Power Plants || INFLUENCE OF TEMPERATURE ON FISH BEHAVIOR

CHAPTER V

INFLUENCE OF TEMPERATURE ON FISH BEHAVIOR1

Jay R. Stauffer^ Jr.

INTRODUCTION

Temperature is among the most important environmental parameters which affect aquatic biota. Fry (1947a) proposed six categories through which an environmental entity can affect an organism: (1) lethal factors which "destroy the integration of an organism"; (2) masking factors which "prevent a second identity from operating on the organism to the extent that it would if the masking factor were not present"; (3) directive factors which "require a response on the part of the organism directed in some relation to a gradient of the factor"; (4) controlling factors which "govern the metabolic rate by operating in the internal medium which is the actual site of metabolism"; (5) limiting factors which "govern the metabolic rate by virtue of their operation within the metabolic chain"; and (6) accessory factors which "impose a metabolic load upon the organism in excess of the rate to which the organism is confined by the factor which is governing the overall metabolic rate". Temperature can function in any of these roles.

The purpose of this chapter is to provide an overview on the effects of temperature on fish behavior. A brief presentation of historical information discusses the lethal effects of temperature, temperature detection and regulation in fishes, and is followed by a comprehensive treatment of temperature selection by fishes and the indirect effects of temperature on fish behavior. Synergistic effects with other parameters associated with power plants are discussed in subsequent chapters.

Reviewers for this chapter: James J. Gift and William W. Reynolds

Copyright © 1980 by Academic Press POWER PLANTS 1 0 3 All rights of reproduction in any form reserved.

ISBN 0-12-350950-5

104 J. R. STAUFFER, JR.

BACKGROUND INFORMATION

Lethal Effects

Historically, the lethal effects of temperature on aquatic organisms were studied first (Vernon 1899; Huntsman and Sparks 1924; and Battle 1926). The direct effect of temperature changes on fish mortality has been extensively studied for the past 30 years, and includes work by Alabaster and Downing (1966), Allanson and Noble (1964), Alpaugh (1972), Biahm and McConnell (1970), Brett (1952, 1956), Charlon et al. (1970), Cocking (1959a, 1959b), Coutant (1970a, 1970b, 1970c), Doudoroff (1942), Edsall and Colby (1970), Fry (1947b, 1960, 1964), Fry et al. (1942, 1946), Gibson (1954), Hoff and West-man (1966), Norris (1963), Olson and Foster (1955), Reynolds et al. (1976a), and Tyler (1966).

Death due to temperature intolerance may result from either exposure to cold or high temperatures. For the most part, exposure to cold temperatures is lethal to temperate fishes only when these organisms are acclimated to elevated temperatures and then abruptly exposed to lower temperatures. This phenomena has been termed cold shock, and is usually associated with shutdowns of thermal outfalls during winter months. Cases of mass mortality are documented in the literature (Anon. 1971, 1972; Ash et al. 1974; Clark and Brownell 1973). Becker et al. (1977a) conducted cold resistance (gradual and abrupt cold shock tests) experiments on organisms from the Columbia River. Lower incipient lethal temperatures for several freshwater fishes are reported in Hart (1947). Becker et al. (1977b) proposed a model which would predict the effects of cold shock and also reviewed existing literature. Other physiological effects of low temperature on fishes were reviewed by Umminger (1969) and Prosser et al. (1970).

Death from temperature intolerance results in the ultimate breakdown in the organization of the individual (Brett 1956), although the actual cause of death is debated. Mayer (1914) postulated that at high temperatures there was insufficient oxygen to sustain increased metabolic activity and, therefore, death resulted from asphyxiation. This hypothesis was supported by Weatherly (1970), who showed that the mean lethal temperature of certain species could be increased by supersaturation of the water medium to the equivalent of five atmospheres. Other phenomena which have been postulated as the cause of death with elevated temperatures include: syn-aptic failure occurring in the rnyoneural junctions and smooth muscle peristalis (Fisher 1958); denaturation of body cells and inactivation of cholinesterase (Kusakina 1963); changes in enzyme systems due to very high concentrations of lactic acid

