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In Hrysiological

L969.

C. Clayton Hoff and. i{arvin L. Ried.esel, Eid.itors. University of t{ew Mexico Press. pp. 15-52.

$ystems

in Semiarid Environments.

Sodium-Z2 Retention as a Function of Water Intake by Citellus lateralis'


Gnnv L. BrNrz
Department of Biology, The University of New Mexico, Albuquerque, New Mexico

Nncenvn wATER balance is one aspect of the stress involved in hibernation. Hibernation has been observed to continue uninterrupted as long as 3 weeks by animals maintained in a laboratory cold room. Such observations imply that in the field (natural conditions) animals may hibernate, without waking, for periods uP to at least 3 weeks in duration. During a period of hibernation an animal has no water intake, but water loss still occurs. Although water loss is probably slight during hibernation, 3 weeks is a long period to be without
water intake.
Because of the ability of hibernators to tolerate negative water balance for extended periods, perhaps hibernators demonstrate quantitative differences in water metabolism, especially tolerance to negative water balance. If such is the case, the capacity of hibernators to tolerate negative water balance may not be restricted to periods of hibernation, but would be demonstrable during periods of activity of the animals. This hypothesis has been tested and found true by studying responses to negative water balance in nonhibernating animals (laboratory rats) as controls versus animals capable of hibernation (Citel' /zzs sp.) (Riedesel, Klinestiver, and Benaliy, 1964). In subsequent studies (Bintz, 1965; Bintz and Riedesel, 1967) laboratory rats and ground squirrels were deprived of drinking water unti\ 30/o of original body weight had been lost, after which the water content of various tissues was determined. All rat tissues analyzed were dehydrated.

In ground

squirrels, blood only n'as dehydrated, liver had an in-

creased water content, and kidney, heart, lung, and muscle had no change in rvater content. Also, employing weight loss as a criterion

of tolerance, we found that under various conditions (temperature,


'Srpport"a in part by

.'vt,

L . K.'eciegE I l*-ln*i-a-tt' l.f

U.S. Atomic Energy Commission Contract AT-(29'2)-1629.

46

PHYSIOLOGICALSYSTEMS

season, food allotment) ground squirrels were more tolerant to drink-

water requirements in this species. Water requirements cannot be determined simply by measuring rhe animal's ad-lib. consumprion of water in the laboratory. Many animals have a tendency to overhydrate under laboratory conditions (Hudson, 1962; Chew, lg65). In this study a water-soluble isotope, 22Na, was employed to study the water requirements of C. lateralis. During the study the following assumptions were made: (i) excretion of 22Na is directly proportional to water intake and turnover, and (ii) when water requirements of the animal are satisfied, rate of excretion of 22Na will be maximal. It is becoming well known that laboratory-acclimated animals often differ physiologically from rhe same individuals under natural conditions. For example, Young and Riedesel (I967) demonstrated that laboratory-acclimated C. lateralis tend to be more homeothermic than the same animals living under natural conditions. Because of the tendency of animals to overhydrate in a laboratory environment, perhaps the water requirements of laboratory-acclimated animals are difierent from the water requirements of animals living under natural conditions where water is not always readily available. This hypothesis was tested by comparing rhe rates of excretion of 22Na in recently caught and in laboratory-acclimated C. lateralis when all animals were allowed equivalent amounts of water daily.

ing-water deprivation than were laboratory rats. Following experimenrs in studying rolerance of Citellus lateralis to drinking-water deprivation, we became interested in studying

Marrnrarsexo MBrHoos
Animals. Specimens of C. lateralis were captured in Sherman traps in the mountains of northern New Mexico. Animals were dusted with l/o rotenone powder shortly after capture. During the time interval between capture and return to. the laboratory, animals were
fed raw carrot and apple.

IJpon return to the laboratory and until the time of experimentation, animals to be studied as laboratory-acclimated animals were individually caged and allowed free access to drinking water and
Wayne Lab Blox. Acclimation to laboratory conditions (temperature 22-+2 C; relative humidity 30-+5Tot photoperiod, light from 7 err to 7 rlc) was for 6 to 9 months. Animals to be studied as field-adapted animals were individually

SODIUM.22

RETENTION

47

the period prior to ."p"rirrr.rrtution. Experimentation began rvithin I week of the date that animals were brought into the laboratory, thus acclimation ro laborarory conditions (especialry drinking lvater ad lib.) was minimized.
Pre.exP-erimental procedure. one week prior to experimentation a]r animals were weighecr and transferred to metabolic cages located in an environmental chamber (temperature 2r -+0.5 c; relaiive humidity and photoperiod as above). Bohy weights of animals ranged from 170 to 390 g. Feces and urine producecl-were weighed daily through-

dyliig

caged. Food and water consisted of Wayne Lab Blox and raw potato. Fach animal was given daily l0 g of porato per 100 g of body weight

out the experiment for ail animals. Ad_lib. drinking water consumplaboratory-acclimatecl animars was derermi.,Ja auity by weighlio"-bI ing drinking-warer bottles.
Experimental procedure..Animals were given a single ip injection of 22fa in Ringer's solution. Specific acivity of the" r2Na was 0.30

