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NATURAL HISTORY SURVEY
&~
J UN 2 , 1986
Air
ILLINOIS
NATURAL HISTORY
SURVEY
LIBRARY
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THE EFFECTS OF DISSOLVED OXYGEN, TEMPERATURE, AND LOW
STREAM FLOW ON FISHES:
A LITERATURE REVIEW
Aquatic Biology Section
Technical Report
by
D. C. Dowling and M. J. Wiley
NN
Report to Springfield City Water,
Light, and Power Company
L
)logy Technical Report 1986(2)
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A
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Illinois Natural History Survey
Aquatic Biology Technical Report 1986(2)
THE EFFECTS OF DISSOLVED OXYGEN, TEMPERATURE, AND LOW
STREAM FLOW ON FISHES:
A LITERATURE REVIEW
by
D. C. Dowling and M. J. Wiley
to
Springfield City Water, Light, and Power Company
M. J. Wiley, Princip 1 Investigator
\
Robert W. Gorden, Head
Aquatic Biology Section
THE EFFECTS OF DISSOLVED OXYGEN, TEMPERATURE, AND LW
STREAM FLOW ON FISHES: A LITERATURE REVIEW
The chemical composition of rivers is very variable, depending on season,
time of day, place, and depth (Golterman 1975).
Of all the chemical substances in natural waters, oxygen is one of the most significant.
The
annual cycle of oxygen in a stream is closely correlated with
temperature. Studies of large rivers and small streams in warm southern
regions of moderate temperature regimes and in northern climates of broad
temperature fluctuations have shown that the oxygen content of flowing
waters is generally highest in winter and lowest in late summer.
In
small, slow-flowing streams of northern latitudes, decreased day length
and ice and snow cover may inhibit photosynthesis and prevent entry of
oxygen from the air, thereby depressing the oxygen concentration.
Sometimes this oxygen depression becomes quite severe, stressing the
resident fish populations, as was documented in the River Ob in Siberia
(Mosevich 1947, Mossewitsch 1961).
They documented oxygen levels falling
to 14% saturation; the fish left the affected regions and congregated in
a few more tolerable areas.
The vernal decline of oxygen content in slow-flowing streams may be
attributed to the action of spring floods in removing vegetation and to
the increased rate of decay of organic material with increasing water
temperature. Further decreases in oxygen content toward late summer are
due to one or more factors. High temperature water, reaching a maxinum in
late summer, holds less oxygen in solution; decreased discharge results in
diminished physical mixing and reoxygenation; and greater decomposition of
naturally produced organic material uses some of the available oxygen
(Reid 1961).
When rainfall is light, rivers are generally fed by
groundwater.
Often underground water is devoid of oxygen and contains
large amounts of carbon dioxide, because of its exposure to organic matter
and bacterial respiration in the soil;
although in limestone strata,
where underground water usually flows through large solution channels,
oxygen content of the emerging water may be high.
In many streams there is also a diurnal variation in oxygen content. The
diurnal pulse is largely a reflection of temperature fluctuations and
photosynthesis-respiration relationships. This diurnal pulse may become
exaggerated in streams where the discharge is low and algal populations
large. In summer, the diurnal curve is characterized by oxygen production
lagging behind rising temperature, often by 2 or 3 hours, thus imparting
high saturation values even after sundown. Depending upon oxygen demand,
the minimum, and possibly most critical, level usually occurs prior to the
early-morning low temperature.
In streams of this nature, the oxygen
concentration may be actively affected by sunlight intensity.
Autumn leaf fall, in combination with low flow, has been shown to create
In the U.S. Southeast and Midseasonally low oxygen concentrations.
western regions, and doubtless in other areas with similar hot and dry
summers, small streams are reduced to a series of disconnected pools with
little more than seepage between them. During autumn, great quantities of
leaves are shed by riparian trees into the pools. The water acquires an
inky black color, because organic matter leaches from leaves; oxygen
concentrations fall to low levels; and these conditions persist until it
rains (Schneller 1955, Slack 1964). Any organism that survives in such
streams must be either very resistant to low oxygen or have a life history
that ensures that no specimens are active in the water during late summer
and autumn (Hynes 1970).
Documented Effects of Drought
Organisms inhabiting small, warmwater streams of central North America are
normally exposed to severe environmental fluctuations; during periods of
deficient rainfall and reduced stream flow, they are subjected to extreme
physical, chemical, and biological stresses. Constriction of the environment changes competition, predation, and survival among crowded, displaced
organisms. Evaporation, stagnation, and organic decomposition alter the
2
dissolved salts and gases upon which aquatic life depends (Larimore et al.
In Smiths Branch, a small warmwater stream in Vermilion County,
1959).
Illinois, discontinuous flow in the summer of 1953 reduced the aquatic
habitat and exposed fish and invertebrates to desiccation, stagnation, and
predation. Larimore et al. (1959) determined that mid-summer impoundment
of the stream waters was not as detrimental to fish as might be expected.
Even though the supply of well oxygenated riffle waters ceased and extreme
thermal stratification occurred, no fish mortality was seen in July or
August of any year with discontinuous flow. Stagnation was more destructive to aquatic animals during September and early October, as leaves and
other dead plant materials accumulated in the pools, decomposed, and progressively increased oxygen demand.
Most fish withstood extreme drought conditions in at least a few parts of
Re-establishment of the populations began as soon as the
the stream.
stream resumed its flow, and during the following 2 weeks, 21 of 29
regularly occurring species moved into most of the stream course (Larimore
et al. 1959).
Twenty-five of the regular species of fish entered Smiths
Only three species were sigBranch by the end of the first summer.
nificantly absent. The longear sunfish (Lepomis megalotis) was not taken
until fall of 1955, and the blackstripe topminnow (Fundulus notatus) and
the hornyhead chub (Nocumis biguttatus) had not repopulated the stream
when final collections were made in 1957. Invertebrates displayed
remarkable adaptations to drought conditions and repopulated Smiths Branch
soon after flow resumed.
Other researchers have also studied the impact of droughts on stream
systems. Cowx et al. (1984) investigated the effects of the 1976 summer
drought on both fish and invertebrate populations of the Afon Dulas stream
in Wales. The most significant effect was the failure of the 1976 year
class of salmon. They felt that salmon alevins of the 1976 year class,
which are less tolerant than either trout or older salmon par to prolonged
exposure to temperatures around their upper lethal limit, did not survive
high water temperatures in June and July. Some compensation for this loss
to the fish populations was indicated by enhanced recruitment in 1977,
3
i.e., the numbers of juvenile salmon surviving beyond fry stage was
greater in 1977 than in 1975, a year of typical recruitment.
One of the major effects of drought on invertebrates was the overall
decrease in potentially colonizable habitats due to reduced depth and
width of the river (Cowx et al. 1984). A reduction in wetted bed area may
also have long-term implications, because many adult insects lay eggs in
fast-flowing or broken water (Sawyer 1950); if suitable areas are
Follow-up
reduced, subsequent populations may be affected (Hynes 1958).
surveys in 1977 and 1978 indicated that invertebrate fauna in the Ob Dulas
had fully recovered from the drought, both in terms of taxonomic diversity
and abundance, suggesting that invertebrate fauna in rivers can recover
quickly from the effects of drought, usually within several life cycles.
Hynes (1958), working on a small stream in Wales, found that several
species of worms and small crustacea can survive prolonged drought and
that, although most insect nymphs and larvae are killed, eggs may survive.
The number surviving depends on whether the drought coincides, at least
partly, with the normal hatching period. Wickliff (1945) indicated that
crayfish that normally do not burrow, may follow the receding water
table for several inches underground and survive for weeks if the chamber
in pool and riffle headwaters,
into which they retreat remains moist;
small fish are able to exist for months in large numbers in greatly
reduced volumes of suitable water, returning to the riffles when flow
resumes.
