BEHAVIORALBIOLOGY24, 474-480 (1978)

Saline Intake in Hamsters

RODERICK

WONG

AND

WAYNE

JONES 1

Department of Psychology, University of British Columbia, Vancouver, Canada V6T 1W5

Experiment 1 assessed the preference-aversion function of hamsters for water
and 0.08, 0.15, 0.35, and 0.71 M NaC1 solutions in the two-bottle test. Unlike
rats, the hamsters did not show a strong preference for isotonic (0.15 M) saline,
nor did they exhibit any preference for the other solutions. Experiment 2 examined the effects of hydrochlorothiazide on the hamsters" intake of 0.71 M NaC1
solution. The results indicated that this diuretic agent did not cause the hamsters
to increase their saline intake relative to the baseline level. Experiment 3 also
indicated that a high (5 rag) dose of desoxycorticosterone acetate failed to produce
sodium appetite in the hamsters. These results contrast with those found with rats.
The golden hamster (Mesocricetus auratus ) is a rodent native to Syria
that has adapted to the natural selection pressures of a dry, xerophytic,
desert environment (Schmidt-Nielsen, 1964). Although psychologists
have conducted extensive studies on the agonistic, social, sexual, and
open-field field behavior of members of this species, there are much less
data on the hamster's mode of adaptation to fluid requirements. Experimental data on the determinants of fluid ingestion in hamsters would
enable us to evaluate the generality of models about the regulation of
consumption found in rodents from mesophytic niches. Unfortunately,
most of our knowledge about fluid ingestion in rodents has been derived
from rat experiments.
Pioneer studies on species differences in water regulation had originated in K u t s c h e r ' s (1969) laboratory. His research group had demonstrated that following food deprivation, guinea pigs b e c o m e adipsic, rats
show restricted drinking, and gerbils and hamsters develop polydipsic
behavior. Such results certainly indicate that the rat's physiological and
behavioral systems operate on principles different than those upon which
other rodents operate. In another comparative study, Carpenter (1956)
indicated that hamsters differ from rats, cats, and rabbits in their be1 This research was aided by Grant 67-0247 from the National Research Council of
Canada and by the University of British Columbia Natural, Applied and Health Sciences
Grants Committee. We thank Amyra Carsh for her help in running and analyzingthe data of
Experiment 3.
474
0091-6773/78/0244-0474502,00/0
Copyright @ 1978 by Academic Press, lnc,
All rights of reproduction in any form reserved.

SALINE INTAKE IN HAMSTERS

475

havioral responses to salt solutions. Unlike the other species in Carpenter's experiment, hamsters showed no preference for NaC1 solutions over
water in the two-bottle test. Since the time of CarpenteVs study, numerous experiments have shown that rats exhibit greater preference for
isotonic saline than for any other concentration of the solution or of water
(see Young, 1966). In Carpenter's experiment, the hamsters' intake of
hypotonic saline was equivalent to that of water. When the saline was
increased to 0.2 M, the hamsters drank more water than saline solution.
The animals showed progressively greater intake of water and decreased
intake of saline when the tonicity of the NaC1 solution was increased from
0.2 to 1.0 M.
The Carpenter (1956) results with hamsters are particularly interesting
because their NaC1 preference-aversion function was clearly different
from that typically observed in rats. However, there are a number of
methodological features of Carpenter's experiment that may be problematic. Carpenter took only a single reading of the hamsters' intake of a
number of NaC1 solutions varying in concentration. The results of many
saline preference studies in our laboratory with rats (Wong, 1977a; Wong
and Kraintz, 1977) and gerbils (Wong, 1977b) have indicated that a daily
reading of the animals' intake over a substantial number of sessions
provides a more stable estimate of an animal's preference than that
obtained with a "one-shot" test. We also believe that Carpenter should
have presented and analyzed his data in terms of a saline preference score
(percentage of saline intake relative to total fluid intake) in addition to that
based upon absolute intake.
The present experiment was designed to ascertain the preferenceaversion function of the hamster taking into account the methodological
features previously discussed. Of particular importance is the fact that we
tested the animals with isotonic (0.15 M) NaC1 in addition to testing them
with hypotonic and hypertonic solutions. Under normal conditions, rats
invariably prefer isotonic saline over any other solution (Young, 1966).
Unfortunately, Carpenter did not study the hamster's intake of this solution relative to water.
EXPERIMENT

