Sexually inexperienced male Wistar rats (strain WI in our colony) known to very infrequently display spontaneous lordosis behavior (Schaeffer et al., 1990b) were used. A first group was tested four times at 5-day intervals for lordosis with vigorous stimulus males (heterotypic sexual behavior), immediately following testing for masculine sexual activity with highly receptive females (homotypic sexual behavior). A small number of animals displayed lordosis during the first test, but more and more animals displayed this behavior from the first to the fourth test. There was no relationship between the degree of masculine sexual activity—intromission without ejaculation or ejaculation—and the occurrence of lordosis behavior. A second group was tested only once for both masculine sexual activity and lordosis behavior as above and afterwards three times at 5-day intervals for lordosis behavior in the absence of any previous testing for masculine sexual activity. A few animals displayed lordosis during their first test. As compared to the first group, the animals which had not displayed lordosis in the first test never showed lordosis responses in the following tests. It is concluded that both homotypic and heterotypic sexual interactions are required for the display of lordosis behavior in the strain of Wistar rats used in this study.

The purpose of this study was to determine whether the effects of estrogen on lordosis behavior in the male rat were related to the number of progesterone (P) receptors in the mediobasal hypothalamus (MBH) and/or dependent on blood P concentration. Two groups of gonadally intact male rats were given five successive doses of 1.0 or 2.5 micrograms estradiol benzoate (EB) and tested for lordosis behavior with a male stimulus at the end of the treatment. One month later they were again injected with EB and sacrificed under the same temporal schedule, but they were not tested for lordosis so as to prevent any emotionally stressful effects of intermale cohabitation. The males given 2.5 micrograms EB more frequently displayed lordosis responses to male mounts than those receiving 1 microgram EB, with a parallel increase in the number of MBH P receptors. The total number of MBH P receptors also appeared to be higher in the animals that displayed lordosis responses (lordosis group) than in those which did not (no lordosis group). In contrast, the display of lordosis behavior was negatively correlated with blood P concentration. Comparing MBH P receptors and blood P values in the EB treated and in nonhormonally treated gonadally intact animals which had been selected for either ability or inability to spontaneously display lordosis behavior, we observed that (1) EB was capable of increasing the number of MBH P receptors in the male rat; and (2) in the absence of EB treatment blood P values were higher in the animals showing lordosis than in those which did not. These data are discussed with respect to observations made in castrated male rats and in ovariectomized females.

Adult ovariectomized female rats received a testosterone propionate (TP) silastic implant (day 0) and were subsequently tested for mounting behavior. The number of females displaying mounting behavior increased from day 2 (0%) to day 14 (94%) following testosterone implantation. Olfactory bulbectomy (OB) performed on day -7 or on day +14 reduced the number of females that mounted on day +16 as compared to sham counterparts. However, the number of females displaying mounting behavior was significantly higher when OB was performed on day +14 than on day -7. Peripheral anosmia induced by ZnSO4 instillation, either with multiple instillations beginning on day -8 before TP implant or with a single injection on day +14, equally impaired mounting behavior on day +16. These results, which further confirm the role of the olfactory system in facilitating the display of mounting behavior in the female rat, suggest an olfaction-testosterone interaction.

Etude au comportement de lordose chez des rats mâles sexuellement inexperimentes ou apres une experience copulatoire avec des femelles. Cette etude s'interesse aux correlations existant entre le comportement de lordose et les concentrations sanguines de progesterone, œstradiol-17β et testosterone

The aim of this study was to investigate the olfactory mechanisms regulating the display of lordosis behavior in intact Wistar male rats bred in our colony. Gonadally intact males show a low capacity to respond by lordosis to male mounts and were insensitive to manipulations of the olfactory system (exposure to the odor of male urine or accessory bulb removal (AOBR)) which have been previously shown to facilitate the display of lordosis behavior in orchidectomized animals primed with ovarian hormones. Treatment with either estradiol benzoate (EB) or EB and progesterone (P) consecutively did not render these gonadally intact animals sensitive to the effects of AOBR. By contrast exposure to male urine was capable of facilitating the display of lordosis behavior in intact male rats given EB + P consecutively. These results are discussed in the light of previous findings showing that (1) two inhibitory structures, the accessory olfactory bulb and the septal and preoptic areas, are involved in the control of lordosis behavior in the male rat; (2) the effects of olfactory cues on the display of lordosis behavior are dependent on the action of both EB and P in orchidectomized animals.