In general, invertebrates are better
drought than are fish, because of egg,
gence, or burrowing (Canton et al.
following resumption of flow (Hynes
Williams and Hynes 1976a, 1976b, 1977).
able to withstand the effects of
nymphal, or pupal diapauses, emerRecolonization is rapid
1984).
1975, Townsend and Hildrew 1976,
In a Colorado stream in 1978 and 1979, Canton et al. (1984) investigated
Brook trout (Salvelinus fontinalis)
the impacts of low flow in 1978.
collected in 1978 were thin and moribund when handled. Average weights
4
for each age class were lower than
American studies (Carlander 1969),
factor (K) was lower in 1978 than
often indicate environmental stress,
expected on the basis of other North
and as a result, the mean condition
Regenerated scales, which
in 1979.
were found on many fish in 1979. In
a few cases, especially on white suckers (Catostomus camersoni) and brook
trout, only regenerated scales were found. Regenerated scales were not
observed in 1978. Fish population densities were significantly reduced by
After
the drought, with no fish collected in most of the study area.
resumption of normal discharge, fish were abundant throughout the study
area, although the greater abundance of trout at the upper end of the
study area suggested that migration from Manitou Lake into the study area
occurred. This lake probably was a refuge for stream fish during the
drought. The rapid recovery of fish populations from drought has also
been noted in studies by Stehr and Branson (1938) and Starrett (1950).
Canton et al. (1984) discovered through functional feeding group analysis
that collector-gatherers and collector-filterers were relatively more
important during normal flow years, whereas shredders and predators had
greater importance during low flow. The decreased importance of gatherers
and filterers during a drought may be related to a combination of factors,
such as loss of habitat, decreased suspended matter, and increased predation pressure.
However, the large increase in total density in 1979
again indicated rapid recovery of stream invertebrates camnunities from
drought. This recovery was probably a result of survival of eggs in moist
substrate, subsequent oviposition by aerial adults, and invertebrate drift
from the upstream reach when flow resumed (Gray and Fisher 1981, Fisher et
al. 1982). Mayflies appeared to be most severely affected by reduced
discharge and Diptera least affected.
Low discharge seemed to affect
invertebrates primarily through loss of habitat, because temperature was
not significantly affected.
5
When faced with low dissolved oxygen, many fish species will migrate to
more tolerable regions, dramatically altering the local fish community
structure. Tsai (1970), studying the Little Patuxent River in Maryland,
found that fish species sensitive to organic enrichment and low dissolved
They were replaced by
oxygen decreased in abundance or disappeared.
species that were tolerant to organic enrichment and low dissolved oxygen.
Adversely affected species included important game fishes, such as
smallmouth bass (Micropterus dolmieui), largemouth bass (L salmoides,
and black crappie (Pomoxis nigrmaculatus).
Favorably affected species
included less important game fish, such as sunfishes, and coarse fish,
such as the American eel (Anmuilla rostrata) and the creek chubsucker
(Er imyzon oblongs).
Garmmon and Reidy (1981) stressed the important role that tributaries play
during episodes of low dissolved oxygen. They noted that, under normal or
high flow conditions, fish commnunities in tributaries to the middle Wabash
River, Indiana, tended to be smaller, less diverse, and of a different
species composition than fish conmunities in the mainstem. In late July
and early August 1977, dissolved oxygen concentrations declined to a low
of 1.0 mg/liter in a 35-mile stretch of the river. During that period,
pollution-sensitive fishes moved into the mouths of tributaries,
increasing several caununity indices.
Such movements to avoid stress
conditions in the Wabash mainstem have been hypothesized by Gammon (1976) .
Previous work by Cross (1950), Minckley (1963), and Krumholz and Minckley
(1964) documented this phenomenon in other rivers.
Fish often seem to avoid lethal levels successfully when better oxygenated
water is accessible (Doudoroff and Shumway 1970). Concentrations below 45 mg/liter apparently interfere with upstream migration of adult salmonids
(Sams and Conover 1969), but migration of these and other anadromous fish
through waters of lower dissolved oxygen concentrations of 2-3 mag/liter
has been observed. Tagatz (1961) observed that young American shad
ceased schooling when the dissolved oxygen concentration was reduced
slowly to 1.4 mg/liter or rapidly to 2.4 mg/liter. However, sublethal
effects of hypoxia on young shad, indicated by refusal to eat, rapid
gulping, and breaking up of schools, seemed to begin at an oxygen concentration of 4.5 mg/liter (Chittenden and Westman 1967).
Time of year may delay recolonization of streams, even if they are fully
recovered in terms of water quality and flow.
Larimore et al. (1959)
found that fish moved back into Smiths Branch quickly during April and
May, but re-invasion was slow for several weeks during late February and
early March, after stream flow resumed.
Katz and Gaufin (1953) also
observed a delay in re-invasion during winter; fish in winter did not
move into a previously unacceptable area of a polluted Ohio stream, even
though it was judged to be free of harmful pollution.
As indicated previously, many different grades of tolerance to low dissolved oxygen exist between stream fishes. Matthews and Styron (1981)
investigated the tolerance of headwater versus mainstream fishes to abrupt
physicochemical changes.
They found that the mountain redbelly dace
(Phoxinus
)ore),
the most abundant cyprinid in the intermittent head-
waters of Mason Creek, Virginia, to be significantly more tolerant of
abrupt changes in dissolved oxygen concentration, temperature, and pH than
were three cyprinid species characteristically found in the more
physicochemically stable Roanoke River and its larger tributaries. Their
results suggest a correspondence between physicochemical tolerances of
fish species and the upstream limits of their distributions in a stream
with environmentally unstable headwaters.
Fishes in other physically
harsh environments have wide physicochemical tolerance limits, which can
be related to their distributional limits. Cross and Cavin (1971) found
the red shiner (Notropis lutrensis), which is highly successful in
intermittent prairie streams, to be more tolerant of low oxygen conditions
that the bluntface shiner (L. camurus), which is restricted to permanently
flowing upland streams.
7
Matthews and Styron (1981) also reported that a greater portion of head-
water than mainstream fantail darters (Etheostma flabellare)
survived
brief periods of oxygen stress. Small intraspecific differences in
tolerance could be important, if over time they resulted in ecotypes
Intraspecific
adapted to specific parts of a heterogeneous watershed.
differences in physiological tolerances among populations in a single
watershed could influence local distribution patterns and help isolate
populations, comprising an initial step in the speciation process.
Burdick et al. (1954) noted that smallmouth bass in Bebe Lake, New York,
were less sensitive than populations in Deer River to dissolved oxygen
(Table 1).
Oxygen acclimatization in a lake might be to a lower level
than that in a rapidly flowing, relatively unpolluted stream.
Matthews and Hill (1979) looked at age-specific differences in the distribution of red shiners over physicochemical ranges and found that juveniles
occupied warmer water than did adults in February, May, and August, higher
oxygen concentration in February and October, and higher pH conditions in
February and August. Juveniles occupied shallower water than did adults
in February and August, locations with less shade in May, and less shelter
Both groups occupied locations
in February, May, August, and October.
with very slow current. The mean habitat breadth of adults changed more
as environmental conditions changed than did that of juveniles. Stress on
red shiners was probably greatest in October, near the end of a lengthy
drought, and in December-January when it was unusually cold.
Temperature and oxygen tolerances of four cyprinid species in a southwestern river coincided with their relative success during drought
(Matthews and Maness 1979).
The emerald shiner (Notropis atherinoides),
which was least tolerant of temperature and oxygen, decreased in abundance
in the sumner and apparently failed to spawn in 1976. The Arkansas River
shiner, which was highly tolerant of low oxygen, was more successful than
the plains minnow (fHybonathus placitus) or the red shiner from August to
October, although substantial algal blooms during that time caused low
nocturnal oxygen concentrations. Oxygen probably exerted less selection
pressure than did temperature. The critical thermal maxima (CTM), which
8
Table 1. Summary of data on lethal oxygen concentrations
(in parts per million) for smallmouth bass (Burdick et al. 1954).
