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WATERBATH STUDItrS ON FIELD AND CAPTIVB CITtrLLUS LATERALIS1

J. J. YOUNG AND M. L. RIEDESEL Department of Biology, University of New Mexico, Albuquerque
INTRODUCTION

ATERBATTT

experiments were

conducted under field and laboratory conditions to investigate the extent to which physiological

importance and characteristics of acclimatization in mammalian hibernators are ill defined (Mayer, 1953; Deane and

Lyman, 1954).
MATERIAL AND METHODS

factors are involved in acclimatization of field animals to laboratory animal quarters. The waterbath technique provides a uniform surface temperature that facilitates quantitative measurement of the capacity of an animal to limit the flow of heat from the site of production, or core, to body surface. The tolerance of Citellus lateral'is to immersion in cold water has been demonstrated to be primarily determined by the extent of peripheral vasoconstriction and metabolic heat production (Yelverton, 1964). The lack of acclimatization following waterbath exposures and other advantages of this technique have been reported by Adolph and Richmond (1956). Acclimatization or preconditioning of animals very often is the most important factor in determining the tolerance of animals to environmental stress. Preparation of animals for hibernation represents an extreme example of acclimatization as hibernation involves changes in essentially all physiological processes (Lyman and Chatfield, 1955). Definition of mechanisms involved in acclimatization of mammalian hibernators would facilitate understanding hibernation as well as acclimatization. However, the
l Work supported by National Science Foundation grant GB2l6.

The experimentai animals were goldenmantled ground squirrel s, Citellus lateralis

(Say). Collections were made in a 12sq-mile area, 10 miles east of Cuba on

Route 126, Rio Arriba County, New Mexico, at 8,300-ft elevation. Four collecting trips were made to the above
The term "field condition" refers to the data collected from animals 15 min after capture; the term "captive" refers to the same animals captive 14-62 days. In each series of experiments animals were first exposed to a given waterbath temperature in the field. Animals were weighed to the nearest gram, and rectal probe inserted to 6-cm depth prior to immersion in water for 30 min. The waterbath chamber had inside dimensions of 32 X 32 X 26 cm and was placed in a 25-gal tank of water to facilitate maintenance of the specified waterbath temperature. After completion of the field experiments, the animals were placed in individual wire cages with water and food availablc, and transported within 48 hr to the air-conditioned laboratory, 24 + 3 C. Later, each animal was exposed to the same procedure in the laboratory. Each series is described as follows: Series I: four animals exposed
area between June 28 and JuIy 31,1964.

189

190
15

J. J, YOUNG AND

},T.

L. RItrDESEL

min, 14 days and 48 days after capture to 25 C water; Series II: five animals exposed 15 min, 21 days and 62 days after capture to 25 C water; Series III: two animals exposed 15 min, 14 days and 50 days after capture to 33 C water; Series IV: six animals exposed 15 min and 28 days after capture to 35 C water. Ail waterbath exposures were of 30-min duration. Recordings of the rectal, water, and air temperatures were made at 2-min intervals for the first 10 min and at 10min intervals for the next 20 min with a telethermometer, + 0.1 C. During the immersion, animals were confined in a cone of f-inch mesh wire. Suspending the cone from a tripod provided constant depth of immersion, water level approximately 0.5 cm below the ears. The term "body-heat loss" refers to the change in the heat content, or stored heat, of the animal during the time indicated and was calculated as described by Burton and Edholm (1955). The total heat loss includes the metabolic heat plus the body-heat loss. The mean metabolic rate of C. lateralis in confined experiments with waterbath temperatures of 25-35 C was 5.3 kcal per hour with 1.1 sn (Yelverton, 196+).

