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THE EFFECTS OF DISSOLVED OXYGEN, TEMPERATURE, AND LOWSTREAM FLOW ON FISHES: A LITERATURE REVIEW

Aquatic Biology SectionTechnical Report

by

D. C. Dowling and M. J. Wiley

Report to Springfield City Water,Light, and Power Company

)logy Technical Report 1986(2)

NATURAL HISTORY SURVEY

J UN 2 , 1986

ILLINOIS LIBRARY- NATURAL HISTORY -

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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

Robert W. Gorden, HeadAquatic Biology Section

M. J. Wiley, Princip 1 Investigator

\

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 sub-stances in natural waters, oxygen is one of the most significant. Theannual 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 broadtemperature fluctuations have shown that the oxygen content of flowingwaters is generally highest in winter and lowest in late summer. Insmall, slow-flowing streams of northern latitudes, decreased day lengthand ice and snow cover may inhibit photosynthesis and prevent entry ofoxygen from the air, thereby depressing the oxygen concentration.

Sometimes this oxygen depression becomes quite severe, stressing theresident fish populations, as was documented in the River Ob in Siberia(Mosevich 1947, Mossewitsch 1961). They documented oxygen levels fallingto 14% saturation; the fish left the affected regions and congregated ina few more tolerable areas.

The vernal decline of oxygen content in slow-flowing streams may beattributed to the action of spring floods in removing vegetation and tothe increased rate of decay of organic material with increasing watertemperature. Further decreases in oxygen content toward late summer aredue to one or more factors. High temperature water, reaching a maxinum inlate summer, holds less oxygen in solution; decreased discharge results indiminished physical mixing and reoxygenation; and greater decomposition ofnaturally produced organic material uses some of the available oxygen(Reid 1961). When rainfall is light, rivers are generally fed bygroundwater. Often underground water is devoid of oxygen and containslarge amounts of carbon dioxide, because of its exposure to organic matterand 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 impartinghigh 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 createseasonally low oxygen concentrations. In the U.S. Southeast and Mid-

western regions, and doubtless in other areas with similar hot and drysummers, 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 aninky 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 summerand 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 environ-

ment 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.

1959). In Smiths Branch, a small warmwater stream in Vermilion County,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 destruc-

tive to aquatic animals during September and early October, as leaves and

other dead plant materials accumulated in the pools, decomposed, and pro-

gressively increased oxygen demand.

Most fish withstood extreme drought conditions in at least a few parts of

the stream. Re-establishment of the populations began as soon as thestream 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

Branch by the end of the first summer. Only three species were sig-

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 streamin 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

reduced, subsequent populations may be affected (Hynes 1958). Follow-up

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

into which they retreat remains moist; in pool and riffle headwaters,

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 able to withstand the effects of

drought than are fish, because of egg, nymphal, or pupal diapauses, emer-

gence, or burrowing (Canton et al. 1984). Recolonization is rapid

following resumption of flow (Hynes 1975, Townsend and Hildrew 1976,

Williams and Hynes 1976a, 1976b, 1977).

In a Colorado stream in 1978 and 1979, Canton et al. (1984) investigated

the impacts of low flow in 1978. Brook trout (Salvelinus fontinalis)

collected in 1978 were thin and moribund when handled. Average weights

4

for each age class were lower than expected on the basis of other North

American studies (Carlander 1969), and as a result, the mean condition

factor (K) was lower in 1978 than in 1979. Regenerated scales, which

often indicate environmental stress, 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

the drought, with no fish collected in most of the study area. After

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 pre-

dation pressure. However, the large increase in total density in 1979again 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 wasnot significantly affected.

5

When faced with low dissolved oxygen, many fish species will migrate tomore 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

oxygen decreased in abundance or disappeared. They were replaced byspecies that were tolerant to organic enrichment and low dissolved oxygen.

Adversely affected species included important game fishes, such assmallmouth bass (Micropterus dolmieui), largemouth bass (L salmoides,and black crappie (Pomoxis nigrmaculatus). Favorably affected speciesincluded 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 Julyand early August 1977, dissolved oxygen concentrations declined to a lowof 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 oxygenatedwater is accessible (Doudoroff and Shumway 1970). Concentrations below 4-

5 mg/liter apparently interfere with upstream migration of adult salmonids(Sams and Conover 1969), but migration of these and other anadromous fishthrough 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 concen-

tration of 4.5 mg/liter (Chittenden and Westman 1967).