,.:/"tJ. Each injectior y^: O.Z miTtOO g body weight. As soon as possi_ ble after injection whole-body radioaciivity was counred with a packard Auto-Gamma model 410 A single-channel spectrometer and an Armac deep-well, exrernal liquid scintillator model 440. For each radioactivity was counted at least once daily untit b\/o of 1ni1al the isotope was excreted. Experimental animals were dividea'into the following groups:
Laboratory-acclimated animals group a ("-6), drinking water ad lib. group b (n-8), no drinking water or potaro group c ("-8), b ml rvater/ 100 g body weight per day group d (n-8), l0 ml water/100 g body weighi per day Field-adapted animals group e (n-8), no drinking water or porato group f (n-8), 5 ml warer/100 g bocly weight per day group S (n-8),warer from poraro ad lib.

In all groups where b or l0.ml water/100 g body weight per day were allowed, the water was given in raw potato. some animars did
not eat all the potaro provided. Only animals that readily ate rhe potato were included in the experiment.

48

PH''SIOLOGICALSYSTEMS

Rnsurrs Laboratory-acclimated, animals. Mean biological half-life values (To) and standard errors for four groups (a, b, c, d) of Citellus lateralis are presented in the upper part of Table l. In all four experimental groups excretion of the isotope was rapid during the first 24 hr following injection. Mean values ranged from l6 to 22/o of total isotope injected Fig. t). Animals of the three groups that had nrater available
C, LATERALIS
22

z
F F UJ (r F t
trJ
UJ

z u
z.
C)

90
80

Temp.=

, sinqle i.p. lnjeclion 2lt o.5"c, Rel Humid = 30l 57' Photoperiod, Light 7A.M.-7P.M.
No

o= Lob Acclimoted, Drinking Woter od lib , n=5 O: , lOml HaOllOOg BodyWeight/Dov, n=8 A: ,5mlHaO/lOOqBodyWeiqht/Dov'n=8 a: , No Drinking Woler, n = 8

(L

z
U

24

48
HOURS

72 ,

96

t20

t44

POST INJECTION

Ftcunr l Nlean percentage retention of injected "Na by laboratory-acclimated Citellus lateralis allowed different amounts of water'

continued to lose isotope rapidly after the first 24 hr (Fig. l)' In fact, rate of excretion of the isotope was nearly linear with respect to time, at least until biological half-life was reached' On the other
hand, rate of 22Na excretion in animals with no drinking water available was considerably less afrer the first 24 hr than during the first
24

hr following injection. Statistically the biological half-life for those animals allowed no drinking water difiered from the biological half-life values for those animals allowed 5 ml or more of water/100 g body weight per day (p 4 0.01). None of the biological half-life values for animals of the three groups given water differed from each other.

SODIUAI.22

RETENTION

49

L'ield-adapted animals. Rate of excretion of the isotope in the two groups of animals allowed 5 ml water/100 g body weight per day (group f) and potaro ad lib. (group g) was rapid during the first 24 hr following injection. Mean values ranged from 17 to 20/o of the total isotope injected Fig. 2). Rate of isotope excretion continued
C LATERALIS
?2No

, Single i.p. lnj.ction

, Tcmp..

21

0.5 .

C .

Rol.Homid..30: 5%, Pholoperiod, Lighl 7A.lr.-7PM.

z
o
F-

90
80

oeFiald Anlmols, NoDrinkihg Wotcr, n. 4 , 5ml H2O /lOO9 Body W.ighl/Doy, n.8

F t! F

E,

z
c

U
ru

o
E.