Oxygen concentrations
Number ; Temperature
of fish
in degrees F.
]imum
found lethal
M aximum
m um
mum I Median I'Mean
Standard Standard
deviation
error
Deer River
29
54.0
0.95
0.58
0.72
0.74
±0.083
±0.015
20
60.0
0.98
0.63
0.87
0.83
±0.121
±0.028
14
70.0
1.23
0.77
0.91
0.96
±0.138
±0.037
9
80.0
1.32
0.99
1.17
1.15
±0.132
±0 044
Beebe Lake
20
52.5
0.85
0.48
0.63
0.63
±0.142
±0.032
15
60.0
1.10
0.48
0.73
0.73
±0.145
±0.037
20
70 0
1.19
0.61
0.86
0.87
1±0.133
+0 030
20
80.0
1.56!
0.72
1.03
1.08
±0.221
±0 050
1
9
is the thermal point at which locomotry activity becomes disorganized and
the animal loses its ability to escape from conditions that will cause its
death (Mihurksy and Kennedy 1976), were more closely related to field
The red shiner and the plains
success than was low oxygen tolerance.
minnow, with higher CT' s, exhibited greater population increases from May
to October than did the other two species.
Fish may use other mechanisms to avoid the stress of deoxygenated waters.
Gee et al. (1978) studied the reactions of 26 species of Great Plains
fishes from eight families to progressive hypoxia.
He noted that when
freshwaters become hypoxic some fishes respond by breathing air facultatively. Others remain aquatic breathers, rising to the surface where
they irrigate their gills with relatively oxygen-rich surface film. All
species were able to use oxygen in the surface film, except Salmonidae
(three species) and the walleye (Stizostedion vitreum)i.
The concentration
of dissolved oxygen at which this reaction occurred was found to be
strongly temperature dependent in the fathead minnow (Pimephales
promelas). Lowe et al. (1967) stated that, on the whole, minnows are more
advantageously adapted structurally and behaviorally than are suckers to
competition for surface-layer oxygen.
Connell (1975), Wiens (1977), and Diamond (1978) suggest that competition
or predation pressures may be less intense, or at least occur less frequently, in harsh, fluctuating environments and some species avoid
competition or predation through evolution of wide physicochemical
tolerance and use of rigorous environments. Whether this is true remains
to be seen, but it is readily apparent that a fish is affected by numerous
biological factors: through its senses, by its associates, its school,
predators, prey, algae, and aquatic plants, in addition to the changing
physical and chemical characteristics of the water, which vary with depth
and season (Sondheimer and Simeone 1970).
It is this array of factors
that actually constitutes what chemical concentrations a fish will survive
and where it will reside. However, the effects of the combined influence
of current, oxygen, and temperature on lotic species has not been accuIndividual laboratory experiments have
rately assessed (Brown 1971).
10
shown how some of these factors may impact fish in their lotic
environment.
Physiological Responses
The rate of oxygen consumption in freshwater fishes is not constant
(Clausen 1936); it varies in the same fish from hour to hour, and it
Daily fluctuations are
varies between individuals of the same species.
probably part of larger rhythms that harmonize with the physiological
cycles of the animal. Wells (1914) pointed out differences in the resistance of fishes during breeding and later in the summer. Beamish (1964)
investigated the standard rate of oxygen consumption (when the fish is
nearly or completely resting) and the active rate of oxygen consumption
(when the fish is stimulated to continuous activity) for brook trout,
carp, and goldfish. These data indicate that, over a range of relatively
high partial pressures of oxygen, the standard rate of consumption remains
constant while the active rate increases with increases in the partial
pressure (Fig. 1). Basu (1959) found that, when the oxygen concentration
of water falls below a critical level, the fish is unable to satisfy its
need for oxygen and oxygen consumption becomes dependent on the oxygen
concentration of the water. For many fish species, oxygen concentrations
below which activity is limited have been determined (Fry and Hart 1948,
Graham 1949, Gibson and Fry 1954, Job 1955).
The classical concepts of Fry (1957) are embodied in Fig. 2. The point at
which available oxygen coincides with oxygen needs for bare maintenance is
termed the incipient lethal tension. Below this level, organisms resist
for a time but eventually die. Standard metabolic rate was defined by
Brett (1962) as the minimal metabolic rate accompanying the energy cost of
It is close to, but
maintenance, measured under laboratory conditions.
slightly higher than, the thermal basal metabolic rate used by others.
The point at which metabolic rate ceases to be dependent on available
oxygen is termed the incipient limiting tension, or also the no-effect
In
oxygen threshold, which affords good protection for fish species.
animals tolerant of low oxygen, the curve in Fig. 2 shifts to the left,
11
E
cb
C
0
I
E
ro
0
o
-
280
-
c
-
o 200
120
40
0
20
Partial
19EON
1 l
%d
60
Pressure
,
100
140
180
of Oxygen, mm Hg
Fig. 1. Relation between standard and active oxygen consumption for carp. Solid symbols represent values for fish
acclimated to air saturation.
The standard curves have been
taken from Fig. 2. The broken line represents the results for
carp acclimated to the various partial pressures of oxygen
applied. The weight range of carp used at the active level
was 36 to 750 g at 10 C and 140 to 540 g at 20 C (Beamish 1964).
I .
12
ZONE
I
I
ZONE OF RESPIRATORY DEPENDENCE
RESPIRATORY
OF
REGULATION
ZONE OF
L RESISTANCE
ZONE OF TOLERANCE
I4
hi
INCIPIENT LMITING TENSION
4
B.
z
hi
0
K
0
Ii.
0
STANDARD METABOLIC RATE
hi
4
INCPIET
INCIPIENT
ENSION
ETHL
LETH
AL
TENSION
ENVIRONMENTAL
0 2
Fig. 2. Relation between standard and active (Maximum) oxygen
uptake rates at different environmental oxygen concentrations (from
Hoar 1966).
13
while in animals requiring higher levels of oxygen the curve shifts to the
Thus, the oxygen tension that produces respiratory dependence
right.
varies with species (Davis 1975).
Ventilatory and circulatory changes have been observed in a number of fish
species in response to hypoxia. Rainbow trout (Salm~ gairdneri) (Holeton
and Randall 1967a, 1967b) and carp (Cyprinus car.piQ) (Garey 1967) show
elevations in both rate and amplitude of breathing and decreased heart
rate in response to low oxygen. In trout, the stroke volume of the heart
increases, thus maintaining cardiac output as the heart rate decreases.
The gill transfer factor for oxygen (a measure of the ability of the gills
to transfer oxygen per unit gradient) increases, water flow through the
gills goes up, and venous oxygen tension drops. All of these factors
operate to maintain oxygen uptake with reduced oxygen availability.
These responses of fish to hypoxia represent compensation or adjustment of
bodily processes. Fish can only resist or tolerate reduced oxygen levels
for short periods. The success of tolerance depends on species, oxygen
levels, and environmental factors, particularly temperature. Davis (1975)
summarized temperature and oxygen levels at which many fish species are
affected and the degree to which they are affected (Table 2).
Effects n Growth and Fecudit
Use of regulatory or compensatory mechanisms requires energy expenditure
and reduces energy reserves for swinming, feeding, predator avoidance, and
other activities.
Stewart et al. (1967) investigated the influence of
Its
oxygen concentrations of the growth of juvenile largemouth bass.
growth and food consumption rates increased with increases in dissolved
oxygen to levels near saturation and declined with further increases of
oxygen concentration (Fig. 3 and 4).
