His results show that
fish prestressed with high nitrogen saturation levels (115 percent)
suffered higher mortalities from a thermal shock than did fish
subjected to the thermal shock at normal saturation values. However,
since the rate of passage through the plume is relatively rapid
(0.5-1.0 meters per second), he concludes that the juvenile migrants
would be exposed to temperature increases greater than 5.4 deg. F
(3 deg. C) for only a few minutes, resulting in sublethal doses.
Coutant concludes from his lab studies that fish passing through the
Hanford plumes would not experience stresses from temperature-induced
supersaturation sufficient to cause significant mortalitites, either
immediate or thirty days after the experience.
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72
Studies conducted by Ebel et al. (ms 13) verify the detrimental
effects of nitrogen supersaturation and temperature conditions. The
data indicate:
1. Supersaturation of nitrogen drastically affects the tolerance
of juvenile coho, chinook and steelhead to temperature increases.
2. Acclimation to higher temperatures is of some value to the
fish in withstanding nitrogen supersaturation. However, the 50
percent mortality level will be reached in less than 18 hours at all
acclimation temperatures when nitrogen supersaturation levels are
125-130 percent.
3. When supersaturation of nitrogen is present, depth is an
important compensating factor when fish are able to "sound".
4. During periods of nitrogen supersaturation, any increase in
temperature will be detrimental to migrating juvenile salmonids.
The concern of the fisheries biologists is substantiated that
supersaturation of nitrogen is a significantly detrimental factor
weighing against successful downstream migration of juveniles and
that the effects of supersaturation are accentuated by temperature
increases. At saturation levels over 115 percent, any increase in
temperature could be damaging due to temperature-gas solubility
relationships.
Toxicity
The relationship of temperature to metal toxicity was shown by
Bouck et al. (ms 1) in laboratory bioassay studies. Juvenile steel-
head and rainbow trout were exposed to toxicant concentrations
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73
ranging from existing (control) levels in Willamette and Columbia
River waters to 100 percent concentration of the allowable levels of
ten toxicants. Results show the 96 hour TLm (tolerance limit median)
concentrations were inversly related to increasing temperatures. As
the test temperature rose from 5 C to 20 C the 96 hour TL dropped
to about half the former value.
Beneficial Effects
The beneficial effects of rearing catfish, shrimp, and oysters
in thermal effluents have been demonstrated in several small scale
experiments but only two references to rearing salmonids in heated
water are reported (Matthiason, in press; Wallis, in press). Carton
and Chriastianson, (in press) and Coutant (in press) pointed out
some problems to the use of waste heat for aquaculture including
demonstration of economic feasibility, design of a system to maintain
proper temperatures, and public acceptance of a product cultured in
an environment containing traces of radioactivity.
-------
VI - THERMAL EFFECTS ON NON-SALMONID FISH
While salmonid fish were the prime target of the biological
effects studies and review as a part of the Columbia River Thermal
Effects Study, other species of cold-water fish are found in the
Columbia River which would be pertinent to th.e over-riding concern
for water quality standards to protect and enhance aquatic life. The
following discussion presents a brief summary of the limited infor-
mation available on the temperature requirements and tolerances of
these non-salmonid fish.
The National Marine Fisheries Service, in studies of the
occurrence of fish in the vicinity of its research barge on the
Columbia River (RM 68), found twenty-seven species of fish CMcConnell
and Snyder, ms 12). Twenty-one of these species were non-salmonid
and accounted for 43 percent of their total catch. The non-salmonid
species which were captured throughout the year included three-spine
stickleback, Columbia River chub, and starry flounder. The non-
salmonid fish of commercial and sport value were American shad, white
sturgeon, and eulachon (smelt).
In temperature studies on the eulachon, Smith and Saalfeld (1955)
reported the fish entered the Columbia River when the temperature was
between 33.8 and 50 F (2 and 10 C) but they migrate up to and beyond
the Cowlitz River (RM 68) when the Columbia is approximately 40 F
(4.4 C). The smelt run was delayed five weeks from entering the
Cowlitz River because of low water temperatures during December 1968
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76
and January 1969 (Snyder, ms 27). Eulachon eggs appear to be more
tolerant than adults to temperature increases. The eggs can with-
stand a temperature of 25.2 F (14 C) from a base temperature of 39.2
to 46.4 F (4 to 8 C) without appreciable mortalities (Parente and
Ambrogetti, ms 26), but a 5.4 deg. F (3 deg. C) increase halts
maturation of adult females. In tests in 1968 and again in 1969,
it was observed that female smelt exposed to water heated 7 deg. F
(3.89 deg. C) above river temperatures were reluctant to spawn.