V INFLUENCE OF TEMPERATURE ON FISH BEHAVIOR 105

(Pegel and Remorov 1961); partial blood coagulation in branchial capillaries followed by rupture of vessels in respiratory system (Agersborg 1930); and failure of coordinating mechanisms of central nervous system (Orr 1955). Heinicke and Houston (1971) demonstrated osmoregulatory breakdown during heat shock, but indicated that this was not the sole cause of death. Other workers have attempted to correlate cellular studies with upper lethal temperatures. Timet (1963) demonstrated that high temperatures caused injuries to living protoplasm and decreased oxygen consumption in marine fishes; while, Wolf and Quimby (1962) showed that the upper ultimate lethal temperature (26 C) of cultured cells of rainbow trout agreed with estimates of the lethal temperature of the intact organism. Moreover, Battle (1926) showed close agreement between the lethal temperature of tissues and lethal levels of complete organisms. The failure of a number of biological systems at temperatures above 30 C was discussed by Drost-Hansen (1965), who indicated that although an organism may be alive at 33 - 34 C, vital bodily mechanisms may be irreversibly damaged. Dean and Coutant (1968) showed that 85% of juvenile chinook salmon which lost equilibrium because of thermal shock died when returned to non-lethal temperatures. This concept of a "point of no return" is taken one step farther with the development of the term critical thermal maximum. Gift (1970) based on Mihursky and Kennedy (1967) defined critical thermal maximum as the "thermal point at which loco-motor activity becomes disorganized and the animal loses its ability to escape from conditions that will soon cause its death". Such a phenomenon would result in the immobilization of fish in heated areas if the fish did not avoid these areas. Gift (1970) further stated that the organism is "ecologically dead" at its critical thermal maximum. Therefore, a thermal dose which induces equilibrium loss is probably just as important as one which causes death (Becker 1973).

The fact that heat death is rarely observed in nature appears to contradict the fact that many organisms are killed at temperatures only slightly above those at which they normally live or prefer (Gunter 1957 ;Kerr 1953; Naylor 1965; Reynolds and Thompson 1974). It would seem that the ability to detect lethal waters and the fishes' subsequent behavior is undoubtfully responsible for the relative scarcity of fish kills (Gammon 1971). On the other hand, Jensen (1970) reported on a fish kill in the vicinity of a power plant and stated that there was no evidence that heat per se was the direct cause of death; however, he postulated that the fish would not have been in the vicinity of the plant were they not attracted by the heated discharge.

Therefore, the remainder of this chapter will focus on temperature detection, thermal regulation and the behavioral


responses of fish to temperature. For a comprehensive review of the other effects of temperature on aquatic organisms, the following symposia, reviews, and bibliographies should be consulted: Beltz et al. (1974), Becker (1973), Bullock (1955), Coutant (1968, 1969, 1970d, 1971, 1973), Coutant and Goodyear (1972), Coutant and Pfuderer (1973), Coutant and Talmage (1976), Fry (1947a, 1957), Mihursky and Kennedy (1967), Kinne (1963) Krenkel and Parker (1969), Parker and Krenkel (1969), Raney and Menzel (1967, 1969), and Raney et al. (1973).

Temperature Detection and Thermal Regulation

Fish are predominately ectothermic, with the exception of lamnid sharks and scombrid (tunas, etc.) species which have achieved a limited degree of endothermy (Carey and Teal 1969). The development of endothermy requires temperature control at the site of respiration, or of the circulatory fluid. Insulation of the gills would defeat the primary purpose of gas exchange; however, Reynolds (1977a) and Reynolds et al. (1976) observed that gill perfusion may be restricted following temperature change as a method to reduce heat exchange. In tuna, there is a countercurrent heat exchange in the form of a rete mirabile interposed in the vasculature between the gills and the muscles (Hammel et al. 1972), thus minimizing heat loss through the gills.

Other fishes have a body temperature which does not differ greatly from the static environmental temperature, except for minor instances. Brett (1956) noted that heat production from excessive activity which may prevent normal respiration could result in increased body temperature. Brett (1956) further notes that rapid passage of fish through stratification layers may also result in the body temperature being different than environmental temperature as shown by Morrow and Mauro (1950). Therefore, since tissues of most fish are constantly brought into equilibrium with the aquatic environment, they, like other ectotherms, are obliged to rely almost exclusively on behavioral responses to regulate body temperature. In general, temperature equilibrium is established more quickly in smaller fishes than large forms (Simpson 1908; Pearse and Hall 1928; Neilsen 1938; Gunn 1942; Davis 1955; Spigarelli et al. 1974). However, Stauffer et al. (1975a) found no positive correlation between length or weight and the time for body temperature to reach equilibrium with environmental temperature. Fish have been shown to prefer a certain range of temperatures when placed in a thermal gradient (e. g., Meldrim and Gift 1971; Stauffer et al. 1975b, 1975c, 1976a; McCauley and Pond 1971; McCauley and Read 1973; Mc-Cauley and Tait 1970; and Cherry et al. 1975, 1977, among


others) and are capable of learning a behavior response which will alter their thermal environment (Rozin and Mayer 1961). The fact that fish can select certain temperatures and thereby regulate body temperatures implies that the organism can sense body temperature and compare it with some optimal or preferred condition (Hammel et al. 1972).