temperature occurred during the first 10 min of immersion in the water. The mean decrease in rectal temperature between the 10th and 20th min of immersion was 1.6 C with 1.3 sn. The mean decrease in rectal temperature between the 20th and 30th min of immersion was 0.3 C with 0.4 sn. The mean decreases in rectal tempera-

ture observed during the 30-min exposures are given in Figure 1. In Series I (25 C water) the decreases in rectal
temperature of field and 14-day captive

animals were similar, P ) .3 (Fig. 1). However, the 48-day captive animals lost less heat (p < .001). In Series II (25 C water) the decrease in rectal temperature of.2I- and 62-day captive animals was less than recorded for field animals (2 < .OOt). The rectal temperature data on 48- and 62-daY caPtive animals were similar (2 > .10). In Series III (33 C water) the 14- and 50-day captive animals were able to maintain body temperatures higher than field animals (2 < .05). In Series IV, the 35 C water represented a cold stress for field animals, 1.6 C decrease in rectal temperature, whereas 28-day captive animals had a mean decrease in rectal temperature of 0.6 C (p < .01). The animals consistently gained weight RESULTS during captivity; however, the correlaA constant rectal temperature indi- tion between body weight and decrease cates a balance between heat gain and in rectal temperature was poor, - '377 heat loss and implies the existence of correlation coefficient. Most of the weight temperature regulation. During the 30- differences between test groups can be acmin waterbath exposures, the body-heat counted for by the rapid gain in weight loss in the "field condition" was greater of young adults, 140-160 g, in the labothan the heat loss of the same animals ratory. The effect of the body weight on the amount of heat lost by animals in in the laboratory. The decrease in rectal temperature the waterbath was minor, for instancel was the principal criterion for interpret- in the field, the rectal temperature of a ing the capacity of the animals to tolerate 151-9 animal decreased I2.5 C during the cold water. In all exposures, field and 30-min immersion, and the rectal temlaboratory, the greatest decrease of rectal perature of a 210-9 animal decreased

WATERBATH STUDIES

t91

13.0 C under the same conditions. Atrthough some field animais weighed more than captive animals, the drop in rectal temperature of fie1d animais was alr,vays greater than the acclimated captive animals.

or 14-day captive anirnals, but vigorous shivering occurred in 25 and 33 C water by animals captive 2l days or longer.
DISCUSSION

The rectal-temperature measurements prior to each waterbath exposure demonstrate the range of rectal temperature of field animals to be greater than captive animais (Table 1). Shivering was never observed in fietd

The data inCicate limited body-temperature regulation by Citellus lateralis collected during summer months. Ilibernation and the accompanying lowering of body ternperature and conservation of body stores of energy are recognized to be important facets of mammalian

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Frc. 1.-Thermal responses of Citellu,s laterali,s to immersion tn 25, 33, and 35 C water

192

J. J. yOU\TG AND M. L. RIEDBSEL namely, hypothermia in response to cold environments and homeothermia during moderate conditions, facilitate conservation of energy and a high rate of activity during the short summer period available for growth and reproduction.

hibernation. In contrast, nurnerous laboratory experiments have demonstrated t'he capacity of mammalian hibernators to have homeothermic temperature regulation in various environments (Kayser, 1961; Suomalainen, 196+). The water-

bath experiments clearly demonstrate lability of body temperature of field animals in contrast to homeothermic temperature regulation by animals maintained in a neutral thermal environment.
TABLE
1

IJnder given conditions, a heterothermic hibernator may bc more capable of maintaining a higher core temperature

than a non-hibernator. Earlier studies

have demonstrated C. spilosoma, a hiber-

nator, maintained a higher rectal temperature than C.lewcuyis and Dipod,ontys ord;i, non-hibernators (Yelverton, 196+) . Apparently, acclimatization and preconditioning are as important in defining limits of temperature regulation as the genetic characteristics. Kayser in a discussion of awake hibernators during summer (Mayer and Van Gelder, 1965)
have chemical thermoregulation (heat production), but their physical thermoregulation (heat-Ioss regulation) is insufficient. The responses to acute cold stress, 25 C water, suggest that both chemical and

RECTAT- TEMPERATURE op "CtTBLLus LATERALrs" MEASURED pRIoR To WATERBATH EXPERIMENTS

Days after Capture

DifferMean
Range ences in

Extremes

14...
21

0...

114
10
17

39.9

37

.6-42.8
2.2 2.3
a,

28... 48...
62.