The studies on smelt indicate that those fish have a lower lethal
temperature limit than do the salmonids and a lower optimum temper-
ature preferendum. Adult female smelt are less tolerant of temperature
changes than other fish.
Thermal tolerance tests on yellow perch showed a 50 percent
mortality level at 89.6 F (32 C) in 2.4 hours (Blahm and Parente,
ms 10). A chill period at about 42.8 F (6 C) for an undefined
period of time is important to induce reproductive development and
o /
spawning in yellow perch.' Thermal tolerance studies on three spine
stickleback indicate a 50 percent mortality level at 78.8 F (26 C)
in about 6 days (Blahm and Parente, ms 10). Thus, both yellow perch
and three-spine stickleback have a higher thermal tolerance than do
the salmonids.
J5/ Mr. Bernard Jones at the National Water Quality Laboratory,
Duluth, Minnesota.
-------
VII - SECONDARY PRODUCTION ORGANISMS
Limited studies in other areas of the world have shown the
potential for adverse effects of a large thermal discharge upon
elements of the food chain. In terms of the ecology of the river
system, disruption of the food chain through destruction of the
smaller organisms could have serious consequences to the fish
resources of the Columbia River. Little information is available
on the secondary food organisms in the Columbia River or the thermal
effects on those organisms.
Specific Foods
Among some fishery biologists, there has been a popular concep-
tion that certain particular foods are required by juvenile salmonids
at various stages of their development (Davidson, 1965). Zooplankton
(Daphnia pulex) was found to be the dominant food organism of
juvenile echo in Lake Merwin, while insects were the dominant food
in Speelyai Creek, a tributary (Hamilton et al., 1970). Insects
formed a major portion of the diet of juvenile chinook migrating
through the lower Columbia River in spring and fall months, whereas
zooplankton were of major importance from July through October
(Craddock and Parente, ms 24). Fish were highly selective on the
larger plankton (Daphnia). Availability of insects was not studied
so it was not known if this was a matter of preference or avail-
ability.
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78
Becker (ms 40) found that larval and adult forms of aquatic
insects (Tendipedidae) were the primary food of even the smallest
chinook fry in the Hanford reach of the Columbia River. His opinion,
and that of most investigators, is that salmon are opportunistic
feeders and take whatever food is most readily available to them.
There is reason to believe this holds true throughout their life.
Food requirements are more specific for some non-salmonid fish
residing in the Columbia River estuary (Haertel and Osterberg, 1967) .
Plankton
Plankton populations in the Columbia River vicinity of Prescott,
Oregon, were studied by Craddock and Parente (ms 24) in 1968 and
1969. Daphnia and Bosmina were found to be the dominant genera of
cladocerans, the most abundant plankton organism. Daphnia was most
abundant at the surface and its peak of abundance occurred simul-
taneously with maximum water temperature in August. Bosmina was
uniformly distributed from 5 to 15 meter (16.4 to 49.2 foot) depths
and peak abundance was in June or July, with a second peak in October
and November.
Thermal Effects on Secondary Organisms
From observations of the thermal effects in the Hanford reach of
the Columbia River, Coutant (1968) indicates caddis flies emerge
about two weeks earlier at a point 14 kilometers below the lowermost
Hanford thermal discharge when compared to caddis flies at a point
16 kilometers above the discharges. Temperatures at the two points
were 54.5 and 52.3 F respectively (12.5 and 11.3 C).
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79
Thermal shock studies were conducted on the plankton Daphnia by
Craddock (ms 23). He found that a fifteen minute exposure to 86 F
(30 C) seemed to have little effect on survival of the plankton,
which had been previously acclimated to 59 F (15 C). Continued
exposure caused a fifty percent mortality in about twenty-four hours,
and a hundred percent mortality in about forty-eight hours.
Studies on the opposum shrimp, an important plankton of the
Sacramento River, by Hair (1971) showed that it can stand
a rapid temperature rise of 25 deg. F (13.9 deg. C), provided the
exposure time is short and that the ultimate temperature does not
reach a critical maximum of 87 F (30 C).
From the limited information available, then, it can be
generally concluded that the planktonic organisms are more resistant
to thermal shock than are the fish species that feed upon them.
-------
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*Matthiason, Mathias, "Beneficial Uses of Waste Heat in Iceland."
