AMER. ZOOL., 19:345-356 (1979).

Amphibian Temperature Regulation Studies in the Field and Laboratory

BAYARD H. BRATTSTROM

Department of Biology, California State University, Fullerton, California, 92634

SYNOPSIS. Studies on thermoregulation in the laboratory and field have come a long way
from the early work done between 1940 and 1960. While some physiological studies on
amphibians have progressed at the same rate as those on reptiles, field studies have been far
behind. Laboratory studies have largely delt with thermal acclimation, evaporative water
loss, and thermal and moisture gradient behavior. Field studies, following early summaries
of body temperatures of field animals, have stressed behavioral thermoregulation; yet,
detailed studies on behavioral thermoregulation in amphibians have been completed for
only a handful of species. A few studies have placed behavioral and physiological ther-
moregulation into an ecological or energetic framework; these studies are reviewed, and
suggestions are made for future work.

INTRODUCTION
1975; Mullen and Alvarado, 1976; Walters
The study of temperature regulation in and Greenwald, 1977). These processes
amphibians is complicated by the require- are, of course, intimately tied to the prob-
ment of amphibians to maintain a moist lems of osmo- and thermoregulation
skin or to occur in an aquatic environment. (Reves, 1977). While I argued some years
Numerous studies have dealt with prob- back, "Pity the poor frog, nobody studies
lems of amphibians in maintaining body his physiology," I would now suggest that
water, avoiding dehydration and adapta- the situation, "Pity the poor frog, his be-
tions of anurans to the desert (Seymour and havioral and physiological problems are so
Lee, 1974; McClanahan, 1975; Shoemaker complicated and interrelated, it is amazing
and McClanahan, 1975). While these that we can understand them and that he is
studies have been concerned with water alive at all!!" Since I have recently reviewed
conservation, they are intimately tied to temperature regulation in amphibians
problems of temperature regulation. In an (Brattstrom, 1970a), I propose here to up-
amphibian involved in avoiding dehy- date the field to expose some problems in
dration, these responses may take prece- our approach, and to suggest some direc-
dence over thermoregulatory ones (Tracy, tions for future research.
1975). Thermoregulation may be com-
promised by demands of hydroregulation,
BEHAVIORAL THERMOREGULATION IN THE FIELD
while in other situations, thermoregulatory
demands may predominate. In a review of body temperatures of am-
The amphibian's integument also func- phibians, I confirmed the old notion that
tions in osmoregulation and respiration many amphibians are poor thermoreg-
(Houston, 1971; Wakeman and Ultsch, ulators (Brattstrom, 1963). For many of
these species, water conservation, or re-
striction to aquatic environments, forces
I thank Dr. W. W. Reynolds for organizing, and
inviting me to this symposium. Page charges for publi- them into an essentially non-thermoregu-
cation of this paper were supported by N.S.F. grant latpry mode (Fig. 1). The best some aquatic
No. PCM-78-05691 to W. W. Reynolds. The figures amphibians may be able to do is to seek out
were prepared by Mark Zolle supported by a grant warmer or cooler portions of their environ-
from the Department of Biology, California State Uni- ment. This thermoregulation behavior
versity, Fullerton. I wish to thank Mark for his drawing
and W. W. Reynolds, Victor Hutchison, and Lon may push them into environments which
McClanahan for comments and remarks on the man- conflict with their respiratory and osmo-
uscript. regulatory demands {e.g., a warm area of a

