Regulation of sexual odor preference by sex steroids in the posterodorsal medial amygdala in female rats

Our previous study in male rats demonstrated that bilateral administration of flutamide, an androgen receptor (AR) antagonist, into the posterodorsal medial amygdala (MePD) increased the time sniffing male odors to as high as that sniffing estrous odors, eliminating the preference for estrous odors over male odors. This made us speculate that under blockade of AR in the MePD, testosterone-derived estrogen acting on the same brain region arouses interest in male odors which is otherwise suppressed by concomitant action of androgen. In cyclic female rats, endogenous androgen has been thought to be involved in inhibitory regulation of estrogen-activated sexual behavior. Thus, in the present study, we investigated the possibility that in female rats the arousal of interest in male odors is also normally regulated by both estrogen and androgen acting on the MePD, as predicted by our previous study in male rats. Implantation of either the estrogen receptor blocker tamoxifen (TX) or a non-aromatizable androgen 5α-dihydrotestosterone (DHT) into the MePD of ovariectomized, estrogen-primed female rats eliminated preference for male odors over estrous odors by significantly decreasing the time sniffing male odors to as low as that sniffing estrous odors. The subsequent odor discrimination tests confirmed that the DHT and TX administration did not impair the ability to discriminate between male and estrous odors. These results suggest that in estrous female rats estrogen action in the MePD plays critical roles in the expression of the preference for male odors while androgen action in the same brain region interferes with the estrogen action.     

PSA-NCAM in the posterodorsal medial amygdala is necessary for the pubertal emergence of attraction to female odors in male hamsters

During puberty, attention turns away from same-sex socialization to focus on the opposite sex. How the brain mediates this change in perception and motivation is unknown. Polysialylated neural cell adhesion molecule (PSA-NCAM) virtually disappears from most of the central nervous system after embryogenesis, but it remains elevated in discrete regions of the adult brain. One such brain area is the posterodorsal subnucleus of the medial amygdala (MePD). The MePD has been implicated in male sexual attraction, measured here as the preference to investigate female odors. We hypothesize that PSA-NCAM gates hormone-dependent plasticity necessary for the emergence of males' attraction to females. To evaluate this idea, we first measured PSA-NCAM levels across puberty in several brain regions, and identified when female odor preference normally emerges in male Syrian hamsters. We found that MePD PSA-NCAM staining peaks shortly before the surge of pubertal androgen and the emergence of preference. To test the necessity of PSA-NCAM for female odor preference, we infused endo-neuraminidase-N into the MePD to deplete it of PSAs before female odor preference normally appears. This blocked female odor preference, which suggests that PSA-NCAM facilitates behaviorally relevant, hormone-driven plasticity.     

Photoperiod-dependent response to androgen in the medial amygdala of the Siberian hamster,Phodopus sungorus

The medial nucleus of the amygdala (MeA) is a steroid-sensitive region that has been implicated in the expression of behaviors such as mating and aggression. The male Siberian hamster (Phodopus sungorus) uses light cues to regulate its reproductive neuroendocrine system, reducing androgen synthesis in the autumn and increasing it in the spring. There is also evidence that short photoperiods reduce the sensitivity of the brain to the behavioral effects of androgen. The authors tested the hypothesis that MeA neurons are less responsive to androgen in short photoperiods by comparing the regional volume and average soma size of the four MeA subnuclei (anterodorsal [MeAD], anteroventral [MeAV], posterodorsal [MePD], and posteroventral) in adult male hamsters that had been castrated and then implanted with capsules containing either testosterone (T) or nothing. Animals from each group were housed in either long or short photoperiods for 15 weeks. MeAD and MeAV somata displayed photoperiod-dependent responses to androgen, increasing in size after T treatment only in long days. In contrast, the average soma size and the regional volume of the MePD subnucleus were significantly larger in T-treated males regardless of photoperiod. The authors conclude that photoperiod influences the sensitivity of the MeA to androgen.     