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-------
APPENDIX A
MANUSCRIPTS
Manuscript
Number FEDERAL WATER QUALITY ADMINISTRATION
1 Bouck, G., G. Chapman, P. Schneider, D. Stevens, and
J. Jacobson, "Initial Studies of Temperature
Requirements of Adults Sockeye Salmon (Oncorhynchus
nerka), Adult Coho Salmon (Onocorhynchus kisutch),
and Thermal-Chemical Requirements of Juvenile
Steelhead Trout (Salmo gairdneri) in the Columbia
River", October 1970.
NATIONAL MARINE FISHERIES SERVICE
Adult Salmon Behavior in River
2 Monan, Gerald E., Kenneth L. Liscom, and Jim R. Smith,
"Sonic Tracking of Adult Steelhead in Ice Harbor
Reservoir, 1969", July 1970.
Thermal Tolerance of Juvenile Salmonids
3 Blahm, Ted H., Robert J. McConnell, "Effect of Increase
Water Temperatures on the Survival of Spring and
Fall Juvenile Chinook Salmon (Oncorhynchus
tschawytscha) from the Lower Columbia River",
August 1970.
4 Blahm, Ted H. and William D. Parente, "Survival of
Juvenile Chum Salmon Exposed to Elevated Tempera-
tures in Columbia River Water", July 1970.
5 Snyder, George R. and Ted H. Blahm, "Mortality of
Juvenile Chinook Salmon Subject to Elevated Water
Temperatures", August 1970.
6 Snyder George and Donovan R. Craddock, "Effect of
Temperature Increases on Juvenile Steelhead Trout
(Salmo gairdneri) from the Columbia River",
August 1970.
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90
Manuscript
Number Thermal Shock of Juvenile Salmonids
7 Blahm, Ted H. and Robert J. McConnell, "Survival of
Juvenile Coho Salmon Exposed to Sudden Water
Temperature Increases", August 1970.
8 Snyder, George R. and Ted H. Blahm, "Survival Times
of Juvenile Salmonids Exposed to Water Temperatures
Causing Thermal 'Shock'", August 1970.
Thermal Tolerance of Non-Salmonids
9 Blahm, Ted H. and Robert J. McConnell "Mortality of
Adult Eulachon (Thaleichthys pacificus) Subjected
to Sudden Increases in Water Temperature",
July 1970.
10 Blahm, Ted H. and William D. Parente, "The Effect of
Increased Water Temperatures on the Survival of
Adult Threespine Stickleback (Gasterosteus
aculeatus) and Juvenile Yellow Perch (Perca
flavescens) in the Columbia River", August 1970.
11 Blahm, Ted H. and William Parente, "A Record of the
Development of the Shad Egg (Alosa sapidissima)",
August, 1970.
"On Site" Timing of Movement of Juvenile Fish at
Prescott, Oregon
12 McConnell, Robert J. and George R. Snyder, "Occurrence
of Fish in the Vicinity of Proposed Sites of Two
Nuclear Electric Plants on the Lower Columbia
River", August 1970.
Effect of Supersaturation of Nitrogen on Juvenile
Salmonids in the Columbia River
13 Ebel, Wesley J., Earl M. Dawley, and Bruce Monk,
"Thermal Tolerance of Juvenile Salmon in Relation
to Nitrogen Supersaturation", September 1970.
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91
Manuscript
Number Effects of Temperature Acclimation on Resistance to
Thermal Challenge
14 Groves, Alan B., "Maximal Swimming Performance of
Juvenile Coho Salmon (Oncorhynchus kisutch)
Following Abrupt Thermal Change", August 1970.
15 Groves, Alan B. and James L. Mighell, "Thermal Dose
and Equilibrium Loss in Chinook Salmon Fry
(Oncorhynchus tschawytscha) (Summary)",
August 1970.
16 McConnell, Robert J. and Ted E. Blahm, "Resistance of
Juvenile Sockeye Salmon (Oncorhynchus nerka) to
Elevated Water Temperatures", September 1970.
17 Mighell, James L., "Effects of Acclimation on
Resistance of Juvenile Salmonids to High Lethal
Temperatures", September 1970.
18 Sylvester, J. R., "Thermal Dose and Predator
Avoidance in Sockeye Salmon", July 1970.
Limnological Survey of Lower Columbia River
19 Clark, Shirley M. and George R. Snyder, "Limnological
Study of Lower Columbia River, 1967-68",
February 1970.
20 Clark, Shirley M. and George R. Snyder, "Timing
and Extent of a Flow Reversal in the Lower
Columbia River", November 1969.
21 Snyder, George R. and Robert J. McConnell, "Frequency
and Duration of the Reversal of Direction of
River Flow in the Lower Columbia River, April
1968 - March 1970", August 1970.
