Inhibitory centers in sexual behavior in the male rat.

Small lesions placed near the diencephalic, mesencephalic junction, in either the lateral or medial mammillary region, resulted in an increase of copulatory behavior. This increase was expressed both by increased numbers of copulation plugs formed per 14-day interval and by increased percentage of days on which copulation occurred. Inhibitory structures thus form an essential part of the circuitry involved in mediation of sex behavior in the male.

Intracranial sites regulating the biphasic action of progesterone in estrogen-primed golden hamsters.

Diencephalic and mesencephalic neural sites regulating the biphasic effect of progesterone (P) were investigated using the hormone implantation technique in ovariectomized female golden hamsters primed with estrogen. Double barreled cannulae were implanted unilaterally and bilaterally in the medial preoptic area, anterior hypothalamus, ventromedial hypothalamus (VMH), central gray, or interpeduncular nucleus. Testing was conducted using a sequential paradigm; facilitation tests commenced after 44 h of estrogen priming. P-filled cannulae placed in the VMH region facilitated lordosis behavior in 42% and 60% of unilaterally and bilaterally implanted females, respectively. In the anterior hypothalamus, only P implants adjacent to the VMH area effectively promoted receptivity. Lordosis behavior was also observed in 20-36% of females with P implants in the medial preoptic area. P implants in central gray and interpeduncular nucleus regions had no significant facilitating effect on sexual behavior. Tests for inhibition occurred 24 h after facilitation testing and consisted of a pretest, followed by systemic P administration and a behavioral test 4-5 h later. During the pretest for inhibition, females that were receptive in the facilitation test attacked males more rapidly than previously nonreceptive animals and showed decrements in lordosis scores after systemic P delivery. This biphasic effect of P completely inhibited receptivity among several animals in the VMH group. Additional experiments, however, investigating the biphasic effect of P implants in the VMH suggested that the occurrence of copulation in the facilitation test may have been involved in mediating the subsequent increase in aggressive behavior and the suppression of sexual responsiveness in the inhibition test. Nevertheless, a final experiment showed that when P was implanted sequentially in the VMH, facilitation and, more importantly, a later reduction in lordosis behavioral scores occurred even when copulation was eliminated in the facilitation test. P implants in mesencephalic regions exerted no significant inhibitory effect on receptivity. These findings demonstrate that the biphasic action of P in the female hamster is regulated by nerve cells located in the diencephalon, especially in the VMH region.

Diencephalic sites of progesterone action for inhibiting aggression and facilitating sexual receptivity in estrogen-primed golden hamsters.

Diencephalic sites of action of progesterone (P) responsible for inhibiting aggression and facilitating sexual receptivity were examined in ovariectomized golden hamsters primed with Silastic capsules of estradiol. P was applied centrally by inserting a hormone-filled, 27-gauge cannula into a 22-gauge guide cannula that was implanted unilaterally in the medial preoptic area (MPO), the anterior hypothalamus (AH), or the ventromedial hypothalamus (VMH). Control implants consisted of cholesterol-filled cannulae placed within the same regions of the brain. Tests for sexual and aggressive behavior occurred 1, 2, 4, and 6 h after hormone implantation by introducing a sexually experienced male into the home cage of the female. Nine of 20 females with P in the VMH exhibited lordosis in comparison to 1 of 12 females in the MPO group and 5 of 16 animals in the AH group. The induction of sexual responsiveness after P implantation in the VMH was further demonstrated in 6 of 11 ovariectomized-adrenalectomized females, indicating that the observed receptivity was not contingent upon activation of the hypothalamic-pituitary-adrenal axis. In addition to the receptive promoting action of P in the VMH, P implantation in the MPO and VMH but not in AH regions was highly effective in inhibiting female biting attacks upon males. In summary, these findings indicate that P can both facilitate sexual receptivity and inhibit aggressive behavior and that P induces these changes in behavior at different locations in the diencephalon.

Dexamethasone: increased weights and decreased [3H] estradiol retention of uterus, vagina and pituitary in the ovariectomized rat.