Adult ovariectomized female rats received a testosterone-propionate (TP)-filled silastic implant and were tested for mounting behavior seven days later. Exposure to estrous female urine but not to urine from ovariectomized females increased the number of TP-treated females that displayed mounting behavior. Olfactory bulb removal on its own and peripheral anosmia induced by intranasal ZnSO4 application impaired mounting behavior. These results are discussed with respect to the effects of anosmia on copulatory behavior in the male.

The purpose of this study was to determine whether facilitory effects exerted by olfactory cues on lordosis behavior in the male rat involved changes in estradiol receptors at the hypothalamic level. Male rats were orchidectomized as adults. They were given either 25 micrograms estradiol benzoate (EB) alone or 25 micrograms EB and 100 micrograms progesterone (P) sequentially and exposed or not to the odor of male urine. Some of them were tested for lordosis behavior at 8 h after P. The other ones were killed 4 h after P and used for estradiol (E2) and P receptor assay in mediobasal hypothalamus (MBH). Olfactory cues were shown to increase the number of E2 receptors in both the animals given EB or EB + P. Progesterone as such appeared to be capable of increasing the number and the rate of occupancy of E2 receptors. A population of constitutive and estrogen-inducible P receptors was detected in the MBH. Since only the animals given EB + P were shown to be sensible to the facilitory effects of male urine on lordosis behavior, it may be assumed that E2 and P on one hand and olfactory cues on the other exert cumulative effects at the level of the MBH and that both a high level and a high rate of occupancy of E2 receptors are necessary for the olfactory cues to facilitate the display of lordosis behavior in the male rat.

The aim of the experiment was to study the effects of stereotaxic lesions of the anterior and the posterior regions of the corticomedial amygdaloid nucleus (CMN) on the display of lordosis behavior by the male rat. Animals were orchidectomized as adults and given estradiol benzoate and progesterone (P) sequentially. Sexual behavior testing was performed by 9 +/- 1 hr after P injection. Lesions placed into the posterior region of the CMN significantly decreased the proportion of animals showing lordosis behavior as compared to sham-operated and control animals. By contrast lesions in the anterior region of the CMN did not cause any changes in the proportion of animals displaying lordosis but markedly increased the lordosis quotient (LQ) of responding animals. The CMN was then concluded to exert a dual control in the display of lordosis behavior in the male rat with a posterior region regulating the willingness of animals to display lordosis behavior and rostral region subserving inhibitory mechanisms related to the sexual performance (LQ values).

The effects of androgen treatment (TP) from postnatal day 20 (P20) onwards on the accessory olfactory bulbs' (AOB) development were studied in rats orchidectomized at birth (PO) or on P20. TP failed to restore impaired AOB development in the rats castrated at PO but completely restored AOB development in those castrated at P20. Androgen secretion during both the perinatal and prepubescent periods appeared to be critical for male AOB development.

The aim of this study was to investigate the mechanisms involved in the inhibitory action of progesterone on estrogen-induced facilitatory effects of estradiol benzoate on lordosis behavior in the male rat. Intact adult male rats were given 1) 25 μg estradiol benzoate (EB) and 100 μg progesterone (P) at an interval of 42 hr. EB injected animals served as controls 2) EB followed by 3 doses of 400 μg dexamethasone (DEXA) and P as above. EB + DEXA injected animals served as controls. Testing for lordosis behavior was performed by 50±hr after EB injection. A significant decrease in the number of the males displaying lordosis in response to the mounts of stimulus males resulted from P injection following EB treatment as compared to EB controls. DEXA treatment significantly reduced the number of EB animals showing lordosis responses but completely prevented the inhibitory effects of exogenous P to occur. Blood P values appeared to be significantly lower in EB + DEXA males than in their EB counterparts. The results provide evidence that endogenous P is involved in the display of lordosis behavior by EB-treated intact males. They mainly suggest that the effects of exogenous P on estrogen-induced lordosis behavior in the intact male rat result from sequential inhibitory mechanisms involving exposure of the animals to the successive action of endogenous and exogenous P.