345

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 346 BAYARD H. BRATTSTROM

pond that is low in oxygen) (Houston, 1971; Tracy (1976) suggests that such basking
Wakeman and Ultsch, 1975; Mullen and plays less of a role in the behavioral ther-
Alvarado, 1976; Reves, 1977; Shoemaker moregulation of adult Rana pipiens and that
and Nagy, 1977). behavioral thermoregulation in this species
Many other amphibians, however, have is poor; and his field data and computer
been shown to be fairly active thermo- simulations indicate that the body temper-
regulators, interplaying water economy atures of this species are always near am-
and thermoregulation to allow them to be bient. Evaporative water loss is apparently
more active longer. This presumably allows too high for effective basking thermoregu-
time for more feeding, reproduction, and lation especially at high radiant levels, low
growth (Lillywhiteetal., 1973). In my initial relative humidities, high air temperatures,
review of amphibian body temperatures, I and high wind speeds. Tracy (1975) also
presented data on several species of frogs showed that a bullfrog, Rana catesbeiana,
(especially Rana cascadae, pipiens, and placed in the sun in a cage, maintained its
clamitans) observed basking in the sun (Fig. body temperature within 4°C. He argues
1), in water or on moist soil (Brattstrom, that this was not thermoregulation as the
1963). These frogs behaved like typical frog was passive and not shuttling (as it
heliothermic lizards, with body tempera- couldn't do). But, he noted, by noon it be-
tures above ambient. The frogs were ap- came dehydrated and stopped producing
parently utilizing evaporative water loss to urine. I suggest that perhaps the frog was
maintain body temperatures below lethal maintaining its body temperature fairly
levels. Later, Lillywhite (1970, 197 la, 1974) constant by physiological thermoregulation
who monitored body temperatures by (evaporative cooling), as it was restrained
telemetry, demonstrated behavioral ther- from doing any behavioral thermoregula-
moregulation in Rana catesbeiana which in- tion.
cluded diurnal basking complete with pos- Basking has only been reported for a few
turing. Brattstrom (1970a) demonstrated other amphibian species. Valdivieso and
that the basking Australian hylid frog, Tamsitt(1974) showed that the montane
Hyla(=Litoria) caerulea, could maintain a Colombian frog, Hyla labialis, is a ther-
fairly constant body temperature by mophilic heliotherm whose body temper-
evaporative cooling under forced basking, atures are usually higher than ambient due
even when the heat load was increased. The to basking or absorbing solar radiation that
frog apparently could regulate the amount filters through clouds. Adults and sub-
of water passing through its skin, as a func- adults were active in the morning and early
tion of heat load. Another Australian hylid, afternoon. Terrestrial juveniles were active
Hyla Moris, could not do this. Basking anu- at all hours, but were most active at night.
rans use a variety of mechanisms (skin This is in contrast to observations on toads
sculpturing, water movements over the (Seymour, 1972; Lillywhite et al., 1973) in
skin, mucous secretions, activity, and cra- which the juveniles often have higher body
nial co-ossification) to reduce or facilitate temperatures than adults, and spend con-
evaporative water loss and cutaneous gas siderable time basking. Seymour (1972)
exchange during thermoregulatory bask- suggests that the young of the desert toad,
ing (Lillywhite, 19716; Heatwole and Bufo debilis, utilize basking thermoregula-
Newby, 1972; Christensen, 1974; Lil- tion for rapid feeding, accelerated diges-
lywhite and Licht, 1974, 1975; Sievertetal., tion, and rapid deposition of fat prior to
1974). Lillywhite (1975) showed the im- winter dormancy. Lillywhite et al. (1973)
portant role of blood circulation in main- showed that youngB. boreas basked, except
taining levels of skin hydration during when food was withheld. Orientation to a
basking. As long as water can enter the frog heat source could be elicited in starving
and be effectively evaporated from the skin animals by feeding them, suggesting that
(thus precluding desiccation), many anu- one advantage of such behavior is accelera-
rans, especially Rana catesbeiana, can main- tion of the digestive processes. Growth
tain fairly constant body temperatures. studies at a variety of temperatures indi-