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8 8

39.4 38.5
39.
1

38.2-40.4
37 .3-s9.6 38 . 0-40.4 36 .7-39 .5

points out that these animals

38.6
37 .7

36.7-39 .2

2.8 2.5

Definition of pre-experimental environrnental conditions should be recognized in future studies regarding the bodytemperature-regulating capacity of mammalian species. The lack of homeothermic temperature regulation in field animals has been suggested by observation of the wide range of rectal temperature (Muilally, 1953). Assuming a Qrc ol 2, a 5 C drop in body temperature represents a 50/s reduction in energy expenditure. The absence of shivering and apparent inability to limit heat loss by vasoconstriction in 25 and 33 C water confirm the existence of labile body-temperature regulation in field conditions. On the other hand, during periods of moderate environmental temperature (21-27 C)
the animals have homeothermic tempera-

physical thermoregulation are very limited in field animals. Future studies should define the extent of environmental stress which is necessary to produce homeothermic or heterothermic temperature regulation. Acclimatization or changes in thermoregulation in response to cold stress must be a key factor in preparation for hibernation.
SUMMARY

ture regulation which facilitates maximum activity. The temperature-regulation characteristics of the mammalian hibernators during summer months,

air-conditioned animal quarters involves improved homeothermic temperature regulation as evidenced by a smaller drop in rectal temperature when immersed in 25 or 33 C water. 2. Apparently a minimum of 15-21 days is required for acclimation and development of homeothermic temperature regulation.

to

1. Acclimation of Citellus

lateralis

Young, J. J. aad lI. L. Riedesel. L967. Waterbath studles sn field and captive C{tel"1us Latetal"J"s,

Fhysiological 4_qqlpsy 40: 189-193

\\'ATERBAT'}I
3. Ar:elirnntcd animals shivercd during 30-min immcrsir:n in 25 anrl 3.j C rvatcr, whereas shivering rvas not observed in

ST LIDIES

193

ficltl animals.

4. Acclimatization tp therm:rl environment appcars to be an important aspcct in determining the response of mammalian hibernatnrs to ncute cokl stress"

I,ITER]ITURE CITSD
Arcr.Fu, Il, F., ancl J. Rrcuaraxn. 19.56. lltlaptation to cold in golden h*nrsler and ground squirrel rneasured chietly lry rrtes ol body cooling, J. r\ppl. Physiol. e:53-58. BurmN, A, C., anel O. G, E$Eor*e, 1955, Man in a e*ld environment. Edtvar<i Arnold, London. f)rlxn, ll. W., arrd C. P. I-vveu, 1954. Body temper*ture, tbyroirl rnd adrenal c$rlex of hs.msters during coltl exposurc and hibernation, with comparisons to rats, Endocrinology 55r300*31$" K,t vsut, C. 19dI. The ph1*iology of nntura! hibernation. Pergamon, New liork, LvM,tN, C. P., and P. O. Cll,u:nuru. 195.1. Ph-.-'sinlcgy of hitrernation in mammais, Ithysi*1, Rev. 35;403*425.

M*vnt,

1T.

\r,

1953. Acclimatization
Sperrropt;d;/lra".s

grcund *quirrel,

of ihe Barrow *ndwJsL*s. hn*t.

Rec. 120:437*438.

l\{even, W. V., arrd R. C. \i,rru Gnt.nr:n. 196.5. }Iammrlian reactitrns to stressful environnrents. Pp. 180-2?8 d* fhysiolngical m*mmalogy. Yol. 3. Acndcnric, New York. *,{ur"r,elrv, D. P. 1!}53, l{ibernation in the goldenmantled ground squirrel. J. Mammal. 3{:65-73. Suoxar.atr.lux, Paavo. 1$64. Proeeeding of thr $econd lnterns,tionel Symposium on Natural Manrmalian tlibcrnation. ^Ann. Acad. Sci, Fenn" $cr. r\; oV{71l18)" 4.i3 pp. Yntxr:ntox, J. T. l9S{, Tlle water-lrath as s tocl iB evaluating r*sponses of small ms.mmals to tbermal stress, M.S. Thesis. Univ. New !Iexie*.