Gross food conversion efficiencies
were reduced only at concentrations below 4 rag/liter (Fig. 5). Stewart et
al. (1967) also noted that growth of bass subjected alternately to low and
14
Table 2. The incipient oxygen response thresholds for various fish groups and
habitats gathered from the literature with emphasis on Canadian species. The rationales for choosing the reported response and its significance are reported in the
following pages 2301-2311. In most instances, oxygen response thresholds were found
in only one form of terminology (e.g. column 6--mg 02/liter)and the others in the
table were calculated. Calculations for freshwater fish made use of the reported
temperature and the 02 solubility data in Tables 1 and 2 while those for nonanadromous
fish in sea water used the data in Table 3 and an assumed salinity of 28%. It is
stressed that the response levels reported are incipient oxygen levels where effects
first become apparent thus providing a "biological indication" of the onset of hypoxic
stress. A) Freshwater species, B) Marine nonanadromous species, C) Anadromous species,
D) Eggs and larvae (freshwater and marine species) (Davis 1975).
Level of dissolved oxygen
Species
Size
Temp
(C)
P0 2 -mm Hg
ml O 2 liter
mg O 2 /liter
7, Satn.
Response to oxygen
level
Itrema
A. Freshwater species
Arctic char,
Salvelinus
alpinus
brown bullhead,
Ictalurus nebulosus
Rainbow trout,
Salmo gairdneri
carp,
Cyprinus carpio
2±.05
25
1.53
2.18
15.8
9-10
60
4.87-4.76
6.95-6.80
38.0
onset of oxygen dependent oxygen
uptake
(196)
682-1136
g
20
78
3.20
4.59
50
below this level blood
is not fully saturated
with oxygen
Irving et .
(1941)
120-250
g
8.5-15
-
43-127
g
50.7-63.7 circulatory changes
occur, including a
slowing of heart
standard oxygen uptake elevated (depressed at lower
levels)
Randsd
andSIta
Beamilh
10
50
2.51
3.59
31.7
20
80
3.29
4.71
51.3
80
2.74-3.04
5.39-4.34
50.9-51.6 blood ceased to be
fully saturated with
02 below this level
Itazaw
(1970)
35.2-70.4
2.8-1.4
4-2.0
ScShr
(1971)
70.4-96.8
3.85-2.8
5.5-4.0
45.3-22.7 loss of negative phototaxis (light avoidance behavior)
45.3-62.3 increased mobility,
"darting" behavior
2 yr-old
5-9
cm
Brook trout
(speckled trout)
Salvelinus
fontinalis
682-1136
g
17-65
g
13.3-25
22
-
20
56-140
g
682-1136
g
-
77.95
Whitmort
et al.
50
blood ceases to be
fully satd. with 02
below this level
Irvialgetid.
(1941)
onset of Or-dependent
metabolism
reduced cruising speed
onset of Or depen.
dent metabolism
reduced activity, all
temps.
Graham
(1949)
4.5
-
I.I
1.5
-
3.21
4.59
100
5.66
8.09
63.2
8
20
80
154
4.19
6.34
5.99
9.06
50.7
98.8
6.72-4.82
9.6-6.88
4.03-3.63
5.75-5.18
3.21
4.59
50
156.6
6.82
9.74
100
10,15
20
116.9-118.7
80
77.95
(1964)
some indication of
avoidance behavior
marked avoidance
3.15
5
5-20
Rainbow trout,
Salmo gairdnerl
5.18-7.34
30-600 g
Largemouth bass,
Micropterus
salmonides
Brown trout,
Salmo trutta
3.63-5.14
77-350 g
235-785g
Walleye,
Stizostedion vitreum
80-100
"signs of asphyxia
Holetm
and loss of equilibrium" (1973)
75
50.7-51.0 standard oxygen uptake reduced below
this level
13.3±1.4
cm
17± .5
20 mos old
8-10
78.85-79.95
4.16-3.97
5.94-5.67
50
21-23
77.85-77.6
3.15-3.04
4.50-4.34
50
blood ceases to be
fully satd. with O
below this level
(190)
Beamish
(1964)
Irving et aL
(1941)
any reduction in
Downing
(1954)
oxygen led to more
rapid death in cyanide
43% reduction in
maximum swimming
speed
30% reduction in
maximum swimming
Jones
(1971b)
speed
2.3-13
100
6.11-4.71
8.73-6.74
-
15
78.45
3.55
5.08
-
15
3.63-4.53
5.18-6.47
235-510
g
80-100
15
63.1-63.6 blood is not fully
satd. with oxygen
below this level
50
Itazawa
(1970)
altered respiratory
Kutty
quotient, little capacity (1968a)
for anaerobic metabolism below this level
51.0-63.7 changes in oxygen
transfer factor and
effectiveins of O
exchange occur
Randall
et al. (1967)
Table 2 (continued).
Species
Size
Temp
(C)
Level of dissolved oxygen
POa-mm Hg
ml O0/liter
mg O/liter
7%SaIa.
51.0
t
t
400-600
g
13.5
80
3.74
5.35
.
t
approx. 300
g
10,15,
20
80
3.30-4.03
4.71-5.75
1-11
g
17.5
93.8
4.05
5.78
92.3-110.7
3.5-4.2
5-6
Largemouth bass,
Mkropterus
juvenile
25
5-9
cm
-
Response to oxygen
level
breathing amplitude
and buccal pressure
elevated
50.7-51.3 below this level blood
is not fully satd. with
oxygen
60
toxicity of zinc, lead,
copper, phenols inScreased markedly
below this level
59.7-71.6 final (maximum sustained) swimming
salmoldes
Reference
Hughes and
Saunders
(1970)
Cameron
(1971)
Lloyd
(1961)
Dahlberg
et al. (1968)
speed reduced below
this level
Ulvuell,
L/pomis
macrochirus
-
2.1
3.0
-
1.1
1.5
-
some avoidance
behavior
strong avoidance
Whitmore
et al. (1960)
B. Marine nonanadromous species
-4it perch,
Macochilus vacca
adults
11
80
3.19
4.56
50.8
blood ceases to be
fully 02 saturated
below this level
Webb and
Brett
(1972)
Dofish,
Sq"as suckleyl
2.5-6.0
kg
10
115
4.68
6.69
72.92
blood ceases to be
fully 0O saturated
below this level
Lenfant and
Johansen
(1966)
Dogtish,
Scylo inus
caniculas
150-600
g
12-14
80
3.12-2.95
4.46-4.20
50.8-51.0 Oxygen-dependent
metabolism begins
below this level
Hughes and
Umezawa
(1968a)
Dragonet,
Cutieymus lyra"
70-140
g
11-12
125
4.98-4.88
7.12-6.97
79.4
100
3.97-3.90
5.67-5.58
63.5
-
-
-
80-120
Dragnet,
Ca•Meymus lyrea
a1sh5
-
-
80
11
150
5.98
8.54
158.2
7.14
10.20
Ndretegus colliei
Atlantic cod,
Gdus morsua
1.14-2.33
kg
5
Oxygen-dependent
metabolism below
this level
altered heart rate
Hughes and
Umezawa
(1968b)
increased ventilatory
muscle activity below
this level
Hughes and
Ballintijn
(1968)
blood ceases to be
fully Oa saturated
below this level
Hanson
(1967)
100
any reduction in
ambient O0 level
produces rise in ventilatory water flow
Saunders
(1963)
100
available oxygen level
appears to limit active
metabolism and
maximum swimming
Brett
(1964)
95.2
C. Anadromous species
Seilbead,
Selsm gairdnert
-
Sockeyesalmon,
Owerkynchus nerka
Caoe salmon,
oerwhynchus
kisutch
see freshwater species
50 g
20-24
1579 g
13
1.5-1.7
kg
15
15$5.9-154.9
100
78.45
6.42-5.97
9.17-8.53
4.72
6.74
63.6
3.55
5.07
50
-
"summer
temperatures"
-
3.15
4.5
5.1-14.8
cm
12±1
130.8
6.3
9.0
Juvenile
10-20
1 57.7-155.9
7.93-6.42
11.33-9.17
155.9
6.42
9.17
5.6-2.8
8.0-4.0
6.3-11
cm
speed
blood ceases to be
fully 0O saturated
below this level
elevated blood and
Randall
buccal pressure and
and Smith
breathing rate increased (1967)
erratic avoidance
behavior
Whitmore
et al. (1960)
below this level acute
mortality in kraft
pulpmill effluent
Hicks and
DeWitt
(1971)
100
reduction of 02 below
saturation produced
some lowering of
maximum sustained
swimming speed
Davis et al.