Olfactory preference in the male rat depends on multiple chemosensory inputs converging on the preoptic area

Both volatile and nonvolatile molecules are involved in chemosensory communication in rodents. Volatile odors from physically inaccessible estrous females induced increased numbers of c-Fos-positive cells in the preoptic area (POA) and in the cortical nucleus of the amygdala (CoA) of male rats. The numbers of c-Fos-positive cells in the medial nucleus of the amygdala (MeA) increased in response to the nonvolatile odors of bedding soiled with the excreta of estrous females. In an alternate choice paradigm, male rats carrying ibotenic acid lesions in either the MeA or the CoA—or a combination of both—distinguished the odors of estrous females from those of males, although the time spent sniffing the stimuli was diminished. Males with POA lesions showed complete loss of this capability. Males carrying either of the lesions did not detect differences between estrous and anestrous females or between intact and orchidectomized males. Lesions in the POA or MeA severely impaired male sexual behavior, whereas a CoA lesion had no effects. Thus, c-Fos-positive cells in the CoA might be involved in chemosensory transmission relevant to certain social contexts, but not in the execution of male sexual behavior. The POA is indispensable for both olfactory preferences and sexual behavior. The residual olfactory preference in males with MeA or CoA lesions or the combination of both could reflect an additional route for chemosensory transmission from the main olfactory bulb to the POA.     

Androgen implants in medial amygdala briefly maintain noncontact erection in castrated male rats

Castration of male rats causes a rapid loss of their normal erectile response to inaccessible estrous females. Previous studies had demonstrated that these noncontact erections (NCEs), a putative sign of sexual arousal, could be restored by systemic treatment with testosterone (T) or dihydrotestosterone (DHT), but not estradiol (E). We examined whether androgen delivered to the medial amygdala (MeA) of castrated rats would maintain NCE. In Experiment 1, males received bilateral cannulae filled with T, DHT, or E directed at the MeA. Control males had the same hormone-filled cannulae implanted subcutaneously and blank cannulae in the MeA, or they received T in the anterior forebrain. During the 2 weeks after surgery, males were tested twice for NCE and copulation. About half the males with androgens in the MeA had NCEs 1 week after castration, but few responded a week later. Closer proximity of androgen implants to the posterodorsal MeA (MeApd) predicted shorter NCE latencies. No males with subcutaneous androgen had NCEs in either test, and few anterior forebrain-implanted males did. Some males receiving E in MeA or subcutaneously had NCE in each test. In copulation tests, the type of steroid treatment did not affect the incidence of ejaculation or most measures of copulation, and the proximity of cannulae to MeApd predicted only the time from ejaculation to the occurrence of NCE during the postejaculatory interval. Experiment 2 showed that NCEs displayed by males with androgen in MeA occurred in response to estrous females, not spontaneously. The results suggest that androgens, perhaps augmented by estrogen, act in the posterodorsal MeA to facilitate NCE and its associated arousal.     

Sexual incentive motivation, olfactory preference, and activation of the vomeronasal projection pathway by sexually relevant cues in non-copulating and naive male rats

There are some apparently healthy male rats that fail to mate after repeated testing with receptive females. We have previously shown that these "non-copulator (NC)" males show no partner preference for a receptive female when given the opportunity to physically interact with a sexually receptive female or a sexually active male. We also demonstrated that although NC males prefer odors from estrous females to odors from anestrous females, this preference is significantly reduced in comparison to the preference displayed by copulating (C) males. The aim of the present study was to evaluate in NC males sexual incentive motivation, that is, the approach behavior of male rats to either a sexually receptive female or a sexually active male in a test where the subjects can smell, hear, and see the stimulus animal but prevents their physical interaction. In addition, we determined whether NC rats have alterations in their ability to detect odors from conspecifics or odors related to food. In the detection of odors from conspecifics, we determined if these NC males are sexually attracted toward odors from receptive females or sexually active males. For food-related odors, we quantified the time it took the subjects to locate a hidden a piece of apple. Finally, using the induction of Fos-immunoreactivity (Fos-IR) as an index of neuronal activation, we compared the response of the vomeronasal projection pathway (VN pathway) of C and NC male rats exposed to estrous bedding. Males without sexual experience (WSE) were included in all experiments to determine the importance of previous heterosexual experience in the different behavioral tests and in the activity of the VN pathway. In the sexual incentive motivation test, we found that C and WSE male rats have a clear preference for estrous females over sexually active males, whereas NC male rats showed no preference. In odor tests, our results showed that C males had a clear preference for odors from estrous females as opposed to odors from sexually active males. Although NC and WSE male rats showed a preference for estrous female odors, this preference was significantly reduced compared to that shown by C males. No differences were found between WSE, C, and NC males in the detection of stimuli associated with food-related odors. A significant increase in Fos-IR was observed in the mitral cell layer of the accessory olfactory bulb in all groups when exposed to estrous bedding. However, only the C male rats exposed to estrous female bedding showed an increase Fos-IR in all structures of the VN pathway. An increase in Fos-IR was observed in the medial preoptic area (MPOA) of WSE males exposed to estrous bedding. No increases in Fos-IR were detected along the VN pathway in NC male rats. We proposed that NC male rats do not display sexual behavior due to a reduced sexual motivation that could be caused by alterations in the neuronal activity of the VN pathway during the processing of estrous odors.     