Ovariectomized rats given 100 mug dexamethasone per day for 5 days had significantly heavier dry weights for uterus, vagina and pituitary, indicating a growth promoting activity of dexamethasone on these tissues in which estrogen normally promotes growth changes. The dexamethasone treated animals also retained significantly less [3H]estradiol per mug dry weight of tissue for uterus, vagina and pituitary. When[3H]estradiol retention was examined in vitro for the nuclear fraction, a significant decrease in retention was found for uterus, vagina and pituitary but not for hypothalamus or cerebral cortex. The decreased ability to bind [3H]estradiol, shown by the estrogen target tissues of the dexamethasone-treated rats, along with the increased growth of the estrogen target tissues, demonstrates that these tissues were able to show trophic responses even when greater levels were one-third of normal. Dexamethasone-treated animals tested for sexual receptivity in the presence or absence of progesterone priming did not show induction of facilitation of sexual receptivity. However, estrogen plus progesterone injections induced sexual receptivity in the presence of dexamethasone. When dexamethasone was combined with a dosage of estrogen, which by itself did not induce sexual receptivity, there was a significant response with 6 to 10 animals showing a low level of receptivity. Thus, dexamethasone can apparently synergize with estrogen to facilitate sexual receptivity.

In vivo progesterone treatment enhances [3H]estradiol retention by neural tissue of the female rat.

[3H]Estradiol retention was examined for neural tissues of ovariectomized rats as a function of various progesterone pretreatments. Short-term progesterone pretreatment (6 or 24 h) with a 50 mg pellet of progesteron sc resulted in increased in vivo retention of [3H]estradiol when measured at 6 or 24 h following withdrawal of the progesterone source. This increase was greatest in arcuate-median eminence tissue. A less pronounced increase was seen in the preoptic-anterior hypothalamus and cerebral cortex, and no increase was seen in the amygdala or mammilary bodies. Following long term progesterone pretreatment increased [3H]estradiol retention was observed in vivo at 48, 72, 96 or 120 h in the continued presence of the progesterone source for arcuate-median eminence, preoptic-anterior hypothalamus, amygdala and cerebral cortex. Also, when progesterone pretreatment was 24 h, [3H]estradiol retention was still increased at 36, 48, 60 and 72 h following removal of the progesterone source for arcuate-median eminence, preoptic-anterior hypothalamus, amygdala and cerebral cortex. Our demonstration that progesterone pretreatment can significantly increase [3H]estradiol retention by neural tissue suggests a possible mechanism by which progesterone can regulate the timing of ovulation and sex behavior. Furthermore, our observations are in keeping with the finding that progesterone has little or no effect at the organismal or tissue level unless estrogen is present.

Cyclic Djungarian hamsters, Phodopus campbelli, lack the progesterone surge normally associated with ovulation and behavioral receptivity.

Serum levels of 17 beta-estradiol and progesterone were assayed at 4- to 5-h intervals across the estrous cycle in female Djungarian hamsters, Phodopus campbelli. The pattern of secretion for estradiol is similar to that described for the rat and the Syrian hamster, rising slowly from a baseline of 54 +/- 2 pg/ml during the morning of estrus to a peak of 187 +/- 16 pg/ml in the early afternoon of proestrus, then falling rapidly back to baseline levels. The pattern of progesterone secretion is significantly different from that of any estrous cycle previously described for a 4-day cyclic mammal. There is no evidence for a significant peak in serum progesterone levels associated with ovulation and receptivity. The highest levels of serum progesterone occur during the early afternoon of diestrous day 2 (8046 +/- 1063 pg/ml). The lowest levels of serum progesterone occur in the early morning of proestrus (720 +/- 125 pg/ml). During the period of sexual receptivity only 2504 +/- 654 pg/ml progesterone are found in the serum. Ovariectomized females show behavioral receptivity after 48 h of treatment with 50 micrograms estradiol benzoate/kg BW in sesame oil. Progesterone is not necessary for receptivity and will not facilitate receptivity when administered in conjunction with a subthreshold dose of estrogen. The presence of tubular ova confirms the time of ovulation to be a period of approximately 7 h between 2200 h on proestrus and 0500 h on estrus. Histological examination of ovarian sections from each of the 4 days of the estrous cycle shows follicular development to differ from that in the Syrian hamster in that the total number of follicles in the ovary is variable and low, and the ovulated follicles (new corpora lutea) are blood filled. Djungarian hamster primary follicles averaged 83 +/- 2 microns in diameter, and preovulatory follicles averaged 455 +/- 17 microns in diameter. The total number of intact follicles per ovary rose from 21 +/- 5 on the evening of estrus to 73 +/- 29 on the evening of diestrous day 2, then fell to 32 +/- 26 by the evening of proestrus. Preovulatory follicles increased in number from 1.0 +/- 0.6/ovary early on estrus to a maximum of 4 +/- 1 on the evening of diestrous day 2. However, at ovulation a combined total of only 5.1 +/- 0.8 ova are shed.