22 Snyder, George R. and Robert J. McConnell, "Subsurface
Water Temperatures of the Columbia River at
Prescott, Oregon (River Mile 72), 1968-69",
April 1970.
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92
Manuscript
Number Thermal Tolerance of Zooplankton of Lower
Columbia River
23 Craddock, Donovan R., "Thermal Effects on an Important
Zooplankter (Daphnia pulex) of the Columbia
River", July 1970.
24 Craddock, Donovan R. and William D. Parente,
"Utilization of Columbia River Zooplankton
by Juvenile Chinook Salmon", August 1970.
Physiological Effect of Temperature Increase on
Anadromous Fish of Lower Columbia River
25 Anonymous, "The Effect of Water Temperature Increase
on Spawning of Columbia River Smelt", (Working
Paper) , May 1969.
26 Parente, William D. and Walter J. Ambrogetti,
"Survival of Eulachon Eggs (Thaleichthys
pacificus) at Different Water Temperatures",
August 1970.
27 Snyder, George R., "Thermal Pollution and the
Columbia River Smelt", July 1970.
Prediction of Environmental Conditions in John Day
Reservoir
28 Novotny, Anthony J. and Shirley Miller Clark,
"Preliminary Predictions of Water Temperatures
in John Day Reservoir".
General
29 Snyder, George R., Theodore H. Blahm, and Robert
J. McConnell, "Floating Laboratory for Study
of Aquatic Organisms in their Environment".
30 Snyder, George R., Donovan R. Craddock, and Ted
H. Blahm, "Thermal Effects Studies on the
Lower Columbia River, 1968-70", August 1970.
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93
Manuscript
Number ATOMIC ENERGY COMMISSION
Performance of Thermally Shocked Young Salmon
31 Coutant, C. C., "Relative Vulnerability of Thermally
Shocked Juvenile Salmonids to Predation",
January 15, 1970.
32 Coutant, C. C., "Relative Vulnerability of Thermally
Shocked Juvenile Salmonids to Predation. II.
A Dose Response in Rainbow Trout",
February 4, 1970.
33 Coutant, Charles C. and John Mark Dean, "Relationships
Between Equilibrium Loss and Death as Responses
of Juvenile Chinook Salmon and Rainbow Trout
to Acute Thermal Shock".
Gas Bubble Disease of Young Salmonids
34 Coutant, Charles C., "Exploratory Studies of the
Interactions of Gas Supersaturation and
Temperature on Mortality of Juvenile
Salmonids, September 28, 1970.
Sonic Tracking of Adult Salmonids
35 Coutant, C. C., "Behavior of Sonic Tagged Chinook
Salmon and Steelhead Trout Migrating Past
Hanford Thermal Discharges (1967)",
Effect of Thermal Shock from Effluent Discharges on
Young Salmon
36 Becker, C. D., C. C. Coutant, "Experimental Drifts
of Juvenile Chinook Salmon through Effluent
Discharges at Hanford in 1968", September,
1970.
37 Becker, C. D., C. C. Coutant, and E. F. Prentice,
"Experimental Drifts of Juvenile Salmonids
through Effluent Discharges at Hanford,
Part II. 1969 Drifts and Conclusions",
July 15, 1970.
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94
Manuscript
Number Temperature Tolerances of Adult Salmon
38 Coutant, Charles C., "Thermal Resistance of Adult
Coho (Oncorhynchus kisutch) and Jack Chinook
(0. tschawytscha) Salmon, and Adult Steelhead
Trout (Salmo gairdneri) from The Columbia
River", October 1970.
Hanford Chinook Salmon Population Studies
39 Watson, D. G., "Fall Chinook Salmon Spawning in
the Columbia River near Hanford 1947-1969",
October 1970.
Relationship of Thermal Discharge to Food and Feeding
of Juvenile Chinook Salmon
40 Becker, C. D., "Food and Feeding of Juvenile Chinook
Salmon in the Central Columbia River in Relation
to Thermal Discharges and Other Environmental
Features', August 1, 1970.
41 Becker, C. D., "Temperature, Timing and Seaward
Migration of Juvenile Chinook Salmon from the
Central Columbia River", July 1970.
Effects of Temperature on C. Columnaris, Dermocystidium
Salmonis, and A. Salmonicida Fish Disease
42 Fujihara, M. P- and F. P. Hungate, "Seasonal
Distribution of Chondrococcus columnaris
Disease in River Fishes: A Procedure Using
Antibody Synthesis in Fishes as a Survey
Technique", October 1, 1970.