The subjects were eight male and six female golden hamsters (Mesocdcetus am'atus) ranging in weight from 96 to 130 g at the beginning of the
experiment. They were housed in single stainless-steel cages measuring
17.8 x 24.2 17.8 cm. Purina Lab Chow was placed on the wire mesh
floor instead of in the food hopper on the side of the cage because the
animals had difficulties in biting between the wire mesh wall adjacent to
the hopper.
We placed a bottle of water in each cage and maintained their daily
intake for 10 days. Then we tested the two-bottle preferences of the

476

WONG AND JONES

hamsters by placing one bottle of water and one of saline through the front
of the cage. The left-right position of the bottles was alternated from day
to day, and the daily intake from each bottle was recorded around the
same time (0930 hr). The animals were first tested for 10 days with a bottle
each of water and 0.15 M NaC1. Then they were given water only for the
subsequent 3 days before being tested with a bottle of water and one of
0.08 M NaCi for 10 days. Following another 3 days during which water
was the only source of fluid, the hamsters were given a choice between
water and 0.35 M NaC! for another 10 days. After the 3-day water-only
period, the animals were given 10 days to choose between water and 0.71
M NaC1 solution.
During the initial 10-day period, the hamsters' mean water intake was
9.72 ml. Data from the two-bottle test were first analyzed in terms of a
preference score consisting of (saline intake/saline + water intake) x 100.
This score minimizes the error due to individual variation in total fluid
consumption within and between subjects. The saline preference of the
hamsters over the 10 test days for each NaCI concentration is shown in Fig.
1. Sex differences were not evident (P > 0.05), so the data were pooled.
An examination of the data indicates that, unlike rats, hamsters do not
show a strong preference for 0.15 M NaC1 solution. In contrast to the rats'
tendency to show a saline preference of about 70% (Devenport, 1973;

35

3O
e0O

25
oi _

20

. 0 8'
NaCI

.15~

.3'5

.71'

Molarity

FIG. I. Percentage intake of 0.08, 0.15, 0.35, and 0.71M NaCI solutions by hamsters in
the two-bottle test.

SALINE INTAKE IN HAMSTERS

477

Wong, 1977a), the hamsters showed 37.2% preference for isotonic saline.
This figure is even lower than the 53% saline preference shown by gerbils
in the Wong (1977b) experiment. When the hamsters were tested with 0.08
M (0.45%, w/v) NaC1 solution, they showed 36.5% saline preference.
When the hamsters were tested with 0.35 M (2.0%, w/v) NaC1, their saline
preference was 31%, and when tested with 0.71 M (4.0%, w/v) NaC1, their
salt preference dropped to 24.2%.
A repeated-measures analysis of variance where the mean saline preference for each solution was dealt with as a within-subject variable
indicated significant differences arising from the tonicity of the test solutions [F (3, 39) = 4.36, P < 0.01]. When multiple comparisons were
performed between the pairs of test conditions, the results indicated that
the saline preference of the hamsters differed only when they were tested
under the 0.08 and 0.71 M NaCI solutions (Tukey T = 5.14, P < 0.05).
None of the other pairwise differences was statistically significant.
The data on the hamsters" absolute intake of water and saline were also
recorded and analyzed. Their mean intake of 0.15 M NaC1 during the 10
test days was 4.67 ml in contrast to their intake of 9.75 ml of water. When
offered a choice between 0.08 M NaCI and water, the animals' mean
intake of saline was 4.51 ml and that of water was 7.44 ml. When tested
with 0.35 M NaC1 and water, the hamsters drank 4.45 ml of saline and 9.14
ml of water. However, when exposed to 0.71 M NaCI and water, the
hamsters" mean intake of saline was 3.02 ml and that of water was 8.95 ml.
A repeated-measures analysis of variance of the saline intake indicated a
significant effect due to conditions IF (3, 39) = 4.25, P < 0.05]. Multiple
comparisons of the pairs of test conditions indicated that the hamsters'
intake of 0.71 M NaC1 was significantly less than their intake of 0.08, 0.15
and 0.35 M NaCI solutions (P < 0.05). Analyses of the hamsters' water
intake under the various test conditions did not reveal any significant
differences (P > 0.05).
The present results indicate that hamsters show little preference for
NaCI solutions of varying tonicity. Their intake of hypotonic, isotonic,
and 0.35 M hypertonic NaCI saline is at a similar low level. When offered
a highly hypertonic solution (0.71 M NaC1), hamsters show a significant
decrease in saline preference relative to the other concentrations. In some
respects our results with hypertonic solutions are similar to those found
by Carpenter (1956). However, we did not replicate Carpenter's finding
that hamsters show an equivalent intake of hypotonic saline and water.
These interexperimental differences may be due to the different procedures (i.e., estimation of saline preference from a "one-shot" vs a
repeated-measures assessment). At any rate, the preference-aversion
function for NaC1 among hamsters is clearly different from that among
rats and gerbils.