Brief Exposure to Female Odors “Emboldens” Male Mice by Reducing Predator-Induced Behavioral and Hormonal Responses

In rodents, where chemical signals play a particularly important role in determining intersexual interactions, various studies have shown that male behavior and physiology is sensitive to female odor cues. Here we examined the effects of brief (1 min) and more prolonged (60 min) preexposure to the odors of a novel estrous female on the behavioral and hormonal responses of sexually experienced and inexperienced male mice, Mus musculus, to subsequent predator (cat and weasel) odor exposure and potential predator risk. Brief, but not prolonged, preexposure to the odors of an estrous female decreased the aversion and avoidance responses of male mice to cat odor in a Y-maze preference test, with the extent of responses being affected by a males prior sexual experience. Similarly, brief, but not prolonged, preexposure to female odors markedly attenuated the analgesic responses elicited in male mice by weasel odor. Brief exposure to a novel estrous female by itself had no significant immediate effects on either corticosterone or testosterone levels in the males. However, brief, but not prolonged, preexposure to the odors of an estrous female attenuated the marked increase in corticosterone and decrease in testosterone that were induced in males by exposure to weasel odor. The decreases in aversive responses to, and effects of, predator odor exposure that are induced by brief exposure to a novel estrous female may reflect a greater risk taking and boldness in males that could directly facilitate access to an immediately, and possibly transiently, available novel sexually receptive female.     

光周期对雄性布氏田鼠种内个体气味辨别的影响

本文研究了成年雄性布氏田鼠对不同光照周期下(长光照: LD; 短光照: SD) 的陌生雌鼠和陌生雄鼠气味的行为表现。实验发现: 所有被试雄鼠对LD 雄鼠、雌鼠气味比SD 雄鼠、雌鼠气味有更多的社会探究行为。LD 雄鼠比SD 雄鼠对同性或异性陌生个体气味的嗅闻和挖掘行为要多, 而且差异显著。在探究LD 动情雌鼠气味源时, LD 雄鼠比SD 雄鼠表现出更多的嗅闻和挖掘行为; 在探究SD 动情雌鼠气味源时, LD 和SD 雄鼠的行为反应没有明显差别。所有被试雄鼠在LD 雄鼠气味源箱中的停留时间都显著多于在SD 雄鼠气味源箱中的停留时间。LD 雄鼠在LD 雌鼠气味源箱中的停留时间显著多于在SD 雌鼠气味源箱中的停留时间; 而SD 雄鼠在LD 和SD 雌鼠气味源箱中的进入频次和停留时间没有明显差别。同时, LD 雄鼠对LD 动情雌鼠的气味在嗅闻频次和嗅闻时间上多于非动情LD 雌鼠, 而对SD 动情和非动情雌鼠的气味没有表现出明显的偏好; SD 雄鼠对LD 和SD 动情雌鼠的嗅闻行为仅在频次上显著多于对非动情雌鼠的嗅闻, 而在时间上没有显著差别。结果表明: 布氏田鼠嗅觉通讯中的个体气味及其对气味进行辨别过程中的行为反应都随着光照周期的不同而发生变化。动物的个体气味带有季节性信息, 来源于长光照下的气味比短光照下的气味更具有性吸引; 短光照在一定程度上可能抑制了雌鼠的动情和雄鼠的性行为反应。其嗅觉通讯行为的适应意义可能在于: 减少秋季短日照环境下的繁殖活动, 提高雄性个体之间的社会容忍性以利于集群越冬。     

Chemosensory and hormone information are relayed directly between the medial amygdala, posterior bed nucleus of the stria terminalis, and medial preoptic area in male Syrian hamsters