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 AMPHIBIAN THERMOREGULATION 347

cated that energy ingestion, linear growth, tion of lipids onto the integument is fol-
weight increase and gross conversion lowed by a complex wiping movement
efficiencies were all maximal at 27°C and which spreads the secretion over the body
were nearly identical to that of toads al- of the frog. The frogs then remain mo-
lowed to thermoregulate in a photothermal tionless. This lipid material reduces evap-
gradient (25.6°). Diurnal behavior of small orative water loss up to 30°C, above which
toads and of tadpoles, compared with the evaporative water loss begins to increase
more nocturnal adults, may have evolved to and then increases precipitously between
maximize growth rates of younger indi- 35 and 40°C. At 40° ambient temper-
viduals, shortening the time to adult size atures, skin and core temperatures of the
(Brattstrom and Warren, 1953; Brattstrom, frogs remained at 36-37°C. Small in-
1962). creases in water loss resulted in depres-
While Bufo boreas, and most amphibians, sions of surface temperature correspond-
bask only on wet soil, Lilly white et al., ing to the release of clear fluid onto the
(1973) observed some toads basking on dry skin surface (McClanahan et al., 1978).
soil. Perhaps the water storage ability of B. This frog thus uses a skin-surface-protec-
boreas allows it to bask on dry soil, and then tive device to reduce water loss while living
move to wet soil or water to replenish in the desert, yet as ambient temperatures
water loss. rise and the possibility of thermal death
Several species of Australian hylid and approaches, the protective nature of this
leptodactylid frogs bask (Johnson, 1970, substance breaks down and the skin be-
I97la,b). These include montane rapid- comes available for evaporative cooling to
stream-dwelling frogs, diurnal tropical maintain body temperatures below lethal
frogs, and semi-desert diurnal and noc- levels.
turnal frogs. The diversity of frogs that
utilize basking suggests that it may occur in BEHAVIORAL THERMOREGULATION STUDIES IN
different species for different reasons and THE LABORATORY
that the mechanisms of reducing the prob-
lems of dehydration and cutaneous gase- Studies on behavioral thermoregulation
ous exchange may be different. Seymour of adult amphibians in the laboratory have
and Lee (1974) have suggested that such been concerned largely with the role of
xeric Australian frogs as Hyla(=Litoria) ru- evaporative water loss and the behavior of
bella and caerulea may solve water and ther- amphibians in thermal gradients. Thermal
moregulatory problems by excreting ur- gradients are difficult to set up for am-
ates, as in other xeric frogs (Loveridge, phibians because of the necessity to avoid
1970; McClanahan, 1975; Shoemaker and dehydration to the animals. It is also
McClanahan, 1973; Shoemaker and difficult to determine whether an animal is
McClanahan, 1975; Blaylock et al., 1976; responding to a thermal or a moisture
Drewes et al., 1977). gradient. Spotila (1972) in a study of
The tacky and moist-looking skin of plethodontid salamanders in thermal and
many hylid frogs from different deserts relative-humidity gradients, comparing
(such as H. caerulea from Australia, Phyl- gradient studies with field studies, indi-
lomedusa spp. in the New World) elicits cated that these salamanders do avoid ex-
speculation as to its function. The ar- tremes and exhibit thermal preferenda,
boreal, desert-dwelling Argentinian P. which .were species-specific and not signifi-
sauvagei, not only has utilized certain as- cantly affected by acclimation temperature
pects of nitrogen excretion and osmoreg- or photoperiod. Dehydration was greater
ulation to survive in the desert (Shoemaker at higher temperatures, and in apparent
and McClanahan, 1975), but also uses compensation, the salamanders selected
evaporative cooling in a subtle manner. the highest relative humidity in the gra-
The skin of this frog is hydrophobic and dient (Spotila, 1972). Thus, the salaman-
contains numerous alveolar glands con- ders seem to be interplaying thermal and
taining lipids (Blaylock et al., 1976). Secre- moisture responses.