In theory, there are two ways in which an organism can detect temperature variations: (1) temperature changes can excite thermal receptors which send nerve impulses to the central nervous system or (2) it can act on a chemical process in some or all tissues (Sullivan 1954). Sullivan (1954) states "according to Kappers et al. (1936), no complicated sensory terminations are known in skins of vertebrates below reptiles and amphibians". More specifically, cutaneous thermal receptors have not been localized in fish (Prosser et al. 1950). Despite the lack of morphological evidence, the functional existence of skin thermoreceptors has been shown via experimental techniques (Sullivan 1954; Wells 1914). It has been demonstrated that several species can detect temperature changes ranging from .05 to 0.2 C (Bardach and Bjorklund 1956; Breeder 1951; Collins 1952; Shelford and Powers 1915). Bull (1936) reported that some teleosts can detect water temperature changes as small as 0.03 C. Moreover, Breder's (1951) field observations on schools of dwarf herring demonstrated an extremely refined sense of temperature perception. Studies by deSylva (1969) documented that larvae have functional thermal receptors immediately after hatching which can detect slight temperature variations. Based on evidence similar to the above, Sullivan (1954) concluded that both bony fishes and selachians (sharks, rays) have surface thermal receptors.

The neuronal system which controls temperature selection in most bony fishes is similar to that in other vertebrates in that both peripheral and rostral brainstem temperatures provide valuable input (Crawshaw and Hammel 1972; Hammel et al. 1972). It was shown in a series of experiments that warming and cooling the brainstem of selachians (sharks) produced major effects (Crawshaw and Hammel 1973), and the effect of heating the brainstem of brown bullheads increased preference for cooler waters (Crawshaw and Hammel 1972). Sullivan (1954) demonstrated the lesions of the forebrain changed the behavior of brook trout when placed in a thermal gradient. Hammel et al. (1972) hypothesized that neural output from the anterior brainstem coordinates autonomic responses which eventually result in thermoregulatory behavior. They further postulated that these thermoregulatory processes evolved as secondary functions to already existing organ systems. At any rate, ". . . most poikilotherms appear to seek temperatures which match the most efficient operation of their metabolism" (Coutant 1974). In summary, temperature preference experiments do not


attempt to explain physiological mechanisms although they do demonstrate the ability of fish to respond to temperature gradients and temperature (Bardach and Bjorklund 1957). For an excellent review on the physiological reactions of fish to temperature, see Crawshaw (1977).

TEMPERATURE SELECTION

Studies which attempt to determine the temperature preference or avoidance of fish species can be categorized into laboratory or field studies. Laboratory studies in general are more numerous than field studies. Detailed j[n situ investigations have been conducted by Marcy (1971, 1976), Nakatani (1969), Gammon (1971), Neill and Magnuson (1974) and Stauffer et al. (1975b, 1976a, b). However, these have been limited to either freshwater or anadromous species. Field studies with marine fish, which are usually considered to be more exacting in their environmental requirements (Kinne 1963; Hedgpleth 1957), are rare.

Laboratory Studies

As indicated by Stauffer et al. (1975b), many authors have recognized the importance of temperature preference studies and have conducted studies under laboratory conditions (e.g., Doudoroff 1938; Fisher and Elson 1950; Brett 1952; Garside and Tait 1958; McCauley 1958; Moss 1970; McCauley and Pond 1971; Meldrim and Gift 1971; Neill and Magnusson 1974; Stauffer et al. 1976a; and Richards et al. 1977, among others). Laboratory studies control multiple factors which may influence fish behavior such as light, gradient, gas and chemical factors, and biological interactions (Brett 1956). Disadvantages of laboratory investigations include: (1) limitation of time and space, (2) compressed thermal gradients, and (3) numbers and size of individuals which can be tested (Ferguson 1958).

Techniques used to determine temperature selection of fishes range from electronic shuttle boxes in which fish control the water temperature (Neill et al. 1972; Neill and Magnuson 1974; Reynolds and Casterlin 1976; Reynolds 1977a; Reynolds et al. 1976) to vertical gradients (McCauley and Tait 1970; McCauley and Pond 1971; Otto et al. 1976; Muller and Fry 1976) and horizontal gradients (Mendelsohn 1895; Wells 1914; Sayle 1916; Doudoroff 1938; Kruger 1952; Sullivan and Fisher 1953; Zahn 1960; Meldrim and Gift 1971; Stauffer et al. 1976a; Cherry et al. 1977) in which the fish select a prefer-