Relative contributions of oestradiol and progesterone to the maintenance of sexual receptivity in mated female hamsters.

Sexual receptivity was measured as total lordosis duration (TLD) per 10 min test period with mating tests standardized across 24 h by giving the first test at the time when the lights were switched off (0 h) during a schedule of 14 h light : 10 h hardness and re-testing at intervals of 2 h. Long-term (14 days), high-level oestrogen treatment was necessary to achieve TLD scores similar to those found in female hamsters treated with oestrogen plus progesterone for 3 days. Mating of female animals treated with oestrogen only always resulted in a significant decrease in TLD scores at later tests (0 v. 2 h). If progesterone was given after the test at 0 h, a significant increase in TLD score was found at the test at 2 h and this increase was maintained across several tests. By 8 h TLD scores had decreased significantly in progesterone-treated mated female hamsters. Progesterone appeared to have, therefore, at least two roles in the regulation of sexual responsiveness in the female hamster: (a) under normal conditions of oestrogen exposure (1--3 days) progesterone was necessary for the induction of sexual receptivity and (b) progesterone was necessary for the maintenance of sexual responsiveness following short-term (10 min) mating in female hamsters treated with oestrogen only. In the oestrogen- and progesterone-treated female animal which is no longer receptive as a result of previous exposure to progesterone plus having been mated, sexual receptivity can be re-induced by treatment with a larger dose of progesterone. These findings have indicated that progesterone is the primary hormonal agent which regulates sexual receptivity in the mated female hamster. Furthermore, the eventual inhibitory effect of progesterone on sexual receptivity was not absolute even in the mated female animal but rather represent some form of habituation to that particular dosage of progesterone to which the animal had been previously exposed.

Role of the hypothalamic paraventricular nucleus in mediating stress-related litter deficits in the golden hamster.

Abstract Primiparous female hamsters (Mesocricetus auratus) were mated to proven breeders and stressed during early pregnancy. Females were housed singly throughout gestation except for Days 4, 5 and 6 when they were paired for ten-min intervals three times each day with another female matched for age, weight and day of pregnancy. Within each of the pairs, one female was consistently dominant to the other. Controls were exposed to a novel area instead of a conspecific. At parturition, all pups were counted, sexed and weighed. There were no significant differences between control and dominant females' litter sizes or sex ratios (defined as percentage male). Subordinate females produced significantly smaller litters than control or dominant dams and significantly lower sex ratios than control dams. Subordinates produced fewer males than control or dominant dams, but there were no differences in the number of females produced. The paraventricular nucleus (PVN) of the hypothalamus has been implicated as a relay center for the physiological response to stress. Bilateral lesions of the PVN were performed on another group of females and the same protocol described above was followed after the females recovered from surgery. When this 'stress relay center' was lesioned, subordinates did not show the significant deficits in litter size and sex ratio. Sham-operated females showed a similar response to social stress to that of intact females. These results suggest that subordinate dams produce smaller litters via selective resorption of males in utero and that the PVN may be a relay center for the mediation of this response.

Localization of hypothalamic sites for the estrogen-priming of sexual receptivity in female hamsters.

Ovariectomized female hamsters received small unilateral implants of estradiol at a variety of anterior-posterior levels of the medial preoptic area and hypothalamus. The results of an initial experiment using 27-ga. implants showed that females with estradiol implants in the ventromedial hypothalamus (VMN) or nearby anterior hypothalamus consistently showed higher levels of sexual receptivity than did females with implants farther rostral, in the preoptic area, or farther caudal, in the posterior hypothalamus. A second experiment used smaller, 28-ga. implants to compare directly the two areas at which implants were effective in the first experiment. The results confirm the findings of other recent studies of hamsters and rats by identifying the VMN as the most effective hypothalamic site for the estrogen priming of sexual receptivity.

Diencephalic organization of estradiol sensitive sites regulating sociosexual behavior in female golden hamsters: contralateral versus ipsilateral activation.