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 348 BAYARD H. BRATTSTROM
Feder and Pough (1975) studied tem-
perature selection in red-backed salaman-
ders, Plethodon cinereus. Animals acclimated
to low temperatures selected high temper-
atures in the gradient, and those accli-
mated to high temperatures selected lower
temperatures; acclimation of temperature
selection was faster than acclimation of
critical thermal maxima. This suggests that
after exposure to, for example, low tem-
peratures, there is value or at least a be-
havioral response of a salamander to seek
warmer areas. This would facilitate rapid
feeding and digestion following a period
of cold weather and possible starvation.
Hutchison and Hill (1978) showed that
preferred temperatures of bullfrog (R.
catesbeiana) tadpoles varies in a complicated
manner with stage of development and
thermal acclimation. While the mean T b
was 20.7 and the modal T b 21.0°C, there
was a tendency for earlier stages, accli-
mated at lower temperatures, to select
lower preferred temperatures. At later
stages, and especially with tadpoles accli-
mated to high temperatures, there was a
preference for higher temperatures. This
latter response may be associated with the
high temperatures selected near meta-
morphosis, and high temperatures toler-
ated by recently metamorphosed juveniles.
Hutchison and Hill (1978) further suggest
that for those species with plasticity of their
preferred temperatures, with changes in
acclimation temperatures, survival is en-
hanced by avoidance of lethal temper-
atures, and energetic efficiencies are maxi-
mized through maintenance of body tem-
peratures at or near physiological and bio-
chemical optima. Plasticity is thus one
strategy for circumventing serious conse-
quences of rapid temperature change or
thermokinetic extremes.
One problem with thermal gradients
constructed for aquatic organisms is verti-
cal thermal stratification. Reynolds and
Casterlin (1976), Reynolds (1977) and
Casterlin and Reynolds (1977, 1978) have
solved this problem by use of a device
which allows an animal to control water
temperature. The animal controls its own
body temperature by its movements be-
tween chambers of water monitored by
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photocells and associated circuitry which
controls heating and cooling equipment.
Thus by its own behavior an animal can
seek out its preferred temperature by
manipulating the temperature of the water
until it reaches its preferendum. In a study
of R. pipiens tadpoles, Casterlin and
Reynolds (1978) showed that while the
tadpoles had a bimodal aspect of temper-
ature regulation (with highs during day
and again at night and lows during dusk
transitions), the preferred temperatures
and modal preferenda were between 27
and 28°C. These data are similar to results
from typical thermal gradients (Lucas and
Reynolds, 1976) and adult modal field
body temperatures (Brattstrom, 1963). It
seems to me that the development of ef-
fective thermal gradients or use of shuttle-
boxes will provide an opportunity for criti-
cal studies on the roles of photoperiod,
acclimation, and energy metabolism on be-
havioral thermoregulation. The shuttlebox
device (Reynolds, 1977) also may be adapt-
able to studying simultaneous responses to
several factors (i.e.., choices between high
temperature-high oxygen water vs. high
temperature-low oxygen water against a
choice of low temperatures and different
oxygen levels).
Some of the studies of Lilly white et al.
(1973) on B. boreas juveniles were carried
out in a laboratory photothermal gradient.
Behavior of these toadlets differed under
different soil and starvation conditions.
Since it is also clear that acclimation and
photoperiod may affect some amphibians,
it may be important to mention here that
length of time that animals are exposed to
the gradient (number of hours or days in
the gradient), the number of animals used
at any one time, acclimation influences,
gradient size, shape, and thermal consis-
tency, and desiccation levels may also affect
responses of animals to a gradient, so cau-
tion should be taken in construction, ex-
perimental design, and data interpretation
of thermal gradient studies.
PHYSIOLOGICAL THERMOREGULATION STUDIES
IN THE LABORATORY
Most laboratory studies on amphibians