Time of year may delay recolonization of streams, even if they are fullyrecovered in terms of water quality and flow. Larimore et al. (1959)

found that fish moved back into Smiths Branch quickly during April andMay, but re-invasion was slow for several weeks during late February andearly March, after stream flow resumed. Katz and Gaufin (1953) also

observed a delay in re-invasion during winter; fish in winter did notmove 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 dis-solved 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 ofabrupt 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 streamwith environmentally unstable headwaters. Fishes in other physically

harsh environments have wide physicochemical tolerance limits, which canbe related to their distributional limits. Cross and Cavin (1971) foundthe red shiner (Notropis lutrensis), which is highly successful inintermittent prairie streams, to be more tolerant of low oxygen conditions

that the bluntface shiner (L. camurus), which is restricted to permanentlyflowing upland streams.

7

Matthews and Styron (1981) also reported that a greater portion of head-

water than mainstream fantail darters (Etheostma flabellare) survivedbrief periods of oxygen stress. Small intraspecific differences in

tolerance could be important, if over time they resulted in ecotypes

adapted to specific parts of a heterogeneous watershed. Intraspecific

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 distri-

bution of red shiners over physicochemical ranges and found that juvenilesoccupied 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 shelterin February, May, August, and October. Both groups occupied locations

with very slow current. The mean habitat breadth of adults changed moreas environmental conditions changed than did that of juveniles. Stress on

red shiners was probably greatest in October, near the end of a lengthydrought, and in December-January when it was unusually cold.

Temperature and oxygen tolerances of four cyprinid species in a south-

western 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 abundancein the sumner and apparently failed to spawn in 1976. The Arkansas Rivershiner, 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 selectionpressure 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 concentrationsNumber ; Temperature found lethal Standard Standardof fish in degrees F. M axi- deviation error

mum mum Median Mean]imum m um I I'

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 1 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

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

success than was low oxygen tolerance. The red shiner and the plains

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 Plainsfishes from eight families to progressive hypoxia. He noted that when

freshwaters become hypoxic some fishes respond by breathing air facul-

tatively. Others remain aquatic breathers, rising to the surface where

they irrigate their gills with relatively oxygen-rich surface film. Allspecies 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 tocompetition 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 fre-quently, 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 numerousbiological factors: through its senses, by its associates, its school,

predators, prey, algae, and aquatic plants, in addition to the changingphysical and chemical characteristics of the water, which vary with depth

and season (Sondheimer and Simeone 1970). It is this array of factorsthat actually constitutes what chemical concentrations a fish will surviveand where it will reside. However, the effects of the combined influenceof current, oxygen, and temperature on lotic species has not been accu-

rately assessed (Brown 1971). Individual laboratory experiments have

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

varies between individuals of the same species. Daily fluctuations are

probably part of larger rhythms that harmonize with the physiological

cycles of the animal. Wells (1914) pointed out differences in the resis-

tance 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

maintenance, measured under laboratory conditions. It is close to, but

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

oxygen threshold, which affords good protection for fish species. In

animals tolerant of low oxygen, the curve in Fig. 2 shifts to the left,

11

E

Ccb0

Ero0o

280c

o 200

120

40

020 60 100 140 180

Partial Pressure of Oxygen, mm Hg

Fig. 1. Relation between standard and active oxygen con-sumption for carp. Solid symbols represent values for fishacclimated to air saturation. The standard curves have beentaken from Fig. 2. The broken line represents the results forcarp acclimated to the various partial pressures of oxygenapplied. The weight range of carp used at the active levelwas 36 to 750 g at 10 C and 140 to 540 g at 20 C (Beamish 1964).

I .

12

--

-

, 9 EON 1 %d 1 l

I

hi

4

B.

zhi0

K0Ii.0

hi

4

ENVIRONMENTAL0 2

Fig. 2. Relation between standard and active (Maximum) oxygen

uptake rates at different environmental oxygen concentrations (from

Hoar 1966).

13

ZONE OF

I ZONE OF RESPIRATORY DEPENDENCE I RESPIRATORY REGULATION

ZONE OF

L RESISTANCE I4 ZONE OF TOLERANCE

INCIPIENT LMITING TENSION

STANDARD METABOLIC RATE

INCIPIENT LETH AL TENSIONINCPIET ETHL ENSION

while in animals requiring higher levels of oxygen the curve shifts to the

right. Thus, the oxygen tension that produces respiratory dependence

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) (Holetonand Randall 1967a, 1967b) and carp (Cyprinus car.piQ) (Garey 1967) showelevations 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 factorsoperate 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 levelsfor 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, andother activities. Stewart et al. (1967) investigated the influence of

oxygen concentrations of the growth of juvenile largemouth bass. Its

growth and food consumption rates increased with increases in dissolvedoxygen 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 etal. (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 ration-

ales 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 reportedtemperature 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 effectsfirst become apparent thus providing a "biological indication" of the onset of hypoxicstress. A) Freshwater species, B) Marine nonanadromous species, C) Anadromous species,D) Eggs and larvae (freshwater and marine species) (Davis 1975).