43 Fujihara, M. P. and R. E. Nakatani, "Antibody
Production and Immune Response of Rainbow
Trout and Coho Salmon to Chondrococcus
columnaris", October 1, 1970.
44 Fujihara, M. P., P- A. Olson, and R. E. Nakatani,
"Chondrococcus columnaris Disease of Fishes:
Influence of Water Temperature, Age and Size
on Susceptibility", June 15, 1970.
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95
Manuscript
Number Effects of Fluctuating Temperatures on Survival and
Growth of Juvenile Chinook Salmon
45 Olson, P. A. and R. E. Nakatani, "Effects of
Chronic Variable Water Temperatures on
Survival and Growth of Young Chinook Salmon".
46 Olson, P. A., R. E. Nakatani and T. Meekin, "Effects
of Thermal Increments on Eggs and Young of
Columbia River Fall Chinooks".
47 Olson, P. A. and W. L. Templeton, "Effects of
Fluctuating Temperatures on Fall Chinook
Eggs and Young".
48 Olson, P- A. and W. L. Templeton, "Effects of
Temperature Increments on Juvenile Steelhead".
Thermal Effects on Secondary Production Organisms
49 Coutant, Charles C., "Thermal Pollution - Biological
Effects a Review of the Literature of 1969",
February 15, 1970.
50 Coutant, Charles C., "Thermal Pollution - Biological
Effects a Review of the Literature of 1968",
February 17, 1969.
51 Coutant, C. C. and C. D. Becker, "Growth of Columbia
River Limpets, Fisherola Nuttalli (Haldeman),
in Normal and Reactor-Warmed Water", August 15,
1970.
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APPENDIX B
MEMBERSHIP OF THE TECHNICAL ADVISORY COMMITTEE FOR BIOLOGICAL EFFECTS
Columbia River Thermal Effects Study
State Representatives
H. J. Rayner, Ph. D. Oregon State Game Commission
James B. Haas Fish Commission of Oregon
Glen D. Carter Oregon Department of Environmental
Quality
Emanuel H. LeMier Washington Department of Fisheries
Jack Ayerst Washington Department of Game
Roland E. Pine Washington Department of Ecology
Monte Richards Idaho Department of Fish and Game
Robert P. Olson Idaho Department of Health
Ralph W. Boland Montana Department of Fish and Game
Federal Representatives
L. Edward Perry, Ph. D. Bureau of Sport Fisheries and Wildlife,
Department of the Interior
Vernon C. Bushnell, Ph. D. Bureau of Reclamation, Department of
the Interior
Fred Limpert Bonneville Power Administration,
Department of the Interior
Edward M. Mains Corps of Engineers, Department of the
Army
Power Organization Representatives
George Eicher Portland General Electric Company
Roy Hamilton, Ph. D. Pacific Power and Light Company
Sam Billingsley Washington Public Power Supply System
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APPENDIX C
State Water Temperature Standards
The water temperature standards adopted for the Columbia River
by the States of Oregon and Washington are as follows:
Oregon
No waste discharge or activity
will cause any measureable in-
crease when river temperatures
are 68 F. or above, or more
than 2 F. increase when river
temperatures are 66 F. or less.
Washington
No measurable increase shall be
permitted within the waters desig-
nated which result in water temp-
eratures exceeding 68 F. nor
shall the cumulative total of all
such increases arising from un-
natural causes be permitted in
excess of t=110/ (T-15); for pur-
poses here, "t" represents the
permissive increase and "T" rep-
resents the resulting water temp-
erature .
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APPENDIX D
Scientific Names of Fish and Other Aquatic Organisms
Salmon
chinook Oncorhynchus tschawytscha
coho Oncorhynchus kisutch
sockeye Oncorhynchus nerka
pink Oncorhynchus gorbuscha
chum Oncorhynchus keta
Trout
steelhead Salmo gairdneri
speckled Salvelinus fontinalis
Atlantic salmon Salmo salar
Other Species
smelt (Columbia River) (eulachon) Thaleich.th.ys pacificus
shad (American) Alosa sapidissima
sturgeon (white) Acipencer transmontanus
yellow perch Perca flavescens
three-spine stickleback Gasterosteus aculeatus
Plankton
Daphnia pulex
Bosmina sp.
Opossum shrimp Neomysis awatschensis
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102
APPENDIX D (CONT.)
Bacteria
Columnaris Chondrococcus columnaris
Furunculosis Dermocystidium salmonis
Aeromonis salmonicida
Protozoans
Ceratomyxa Ceratomyxa shasta
Insects
Midges Tendipedidae
Caddis fly Tricoptera
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