 AMPHIBIAN THERMOREGULATION
haye dealt with physiological aspects other
than thermoregulation. Considering the
kinds of studies that have been done on
reptiles, it is surprising to find so few
studies on amphibian thermoregulation.
In heating and cooling studies on the
bullfrog,/?, catesbeiana, (Tripp and Lustick,
1974) in water and air, there was no differ-
ence in heating and cooling of frogs in
water, but there was in air. Heart rates
were higher during heating than during
cooling in water, while there was no differ-
ence in heart rates during heating and
cooling in air.
The majority of physiological studies in
amphibians have understandably been
metabolic studies on the relationship of
metabolism to gas exchange, surface area,
and habitat. Most of these studies were
done at a variety of temperatures, and thus
contribute to our understanding of the
physiology of amphibians and the relative
importance of behavioral and physiologi-
cal thermoregulation to enhance or curtail
certain aspects of the animals' physiology
(see the following papers for current
physiological and environmental ap-
proaches to old problems; Clausen, 1973;
Bennett and Wake, 1974; Guttman, 1974;
Turney and Hutchison, 1974; Heath,
1976; Pitkin, 1977; Weathers and Snyder,
1977). In addition, the interplay between
these responses sometimes provides solu-
tions to, or problems for, other physiologi-
cal responses of the animal.
Recent studies have made us aware of
the increasing importance of the role of
anaerobic respiration in stress activity and
in normal behavior. Guttman (1974), for
example, has shown that the toad, B. val-
liceps, can stand anoxia longer than B.
woodhousei. He suggests that this allows B.
valhceps to endure its longer winter dor-
mancy better than B. woodhousei (also see
Armentrout and Rose, 1971). Turney and
Hutchison (1974) have shown in R. pipiens
that of stressed energy, 69 and 73% was
supported anaerobically at 25° and at 15°C,
respectively. They suggest that the ineffi-
ciency of the breathing cycle of this frog,
coupled with limitations of the respiratory
surface and separate gas exchange path-
ways, have placed extreme restrictions on
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349
the capacity of the frog to meet oxygen
demands, thus forcing the animal to incur
a relatively large oxygen debt during
maximal activity. Other studies with an-
aerobic respiration (Bennett and Licht,
1973, 1974; Seymour, 1973; Bennett,
1978) suggest that slow-moving amphib-
ians such as B. boreas produce small
amounts of lactate and do not exhaust. In
contrast, fast-moving jumping forms such
as R. pipiens have high levels of lactate gen-
eration, but are unable to sustain maximal
activity. Aquatic species rely largely on
air-gulping during activity and show little
anaerobiosis. The rate of lactate produc-
tion is also directly correlated with preda-
tory avoidance. Noxious-tasting, aggres-
sive, or cryptically-colored amphibians
have low anaerobic scopes for activity while
others rely upon anaerobiosis for rapid
flight (R. pipiens) or rapid avoidance be-
havior {Batrachoceps attenuatus). In fact,
rapid activity in amphibians appears to be
possible only at the expense of extensive
anaerobiosis (Bennett, 1978). In my opin-
ion, two important lines of research are
developing from these studies. First, is the
question of how the enzymes and enzyma-
tic pathways that are involved in aerobic
and anaerobic respiration (Bennett, 1974,
1978; Baldwin et al., 1977) respond to the
thermoregulatory demands of the animal
and whether the same enzymes and path-
ways are used at all temperatures.
Many organisms, including amphibians,
compensate aspects of their metabolism so
that they are more efficient at specific tem-
peratures than would be predicted other-
wise. The studies on such compensation
mechanisms in amphibians have largely
been restricted to adaptations by amphib-
ian larvae, especially in embryonic temper-
ature adaptations in northern and alpine
frogs (Licht, 1971; McLaren and Cooley,
1972; Packard, 1972; Brown, 1975; Kura-
moto, 1975a,b). A perhaps equally im-
portant aspect of metabolic compensa-
tion is that shown by high elevation popula-
tions of adult salamanders (Fitzpatrick and
Brown, 1975) and anurans (Packard and
Bahr, 1969; Packard, 1971, 1972) though
in recent studies on Pseudacris triseriata, the