Level of dissolved oxygenTemp Response to oxygen

Species Size (C) P0 2-mm Hg ml O 2 liter mg O 2 /liter 7, Satn. level Itrema

A. Freshwater species

Arctic char,Salvelinus

alpinus

brown bullhead,Ictalurus nebulosus

Rainbow trout,Salmo gairdneri

carp,Cyprinus carpio

Walleye,Stizostedion vitreum

Largemouth bass,Micropterus

salmonides

Brook trout(speckled trout)Salvelinus

fontinalis

- 2±.05 25 1.53 2.18 15.8 "signs of asphyxia Holetmand loss of equilibrium" (1973)

43-127 9-10 60 4.87-4.76 6.95-6.80 38.0 onset of oxygen -g dependent oxygen

uptake

682-1136 20 78 3.20 4.59 50 below this level bloodg is not fully saturated

with oxygen

120-250 8.5-15 80-100 3.63-5.14 5.18-7.34 50.7-63.7 circulatory changesg occur, including a

slowing of heart

77-350 g 10 50 2.51 3.59 31.7 standard oxygen up-take elevated (de-

30-600 g 20 80 3.29 4.71 51.3 pressed at lowerlevels)

235-785g 13.3-25 80 2.74-3.04 5.39-4.34 50.9-51.6 blood ceased to befully saturated with02 below this level

2 yr-old 22 35.2-70.4 2.8-1.4 4-2.0 45.3-22.7 loss of negative pho-totaxis (light avoid-ance behavior)

70.4-96.8 3.85-2.8 5.5-4.0 45.3-62.3 increased mobility,"darting" behavior

5-9 - - 3.15 4.5 - some indication ofcm avoidance behavior

I.I 1.5 - marked avoidance

682-1136 20 77.95 3.21 4.59 50 blood ceases to beg fully satd. with 02

below this level

(196)Irving et .(1941)

Randsdand SIta

Beamilh(1964)

Itazaw(1970)

ScShr(1971)

Whitmortet al.(190)Irvialgetid.(1941)

17-65 5 100 5.66 8.09 63.2 onset of Or-dependent Grahamg metabolism (1949)

8 80 4.19 5.99 50.7 reduced cruising speed20 154 6.34 9.06 98.8 onset of Or depen.

dent metabolism5-20 116.9-118.7 6.72-4.82 9.6-6.88 75 reduced activity, all

temps.

56-140 10,15 80 4.03-3.63 5.75-5.18 50.7-51.0 standard oxygen up- Beamishg take reduced below (1964)

this level

Brown trout, 682-1136 20 77.95 3.21 4.59 50 blood ceases to be Irving et aLSalmo trutta g fully satd. with O (1941)

below this level

Rainbow trout, 13.3±1.4 17± .5 156.6 6.82 9.74 100 any reduction in DowningSalmo gairdnerl cm oxygen led to more (1954)

rapid death in cyanide

20 mos old 8-10 78.85-79.95 4.16-3.97 5.94-5.67 50 43% reduction in Jonesmaximum swimming (1971b)speed

21-23 77.85-77.6 3.15-3.04 4.50-4.34 50 30% reduction inmaximum swimmingspeed

235-510 2.3-13 100 6.11-4.71 8.73-6.74 63.1-63.6 blood is not fully Itazawag satd. with oxygen (1970)

below this level

- 15 78.45 3.55 5.08 50 altered respiratory Kuttyquotient, little capacity (1968a)for anaerobic metab-olism below this level

- 15 80-100 3.63-4.53 5.18-6.47 51.0-63.7 changes in oxygen Randalltransfer factor and et al. (1967)effectiveins of Oexchange occur

15

Table 2 (continued).

Level of dissolved oxygenTemp Response to oxygen

Species Size (C) POa-mm Hg ml O0/liter mg O/liter 7% SaIa. level Reference

t t 400-600 13.5 80 3.74 5.35 51.0 breathing amplitude Hughes andg and buccal pressure Saunders

elevated (1970). t approx. 300 10,15, 80 3.30-4.03 4.71-5.75 50.7-51.3 below this level blood Cameron

g 20 is not fully satd. with (1971)oxygen

1-11 17.5 93.8 4.05 5.78 60 toxicity of zinc, lead, Lloydg copper, phenols in- (1961)