Dual ipsilateral, contralateral and bilateral 28-gauge estradiol (E2) filled cannulae were implanted in the medial preoptic area (MPO) and ventromedial hypothalamus (VMH) of ovariectomized female golden hamsters housed in large arenas with male partners. Twenty-four hours after implantation, vaginal scent-marking patterns were significantly and equally elevated in all groups. Forty-four hours after implantation, progesterone was administered and females were tested for sexual receptivity 4-5 h later. Bilateral E2 implants in the VMH as well as dual ipsilateral and contralateral MPO-VMH implants were significantly more likely to facilitate sexual responsiveness than bilateral MPO implants. More importantly, ipsilateral MPO-VMH implants produced significantly longer lordosis duration scores than bilateral VMH and contralateral MPO-VMH implants. After mating, females with bilateral MPO implants attacked their mates more frequently than females with bilateral VMH and dual MPO-VMH implants. Taken together, results suggest that: (1) although MPO and VMH regions are equally sensitive to the vaginal marking promoting effects of E2, these same regions require synergistic ipsilateral activation for the effective priming of sexual responsiveness; (2) the heightened duration of lordosis behavior after ipsilateral MPO-VMH E2 implantation may reflect an anterior diencephalic estrogenic removal of an inhibitory process occurring primarily in the ipsilateral VMH region; and (3) the difference in postcopulatory attacks may reflect variable actions of progesterone on E2-induced progestin receptors in the MPO and VMH.

Dual estradiol action in diencephalon and the regulation of sociosexual behavior in female golden hamsters.

Previously, we found that single implants of estradiol (E2) placed in the ventromedial hypothalamus (VMH) but not the anterior hypothalamus (AH) facilitated precopulatory, i.e. vaginal scent marking, and copulatory, i.e. lordosis, behavior following progesterone administration. However, the duration of lordosis was markedly attenuated in comparison to the duration shown by intact cycling females. This study was designed to examine whether dual implantation of E2 in the diencephalon would facilitate patterns of precopulatory and copulatory behaviors similar to those shown by intact cycling females. One E2 implant was placed in either the medial preoptic area (MPO) or AH and a second E2 implant was placed in the VMH. Control females were tested following E2 application at only the MPO or AH region in conjunction with a cholesterol implant in the VMH. An additional control group was tested with females implanted with cholesterol at both MPO-AH regions and the VMH. During a 2-day postimplantation test period, vaginal marking scores were elevated for both single and dual E2 implanted females and agonistic response patterns toward males declined significantly. In addition, a significant inverse relationship was found between the number of vaginal marks and bites exhibited by females with single E2 implants in the MPO, whereas these two response patterns were positively correlated in females with E2 stimulation occurring only in the AH region. No significant relationship was found between vaginal marking and biting attack for females receiving dual E2 stimulation. Systemic progesterone administration on the third postimplantation day facilitated sexual receptivity in the majority of females with dual E2 implants (greater than 90%). These receptive females displayed lordotic responsiveness that closely matches the full display of sexual receptivity shown by intact cycling females. In contrast, only one female with a single E2 implant in the AH region showed sexual responsiveness. The results demonstrate that: precopulatory vaginal marking and biting attack are mediated by E2 action in the MPO and AH but in a different manner; additional action of E2 in the VMH diminishes the distinctive precopulatory behavioral effects of E2 in the MPO and AH suggesting an influential role of the VMH in regulating sociosexual activities; and E2 action in either the MPO or AH region in conjunction with E2 action in the VMH may be necessary in order to facilitate the species-typical display of lordotic responsiveness.

Effects of sex and reproductive state on acoustic responsiveness in hamsters.

Semi-quantitative [14C]2-deoxyglucose (2DG) autoradiography was used to describe the responses of hamsters to 35 kHz mimics of the "ultrasounds" used for communication during mating. The first study examined the processing of ultrasounds and ambient noise by estrous females, some of which were deafened or hemideafened with plastic ear plugs. These data failed to reveal responses specific to the ultrasounds. However, lateralized responses to the ambient noise were apparent, especially in the hemideafened subjects. For the ventral cochlear nucleus (VCN), 2DG uptake was elevated contralateral to the plug and ipsilateral to the effective stimulus. In contrast, uptake by more rostral structures (dorsal n. of the lateral lemniscus = DNLL; ventral n. of the lateral lemniscus = VNLL; central n. of the inferior colliculus = CIC; medial geniculate n.) was elevated contralateral to the stimulus. A second experiment examined the responses of intact or castrated male and female hamsters to unilaterally presented ultrasounds and ambient noise. As before, relative levels of 2DG uptake differed across hemispheres for structures including the VCN, trapezoid body, VNLL, DNLL, and CIC. More surprisingly, intact females showed more 2DG uptake than males in the DNLL, auditory nerve, and lateral lemniscus. Females also tended to show elevated anterior hypothalamic uptake, but just in the hemisphere contralateral to the stimulus. These results suggest that male and female hamsters differ in acoustic responsiveness, and that this difference is mediated by hormonal effects at several brainstem components of the central auditory system.