compensation may not be in oxygen con-
 350 BAYARD H. BRATTSTROM
sumption but in differences in breeding
seasons, and activity times (Packard, 1971).
A considerable body of literature on am-
phibian physiology has been concerned
with thermal resistances and the effect of
thermal histories on thermal tolerances.
Studies on the rate and range of thermal
acclimation have provided us with infor-
mation on the physiological plasticity of
amphibians, and on genetic limits on ther-
mal tolerances. These data have also been
shown to have zoogeographic implication
(see reviews in Brattstrom, 1970a, b). Re-
cent studies have extended these ideas {e.g.,
Pough and Wilson, 1970; Farrell, 1971;
Fitzpatrick et al., 1971; Holzman and
McManus, 1973; Pough, 1974; Burke and
Pough, 1976; Feder, 1978; Hoppe, 1978).
Importantly, others are looking at acclima-
tion effects on tissues (Shertzer etal., 1975;
Lascano et al., 1976; Lagerspetz, I977a,b;
Ballantyne and George, 1978) and blood
parameters (Weathers, 1975, 1976). In-
terestingly, Ballantyne and George (1978)
have shown that acclimation to cold (from
21 to 5°C over one month) in R. pipiens can
raise muscle mitochondrial content.
Whether this is an adaptation for dormancy
is unknown, but worthy of continued study.
These studies hopefully will contribute to
our understanding of the physiology of the
amphibian in solving its simultaneous
problems of temperature, water, gas, and
ionic regulation (Fig. 1). They are already
giving us a clue to the basis for the marked
seasonal differences seen in some amphib-
ians. Lagerspetz, (19776), for example has
shown that there is a temperature and sea-
sonal effect on CNS activity mediated by the
thyroid via the autonomic nerves. These
seasonal differences may be important for
spring reproductive effort and winter sur-
vival.
PHYSIOLOGICAL STUDIES ON TEMPERATURE
REGULATION IN THE FIELD
Body temperatures taken of amphib-
ians in the field are important, but this is
not studying thermoregulation. As point-
ed out by Heath (1964, 1965) these data
describe only the limits and possible pref-
erenda of the animals. Studying ther-
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moregulation involves monitoring be-
havior at the same time as body tempera-
ture (i.e.., watching for emergence, and
then recording body temperature of
emerging animals). Of all the field body
temperatures recorded (Brattstrom, 1963,
1970a; Lillywhite et al., 1973), behavioral
thermoregulation was sufficiently docu-
mented for only a few species (B. boreas,
Acris crepitans, H. regilla, R. catesbeiana,
Taricha rivularis). A more effective way to
study this type of thermoregulation is by
the use of telemetry. Lillywhite (1970,
1971a, 1974, 1975) has used this and other
techniques to effectively study behavioral
thermoregulation in the field and the lab.
While body temperature is a function of
environmental inputs and of physiological
responses and interactions, in these studies
body temperature was usually the only
physiological parameter measured. It is
important to measure several physiological
parameters of amphibians simultaneously
with modified technology and with patience.
One area of field investigations not yet
touched with amphibians is the construc-
tion of time/activity budgets in the field.
This will be an essential step in the con-
struction of time/activity/energy budgets
and in appreciating the role played by am-
phibians in total community energy budget.
This will be a difficult task due to the secre-
tiveness and nocturnality of many amphib-
ians and will require patience since many
amphibians spend a lot of time being inac-
tive. Yet, this may be an important part of
energy conservation in amphibians.
ENERGY METABOLISM AND ENERGY BUDGETS
Considerable interest has developed re-
cently on the relative roles of energy pro-
duction in amphibians (Hutchison et al.,
1977; Bennett, 1978). We now have studies
on energy metabolism in amphibians in-
volving size, season, fat deposition and
>
FIG. 1. Diagrams of body-environment interactions
of three types of amphibians. Above, a basking
(heliothermic) anuran (partly after Pough, 1974;
Tracy, 1975, 1976). Middle, a cryptic salamander or
largely nocturnal anuran. Below, an aquatic amphib-
ian; (larval or adult).
 AMPHIBIAN THERMOREGULATION 351