red temperature. McCauley (1977), in an excellent review of the different laboratory methods used to determine temperature selection, lists the following generalized groups of techniques: two-chambered devices, radial devices, horizontal tanks, transverse gradients, vertical gradients, electronic shuttleboxes, toroidal gradients, body temperature telemetry and calorimetry. Horizontal troughs have probably been more widely used and modified than the other general types. Probably the simplest units consist of a trough which is progressively heated by an immersion heater (Barans and Tubb 1973) or by a series of heat lamps underneath a long metal trough (Meldrim and Gift 1971; Stauffer et al. 1976a; Cherry et al. 1977). The introduction of cold water at one end of these troughs intensifies the temperature gradient. Unless water level is precisely controlled and kept at a minimum, vertical layering of the water may occur. To alleviate this problem, some investigators have used one or more air curtains (Zahn 1960; McCracken and Starkman 1968; Richards and Ibara 1978). Recently, horizontal troughs have been designed in which hot saturated water is introduced at one end and cooled as it flows down the trough; thus eliminating the potential of a supersaturated condition at the warmer end of the gradient (Ecological Analysts 1978). Other modifications of the horizontal design include separating the trough by a series of baffles (Reutter and Herdendorf 1974). Reynolds and Thomson (1974) used a "selectatron" which consisted of 16 chambers interconnected at the top, while Crawshaw and Hammel (1973) used two chambers connected by a plexiglass door which opened whenever the fish approached. Alabaster (1964) and Alabaster and Robertson (1961) experimented with a 152 m long trough.

Historically, a controversy developed over the best type of unit for determining the temperature a particular species preferred. Javaid and Anderson (1967a) noted a wide discrepancy between their results for rainbow trout using a horizontal gradient and the results of Garside and Tait (1958) who used a vertical gradient. Other investigators have observed similar discrepancies depending upon the particular spacial limitations of certain species and test organisms (Gift, pers. commun.). However, there is evidence which indicates that the geometry of the gradient may not be as important as previously thought (McCracken and Starkman 1968; Fry 1971; McCauley and Pond 1971). For those troughs in which fish control their environmental temperature, Reynolds et al. (1976b) indicates that the results obtained are reproducible and species specific. It would, therefore, appear that the temperature preference of a species is a real and measurable phenomenon. Probably the most important determinant in selecting the type of trough depends upon the species to be tested. Crawshaw and


Hammel (1973) state that horn shark were unable to properly orient in a 9 - 31 C thermal gradient which was easily mastered by brown bullhead catfish. Stauffer et al. (1976a) indicated that the fantail darter performed erratically in a horizontal trough.

In addition to variations in the physical unit, investigators have varied techniques in determining the selected temperature. Because of the type of unit, some investigators have used extensive training periods before conducting their tests (Crawshaw and Hammel 1974). Depending on the techniques and species, authors have allowed their test organisms to habituate to the test chamber for 10 min (Crawshaw 1975a), 40 to 60 min (Stauffer et al. 1976a), 16 h (Otto et al. 1976), or 48 h (McCauley and Pond 1971). In an attempt to minimize the stimulation by the investigator, different authors have utilized closed circuit television and mirrors to view test specimens (Cherry et al. 1977; Meldrim and Gift 1971), while others have blinded their test organisms (Crawshaw and Hammel 1973). Moreover, the temperature selected has been determined by monitoring locations within the experimental unit (McCauley and Pond 1971; Stauffer et al. 1976a), or by measuring body or muscle temperature (Crawshaw 1975a, 1975b).

Another source of variation among different investigators' techniques is the process of temperature acclimation. It is known that thermal acclimation will affect the temperature for optimum activity and the ability of fish to withstand either low or high lethal temperatures (Fisher 1958; Tatyankin 1966). However, the process of acclimation as a physiological phenomenon is not well understood (Kinne 1963). Acclimation temperature may involve alterations in enzymes, cells, organs, metabolic activity, and, as such, the capacity to acclimate depends on genetic background, environmental history and present physiological age and condition (Kinne 1963). Furthermore, it is not clear whether acclimatization of fish under natural seasonal conditions is different from acclimation under laboratory conditions. For example, Meldrim and Gift (1971) and Sullivan and Fisher (1953) found differences in selected temperatures depending upon whether fish were acclimated during rising or falling field temperatures. On the other hand, Cherry et al. (1977) could not distinguish differences for six of nine species when responses during rising field temperatures were compared with those during falling. They attributed this phenomenon to a longer acclimation period in the laboratory; thus fish were acclimated to a more stable condition as opposed to acclimitized to field conditions. Rate of acclimation from collection temperature to test temperature is also debated. It is generally accepted that fish can acclimate to higher temperatures faster than they can to


lower temperatures (Doudoroff 1942, Norris 1963) and that an acclimation rate of 1 C/day is "safe". It has been suggested that large body size may increase tolerance to sudden change in water temperature (Lattimore and Gibbons 1976). Javaid (1972), using three salmonids, showed that the effects of upward acclimation were complete within one week or less. Experiments by Norris (1963) and Crawshaw and Hammel (1974) indicated that the maintenance of fish at a constant acclimation temperature may result in a decrease in accuracy of temperature selection. As a result, some investigators use flow-through holding tanks and keep their fish constantly exposed to natural fluctuations of water temperature (Cherry et al. 1975, 1977).