(1963)
100
as above
83.1
mincreased
to
.
to
t
90 ot
"9
to
20
10
136-67.9
16
Davis
(1973)
87.2-43.6 growth rate proportional to oxygen level
with best growth at
8.0 ms/liter, lowest
at 4.0 mg/liter
Dahlberg
et al. (1968)
Hermann
(1958)
Table 2
(continued).
Level of dissolved oxygen
Size
Species
6.3-11
cm
Chinook salmon,
Oncorhynchus
Ishawytscha
"
"*
Atlantic salmon,
Salmo salar
American shad,
Alosa sapidissima
Rainbow trout,
larvae,
Salmo gairdnerl
Chum salmon eggs,
Oncorhynchus keta
Atlantic salmon,
Salmo salar
Atlantic salmon,
Salmo salar
uveniles
J
87-135
g
-
Temp
(C)
"summer
temps"
"fall
temps"
10-20
15
-
PO-mm Hg
-
ml O/liter
3.15
-
-
4.5
-
157.7-155.9
7.93-6.42
11.33-9.17
69.6
3.15
4.5
1.75-2.1
2.5-3.0
-
2.8
4.0
-
-
-
D. Eggs and larvae
5.03
100
10+1
early
stage
eggs
10
13.9
.7
pre-hatch
eggs
10
>97.4
>4.9
5 day
eggs
8.0-8.2
22.2
(eyed)
85 day
eggs
3.6-4.9
> 60.3
5.5
%7Satn.
4.5
1+8
day
old
larvae
early
eggs
mg Oa/liter
1.17
b
9.62
> 3.5b
7.18
1
> 7.0
1.67
> 5.0
Response to oxygen
level
Refermw
marked avoidance of
this level in summer
little avoidance of
this level in fall
reduction of O
below saturation produced some lowering
of maximal sustained
swimming speed
Salmon stop swimming
at a speed of 55
cm/s at 02 levels
below this. Faster
swimming requires
more oxygen
threshold for migration through
polluted areas
necessary for
spawning areas
Whitmen
et al. (11
63.4
elevated heart and
breathing rates below
this level. Bradycardia (heart rate
slows) sets in at
lower O0 levels
Holetoa
(1971)
8.8
retarded development,
deformities, mortality, respiratory Oadependence
oxygen level required
for normal development and non O2dependent metabolism
Alderdice
et al. (1953)
critical oxygen level
for supply of oxygen
demand and non 02denendent metabhlism
Wickett
(1954)
As above
Lindroth
(1942)
100
44.33
> 61.8
14.07
Davis etd.
(1963)
Kuttysad
Saunden
(1973)
Chittenda
(1973)
> 38.1b
.53
0.76
6.09
4.06
7.0
5.80
10.0
45.31
102.67
2.17
4.97
3.1
7.1
27.36
62.67
near
(from
hatching
hatching
5
17
eyed
hatching
10
10
Northern pike,
Esox lucius
from
fertilization
to
feeding
larvae
15,19
MummichogL
Fundulus
heteroclltus
fertilization
to
hatching
20
Pacific cod,
Gadus
macrocephalus
fertilization
to
hatching
71.7
160.8
43.14
98.83
> 5.18-5.66
101.2
> 2.16-2.35
3.15
> 3.35-3.09
4.5
> 2-3
3-5
*European species - representatives of this family found in Canadian Atlantic waters.
bCalculations based on 4 C.
*Estuarine fish
- calculations based on salinity of 30%,.
17
> 33.0
64.9
Wickett
1954))
critical 02 level for
supply of 02 demand
and non O2 -dependent metabolism
Hayes et al.
(1951)(from
Wickett,
1954)
necessary O2 level for
proper hatching,
survival and
development
Siefert
et al.
(1973)
reduced hatching at
this level compared
to 7.5 mg/liter 0O
Voyer and
Hennekey
(1972)
oxygen level required
for qptimal hatching
at 157,o salinity
Alderdica
and
Forresmt
(1971)
0
w II
Zsc
z
s-
ZU
4
at
r5)
VS)
ri)
NTS)
ys)
WkJ)
I<
hn)
*E
1
4
a
iii,
7
a
9
MILLIGRAMS PER LITER
-,
WD
-i
DISSOLVED OXYGEN CONCENTRATION
--
IN
Fig. 3. Growth of largemouth bass in relation to
dissolved oxygen concentration (Stewart et al. (1967).
18
r-
Ad%^
0%
U.500
0.400
-
3t 0.300
-
4
aJI
>o--
F
a
W
oz
0.200
MENT-I
MENT-2
o 0.100
MENT-4
MENT-5
MENT-6
41O=
•.vvv
l
.
2
I
DISSOLVED
I
.
3
&
.
4
OXYGEN CONCENTRATION
.. I.MA I
5
6
.,,
.
.
.
I
.
7 8 910
20
30
IN MILLIGRAMS PER LITER
Fig. 4. Food consumption rate in relation to dissolved
oxygen concentration (Stewart et al. 1967).
0
0
4
£
I
9
£
hI
O--------
0---0
-----
---
DISSOLVED
OXYGEN
-O EXPERIMENT- I
A EXPERIMENT- 2
EXPERIMENT-3
Q EXPERIMENT-4
-0 EXPERIMENT-5
EXPERIMENT-6
CONCENTRATION IN MILLIGRAMS PER LITER
Fig. 5. Food conversion ratio in relation to dissolved
oxygen concentration. The food conversion ration of those
fish that lost weight was considered to be 0, regardless of
the weight loss (Stewart et al. 1967).
19
high concentrations of dissolved oxygen for either equal or unequal
portions of 24 hours was markedly impaired.
Andrews et al. (1973) studied the influence of dissolved oxygen on the
Growth rates of fish
growth of channel catfish (Ictalurus punctatus).
maintained at 36% air saturation were approximately half that of fish
Food conversion ratios were submaintained at 60 and 100% (Table 3).
stantially poorer for groups maintained at the lower oxygen level. Blood
hematocrit (the packed volume of red blood cells) , hemoglobin levels, and
survival rates (100% in all groups) were not influenced by oxygen levels.
Fish maintained at the two higher oxygen levels were active and fed
vigorously, while those maintained at 36% saturation swam lethargically,
Under ad libutum
fed poorly, and had reduced response to loud noises.
feeding, dissolved oxygen concentrations influenced catfish growth more
than when a constant daily rate of 3% biomass was fed (Table 4). Herrmann
et al. (1962) and Stewart et al. (1967) postulated that food efficiency
was not substantially reduced until the oxygen level was low enough to
reduce food consumption to a level where most of the nutrients were used
to maintain the animal. Stress due to hypoxic conditions did not appear
to enhance disease or parasite problems, because no mortalities occurred
in either experiment.
Chronic effects of low dissolved oxygen concentrations on the fathead
Fathead minnows were exposed to
minnow were studied by Brungs (1971a).
constant dissolved oxygen concentrations (1.0-5.0 rag/liter) for 11 months.