DIURNAL AND
BASKING AMPHIBIAN
'particular

CONVECTIVE THERMAL RADIATION

HEAT LOSS FROM ATMOSPHERE

SCATTERED AND WATER

REFLECTED SUNLIGHT LOSS

THERMAL RADIATION
TO ENVIRONMENT

EVAPORATIVE THERMAL RADIATION

FROM VEGETATION
HEAT LOSS

THERMAL RADIATION
FROM GROUND
HEAT CONDUCTION
TO OR FROM GROUND WATER INPUT
FROM SOIL

SECRETIVE OR
NOCTURNAL AMPHIBIAN
CONVECTIVE HEAT
GAIN AND LOSS

EVAPORATIVE
HEAT LOSS
THERMAL RADIATION
TO ENVIRONMENT

GASEOUS
EXCHANGE

WATER INPUT GASEOUS HEAT CONDUCTION

FROM SOIL EXCHANGE TO OR FROM GROUND

AQUATIC
AMPHIBIAN

BEHAVIORAL RE8PONSE
TO AND FROM
WARM WATER

CONVECTIVE AND
CONDUCTIVE HEAT
GASEOUS TO AND FROM WATER
EXCHANGE

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 352 BAYARD H. BRATTSTROM

utilization, and reproductive effort (Sey-
mour, 1973; Seymour and Lee, 1974; Fitz-
patrick and Atebara, 1974; Beckenback,
1975; Tracy, 1975, 1976; Feder, 1976). Re-
cent studies on feeding behavior and diges-
tion efficiencies have also contributed to
our understanding of the energy utilization
of amphibians (Lillywhite et al., 1973;
Sternthal, 1974; Tracy, 1975; Smith, 1976).
Fitzpatrick (1973) has studied energy
budgets in the salamander Eurycea bis-
lineata, and Smith (1976) has produced an
energy budget (Fig. 2) for the toad/?, terres-
tris. Assuming a digestive assimilation effi-
ciency of 74%, about half of that energy
goes into metabolic costs and half into
production. Of the latter, about half goes
into growth and half into reproductive ef-
fort (Fig. 2). The latter, of course, may be
variously partitioned at different ages and
seasons.
Burton and Likens (1975) have deter-
mined that the energy flow through the
salamander population in a New Hamp-
shire forest is 11,000 KCal/ha/yr, equal to
0.02% of the net primary productivity and
about 20% of the energy flow through
birds and mammals in the forest. On the
other hand, salamanders are 60% efficient
in converting ingested energy into new tis-
sue, and produce more new tissue annually
than do bird populations. If we had some
time/activity budgets for more amphib-
ians, we could begin to develop time/activ-

ENVIRONMENT = 100 °70

INGESTION

26°7C 74°7 C
UNASSIMILATED ASSIMILATED
ENERGY ENERGY

38°7 O 36°7 O
METABOLIC PRODUCTION
COSTS OF TISSUES

o°r. O-36°7o| I O - 3 6 °7 O

GROWTH REPRODUCTION

3O.6°7O f 54°7 n
LEAN DRY FAT
BIOMASS ACCUMULATION
FIG. 2. An energy budget for the toad, Buju ttrre\ln\ (modified after Smith, 1976).