Patterns of 2-deoxyglucose uptake reflect the neural processing of lordosis-inducing somatosensory stimuli in hamsters.

Semi-quantitative [14C]2-deoxyglucose (2DG) autoradiography was used to map the neural responses of female hamsters to lordosis-inducing flank stimuli. Specifically, manual stimulation of one flank was used to maintain estrous females in lordosis for 20 min after an IV injection of 200 muCi/kg of 2DG. Hemispheric differences in 2DG uptake then were sought in brain nuclei implicated in the programming of lordosis, or in the mediation of somatosensory or hormonal influences on this response. The responses to lateralized flank stimulation included reliable contralateral elevations in 2DG uptake in the ventral posterior lateral nucleus of the thalamus (VPL), the dorsal mesencephalic central gray (dCG), and the tectum. Elevated activity on the part of the VPL may not be crucial for lordosis. However, the effects of flank stimulation on 2DG uptake by the dCG and tectum confirm and extend much previous evidence implicating the dorsal midbrain in the mediation of tactile and hormonal effects on sexual responses. For example, these results suggest that somatosensory influences on hamster lordosis are mediated by both the dCG and tectum. In addition, they suggest that these influences are strongly lateralized until at least this stage of sensory processing, leaving for some subsequent element of neural circuitry the task of translating these lateralized inputs into the bilaterally symmetric outputs ultimately required to program the normal, bilaterally symmetric, lordosis response.

Dual progesterone action in diencephalon facilitates the induction of sexual receptivity in estrogen-primed golden hamsters.

In order to identify and characterize the progesterone (P) sensitive neural system that regulates feminine sexual behavior, 28-gauge P-filled cannulae were implanted in the medial preoptic area (MPO), ventromedial hypothalamus (VMH), and central gray (CG) of ovariectomized estrogen-primed golden hamsters. Dual implants of P were placed either ipsilaterally or contralaterally in brain sites consisting of MPO-VMH, MPO-CG, or VMH-CG combinations. Tests for sexual receptivity commenced 44 hr after estrogen priming and consisted of a preimplantation test followed 4.5 to 5.5 hr later by a postimplantation test. In the preimplantation test, stimulus males were attacked when placed into the female's home cage which indicated that the subsequent display of sexual receptivity occurring in the postimplantation test was due to the action of P. Dual implants of P placed either ipsilaterally or contralaterally in MPO-VMH regions were significantly more effective in facilitating lordosis behavior than dual P implants placed in MPO-CG or VMH-CG regions. However, the duration of lordotic responsiveness produced by dual P implants in MPO and VMH regions appears to be shorter than the duration of lordosis typically observed in intact females on proestrus. Results suggest that MPO and VMH regions are sensitive to the lordosis facilitating actions of small dual implants of P.

Organization and expression of agonistic and socio-sexual behavior in golden hamsters over the estrous cycle and after ovariectomy.

Pairs of hamsters were housed in large enclosures that contained separate male and female living areas and observed over the 4-day estrous cycle and after ovariectomy. Agonistic elements exhibited frequently by females included on-back, boxing, lateral posturing, and biting, whereas males engaged frequently in boxing and on-back patterns of behavior. Furthermore, on-back and boxing by females were significantly higher on estrus than on any other day of the estrous cycle. Agonistic acts performed after ovariectomy did not differ in occurrence from those shown by animals on diestrus and proestrus. Vaginal marking increased during diestrus and attained a peak 24 hr prior to sexual receptivity. Both vaginal marking and mating occurred more frequently in the female's than male's home area suggesting that vaginal marking and mating occurred serve to attract males to the nest of females. Males also organized their marking patterns by location as shown by more flank marking in their own than their partner's area, albeit the significance for this difference in location is not known. The results demonstrate that when heterosexual pairs of hamsters are tested in large and partially familiar habitats, a wide range of behavior is exhibited and organized in a manner that is not observed in small and unfamiliar cages.