478

WONG AND JONES

EXPERIMENT 2

The results of Experiment 1 indicate that hamsters show a relatively

low level of acceptance of 0.71 M NaCI under normal, nondeprived
conditions. The present experiment was designed to determine whether
sodium depletion would increase the hamsters' preference for this solution. We believe that this issue is important because of a recent finding by
Salber and Zucker (1974). They reported an absence of salt appetite
among adrenalectomized hamsters. Adrenalectomy produces sodium deficits through a continuous loss of body sodium in urine and other excretions. Thus, we would expect the animal to show a compensatory increase
in available salt.
In the present experiment we injected the hamsters with a diuretic
agent, hydrochtorothiazide, which acts upon the kidneys to produce the
loss of sodium excreted in the urine. This drug has been shown to increase
the spontaneous appetite for NaC1 solution of rats (Fregly, 1967; Fregly
and Kim, 1970). We assumed that with the loss of body sodium, the
hamsters should show a compensatory increase in NaC1 consumption
similar to that shown by rats in the Fregly studies.
With the exception of one female that died between experiments, the
subjects were the same hamsters used in Experiment 1. Eight male and
five female hamsters served in the present experiment. Following 6 days
when their intake of 0.71 M NaC1 and water was assessed, we gave each
hamster daily injections of 3.0 mg of hydrochlorothiazide dissolved in 0.25
ml of water and Tween 80 (Baker) for 3 consecutive days. We recorded
the hamsters' intake of water and 0.71 M NaC1 during this period as well
as during the six postinjection sessions.
During the 6 preinjection days, the mean saline preference score of the
hamsters was 19.0%. During the subsequent 3 days (Days 7-9), there
appeared to be a substantial increase in the hamsters' saline preference ()7
= 31.8%). During the 6 postinjection days (Days 10-15), the hamsters'
saline preference (J = 19.7%) returned to their preinjection baseline
level. A repeated-measures analysis of variance, where the mean saline
preference score during Days 1 to 6, 7 to 9, and 10 to 15 was dealt with as a
within-subject variable, indicated a significant effect IF (2, 24) = 10.96), P
< 0.01].
An examination of the absolute intake of water and saline of the
hamsters during the various phases of the experiment produced the following results. During the 6 preinjection days, the mean saline intake of
the hamsters was 1.97 ml and that of water was 8.55 ml. During Days 7 to
9 the hamsters' mean saline intake was 3.07 ml and that of water was 6.16
ml. During the 6 postinjection days the hamsters showed a mean saline
intake of 2.10 ml in contrast to a mean water intake of 8.49 ml. A
repeated-measures analysis of variance, where the mean saline intake of
the animals during Days 1 to 6, 7 to 9, and 10 to 15 was dealt with as a