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 AMPHIBIAN THERMOREGULATION 353

ity/energy budgets and begin to approach

the problem of the cost of thermoregula-
tion. ~ « LIVER GLYCOGEN
While many workers have noted seasonal • • BLOOD GLUCOSE
II
changes in "winter" and "summer" frogs It ,
1X
t 1
(Brattstrom, 1970a), it has only been in light ^— LIVER LIPIDS a
> 1
FAT BODIES
of renewed interest in energetics that new 1 »
1 t /
/
approaches to seasonal changes in metab- m
t- 1 1 /
olism have been made (Fitzpatrick, 1971; z 1 1 /
O
3 1
Harri, 1973; Harri and Talo, 1975a,b; I<
1 1
1 //
Shertzer, et al., 1975; Weathers, 1975, 1 1 /
1976; Lagerspetz, 1977a,6). In terms of en- V 1 1/
ergy metabolism, and thus the costs for a \ • • *

T\
.' «./-'"
variety of functions, it is also important to
know how an amphibian may be partition-
ing the utilization of its energy with season.
Thus Lillywhite et al. (1973) suggest that
behavioral thermoregulation and energy EMERGENCE BREEDING DORMANCY

partitioning in juvenile toads (B. boreas)

maximize growth, and shorten the time to
reach adult size. This is also apparently true
for B. terrestris (Smith, 1976). In adults, en-
ergy is largely partitioned into reproductive
effort (production of eggs and sperm, and
cost of reproductive behavior). In addition,
amphibians must also prepare for cold
winters or long periods of dormancy un-
derground (Seymour, 1973). Seasonal dif-
ferences in metabolism of adult amphibians
probably represent an interplay between
energy utilization for reproduction and
preparation for winter. Recent studies
(Pasanen and Koskela, 1974; Koskela and
Pasanen, 1975; Byrne and White, 1975)
have demonstrated marked and interesting
changes in liver and muscle glycogen, blood
glucose, and body lipids. In R. catesbeiana
lipid reserves become exhausted from the
time of emergence through the breeding
season (Byrne and White, 1975). Lipid re-
serves then increase prior to and into dor-
mancy, while blood glucose levels rise dur-
ing the breeding season and are lowest
upon emergence from dormancy (Fig. 3).
These may not be the same kind of changes
seen in other amphibians (Reno etal., 1972;
Seymour, 1973; Gehlbach etal., 1973), but
the physiological strategies used are proba-
bly a function of the different seasonal ac-
tivities employed by different amphibians.
Depressed metabolism, fat deposition and
utilization, and ability to endure anoxia (or
have low oxygen demands) probably allow
FIG. 3. Seasonal changes in lipid reserves, and blood
glucose levels in the bullfrog, Rana catesbeiana (Sim-
plified after Byrne and White, 1975).
amphibians to survive harsh periods. At
this time, thermoregulation and its costs are
probably low. The costs and benefits of
thermoregulation are highest during sea-
sonal activity and help in growth, energy
metabolism, and reproductive effort. Much
more needs to be done; modeling efforts
(Tracy, 1975) may be an important next
step.
SPECIAL ASPECTS AND PROBLEMS
Some anurans prefer temperatures ex-
ceeding 30°C (e.g., H. Smithi, rubella, Phyl-
lomedusa sauvagei; Brattstrom, 1970a,b;
Johnson, 1970, 1971a,6; Seymour and Lee,
1974; McClanahan «£«/., 1978). Interesting
physiological processes may occur in frogs
at high temperatures, (Stephenson, 1967;
Harri and Talo, I975a,b) implying the ne-
cessity of measurements at >30°C.
Kluger (1977) demonstrated that H.
cinerea developed a fever (increase of 2°C.in
body temperature) following injections of
killed Gram-negative bacteria (Aeromonas
hydrophila). Casterlin and Reynolds (1977)
demonstrated a similar "behavioral fever"
(significant mean increases in preferred
temperature) in tadpoles of R. catesbeiana
and R. pipiens, following similar injections
with the same bacteria. These studies have
implications both for understanding the

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 354 BAYARD H. BRATTSTROM

adaptive advantage of fever in diseased (ed.), Comparative physiology of thermoregulation, pp.

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