Effects of progesterone on lordotic responses to specific mating stimuli in hamsters.

Mating-induced inhibition of sexual receptivity was examined in ovariectomized, estrogen (E) treated and estrogen plus progesterone (E + P) treated hamsters given 10 min of exposure to male mounting stimulation alone or to mounts, intromissions, and ejaculations at eight hourly intervals. In E + P treated females, no differential effects of exposure to full mating stimulation vs. mounting stimulation alone were observed. In contrast, females given E treatment alone showed a marked differential response. Fully mated, E-treated females showed more lordosis than E-treated females exposed to mounts alone during the initial test. However, total lordosis duration declined precipitiously in the fully mated group by 2 hr and remained significantly below that in other groups during subsequent tests. Levels of receptivity in E-treated females mounted-only remained relatively constant until 8 hr. These results suggest that P reduces the inhibitory effects of vaginocervical cues received during mating without affecting the response to mounting stimulation alone. In addition, vaginocervical stimulation may initially facilitate lordosis in E-treated females.

Estrogen action in anterior and ventromedial hypothalamus and the modulation of heterosexual behavior in female golden hamsters.

Changes in heterosexual patterns of agonistic, marking, and sexual behavior were examined in female hamsters over a three day period following implantation of either estrogen or cholesterol in the hypothalamus. Animals were habituated to large arenas that permit the display of a wide range of behavior. Estrogen implants located in the ventromedial (VMH) but not the anterior (AH) portion of the hypothalamus were effective in facilitating the occurrence of vaginal marking, over a two day estrogen priming period. During this two day period, the exhibition of agonistic behavioral patterns declined significantly. Systemic administration of progesterone elicited sexual receptivity in 75% of the females in the VMH group, in contrast to only 25% in the AH group. Females with hypothalamic implants of cholesterol remained unreceptive following progesterone injections. The results provide important information on the estrogen sites of action at the hypothalamus in mediating heterosexual interactions.

Changes in responsiveness to newborn pups in pregnant, nulliparous golden hamsters.

Virgin female hamsters were mated and tested once daily for maternal retrieving behavior beginning on days 0, 5, 9, 13, 15, of the 16 day gestation period to determine if responsiveness toward newborn pups changes as pregnancy proceeds. Upon initial exposure to 3 newborn pups, only a small percentage of early-to-mid-pregnant females exhibited maternal retrieving behavior spontaneously. In contrast, over half of the 15 day pregnant females displayed retrieving during the first test. Despite the high frequency of initial pup-directed aggression and cannibalism, maternal retrieval was induced in the majority of the females in all groups by repeated daily exposure to 3 newborn pups. However, no significant differences were observed in the number of pup exposure periods required to induce maternal retrieving in 0, 5, and 9 day pregnant females. It is concluded that the high level of maternal responsiveness observed in the parturient hamster develops somewhat abruptly during late pregnancy. In this respect, the pattern observed in the hamster differs from the more gradual increase in maternal responsiveness reported in mid-to-late-pregnant mice and rats.

Differential effects of paternal presence on pup survival in two species of dwarf hamster (Phodopus sungorus and Phodopus campbelli).

The effect of the presence or absence of the male, and of decreased ambient temperature (21 degrees C vs. 4 degrees C) on litter survival, pup survival and pup growth was measured from birth through day 18 after birth in Siberian (P. sungorus) and Djungarian (P. campbelli) hamsters. Siberian hamsters were not significantly affected by the experimental manipulations. In contrast, whereas 100% of litters and 95% of pups were successfully raised to weaning at 21 degrees C by paired Djungarian hamsters, survival fell to 47% when the mate was absent and even further, to 32%, when the ambient temperature was lowered. No significant differences in litter size or pup weight at birth were detected between species at the warmer temperature. However, P. sungorus pups gained weight significantly faster through day 12 after birth (while dependence upon the mother for food was absolute) than P. campbelli pups under all experimental conditions. Although the species are closely related, these data show that male Djungarian hamsters are essential to offspring survival under conditions where Siberian hamsters do not require conspecific help. Species differences in metabolism and thermoregulation may account for the differential pup survival.