Screased markedlybelow this level

Largemouth bass, juvenile 25 92.3-110.7 3.5-4.2 5-6 59.7-71.6 final (maximum sus- DahlbergMkropterus tained) swimming et al. (1968)

salmoldes speed reduced belowthis level

Ulvuell, 5-9 - - 2.1 3.0 - some avoidance WhitmoreL/pomis cm behavior et al. (1960)

macrochirus 1.1 1.5 - strong avoidance

B. Marine nonanadromous species

-4it perch, adults 11 80 3.19 4.56 50.8 blood ceases to be Webb andMacochilus vacca fully 02 saturated Brett

below this level (1972)

Dofish, 2.5-6.0 10 115 4.68 6.69 72.92 blood ceases to be Lenfant andSq"as suckleyl kg fully 0O saturated Johansen

below this level (1966)

Dogtish, 150-600 12-14 80 3.12-2.95 4.46-4.20 50.8-51.0 Oxygen-dependent Hughes andScylo inus g metabolism begins Umezawa

caniculas below this level (1968a)

Dragonet, 70-140 11-12 125 4.98-4.88 7.12-6.97 79.4 Oxygen-dependent Hughes andCutieymus lyra" g metabolism below Umezawa

this level (1968b)100 3.97-3.90 5.67-5.58 63.5 altered heart rate

Dragnet, 80-120 - 80 - - - increased ventilatory Hughes andCa•Meymus lyrea muscle activity below Ballintijn

this level (1968)a1sh5 - 11 150 5.98 8.54 95.2 blood ceases to be Hanson

Ndretegus colliei fully Oa saturated (1967)below this level

Atlantic cod, 1.14-2.33 5 158.2 7.14 10.20 100 any reduction in SaundersGdus morsua kg ambient O0 level (1963)

produces rise in ven-tilatory water flow

C. Anadromous speciesSeilbead,Selsm gairdnert - see freshwater species

Sockeye salmon, 50 g 20-24 15Owerkynchus nerka

1579 g 13

1.5-1.7 15kg

Caoe salmon, 6.3-11 "summeroerwhynchus cm tempera-

kisutch tures"

5.1-14.8 12±1cm

to . Juvenile 10-20 1

t o t "9 20

90 ot to 10

$5.9-154.9 6.42-5.97 9.17-8.53 100 available oxygen level Brettappears to limit active (1964)metabolism andmaximum swimmingspeed

100 4.72 6.74 63.6 blood ceases to be Davisfully 0O saturated (1973)below this level

78.45 3.55 5.07 50 elevated blood and Randallbuccal pressure and and Smithbreathing rate increased (1967)

- 3.15 4.5 - erratic avoidance Whitmorebehavior et al. (1960)

130.8 6.3 9.0 83.1 below this level acute Hicks andmortality in kraft DeWittpulpmill effluent (1971)mincreased

57.7-155.9 7.93-6.42 11.33-9.17 100 reduction of 02 below Davis et al.saturation produced (1963)some lowering ofmaximum sustainedswimming speed

155.9 6.42 9.17 100 as above Dahlberget al. (1968)

136-67.9 5.6-2.8 8.0-4.0 87.2-43.6 growth rate propor- Hermanntional to oxygen level (1958)with best growth at8.0 ms/liter, lowestat 4.0 mg/liter

16

Table 2 (continued).

Species

Chinook salmon,Oncorhynchus

Ishawytscha

" "* J

Atlantic salmon,Salmo salar

American shad,Alosa sapidissima

Rainbow trout,larvae,

Salmo gairdnerl

Chum salmon eggs,Oncorhynchus keta

Level of dissolved oxygenTemp(C) PO-mm Hg ml O/liter mg Oa/liter %7 Satn.Size

Response to oxygenlevel Refermw

6.3-11 "summer - 3.15 4.5 - marked avoidance of Whitmencm temps" this level in summer et al. (11

"fall - - 4.5 - little avoidance oftemps" this level in fall

uveniles 10-20 157.7-155.9 7.93-6.42 11.33-9.17 100 reduction of O Davis etd.below saturation pro- (1963)duced some loweringof maximal sustainedswimming speed

87-135 15 69.6 3.15 4.5 44.33 Salmon stop swimming Kuttysadg at a speed of 55 Saunden

cm/s at 02 levels (1973)below this. Fasterswimming requiresmore oxygen

- - - 1.75-2.1 2.5-3.0 - threshold for migra- Chittendation through (1973)polluted areas

2.8 4.0 - necessary forspawning areas

1+8dayold

larvae

earlystageeggs

pre-hatcheggs

5 dayeggs

10+1 100

10 13.9

10

8.0-8.2

>97.4

22.2

D. Eggs and larvae5.03

.7

>4.9

1.17

7.18 63.4 elevated heart and Holetoabreathing rates below (1971)this level. Brady-cardia (heart rateslows) sets in atlower O0 levels

1 8.8 retarded development, Alderdicedeformities, mortal- et al. (1953)ity, respiratory Oa-dependence