The number of eggs produced per female was reduced at 2.0 mg/liter and no
spawning occurred at 1.0 mg/liter (Table 5) . Fry growth was reduced sigFry
nificantly at all concentrations below the control (7.9 mg/liter).
survival was reduced at 4.0 ramg/liter and lower dissolved oxygen concenEighteen percent of the survivors at 4.0 mg/liter
trations (Table 6).
were deformed. The time required for hatching was increased at successively lower oxygen concentrations by as much as 50%, from 5.0 days under
No effect on percent
control conditions to 7.8 days at 2.0 rag/liter.
hatch was observed. Fry growth, therefore, was the most sensitive indi-
20
Table 3. Experiment 1--Groups of catfish were
maintained at three oxygen levels and fed at a constant rate of 3% of their biomass daily (Andrews et
al. 1973).
Dissolved
oxygen
level
(% of air
saturation)
Average
gains
(g)
Feed
conversion
(g feed/
g gain)
Hematocrita
(%)
Hemoglobin"
(g/100 ml)
100
60
36
294a
287a
151b
1.23
1.30
1.78
33a
35a
32a
8.7a
9.9a
8.4a
a Values within a given column followed by the same
letter are not statistically different (P > 0.01, Duncan,
1965).
b Feed conversion was calculated on the basis of the
total amount of feed offered the fish during the experimental period. No attempt was made to determine what
portion was actually consumed by the fish.
Experiment 2--Groups of catfish were
Table 4.
maintained at three oxygen levels and fed three
times a day ad libitum (Andrews et al. 1973).
Dissolved
oxygen
level
(% of air
saturation)
100
60
36
Feed
Average
gaina
(g)
consumption
(% of
biomass/
day)
Feed
Conversion
(g feed/
g gain)
Hematocrit"
(%)
159a
124b
65c
3.3
2.9
2.1
1.12
1.18
1.47
37a
34a
35a
* Values within a given column followed by the same
letter are not statistically different (P > 0.01, Duncan,
1965).
21
Table 5. Spawning and egg production in the fathead minnow
reared from fry at various oxygen concentrations (Brungs 1971a),
No. of
Mean measured
dissolved oxygen
(mg/liter)
females
1.06
1.99
3.00
3.94
4.91
7.92 (Control)
4"
5
6
4
7
8
males
spawnings/
female
eggs/
spawning
eggs/
female
3'
6
4
5
5
6
0
5.4
10.0
13.3
16.3
10.6
0
171
176
150
146
127
0
925
1763
1992
2377
1349
*Three of the females and two of the males were immature at the end of the test.
Table 6.
Survival and growth during 30 days of fathead
minnow fry spawned and reared at
concentrations
Mean measured
dissolved oxygen
(mg/liter)
2.02
3.08
4.02
5.01
7.26 (Control)
various dissolved oxygen
(Brungs 1971a).
No.
replicates
No. fry
at start
Survival
(%)
Mean length
after 30 days*
(mm)b
3
4
2
2
2
70,38,65
70,42,94,91
70,96
70,73
70,70
0,0,0
10,5,3,5
30,20
64,68
50,50
8.7.9.3a
9.0,9.3
9.0,10.3
12.2,11.1
*Thirty-day period includes the time required for hatching.
bMean length of 143 newly hatched fry from a control dissolved
oxygen concentration was 5.3 mm.
eOnly two of the four replicates were continued for 30
days.
22
cator of sublethal effects of dissolved oxygen with fry survival nearly as
sensitive. Reproduction was least sensitive.
A continuous 12-month exposure of fathead minnows to elevated water
temperatures (26-34 C) showed that reproduction was more sensitive than
survival (Table 7), growth, or egg hatchability in assessing the effect of
The number of eggs produced per female, the
temperature (Brungs 1971b).
number of eggs per spawning, and the number of spawnings per female were
each gradually reduced at successive temperatures above the control (23.5
C) (Table 8).
No spawning or mortality occurred at 32 C, which was the
Male secondary
lowest temperature where growth was apparently reduced.
sexual characteristics were less developed at 30 C than at lower temperatures. Brett (1944) demonstrated that acclimation to higher temperatures
for the fathead minnow increased the upper lethal temperatures. He
reported a maximum lethal temperature of 34 C. Hart (1947) estimated the
upper incipient lethal temperatures to be 28.2 C after 2000 minutes for
fish acclimated at 10 C, 31.7 C after 1800 minutes with acclimation at 20
C, and 33.2 C after 9000 minutes when fish were acclimated at 30 C.
Andrews and Stockney (1972) studied the interactions of feeding rates and
environmental temperature on growth, food conversion, and body composition
of channel catfish. Daily feeding rates were 2, 4, and 6% of total biomass of fish in each aquarium. The resulting data indicated that greater
growth and lower conversions were obtained at 30 C than at either 26 or 34
C (Fig. 6 and 7), which agrees with West (1966) who found that the optimal
temperature for growth was between 29-30 C, with the maximum food conversion efficiency at 28.9 C. Shrable et al. (1969) also suggested that the
most rapid rate of digestion was at 26.6-29.4 C and Kilambi et al. (1970)
found that the optimum condition for raising channel catfish was 32 C with
a 14-hour photoperiod.
23
Table 7. Fry survival and length at three temperatures
(90 days) 1 (Brungs 1971b).
Chamber
4C
5C
6C (Control)
Mean
measured
temperature
(C)
Number
of fry
at 45
days
27.3
25.9
21.5
40
40
40
Male
Female
Survival
(%)
Mean
length
(mm)
Standard
deviation
Range
(mm)
Meanlength
(mm)
98
100
100
43(23)2
46(19)
44(26)
3.4
2.9
3.0
37-50
40-51
39-51
39(16)
41(21)
39(14)
Standard Range
deviation ( mm)
2.5
2.3
2.1
35-43
36-44
35-43
I Spawning in 30 C chamber ceased before the fry survival and growth studies were initiated.
2.Numbers in parentheses indicate number of fish.
Table 8. Egg production data and percent of hatch
six temperatures (Brungs 1971b).
Chamber
1A
1B
2A
2B
3A
3B
4A
4B
5A
5B
6A (Control)
6B ( Control)
Mean
measured
Mean
temperature Number of Number of spawnings/
(C)
spawnings females
female
33.7
34.0
31.9
31.8
29.8
30.0
27.8
27.9
26.1
26.2
23.5
23.5
0
0
0
0
21
24
109
67
150
89
155
144
0
0
7
3
7
7
7
7
8
6
6
8
0
0
0
0
3.0
3.4
15.6
9.6
18.8
14.9
25.8
18.0
1All eggs were hatched in the "A" side of the test chambers.
24
at
Number
of eggs
Mean
eggs/
spawning
Mean
eggs/
female
Percentage
hatch
0
0
0
0
1,414
1,064
13,209
7,942
25,131
14,199
29,570
26,983
0
0
0
0
67
44
121
119
168
160
191
187
0
0
0
0
202
152
1,887
1,135
3,590
2,367
4,928
3,373
0
0
0
0
67
80
90
88
95
96
87
94
FEEDING RATES
800
A
6 percent
*
4 percent
18
22
2
600
0
4c
Iw
qc
400 -
'U
I'
200
0
.
26
34
30
TEMPERATURE
(C)
Fig. 6. The relationship between environmental
temperature and average percentage gain in 12 weeks
of channel batfish fingerlings fed at three feeding
rates.
Each point represents average values from
duplicate aquaria each containing twenty fish (Andrews
and Stickney 1972).
2
I-.
10
2
0
I
8
6
U
0
0
0
4
Is.
"18
22
26
TEMPERATURE
30
34
MC)
Fig. 7. The relationship between environmental
temperature and food conversion ratio (g feed/g gain)
for channel catfish fed at three feeding rates. Each
point represents combined data from duplicate aquaria
each containing twenty fish (Andrews and Stockney
1972).