It is generally agreed that there is an effect of acclimation temperature on acute (short term 1 24 h) preferenda. This effect has been expressed by a variety of relationships. Most commonly, temperature preference is plotted against acclimation temperature. Fry (1947a) used this relationship to estimate the final temperature preference for a particular species. If both axes are represented by the same units, then the point at which the temperature preference-acclimation temperature curve intersects a line which passes through the origin and has a slope of 1.0 is deemed the final temperature preference (Fry 1947a). Theoretically, this should represent the temperature which a species will ultimately select given an expanded temperature gradient and an indefinite amount of time. There is presently some debate as to whether or not this relationship should be predicted with various mathematical models or estimated by eye. In a recent symposium Coutant (Richards et al. 1977) suggested that a visual construction line would be preferable to using the simple model P = a A + B, where P = preference temperature and A = acclimation temperature, since the slope of preference/acclimation curve decreases as the acclimation temperature approaches the final preferred temperature (i.e., the point where P = A). However, other models which more accurately depict the relationship between preference and acclimation temperature are needed when statistically comparing the responses of different species or several races of the same species (Hall et al. 1978). Analysis of acute temperature preference data seems to indicate that the quadratic equation, Ρ = α A + a A + β, most often provides the best fit when compared with higher order polynomials and log transformations (Stauffer ms).

Data from temperature preference studies have also been used to characterize the range of temperature which a species will select. Reutter and Herdendorf (1974) used the standard deviation about the final preferendum as an estimate of how dependent a species was on temperature. It may be, however,


that this range is more dependent on inherent variations among individuals and the response of individuals to the test apparatus rather than an indicator of temperature dependence. Zahn (1962) postulated that if two species share the same final temperature preference, the one which generated a curve with a greater angle between itself and the theoretical line with a slope of 1.0 is considered to be more stenothermal. McCauley and Tait (1970) also indicated that the shape of the preferred temperature-acclimation temperature curve was useful in the physiological description of a species.

Despite the general relationship between temperature selection and acclimation temperature, several salmonids demonstrated a temperature preference relationship which appeared to be independent of acclimation temperature: salmon species ( Oncorhychus) (Brett 1952), rainbow trout (Brett 1952; McCauley et al. 1977), brook trout (McCauley and Tait 1970), and certain salmonid hybrids (Goddard and Tait 1976).

Recently the occurrence of febrile responses has been reported in aquatic ectothermic vertebrates and invertebrates (Casterlin and Reynolds 1977a, b, 1978; Reynolds 1977b; Reynolds et al. 1978a,b; Reynolds and Covert 1977; Covert and Reynolds 1977). Since ectotherms are for the most part dependent upon behavior to control body temperatures, these fever responses are manifested by an increase in the preferred temperature. Survival value of fever in fish is discussed by Covert and Reynolds (1977).

In addition to the concept of temperature preference, several studies have investigated temperature avoidance behavior (Reynolds 1977c). Gift (1970) defined upper avoidance temperature as that "... which creates sufficient stress to cause temperature to be a directive factor". Brett (1956) suggests that the inherent ability to perceive fine temperature gradients is caused by internal drives or environmental stress, which would distinguish between temperature as a directive factor and a preference response. Probably the avoidance unit most commonly used today is one patterned after Meldrim and Gift's (1971), modified from designs of Shelford and Allee (1913), Jones (1952), Sprague (1964) and Hill (1968). This unit consists of two subtroughs in which water enters at opposite ends and drains at the center. This arrangement allows the investigator to provide the test organism with a choice of two temperatures. To correct for positional effects, water from a temperature control unit which supplies one end of the first subtrough, supplies the opposite end of the second subtrough. In practice, a fish or group of fish are placed in each subtrough, and water flows from either end of each subtrough at the acclimation temperature. A temperature differential is created between halves of each subtrough


after the orientation period. The water temperature of each half of each subtrough is raised stepwise, as the experiment progresses until the organisms avoid the warmer water. Results from this type of test generally indicates that the upper avoidance temperature is higher than the preferred temperature but lower than the upper lethal temperature.

Field Studies

Several methods have been used to estimate fishes' temperature preference or avoidance responses from field data. Neill and Magnuson (1974) and Stauffer et al. (1974, 1975b, c, 1976) estimated temperature preference responses from field data. Stauffer et al. (1976a) used a block net and rotenone to sample areas with different temperatures. It has been hypothesized that a bias might be introduced by a possible effect of temperature on fish susceptibility to rotenone (Reynolds 1977c). However, repeated sampling with seines and additional rotenone indicated that, if done correctly, fishes can be effectively sampled at all temperatures above 15 C. However, due to the qualitative nature of most types of gear, it is difficult to predict temperature selection from field data. Probably one of the most exact methods of determining field temperature preference is through the use of acoustic census, echo location techniques, and telemetry (Kelso 1974, Kelso and Minns 1975; Coutant 1974). Because availability of this type of data is limited, most field temperature selection data are based on observations, rather than on specifically controlled experiments. The fact that both freshwater and marine fish are attracted to warm water discharges throughout the colder months of the year has been well documented (Allen et al. 1970; Barkley and Perrin 1972; Dryer and Benson 1954; Elser 1965; Fairbanks et al. 1971, Galloway and Strawn 1971; Gibbons et al. 1972; Grimes and Mountain 1971; Landry and Strawn 1972; Marcy 1976; Marcy and Galvin 1973; Moore and Frisbie 1972; Hatch 1973). Kinne (1963) stated that cases of the opposite type of migration (i.e., attraction to cold areas) are known from glacial relicts in arctic areas.