> 7.0 > 61.8 oxygen level requiredfor normal develop-ment and non O2-dependent metabolism

1.67 14.07 critical oxygen level Wickettfor supply of oxygen (1954)demand and non 02-denendent metabhlism

(eyed) 3.6-4.9 > 60.3b

> 3.5b > 5.0 > 38.1 b85 dayeggs

Atlantic salmon, early 5.5 9.62 .53 0.76 6.09 As above LindrothSalmo salar eggs (1942)

near (fromhatching 5 71.7 4.06 5.80 45.31 Wicketthatching 17 160.8 7.0 10.0 102.67 1954))

Atlantic salmon, eyed 10 43.14 2.17 3.1 27.36 critical 02 level for Hayes et al.Salmo salar hatching 10 98.83 4.97 7.1 62.67 supply of 02 demand (1951)(from

and non O2 -depen- Wickett,dent metabolism 1954)

Northern pike, from 15,19 > 5.18-5.66 > 2.16-2.35 > 3.35-3.09 > 33.0 necessary O 2 level for SiefertEsox lucius fertilization proper hatching, et al.

to survival and (1973)feeding developmentlarvae

fertilization 20to

hatching

fertilization 3-5to

hatching

101.2 3.15

*European species - representatives of this family found in Canadian Atlantic waters.bCalculations based on 4 C.*Estuarine fish - calculations based on salinity of 30%,.

4.5 64.9 reduced hatching at Voyer andthis level compared Hennekeyto 7.5 mg/liter 0O (1972)

> 2-3 oxygen level required Alderdicafor qptimal hatching andat 157,o salinity Forresmt

(1971)

17

MummichogLFundulus

heteroclltus

Pacific cod,Gadus

macrocephalus

0w II

Z scs-

z

ZU

at4

I<r5)ri)VS)

NTS)ys)WkJ)hn)

- iii, -,1 4 a 7 a 9 -i WD --

DISSOLVED OXYGEN CONCENTRATION INMILLIGRAMS PER LITER

Fig. 3. Growth of largemouth bass in relation todissolved oxygen concentration (Stewart et al. (1967).

18

*E

I 2 3 4 5 6 7 8 910

DISSOLVED OXYGEN CONCENTRATION IN MILLIGRAMS PER LITER

Fig. 4. Food consumption rate in relationoxygen concentration (Stewart et al. 1967).

0

4£

I9£hI

to dissolved

0

O--- -O EXPERIMENT- I------ A EXPERIMENT- 2

0---0 EXPERIMENT-3----- Q EXPERIMENT-4

--- -0 EXPERIMENT-5EXPERIMENT-6

DISSOLVED OXYGEN 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

U.500

4 0.400

aJI

>o--

3t 0.300

a

0.200W

ozo 0.100

41-

MENT-IMENT-2

MENT-4MENT-5MENT-6

20 30l I & .,, I I . . . . I. . . .. .MA

0% Ad%^ -r-

O= •.vvv

-

-

F

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 of channel catfish (Ictalurus punctatus). Growth rates of fish

maintained at 36% air saturation were approximately half that of fish

maintained at 60 and 100% (Table 3). Food conversion ratios were sub-

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,

fed poorly, and had reduced response to loud noises. Under ad libutumfeeding, dissolved oxygen concentrations influenced catfish growth more

than when a constant daily rate of 3% biomass was fed (Table 4). Herrmannet 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

minnow were studied by Brungs (1971a). Fathead minnows were exposed to

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 sig-

nificantly at all concentrations below the control (7.9 mg/liter). Fry

survival was reduced at 4.0 ramg/liter and lower dissolved oxygen concen-trations (Table 6). Eighteen percent of the survivors at 4.0 mg/liter

were deformed. The time required for hatching was increased at succes-

sively lower oxygen concentrations by as much as 50%, from 5.0 days under

control conditions to 7.8 days at 2.0 rag/liter. No effect on percent

hatch was observed. Fry growth, therefore, was the most sensitive indi-

20

Table 3.maintained atstant rate ofal. 1973).

Experiment 1--Groups of catfish werethree oxygen levels and fed at a con-3% of their biomass daily (Andrews et

Dissolvedoxygen Feed

level con-(% of air Average version Hema- Hemo-

satu- gains (g feed/ tocrita globin"ration) (g) g gain) (%) (g/100 ml)

100 294a 1.23 33a 8.7a60 287a 1.30 35a 9.9a36 151b 1.78 32a 8.4a

a Values within a given column followed by the sameletter are not statistically different (P > 0.01, Duncan,1965).

b Feed conversion was calculated on the basis of thetotal amount of feed offered the fish during the experi-mental period. No attempt was made to determine whatportion was actually consumed by the fish.