SALINE INTAKE IN HAMSTERS

479

within-subject variable, did not yield any significant effects (P > 0.05). A
similar analysis of the animals' water intake during the three phases of the
experiment yielded a significant effect IF (2, 24) = 10.52, P < 0.01].
Multiple comparisons of the pairs of test conditions indicated that the
hamsters drank less water during Days 7 to 9 than they did during the
periods before and after the injections.
The results of Experiment 2 are interesting in that sodium-depleted
hamsters appeared to show an increased preference for hypertonic saline
even though they really did not show a compensatory increase in saline
consumption. The apparent change in saline preference arising from the
treatment was due to an artifact. The hamsters showed a decrease in
water intake during the injection days but maintained their saline intake at
the level they had exhibited during the pre- and postinjection periods. One
might wonder if we had administered the appropriate dose of hydrochlorothiazide to the hamsters. The amount of the diuretic (3 mg/100 g
body weight) used in the present experiment was comparable to the
amount (2 rag/100 g body weight) used in the Wong and Wilson (1973) and
Wot~g and Hardisty (1974) studies that had demonstrated sodium appetite
in rats.
On the basis of these results one may also question whether hamsters
respond to diuretics like rats by showing a significant sodium loss in their
urine. A study in progress (Kraintz et al., 1978) indicates that hamsters
show a significant increase in urine volume but no significant change in
sodium output following diuretic injections. In view of such results, the
absence of sodium appetite in hamsters following diuretic injections is
not surprising.
EXPERIMENT 3
In addition to studying the effects of adrenalectomy on the hamster's
saline intake, Salber and Zucker (1974) also assessed the effects of
mineralcorticoid injections on these animals. They found an absence of
sodium appetite when the animals were injected with 2.5 mg of desoxycorticosterone acetate (DOCA). In the present experiment we assessed the effects of a DOCA injection that was twice the dose used by
Salber and Zucker.
This experiment was conducted 3 months after Experiment 2 had been
completed and the subjects were the 11 surviving members (seven males
and four females). Following 3 days during which their intake of 0.71 M
NaCI and water was assessed, we gave each hamster daily injections of
5.0 mg of DOCA dissolved in 0.25 ml of peanut oil over 3 consecutive
days. We recorded the hamsters' intake of water and 0.71 M NaCI during
this period as well as during the three postinjection sessions.
In general, the results indicated no significant effects of DOCA injections on either the saline preference or the absolute saline intake of the

480

WONG AND JONES

h a m s t e r s ( P > 0.05). T h e s e r e s u l t s s u p p o r t S a l b e r a n d Z u c k e r ' s (1974)

f a i l u r e to p r o d u c e s o d i u m a p p e t i t e in t h e a n i m a l s w i t h a l o w e r d o s e o f
DOCA. These experimenters had suggested that the ancestors of the
present laboratory hamsters had inhabited a desert environment where
salt w a s r e a d i l y a v a i l a b l e . F o r this r e a s o n h a m s t e r s d i d n o t e x p e r i e n c e salt
deficiencies that would select for those who evolve adrenal salt-retaining
c a p a b i l i t i e s . E v e n w h e n d e p l e t e d o f b o d y s o d i u m ( E x p e r i m e n t 2),
h a m s t e r s d o not s h o w t h e a d a p t i v e r e a c t i o n o f s e e k i n g a n d c o n s u m i n g
h y p e r t o n i c saline t h a t is c h a r a c t e r i s t i c o f m o s t m a m m a l s .

REFERENCES
Carpenter, J. A. (1956). Species differences in taste preference. J, Comp. Physiol. Psychol.
49, 139-144.
Devenport, L. D. (1973). Aversion to a palatable saline solution in rats: Interactions of
physiology and experience. J. Comp. Physiol. Psychol. 83, 98-105.
Fregly, M, J. (1967). Effect of hydrochlorothiazide on preference threshold of rats for NaC1
solutions. Proc. Soc. Exp. Biol. Med. 125, 1079-1082.
Fregly, M. J., and Kim, K. J. (1970). Specificity of the sodium-chloride appetite ~ hydrochlorothiazide treated rats. Physiol. Behav. 5, 595-599.
Kraintz, L., Kraintz, P. W., and Wong, R. (1978). "Effects of Diuretics and AldosteroneInhibiting Substances on Urinary Excretions of Hamsters." In preparation.
Kutscher, C. L. (1969). Species differences in the interaction of feeding and drinking. Ann.
N.Y. Acad. Sci. 157, 539-552.
Salber, P., and Zucker, I. (1974). Absence of salt appetite in adrenalectomized and DOCAtreated hamsters. Behav. Biol. 10, 295-311.
Schmidt-Nielsen, K. (1964). "Desert Animals." Oxford: Clarendon Press.
Wong, R. (1977a). Adaptation of long-term saline consumption and later saline preference.
Behav. Biol. 19, 389-393.
Wong, R. (1977b). Saline intake in gerbils (Meriones unguiculatus). Physiol. Psyehol. 5,
225-229.
Wong, R., and Hardisty, R. (1974). Sodiurfi intake induced by aldactazide and formalin
injections. Behav. Biol. 11, 127-130.
Wong, R., and Kraintz, L. (1977). Desalivation and saline ingestion in rats. Behav. Biol. 19,
130-134.
Wong, R., and Wilson, C. S. (1973). Aldactazide-induced sodium appetite in rats. Behav.
Biol. 8, 285-289.
Young, P. T. (1966). Hedonic organization and regulation of behavior. Psychol. Rev. 73,
59-86.