Furthermore, it may be that periods of torpor can accrue physiological and ecological advantages to animals (Brett 1971). Therefore, voluntary orientation to cold locations could reserve energy sources during periods of low food availability and aid in predator avoidance (Crawshaw 1975b). Stauffer et al. (1976b) reported on fishes avoiding the area of a thermal discharge in the winter months, but attributed this to elevated levels of free chlorine concentrations rather than a temperature directed response. Moore and Frisbie (1972) and Marcy and Galvin (1973) reported increased sport


fishing in heated effluents during the winter months. Mihursky (1969) documented summer movement away and winter attraction of fishes to a heated effluent at Chalk Point, MD.

Several studies have shown specific responses for certain species. Kelso (1976) reports that a thermal discharge caused yellow perch and white suckers to orient into a current produced by the Nanticoke Generating Station. Howe and Coates (1975) performed a series of studies which indicated that movement of winter flounder was related to temperature. Jammes (1931) showed that brown trout preferred 12 C in a lake and dwarf herring avoided 30 C in the field (Breder 1951). However, some individuals of sea catfish apparently did not avoid lethal temperatures under field conditions, although the general reaction of the species was one of avoidance of lethal temperatures (Galloway and Strawn 1971).

INDIRECT EFFECTS OF TEMPERATURE ON FISH BEHAVIOR

In addition to the direct effect of temperature on fish behavior in the form of a directive factor, temperature may affect behavior by influencing activity (Sullivan 1954; Hela and Laevastu 1962; Peterson and Anderson 1969). Activity is usually measured by testing swimming speed or metabolic rate (see Chapter VI). Fry (1947a) reported that the scope of activity increased with increasing temperature over the complete biokinetic range of brown bullhead catfish, while the metabolic scope for activity peaked at 19 C for brown trout (Fry 1947b; Graham 1949). The peaking of activity below the lethal temperature was also reported for yellow bullhead catfish (McLarney et al. 1974). Brett (1956) hypothesized that survival value may be linked with optimum activity level and that an increase in lethal temperature without increase in scope of activity would have little survival value. For example, both Sylvester (1972) and Yocum and Edsall (1974) reported an increased predation vulnerability of fish subjected to thermal stress.

Other changes in behavior at sublethal temperatures include the increase of aggression. McLarney et al. (1974) reported that the ratio of damaging aggressive units per warning aggressive units increased significantly when yellow bullhead were subjected to temperatures greater than 30 C. They suggested that one possible interpretation is that the change in behavior was due to the loss of the ability of the fish to distinguish individuals of its own species which may ultimately reduce the carrying capacity of a body of water. Moreover, Nyman (1972) reported an increase in intraspecific aggression


at high temperatures for the European eel. Nyman (1972), also stated that the behavior of European eel was affected in the following ways: (1) changed substrate selection at different temperatures, (2) crowding occurred at temperatures below 8 C, and (3) foraginq was initiated by a rise in temperature.

Marcy (1976) showed that ovaries of white catfish and brown bullheads inhabiting a heated discharge developed earlier than those of fishes living under ambient conditions. Other instances of accelerated gonadal development cited by Marcy (1976) included Adair and DeMont (1971), Langford (1972), Trembley (1960) and Witt (1971).

Migrations also appear to be controlled in part by temperature. Brett (1956) thought it improbable that temperature could trigger precise migrations based on laboratory studies which showed a relatively high range of preference temperatures. However, other authors have showed that mass migrations have been correlated with temperature changes (Chittenden 1969, 1972; Foerster 1937; Kennedy 1940; Leggett and Whitney 1972). Thermal discharges have been shown to affect movement, especially in the turbulent mixing zones (Kelso 1974).

Temperature may affect feeding behavior of fishes. Latti-more and Gibbons (1976) showed that stomach contents of fish were more herbivorous under ambient conditions than fish located in a thermal effluent, while Massengill (1973) reported a shift from an invertebrate to a piscivorous diet for brown bullheads inhabiting a heated affluent. Peters et al. (1972) showed that the meal size of pinfish, spot and Atlantic silverside decreased as temperatures deviated from optimum. Bennett and Gibbons (1972) also noted that fish contained less food in their stomachs at high temperatures, but attributed this to the possibility of a higher digestion rate at increased temperatures. In conjunction with these observations, several authors noted that fish living at elevated temperatures are in poorer condition than those living at ambient temperatures (Massengill 1973; Stauffer et al. 1974; Marcy 1976). Shcherbukha (1972) reported conflicting results in that three species overwintering in the USSR improved in condition while two species declined.