Table 4. Experiment 2--Groups of catfish were

maintained at three oxygen levels and fed three

times a day ad libitum (Andrews et al. 1973).

FeedDissolved con-

oxygen sumption Feedlevel (% of Conver-

(% of air Average bio- sion Hema-satu- gaina mass/ (g feed/ tocrit"

ration) (g) day) g gain) (%)

100 159a60 124b36 65c

* Values within a givenletter are not statistically1965).

3.3 1.12 37a2.9 1.18 34a2.1 1.47 35a

column followed by the samedifferent (P > 0.01, Duncan,

21

Table 5. Spawning and egg production in the fathead minnowreared from fry at various oxygen concentrations (Brungs 1971a),

No. ofMean measured

dissolved oxygen spawnings/ eggs/ eggs/(mg/liter) females males female spawning female

1.06 4" 3' 0 0 01.99 5 6 5.4 171 9253.00 6 4 10.0 176 17633.94 4 5 13.3 150 19924.91 7 5 16.3 146 23777.92 (Control) 8 6 10.6 127 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 fatheadminnow fry spawned and reared at various dissolved oxygenconcentrations (Brungs 1971a).

Mean measured Mean lengthdissolved oxygen No. No. fry Survival after 30 days*

(mg/liter) replicates at start (%) (mm)b

2.02 3 70,38,65 0,0,0 -3.08 4 70,42,94,91 10,5,3,5 8.7.9.3a4.02 2 70,96 30,20 9.0,9.35.01 2 70,73 64,68 9.0,10.37.26 (Control) 2 70,70 50,50 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 con-centration 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

temperature (Brungs 1971b). The number of eggs produced per female, thenumber of eggs per spawning, and the number of spawnings per female were

each gradually reduced at successive temperatures above the control (23.5C) (Table 8). No spawning or mortality occurred at 32 C, which was the

lowest temperature where growth was apparently reduced. Male secondary

sexual characteristics were less developed at 30 C than at lower tempera-

tures. Brett (1944) demonstrated that acclimation to higher temperaturesfor 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 forfish acclimated at 10 C, 31.7 C after 1800 minutes with acclimation at 20C, and 33.2 C after 9000 minutes when fish were acclimated at 30 C.

Andrews and Stockney (1972) studied the interactions of feeding rates andenvironmental temperature on growth, food conversion, and body composition

of channel catfish. Daily feeding rates were 2, 4, and 6% of total bio-

mass 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 conver-

sion 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 witha 14-hour photoperiod.

23

Table 7. Fry survival and length at three temperatures(90 days) 1 (Brungs 1971b).

Male FemaleMean Number

measured of fry Mean Mean-temperature at 45 Survival length Standard Range length Standard Range

Chamber (C) days (%) (mm) deviation (mm) (mm) deviation ( mm)

4C 27.3 40 98 43(23)2 3.4 37-50 39(16) 2.5 35-435C 25.9 40 100 46(19) 2.9 40-51 41(21) 2.3 36-446C (Control) 21.5 40 100 44(26) 3.0 39-51 39(14) 2.1 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 atsix temperatures (Brungs 1971b).

Meanmeasured Mean Mean Mean

temperature Number of Number of spawnings/ Number eggs/ eggs/ PercentageChamber (C) spawnings females female of eggs spawning female hatch

1A 33.7 0 0 0 0 0 0 01B 34.0 0 0 0 0 0 0 02A 31.9 0 7 0 0 0 0 02B 31.8 0 3 0 0 0 0 03A 29.8 21 7 3.0 1,414 67 202 673B 30.0 24 7 3.4 1,064 44 152 804A 27.8 109 7 15.6 13,209 121 1,887 904B 27.9 67 7 9.6 7,942 119 1,135 885A 26.1 150 8 18.8 25,131 168 3,590 955B 26.2 89 6 14.9 14,199 160 2,367 966A (Control) 23.5 155 6 25.8 29,570 191 4,928 876B ( Control) 23.5 144 8 18.0 26,983 187 3,373 94

1 All eggs were hatched in the "A" side of the test chambers.

24

2

w04cqcI-

'U

I'

800

600

400 -

200

0 .

FEEDING RATES

A 6 percent

* 4 percent

18 22 26 30 34

TEMPERATURE (C)

Fig. 6. The relationship between environmentaltemperature and average percentage gain in 12 weeksof channel batfish fingerlings fed at three feedingrates. Each point represents average values fromduplicate aquaria each containing twenty fish (Andrewsand Stickney 1972).

2I-.

20

IU

000Is.