70

^____-'r_
60

z
u

,)---1 ,) 240 336


HOURS, POST INJECTION

Frcunr 2. Mean percentage rerention of injecred ",Na by field-adapted Cileilus lateralis allowed different amounts of water.

essentially in a linear fashion with respect to time after the first 24 hr following injection and ar least until biological half-life values were reached (Fig. 2). Biological half-life values are given in the lower part of Table l Field-adapted animals of group e were allowed no drinking water
and biological half-life was reached by only one animal. Three of the eight animals died before reaching the biological half-life. Isorope excretion by animals in this group was slow during the first 24 hr following injection and conrinued to be very slow until 836 hr post-

injection, at which time four of the animals were sacrificed because it appeared that they would die before reaching the biological halflife. Statistically the amount of isotope excreted by animals that were allowed no water was different borh at 24 hr (P40.01) and at 48 hr

50

PHYSIOLOGICAL SYSTEMS

WATER CONSUMPTION AND BIOLOGICAL HALF-LIFEVALUES (Tb) OF PNA IN LABORATORY-ACCLIMATED ANIMALS AND IN FIELD-ADAPTED ANIMALS

TABLE I

--:=
a

Group

Water intake in ml per 100 gbodY

To in hr

Mean body

weight per day

Mean-+SE

Range

weightl
98 76 93 98

b
c

Laboratory-acclimated animals 57-r44 83+ 11.8 Unlimited r08-272 169+-14.7 None 64-t46 I07 -F I2.0 5 ml in raw potato 69-128 84-+6.9 l0 ml in raw potato Field-adapted animals 68-96 81-+-3.02 5 ml in raw potato

Potato ad lib.
of injection.

80-F3.02 as percentage

65-93

100

of body weight at time

(f a0.0l) from the amount of isotope excreted by the animals in both groups allowed water. Biological half-life values of isotoPe in the two groups of animals allowed water were not different (P>0.8)'
Fietd-adapted us. Iaboratory-acclimated animals. With respect to the rate of excretion of 22Na, laboratory-acclimated animals demonstrated a greater variability from animal to animal (within the same group) than did the field-adapted animals. In the two groups of animals allowed no water, excretion of the isotope during the first 24 hr following injection was rapid (22/" of total isotope) in the laboratoryacclimated animals and slow (7f" of total isotope) in the field-adapted animals. Statistically biological half-tife values of the isotope in fieldadapted and laboratory-acclimated animals allowed water ad lib. did not differ (P>0.7). Also, biological half-life values of 22Na in fieldadapted and laboratory-acclimated animals allowed 5 ml of water/ 100 g body weight per day did not difier (P20.05). The lack of statistical difference in the latter comparison was probably due primarily to variability of rate of isotope excretion within each group, expecially in laboratory-acclimated animals.
DrscussroN

In studies concerned with

water intake of animals, one must be

SODIUM.22

RETENTION

51

food.

In other words, the animals were obtaining about 1.5 ml of free water for every 20 g of food eaten. Metaboric water may be greater in amount than free warer. Lab Blox has a composition of afproxi_ mately 24/o pr-otein, 4/o fat, b/o crude fiber, antl up to l0fo ash. Assuming that the remaining b\fo is carbohydrate, metabolic'water gained from oxidation of 20 g of Lab Blox to carbon dioxide and water would be approximatery 8 ml, with an intake of 20 g of food per day, an animal would be obtaining approximately l0 miof water from food alone. Those animars alrowed no water ieither drinking or potato) ate very little, if any, food after the first 2 to 3 days follow_ ing water deprivation.
Groups of laboratory animals allowed drinking water ad lib. or allowed l0 ml warer/100 g body weight per day excrered 22Na ar similar rates (To-83 and 84 hr, respectively), whereas animals al_ Iowed 5 ml water/100 g body weight per clayexcrered isorope some_ what more slowly (Tb-107 hr). It *o,rtd seem, therefore, that in the case of laboratory-acclimared animals, l0 ml water/100 g body weight Pfr-day is not limiting and may approximate the water"requirements Groups

aware not only of the drinking water but also of water in the animal's In the present study, animars alrowed at reast some water ate l5 to 20 g of Lab Blox per day. Lab Blox has a warer contenr of 7_g/..

of these animals.

weight per day approximates the drinking_warer reqriremJnts. It is interesting that the maximum rate of e*cietion of the isotope was very similar in laboratory-acclimated animals alrowed drinking water ad lib. or allowed l0 ml warer/ 100 g body weight per day und in n"ld_ adapted animals allowed porato ad lib. o, utt*.d 5 mi water/I00 g body weight per day, even though several laboratory_acclimated ani rnals drank over 100 ml of warer per day. Apparently, above a mini_ mal-required water intake, Na excretion ii-independent of water excretion. Although laboratory-acclimated animals have a high water intake, the osmotic control mechanism keeps Na excretion similar to the rate observed in field-adapted animals. under conditions of no water, field-adapted animars were better able to conserve body sodium than were raboratory-acclimated ani-

of recently trapped animals allowed water ad lib. and those allowed 5 ml water/100 g body weight per day excreted the isotope at similar rares, as biological half-rie oil,r., were g0 and gl hr,.respectively. In the case of field animals, b ml water/100 g body

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