25
Accliation
The phenomenon of acclimation may change the point at which a fish is
affected by dissolved oxygen or temperature. A clear shift in the lethal
level of oxygen in relation to acclimation was shown by Shepard (1955) for
the eastern brook trout and for three warmwater species by Moss and Scott
(1961).
Shepard (1955) suggested that adjustments might be made in the
oxygen carrying capacity of the blood, thus increasing its ability to
extract oxygen from the water. Prosser et al. (1957) found this to be
true for goldfish. MacLeod and Smith (1966) found that fathead minnows
increased their hematocrit in response to low oxygen, thus making more
hemlobin available for oxygen transport in the blood.
As well as resulting in blood oxygen capacity changes, acclimation can be
behavioral in nature (Davis 1975). Considerable energy can be expended by
a nonacclimated fish reacting behaviorally to low oxygen conditions.
Acclimation lowers the magnitude of these reactions and enables energy
conservation during oxygen stress.
Any such ability to adapt to low
oxygen should be useful, especially if the transition to a low oxygen
regime is slow enough to enable acclimation to occur without severe
physiological stress. However, this mechanism would be of little help to
fish suddenly encountering low levels of oxygen. The ability to acclimate
can be considered as a built-in safety factor for species encountering low
oxygen conditions.
Acclimation temperature has a direct positive effect upon avoidance
temperatures of species tested (Mathur et al. 1983).
Fishes avoided
temperatures higher than their acclimation temperatures (Fig. 8).
The
differences between avoidance and acclimation temperatures decreased as
acclimation temperatures increased.
Strong evidence of similarities
rather than differences between populations and among species within a
family was demonstrated. Moss and Scott (1961) studied the differences in
survival between fish exposed to rapid drops in dissolved oxygen to a
critically low point from near saturation (shock test) and fish exposed to
dissolved oxygen declines gradually over a period of time (acclimation
26
L
OU
%MO
0.
E
4)
4)
#I
V
*0
0
>
<
Acclimation
Temp.(C)
Fig. 8. Plot of the published data on the relationship between
avoidance temperature and acclimation temperatures of fishes within
a family.
The 90% confidence intervals of the individual predicted
values are shown. 'Cherry et al. (1975, 1977); -, Muddy Run-Conowingo (Mathur et al. 1983).
27
test). The minimal dissolved oxygen survived by fish in acclimation tests
was lower than that survived in shock tests at any given temperature
(Table 9). The metabolic mechanisms of the species considered appear to
be highly adaptive. In addition, data from the starved and obese groups
of channel catfish used in shock tests at 30 C suggest that excessively
fat fish are more sensitive to low levels of dissolved oxygen than are
fish that have been normally fed.
Allen and Strawn (1971) noted that juvenile channel catfish, changed from
lower to higher temperatures, were nearly reacclimated in 1-3 days, measured in terms of change in heat resistance. However, fish changed from
high to low temperatures required 4-14 days to approach acclimation. In
both changes, complete reacclimation required at least 12 days, indicating
that tempering for a few hours would increase heat tolerance to same
extent, but almost 1 day would be required to significantly increase it.
Channel catfish could be held in cold water for 1-2 days with only a
partial loss of heat tolerance. Fish exposed to fluctuating temperature
should retain a relatively high heat tolerance.
Diana (1983) found that largemouth bass acclimated to a thermocycle
exhibited no stress during rapid temperature changes, and the metabolic
rate at each end point in temperature was similar to fish acclimated to
that temperature. A major reason for stress responses to altered temperature found in earlier studies may be that the fish had been exposed only
to constant temperatures in the lab. Diana's (1983) results make sense in
that largemouth bass are considered temperature eurytherms (Magnuson et
al. 1979) and regularly undergo seasonal temperature extremes from 1 to 32
C.
Their response to temperature fluctuations is more variable than
stenotherms.
Venables et al. (1977) determined that bass were metabolically acclimated to a new constant temperature after only 3 days, much
less than the usual 2-week acclimation period recommended for most fish
(Beamish 1964).
Fish do not appear to handle fluctuations in oxygen as well as they do
those in temperature.
Exposure to cycling oxygen levels appeared to
28
Table 9.
Critical levels of dissolved oxygen (p.p.m.)
as determined in shock and acclimation tests (Moss and Scott
1961).
Type of test and species
Temperature
(degrees centigrade)
(degrees centigrade)
35
30
25
Shock tests
Bluegill
Largemouth bass
Channel catfish
0.75
0.92
0.95
1.00
1.19
1.03
1.23
1.40
1.08
Acclimation tests
Bluegill
Largemouth bass
Channel catfish
0.70
0.83
0.76
0.80
0.83
0.89
0.90
1.23
0.92
29
reduce the growth of largemouth bass juveniles at 26 C (Stewart et al.
Similarly, Whitworth (1968) observed weight loss in brook trout
1967).
exposed to daily fluctuations in oxygen.
Behavioral Responses
A number of other behavioral responses to hypoxia have been reported.
Scherer (1971) described loss of normal avoidance of high light intensities (negative phototaxis) in walleye when dissolved oxygen fell to 2-4
mg/liter. It has also been shown that fish generally become more active
in hypoxic water and attempt to move away from the low oxygen region
(Randall 1970).
Avoidance behavior has been reported for a number of
species, although the reported dissolved oxygen threshold for a given
It is not clear whether avoidance
response often varies considerably.
behavior in response to low oxygen constitutes a highly directed form of
behavior.
It may result simply from increased locomotor activity with
more random movement, which is satisfied by discovery of improved oxygen
conditions. Avoidance behavior in low oxygen would have survival value
and the presence of this behavioral pattern suggests that fish periodiImplicit in a behavioral response to low
cally may benefit from it.
oxygen would be sane means of detecting low oxygen concentrations. The
physiological responses to hypoxia are rapid, consisting of circulatory
and ventilatory changes in trout and tench within a few seconds of the
Such rapid
onset of hypoxia (Eclancher 1972, Randall and Smith 1967).
responses suggest the presence of an oxygen receptor system on or near the
gills.
Whitmore et al. (1960) investigated the avoidance reactions of salmonid
and centrarchid fishes to low oxygen concentrations. Largemouth bass and
bluegill (Lepomis macrochirus) markedly avoided concentrations near 1.5
mg/liter, but they showed little or no avoidance of the higher concentrations. Only bass showed any avoidance of concentrations near 4.5 mg/
liter. Avoidance by fish of water with low dissolved oxygen concentrations was shown not only by shorter excursions in distance and time into
the experimental channels but also by fewer entries into the low oxygen
30
channels. Avoidance revealed by periodic counts of fish in the channels
could be the result of gradually produced discomfort and stimulation at
Shelford and Allee (1913) found that
reduced oxygen concentrations.
several fish species avoided water with low oxygen content in a gradient
tank by turning back into water with more favorable concentrations.
Ironically, the mechanism necessary to move fish fran hypoxic areas, the
locomotory system, is also influenced by low dissolved oxygen. Katz et
al. (1959) reported that 6-cn bass could resist a water current of 25 cm/
second at 25 C in September at 2.0 mg/liter oxygen. However, in December
at 15-17 C, they were unable to do so even when oxygen levels were 5.0 mg/
liter. Dahlburg et al. (1968) studied the influence of dissolved oxygen
and carbon dioxide on the swimming performance of largemouth bass and coho
salmon (Oncorhynchus kisutch). They concluded that concentrations of free
carbon dioxide even above 40 mg/liter, which is rarely found in nature,
had virtually no adverse effect on the swimning performance of largemouth
The swimming speed of largemouth bass at low carbon
bass (Fig. 9).
dioxide concentrations decreased with dissolved oxygen concentrations
At concentrations above 6.0 mg/liter, the
below 5-6 mg/liter (Fig. 10).
performance of bass apparently was independent of oxygen concentration.