Pentobarbital lengthens the estrous cycle when injected on diestrus 1 afternoon in the 4-day cyclic female rat. The present study was undertaken to elucidate the mechanisms involved. Bromocriptine (2.5 mg/kg sc) given 30 min before pentobarbital (30 mg/kg ip) prevented the effects of pentobarbital on both cycle lengthening and progesterone levels in diestrus 2. Serum prolactin concentrations rose 10 min after pentobarbital injection, peaked at 30 min, and returned to control values after 60 min. Our results suggest that pentobarbital-induced cycle lengthening is due to a high progesterone production on diestrus 2 morning related to the luteotropic action of prolactin.

Previous studies showed that different amygdaloid nuclei are involved in the control of lordosis behavior in female rats. The objective of the present study was to determine whether these nuclei played a role in the control of lordosis behavior in the male rat. Lesions were placed into different amygdaloid nuclei in male rats castrated as adults and primed with ovarian hormones. Lesions in the corticomedial amygdaloid nucleus completely suppressed lordosis behavior as expressed by the number of animals displaying lordosis responses to male mounts. By contrast extended lesions placed into the lateral amygdaloid nucleus (LN) remained without effects. Lesions placed in the very posterior part of the LN produced “hypersexuality” with a rise in the number of animals displaying lordosis responses and high LQ values. Lesions in the anterior part of the LN induced a decrease in the number of animals showing lordosis responses. The amygdala was then concluded to represent a functionally heterogeneous structure with different regions exerting opposite effects on the display of lordosis behavior in the male rat.

The study was undertaken to examine LH, prolactin (Prl) and estradiol involvement in the control of corpus luteum function during 4-day cycle in the female rat. A potent LHRH antagonist (Antag) and bromocriptine (BRC) were used to induce LH and Prl deprivation in 4-day cyclic rats. Both drugs were administered after the occurrence of LH and Prl release on the afternoon of proestrus. Blood progesterone (P) concentration on diestrus 1 at 24:00 hr was similar in Antag, Antag + BRC and BRC-treated animals and did not differ from P values in control females. It was then concluded that neither LH nor Prl were necessary for the corpus luteum to achieve its stage of maximal activity during the night from diestrus 1 to diestrus 2. By contrast higher P values were observed on diestrus 2 at 12:00 hr in Antag-treated rats than in those given Antag + BRC, BRC or in the controls. When given on diestrus 1 evening estradiol benzoate completely prevented this effect of Antag to occur in LH and Prl deprived females. These results allow to hypothesize that LH exerts luteolytic effects during the diestrous period of the cycle and also suggest a possible involvement of estradiol in the regression phase of the corpus luteum.

The aim of this investigation was 1) to compare the development of AOB in male and female rats before birth, early after birth, and at later stages of sexual immaturity and maturity 2) to examine the effects of early and delayed castration after birth on AOB development in the male rat. A significant increase of the surface area of AOB was observed in both sexes from birth until postnatal day 7, but AOB was found to be larger in males than in females. From the end of the first postnatal week AOB stopped growing until day 40 in both males and females. After this time AOB resumed its development until day 60 in males, while no changes occurred in females over the same period. Castration of males at birth or six hours later impaired development of AOB. Castration of males on postnatal days 20 or 30 also impaired AOB development until day 60. The results strongly suggest that the development of AOB in the male rat is dependent on the two well known "perinatal" and "adult" phases of the endocrine activity of the testes.