10

8

6

4

"18 22 26 30 34

TEMPERATURE MC)

Fig. 7. The relationship between environmentaltemperature and food conversion ratio (g feed/g gain)for channel catfish fed at three feeding rates. Eachpoint represents combined data from duplicate aquariaeach containing twenty fish (Andrews and Stockney1972).

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 lethallevel of oxygen in relation to acclimation was shown by Shepard (1955) forthe eastern brook trout and for three warmwater species by Moss and Scott(1961). Shepard (1955) suggested that adjustments might be made in theoxygen carrying capacity of the blood, thus increasing its ability toextract oxygen from the water. Prosser et al. (1957) found this to betrue for goldfish. MacLeod and Smith (1966) found that fathead minnows

increased their hematocrit in response to low oxygen, thus making morehemlobin available for oxygen transport in the blood.

As well as resulting in blood oxygen capacity changes, acclimation can bebehavioral in nature (Davis 1975). Considerable energy can be expended bya nonacclimated fish reacting behaviorally to low oxygen conditions.

Acclimation lowers the magnitude of these reactions and enables energyconservation during oxygen stress. Any such ability to adapt to low

oxygen should be useful, especially if the transition to a low oxygenregime is slow enough to enable acclimation to occur without severephysiological stress. However, this mechanism would be of little help tofish suddenly encountering low levels of oxygen. The ability to acclimatecan be considered as a built-in safety factor for species encountering lowoxygen conditions.

Acclimation temperature has a direct positive effect upon avoidance

temperatures of species tested (Mathur et al. 1983). Fishes avoidedtemperatures higher than their acclimation temperatures (Fig. 8). Thedifferences between avoidance and acclimation temperatures decreased asacclimation temperatures increased. Strong evidence of similarities

rather than differences between populations and among species within afamily was demonstrated. Moss and Scott (1961) studied the differences insurvival between fish exposed to rapid drops in dissolved oxygen to acritically low point from near saturation (shock test) and fish exposed todissolved oxygen declines gradually over a period of time (acclimation

26

LOU

%MO

0.

E4)

4)

V

*0

0>

<

Acclimation Temp.(C)

Fig. 8. Plot of the published data on the relationship betweenavoidance temperature and acclimation temperatures of fishes withina family. The 90% confidence intervals of the individual predictedvalues are shown. 'Cherry et al. (1975, 1977); -, Muddy Run-Cono-wingo (Mathur et al. 1983).

27

# I

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 tobe highly adaptive. In addition, data from the starved and obese groupsof 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 fromlower to higher temperatures, were nearly reacclimated in 1-3 days, mea-sured in terms of change in heat resistance. However, fish changed fromhigh to low temperatures required 4-14 days to approach acclimation. In

both changes, complete reacclimation required at least 12 days, indicatingthat tempering for a few hours would increase heat tolerance to sameextent, 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 apartial loss of heat tolerance. Fish exposed to fluctuating temperatureshould 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 metabolicrate at each end point in temperature was similar to fish acclimated tothat temperature. A major reason for stress responses to altered tempera-ture found in earlier studies may be that the fish had been exposed onlyto constant temperatures in the lab. Diana's (1983) results make sense inthat largemouth bass are considered temperature eurytherms (Magnuson etal. 1979) and regularly undergo seasonal temperature extremes from 1 to 32C. Their response to temperature fluctuations is more variable thanstenotherms. Venables et al. (1977) determined that bass were meta-bolically 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 Scott1961).

Temperature(degrees centigrade)

Type of test and species (degrees centigrade)25 30 35

Shock testsBluegill 0.75 1.00 1.23Largemouth bass 0.92 1.19 1.40Channel catfish 0.95 1.03 1.08

Acclimation testsBluegill 0.70 0.80 0.90Largemouth bass 0.83 0.83 1.23Channel catfish 0.76 0.89 0.92

29

reduce the growth of largemouth bass juveniles at 26 C (Stewart et al.

1967). Similarly, Whitworth (1968) observed weight loss in brook trout

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 inten-

sities (negative phototaxis) in walleye when dissolved oxygen fell to 2-4

mg/liter. It has also been shown that fish generally become more activein 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

response often varies considerably. It is not clear whether avoidance

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 periodi-cally may benefit from it. Implicit in a behavioral response to lowoxygen would be sane means of detecting low oxygen concentrations. Thephysiological responses to hypoxia are rapid, consisting of circulatory

and ventilatory changes in trout and tench within a few seconds of the

onset of hypoxia (Eclancher 1972, Randall and Smith 1967). Such rapid

responses suggest the presence of an oxygen receptor system on or near the

gills.