Effects on Reproductive Success
The adult life stage of fish is not the only portion of their life cycle
impacted by low dissolved oxygen. Developing fish eggs and larvae have a
number of responses to low oxygen, including respiratory dependence,
retarded growth, reduced yolk sac absorption, developmental deformities,
and mortality. As developnent proceeds, oxygen requirements of eggs and
larvae increase. Brungs (1971a) found larvae to be the most sensitive
stage in the life cycle of the fathead minnow, when exposed to reduced
oxygen levels of 1.0-5.0 mg/liter.
Carlson et al. (1974) studied the effects of lowered dissolved oxygen
concentrations on channel catfish embryos and larvae. The effects of low
oxygen concentrations on embryos and larvae were evident at all reduced
31
U
w
cc
w
0
.J
z
w
nON)
RE)
TION)
z
I
DISSOLVED OXYGEN CONCENTRATION IN
MG PER LITER
Fig. 9. Mean final swimming speeds, expressed in
L/sec, of largemouth bass at various concentrations of
dissolved oxygen and carbon dioxide. The curve relating
swimming speed -to oxygen concentration is the curve
that was fitted to all data from tests at the low concentration of free carbon dioxide (probably <3 mg/liter)
only, most of which (those from early tests) have not been
plotted here. The stated free carbon dioxide levels of 48,
21, and 38 mg/liter are means of observed values for
individual tests ranging from 44 to 55 mg/liter, 17 to
23 mg/liter, and 24 to 38 mg/liter, respectively (Dahlburg
et al. 1968).
U 'M
w6
9)
IL5
w
-J
Z
4
Q
w
hi
w 3
z
2
ii
0A
Ln
DISSOLVED
OXYGEN CONCENTRATION
MG PER LITER
IN
Fig. 10.
Relationship between the mean final swimming
speed of largemouth bass, expressed in L/sec, and the
dissolved oxygen concentration at nearly natural, low concentrations of carbon dioxide (<3 mg/liter) and temperatures
near 25 C (Dahlburg et al. 1968).
32
concentrations tested at 25 C. Embryo pigmentation was progressively
lighter as the oxygen concentration decreased. At reduced oxygen concentrations, hatching was prolonged by 0.5-2.0 days and feeding was delayed
by 1-3 days. Growth of survivors was reduced at all lowered concentrations so that average lengths attained were 0.9-2.8 mm less than those of
controls. Survival was also significantly less at 30 and 50% saturation.
No hatching occurred at 20% saturation at 25 C. Only three fish survived
at 30% saturation and 28 C. Channel catfish embryos and larvae were
sensitive to all reductions in dissolved oxygen, although survival was
Siefert and Spoor
only slightly affected at 60 and 70% saturation.
(1974), using similar testing methods, found that early-stage white
suckers and walleye were not harmed at 50% saturation.
Dudley and Eipper al. (1975) investigated the survival of largemouth bass
embryos at low dissolved oxygen concentrations. Oxygen concentrations that
caused mortalities during hatching were 2.0, 2.1, and 2.8 mg/liter at
incubation temperatures of 15, 20, and 25 C, respectively (Fig. 11), although some embryos developed and hatched at oxygen concentrations as low
as 1.0, 1.1, and 1.3 mg/liter.
The increasing embryo biomass requires
more oxygen as it develops to the hatching point. Laurence (1969)
reported that oxygen consumption of largemouth bass embryos incubated at
20 C increased from 0.06 microliters during the first 24 hours to 1.92
microliters during the last 24 hours of the 96-hour prehatching period.
During hatching, largerouth bass embryos move vigorously (Carr 1942) .
Laurence (1969) noted a possible peak in oxygen consumption during
hatching.
The increased oxygen consumption coincident with hatching
of largemouth bass embryos undoubtedly increases oxygen needs above that
which some conditions can supply.
Like fish, eggs also show respiratory dependence (Fry 1957). Alderdice and
Brett (1957) reported a gradual increase in the dissolved oxygen requirements of chum salmon (Qncorhynchus kA) eggs as development progressed.
Eggs at early stages of incubation required about 1 mg/liter oxygen, while
those about to hatch required over 7 rag/liter. Eggs held at low oxygen
tensions experienced reduced growth and retarded development. Deformities
33
EXPNM
--.
-----
*
EXPE
5c
a
20 C
0
25C
A
Survival at 90%
Oxygen Saturation
m.--
6
(I)
0
a
cu
s_-
SO-
/
/ /
/
f
0
/
nA°
/
A
/
/
AA/
An
A
Oxygen Concentration (mg/I)
Fig. 11. Relationships between embryo survival and oxygen concentration
at three incubation temperatures. Each point represents the percent of 40
embryos (Experiment II) or 150 embryos (Experiment III) which survived to
completion of hatching. Curves were fitted by eye to Experiment III data,
Although at 20 C survival in Experiment II agreed with that found in Experi'
ment III, survival was higher in Experiment III than in Experiment II at both
15 C and 25 C (Dudley and Eipper 1975).
34
in embryos exposed to hypoxia were observed, with mortality occurring at
Others have reported
the stage when the circulatory system develops.
similar increased oxygen requirements of eggs as development proceeds
(Lindroth 1942, Hayes 1949, Wickett 1954, Guilidov 1969).
Minimum Oxen Standard
Setting a minimum allowable summer dissolved oxygen concentration for
water quality standards in warmwater systems would be extremely difficult.
In his classic work, Ellis (1937) concluded that a minimum summer
dissolved oxygen concentration of 5 mg/liter was necessary to support
good, mixed fish faunas. Coble (1982) looked at fish populations in a
Wisconsin river in relation to dissolved oxygen and found sport fishes and
a variety of other species in water with less than 5 mg/liter oxygen.
However, the quantitative measure, percent sport fish 2100 mn in samples,
revealed that the proportion of a fish community composed of sport fishes
was larger in waters, with higher levels of dissolved oxygen. Moreover,
with a measure of dissolved oxygen concentration of daytime or averaged
values, the level of 5 mg/liter could be identified as a point of
departure between good and poor fish populations.
Davis (1975)
established critical oxygen and temperature criteria for Canadian aquatic
life (Fig. 12).
His criteria contained three levels to assess potential
risk: Level A provides a high safeguard for very important aquatic
populations; with Level B, there is a possibility of moderate risk to
aquatic organisms with depressed oxygen concentrations; and with Level C,
a large portion of a given population or community may be severely
affected by low oxygen levels, especially on a chronic exposure basis.
35
A
100
4
0oa
60
0
40
I
. ..
-AA-I*0O SA.
Ow---*
O
I
20-
a
10
5
SLASONAL
15
TEMPERAITU€
20
ZS
MAXIMUM
- C
B
I?
**9091"eI
N
hi
ao
**eg
**~.*fP*.O
s«O
MI*CMI*
0
O
*
J
0*°
S
a
Ft
I
e..
A
W mmm
--- *
N
I
o
**
S
K
a-
0 1%
a
1
StASONAL
Ti
aA
1
I
TIMPEOATULI
A
4 f
U
MAXIMUM
A,-•
0L
-C
Fig. 12. A, Oxygen criteria, expressed as seasonal
minima in percent saturation, at various seasonal temperature
maxima. Three levels of protection, A, B, and C, are given
as defined in the text. Values used in derivation of the
criteria are summarized in Table 10; B, Oxygen criteria,
expressed as oxygen minima in mg 02 /liter (ppm), at various
seasonal temperature maxima. Three levels of protection, A,
B, and C, are given as defined in the text. These values
were calculated from the percentage saturation values illustrated in (A) and summf4xrzed in Table 10 (Davis 1975),
36
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