INTERPRETIVE ANALYSIS

Despite differences in investigators, techniques, and data interpretation, several generalizations can be drawn from both laboratory and field studies involving temperature selection. Temperature selection appears to be species specific. Evidence of genetic adaptation is inferred not only from


thermal behavior studies, but also from the fact that fish live in a variety of latitudes, climatic zones and depths (Kinne 1963). As stated previously, acclimation temperature probably has the greatest influence on an acute temperature selection, with the exception of a few species, thus, demonstrating a nongenetic adaptation to temperature. Additionally, the thermal history (exposure to varying temperatures in developmental stages and prior to present acclimation regime) of the organism appears to play an important role, but the relationships are not entirely clear (Fry and Hochachka 1970: In Crawshaw 1975a). However, cold exposure did not affect the thermoregulatory behavior of bluegill (Beitinger 1974). Season (Sullivan and Fisher 1954; Hoar 1955; Tyler 1966; Zahn 1963; Barans and Tubb 1973), photoperiod (Hoar and Robinson 1959), diet (Fisher 1958), hormone levels (Hoar 1959; Baslow 1967; Parvatheswaro 1969; Pequingnot et al. 1969), light intensity (Meldrim and Gift 1971; Reynolds and Thomson 1974), salinity (Meldrim and Gift 1971), age (Reutter and Herdendorf 1974), exposure to toxic substances (Ogilive and Anderson 1965; Opuszynski 1971; Peterson 1973, 1976), pathogens (Covert and Reynolds 1977), and size (Meldrim and Gift 1971; Gift 1970) have all been suggested as variables which may influence temperature preference and avoidance of a species, although some of the results are ambiguous.

Javaid and Anderson (1967b) reported that rainbow trout and brook trout selected cooler water, while Atlantic salmon preferred warmer water during periods of starvation. Brett (1971) suggested that the selection of a lower temperature, and, hence, lower metabolism, was a response to limited food supplies. Lillywhite et al. (1973) showed that when food was readily available newly metamorphosed toads (Bufo boreas) preferred higher temperatures than they selected under starvation conditions.

The relationship between temperature avoidance responses and light levels for white perch varied, depending upon whether tests were conducted under rising or falling field temperatures; Atlantic silversides showed an inverse temperature avoidance response with varied light intensities (Meldrim and Gift 1971). Temperature preference responses override light intensity preferences in grunion (Reynolds and Thompson 1974).

Of particular interest is the fact that size was a significant variable in temperature avoidance studies with white perch but not in temperature preference studies (Meldrim and Gift 1971). Size also affected the avoidance responses of winter flounder (Gift 1970). In general, smaller fish and juveniles prefer a warmer temperature than adults of the same species (McCracken and Starkman 1968; Barans and Tubb 1973;


McCauley and Read 1973; Coutant 1974; Otto et al. 1976). Reutter and Herdendorf (1974) noted that the final preferendum for adults of some species was higher than for young-of-the-year fish.

There may be a critical age in development when larval fishes are more susceptible to high temperatures. Altman and Dittmer (1966) showed that larval stages had experimentally lower upper incipient lethal levels than adults while Huntsman and Sparks (1924) showed that the young of several freshwater fish species were more tolerant to high temperatures than adults. Regardless of experimentally derived values, juveniles may be more susceptible to lethal temperature in the field because of an inability to swim to cooler waters (Alabaster 1964; Alabaster and Downing 1966). As stated previously, larval forms have functional thermal receptors immediately after hatching (deSylva 1969), however, the phenomena of low thermal responsiveness appears to be correlated with size (Meldrim and Gift 1971) or age (Reynolds and Thompson 1974). Smaller fish may be more susceptable to this phenomenon since internal body temperatures may reach equilibrium more rapidly than internal temperatures of larger fish (Beitinger and Magnuson 1976). Meldrim and Gift (1971) defined low thermal responsiveness as the inability of a fish to avoid areas in a thermal gradient which produces stressful conditions. Low thermal responsiveness has been characterized in laboratory observations by death, acute thermal stress followed by recovery, or no observable stress, even though test fish frequented temperature extremes in the experimental gradient (Meldrim and Gift 1971). Low thermal responsiveness in the laboratory may also be more prevalent during periods of the year with variable ambient water temperatures or low water temperatures, as well as be related to fish size. Moreover, it may be an artifact of laboratory experiments in which fish are exposed to artificially compressed thermal gradients (Gift, pers. commun.).