Rats were orchidectomized (ORCH) as adults and given successive doses of 0.5 microgram or 1.0 microgram of oestradiol benzoate (EB) combined or not with dexamethasone (DEXA) at the end of oestrogen treatment; successive doses of olive oil. They were presented to stimulus males on the day of the last EB injection and decapitated immediately after the behavioural test. Animals given EB or olive oil only served as controls. There was a significant rise in blood progesterone concentration in animals given 0.5 and 1.0 microgram EB as compared with oil-treated animals. A higher blood progesterone concentration was observed at the end of the behavioural session in oil-treated and 0.5 microgram EB-treated ORCH rats than in their isolated counterparts, an effect which appeared not to dependent on lordosis responses to mounting attempts of the males. DEXA completely suppressed the rise in blood progesterone concentration in oestrogenized ORCH rats presented to stimulus males. Presentation of ORCH rats to stimulus males was then concluded to constitute a stressful condition capable of inducing adrenocortical progesterone secretion in ORCH animals.

Previous observations showed that exposure to the odor of male urine prior to mating could enhance the display of lordosis behavior in male rats feminized with ovarian hormones. This study was performed to determine in feminized male rats whether the control of lordosis behavior by the olfactory system was mediated by the ventromedial nucleus (VMN) of the hypothalamus. Male rats were orchidectomized (ORCH) as adults and primed with 25 micrograms estradiol benzoate (EB) and 150 micrograms progesterone (P) 40 hr apart. Lordosis behavior was tested 9 +/- 1 hr after P injection. VMN lesions were shown to completely suppress the display of lordosis behavior as compared to sham VMN operated and dorsomedial nucleus (DMN) lesioned animals. Exposure of feminized rats to the odor of male urine by 9 +/- 1 hr before mating significantly increased the proportion of ORCH rats that displayed lordosis behavior in response to male mounts. This effect was abolished by VMN lesions but was maintained in the sham VMN operated and DMN lesioned animals. These results were discussed in the light of the present knowledge on the neuroendocrine and olfactory structures which mediate lordosis behavior in the male rat.

The role of pheromones in the process of ovulation and/or luteinization induced by coitus was studied in female rats primed with estradiol benzoate (EB) and early mated during 4-day cycles. The effects of coitus were evaluated by using 1) the proportion of females displaying postovulatory corpora lutea (POCL) and/or luteinized unruptured follicles (LUF), and 2) the ovulation coefficient (OC) computed in each female by dividing the number of POCL by the total number of POCL and LUF. A greater proportion of females displayed ovulation and/or luteinization following coitus than those given EB only. This proportion was slightly but significantly increased when females were exposed to bedding soiled with 5 ml of male urine prior to mating, an effect prevented by complete olfactory bulbectomy. However, in the females given EB only, the pheromonal stimulus remained inefficient. Weak ovulatory effects of coitus, as assessed by low OC values, were noted whether the females were exposed or not to male urine. An increase in the frequency of lordosis occurred after olfactory bulbectomy, but no significant changes in OC values were observed in this experimental group. It was concluded that pheromones, act as a primer on the neuroendocrine mechanisms governing ovulation in the cyclic female rat, as in other species of rodents.

Orchidectomized rats were given estrogen and progesterone and tested for feminine behavior in the presence of a mounting male after accessory olfactory bulb removal (AOBR). Complete AOBR caused a rise in the number of estrogen-progesterone-treated male rats responding by lordosis behavior to male mounts as compared to controls and sham-operated animals. By contrast, LQ scores did not appear to differ in these three groups of animals. The results are discussed in terms of involvement of the main and the accessory olfactory systems in the regulation of feminine behavior in the male rat.

The relation between testosterone-induced oestrous cycle disruption and prolactin (Prl) secretion was studied in the female rat. Testosterone propionate (TP) injection on dioestrus 2 (day 0) of 4-day cycles induced cycle interruption for 12 days. A diurnal pattern of Prl secretion was observed from day 1 to day 6 with high levels in the afternoon and low levels in the morning. Cycle interruption also occurred in TP treated females given a dose of bromocriptine (BRC) known to completely block Prl secretion. Impairment of follicular growth resulted from TP treatment and from TP and BRC treatment as well. Until resumption of prooestrous, the follicular size did not exceed that noted during the dioestrous period of the cycle. Despite resumption of follicular growth from day 9 until day 14, ovulation only took place in a small number of animals. These results indicate that the antigonadotropic effects of Prl were not responsible for cycle disruption in our experimental model and that other mechanisms were in itself sufficient for the disruption. This was discussed in the light of previous findings in our laboratory.