Whitmore et al. (1960) investigated the avoidance reactions of salmonidand centrarchid fishes to low oxygen concentrations. Largemouth bass andbluegill (Lepomis macrochirus) markedly avoided concentrations near 1.5

mg/liter, but they showed little or no avoidance of the higher concen-trations. Only bass showed any avoidance of concentrations near 4.5 mg/

liter. Avoidance by fish of water with low dissolved oxygen concentra-tions 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

reduced oxygen concentrations. Shelford and Allee (1913) found that

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

bass (Fig. 9). The swimming speed of largemouth bass at low carbon

dioxide concentrations decreased with dissolved oxygen concentrations

below 5-6 mg/liter (Fig. 10). At concentrations above 6.0 mg/liter, the

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

Uw

0ccw

.Jz

I

w

z

nON)RE)TION)

DISSOLVED OXYGEN CONCENTRATION INMG PER LITER

Fig. 9. Mean final swimming speeds, expressed inL/sec, of largemouth bass at various concentrations ofdissolved oxygen and carbon dioxide. The curve relatingswimming speed -to oxygen concentration is the curvethat was fitted to all data from tests at the low concen-tration of free carbon dioxide (probably <3 mg/liter)only, most of which (those from early tests) have not beenplotted here. The stated free carbon dioxide levels of 48,21, and 38 mg/liter are means of observed values forindividual tests ranging from 44 to 55 mg/liter, 17 to23 mg/liter, and 24 to 38 mg/liter, respectively (Dahlburget al. 1968).

U 'Mw6

9)w 5IL-JZ 4

Qwhiw 3

z 2

ii0A

DISSOLVED OXYGEN CONCENTRATION INMG PER LITER

Fig. 10. Relationship between the mean final swimmingspeed of largemouth bass, expressed in L/sec, and thedissolved oxygen concentration at nearly natural, low con-centrations of carbon dioxide (<3 mg/liter) and temperaturesnear 25 C (Dahlburg et al. 1968).

32

Ln

concentrations tested at 25 C. Embryo pigmentation was progressively

lighter as the oxygen concentration decreased. At reduced oxygen concen-

trations, 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 concentra-

tions 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 weresensitive to all reductions in dissolved oxygen, although survival wasonly slightly affected at 60 and 70% saturation. Siefert and Spoor

(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 thatcaused 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), al-though some embryos developed and hatched at oxygen concentrations as lowas 1.0, 1.1, and 1.3 mg/liter. The increasing embryo biomass requiresmore 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 1.92microliters 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 duringhatching. The increased oxygen consumption coincident with hatchingof largemouth bass embryos undoubtedly increases oxygen needs above thatwhich 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 require-ments of chum salmon (Qncorhynchus kA) eggs as development progressed.Eggs at early stages of incubation required about 1 mg/liter oxygen, whilethose about to hatch required over 7 rag/liter. Eggs held at low oxygentensions experienced reduced growth and retarded development. Deformities

33

EXPNM EXPE

--. 5 c a

--- * 20 C 0

--- 25C A

Survival at 90%Oxygen Saturation

m.--

/ /

/ /0

A // /

nA° / AA/An A

Oxygen Concentration (mg/I)

Fig. 11. Relationships between embryo survival and oxygen concentrationat three incubation temperatures. Each point represents the percent of 40embryos (Experiment II) or 150 embryos (Experiment III) which survived tocompletion 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 both15 C and 25 C (Dudley and Eipper 1975).

34

6

(I)

0

cs_-a SO-u

f

in embryos exposed to hypoxia were observed, with mortality occurring at

the stage when the circulatory system develops. Others have reported

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 forwater quality standards in warmwater systems would be extremely difficult.In his classic work, Ellis (1937) concluded that a minimum summerdissolved oxygen concentration of 5 mg/liter was necessary to supportgood, mixed fish faunas. Coble (1982) looked at fish populations in a

Wisconsin river in relation to dissolved oxygen and found sport fishes anda 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 fisheswas 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 severelyaffected by low oxygen levels, especially on a chronic exposure basis.

35

A

4

a

0

I

a

100

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60

40

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I Ow---* .. O -AA-I*0 . O SA.

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SLASONAL TEMPERAITU€

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aN

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I?

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StASONAL TIMPEOATULI MAXIMUM - C

Fig. 12. A, Oxygen criteria, expressed as seasonalminima in percent saturation, at various seasonal temperaturemaxima. Three levels of protection, A, B, and C, are givenas defined in the text. Values used in derivation of thecriteria are summarized in Table 10; B, Oxygen criteria,expressed as oxygen minima in mg 02/liter (ppm), at variousseasonal temperature maxima. Three levels of protection, A,B, and C, are given as defined in the text. These valueswere calculated from the percentage saturation values illus-trated in (A) and summf4xrzed in Table 10 (Davis 1975),

36

K a - 0 - - A A,-•Ft a 1% aA 4 f

W ---mmm *

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