Sullivan and Fisher (1953) demonstrated a seasonal change in selected temperature which was distinct from the effect of acclimation temperature. Fry and Hochachka (1970: In Crawshaw 1975a) hypothesized that changes in thermoregulation with season are caused by anticipatory thermoregulation. Differences between temperature responses during periods of risinq and falling acclimation temperatures (Meldrim and Gift 1971) may be a response caused by season or due to different field acclimatization states of fishes which are collected during different seasons. In addition to seasonal changes, fishes may prefer different temperatures on a diel cycle (Barlow 1968; Brett 1971; Coutant 1974).

Biological interactions also influence the temperature


responses of fish. Hatch (1973) reported that the presence of largemouth bass in a heated effluent may be a response to forage fish movement rather than a response to a preferred temperature. Gift (1970) reported that Atlantic silversides remained in water warmer than the normal avoidance temperature when placed in a tank with a predatory species. Moreover, Beitinger and Magnuson (1975) demonstrated that the presence of a socially dominant adult bluegill affected the behavior of smaller fish. Therefore, certain aspects of the life history of an organism should be noted when conducting temperature selection studies. For example, territorial fish should probably be tested singularly while schooling fish may respond more accurately if a group is tested (Richards et al. 1977).

The fact that one of the most important sublethal effects is the behavior of an organism in the presence of a thermal effluent (Meldrim and Gift 1971) has been responsible for generation of much of the above data. It has been well documented that fish will congregate within a particular range both in the laboratory and under field conditions (Brett 1956). However, due to the difficulty in establishing temperature preference in nature, the relationship between laboratory and field derived data is poorly known. Reynolds (1977c) discussed this general topic and stated that determination of preferenda under laboratory conditions is for the most part species specific and reproducible, but that "numerous non-thermal stimuli" are present in nature. Stauffer et al. (1976a) recognized this and mathematically corrected for gradient, photoperiod, interspecific interrelationships, flow, and power plant chlorination schedules. Altitude and stream order were not mathematically corrected because of the close proximity of the stations. Regardless of the difficulties encountered in deriving field temperature preference data, there have been reasonably close correlations between laboratory and field data for most species when data have been obtained on the same populations (Ferguson 1958; Neil and Magnuson 1974; Stauffer et al. 1976a). In addition, McCauley and Pond (1971) found that the temperature preference of juvenile rainbow trout in vertical and horizontal gradients agreed reasonably well with the mid-summer vertical distribution of adults found by Horak and Tanner (1964). Contrary to the above observations, Doudoroff (1942) found that opaleye more frequently selected a water temperature of 26 to 27 C, which was not consistant with the normal habitat temperature (Brett 1956). Brett (1944, 1946) attributed this apparent contradiction in part to possible summation of acclimation temperature. Stauffer et al. (1976a) also noted that the spotfin shiner demonstrated a temperature preference in the laboratory, but showed no definable preference temperature under


field conditions. Richards and Ibara (1978) found that movement of brown bullheads into a thermal discharge during the fall could be predicted with laboratory thermal preference data, but that emigration during the spring could not be predicted.

Because of the species specific nature of temperature responses, Hart (1952) and McCauley (1958) investigated the possibility of differences in temperature responses of various races and subspecies. Both McCauley (1958) and Hart (1952) concluded that differences in thermal responses were found among subspecies but not withi.i physiological races. However, work by Hall et al. (1978) suggests that there are differences in temperature behavior responses among races of white perch.

Temperature preference and avoidance data have been generated for many species using a variety of laboratory and field techniques. In general, there is much more complete information available for freshwater fishes. Behavior information for marine species is limited for the most part to field observations or one or two laboratory tests. Little data are available which correlate the interaction between temperature and salinity (Terpin et al. 1977; Wyllie et al. 1976). For a summary of fish responses refer to Brown (1974) and Coutant (1977).

In summary, temperature is the dominant factor which controls the behavior and activities of aquatic pokilotherms (Becker 1973)· As a result, aquatic organisms demonstrate both genetic and non-genetic adaptation to different thermal conditions (Kinne 1963). Studies have shown that changes in temperature can be perceived with cutaneous thermal receptors and this information coordinated with the central nervous system. Both laboratory and field studies indicate that fish select and avoid certain temperature regimes. In general, fish prefer higher temperatures than their acclimation temperature until their final temperature preferendum is attained, although there have been studies which indicate that acclimation temperature does not affect preference responses for particular species. The occurrence of behavioral fever may have important implications for field evaluations of the effects of thermal outfalls. If, in fact, diseased fish or fish in poor condition are collected in artificially heated areas, care should be exercised in determining whether this condition was caused by the effluent or whether the diseased fish were attracted to the effluent due to a febrile response. Thermal history, season, diet, hormone levels, light intensity, salinity, size, biological interactions and age have all been suggested as possible influences of temperature selection. One of the most obvious effects of temperature is the exclusion of


organisms from areas with either unsuitable or non-preferred temperature conditions (Kinne 1963) and replacement by other species. Less obvious are the effects on activity, feeding behavior, reproduction, substrate selection and aggression.

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