In many rodent species, including Syrian hamsters, the expression of appropriate social behavior depends critically on the perception and identification of conspecific odors. The behavioral response to these odors is mediated by a network of steroid-sensitive ventral forebrain nuclei including the medial amygdala (Me), posterior bed nucleus of the stria terminalis (BNST), and medial preoptic area (MPOA). Although it is well-known that Me, BNST, and MPOA are densely interconnected and each uniquely modulates odor-guided social behaviors, the degree to which conspecific odor information and steroid hormone cues are directly relayed between these nuclei is unknown. To answer this question, we injected the retrograde tracer, cholera toxin B (CTB), into the BNST or MPOA of male subjects and identified whether retrogradely-labeled cells in Me and BNST 1) expressed immediate early genes (IEGs) following exposure to male and/or female odors or 2) expressed androgen receptor (AR). Although few retrogradely-labeled cells co-localized with IEGs, a higher percentage of BNST- and MPOA-projecting cells in the posterior Me (MeP) expressed IEGs in response to female odors than to male odors. The percentage of retrogradely-labeled cells that expressed IEGs did not, however, differ between and female and male odor-exposed groups in the anterior Me (MeA), posterointermediate BNST (BNSTpi), or posteromedial BNST (BNSTpm). Many retrogradely-labeled cells co-localized with AR, and a higher percentage of retrogradely-labeled MeP and BNSTpm cells expressed AR than retrogradely-labeled MeA and BNSTpi cells, respectively. Together, these data demonstrate that Me, BNST, and MPOA interact as a functional circuit to process sex-specific odor cues and hormone information in male Syrian hamsters.     

Behavioral characterization of non-copulating male mice

Non-copulating (NC) males are those animals that do not mate in spite of repeated testing with sexually receptive females. They have been observed in several species including rats and mice. The present experiment was designed to perform a detailed behavioral characterization of NC male mice. Thus, we evaluated their sexual incentive motivation for a sexually receptive female or a sexually active male, olfactory preference for volatile and non-volatile odors from females or males, and olfactory discrimination between female and male volatile odors and food related odors (milk versus vinegar). We compared the activity of the accessory olfactory system (AOS) in copulating (C) and NC males in response to estrous bedding using the induction of Fos-immunoreactivity (Fos-IR) as a measure of neuronal activation. We also determined if estradiol or dopamine treatment could induce sexual behavior in NC males. Finally, we compared the testis weight and the number of penile spines in C, NC, and gonadectomized males. In the sexual incentive motivation test C males spend significantly more time in the female incentive zone than in the male incentive zone. On the other hand, NC males spend the same amount of time in both incentive zones. In tests of olfactory preference, NC males spent less time investigating estrous odors than C males. As well, NC males discriminate urine from conspecifics but they spend less time smelling these odors than C males. In addition, no increase in Fos expression is observed in NC males when they are exposed to odors from estrous females. Our data also suggest that the deficits observed in NC males are not due to lower circulating levels of gonadal hormones, because estradiol supplementation does not induce sexual behavior in these animals, and their testis weight and the number of penile spines are normal. The results suggest that NC males are not sexually motivated by the receptive females and their odors.     

Non-neural androgen receptor promotes androphilic odor preference in mice

In mice, male-typical preference for female olfactory cues results largely from sexually differentiated testosterone production. It is currently unclear on which cells and tissues testosterone acts to produce male-typical preference for female olfactory cues. To further address the site of androgen action on olfactory preference, we have developed a loxP-based transgenic mouse that overexpresses androgen receptors (AR) only when activated by Cre. We used this transgene to overexpress AR globally in all tissues using a CMV-Cre driver and a Nestin-Cre driver to overexpress AR selectively in neural tissue. We then examined olfactory preference in transgenic and wildtype (Wt) littermates by simultaneously exposing animals to female-soiled, male-soiled and clean bedding. Ubiquitous overexpression of AR in CMV-AR mice increased preference for male bedding, whereas neural-specific AR overexpression in Nestin-AR transgenic mice did not differ from wildtype siblings in olfactory preference. Neural activation of olfactory brain areas in response to female-soiled bedding was also evaluated in these mice by measuring FOS immunoreactivity. This revealed a decrease in neural activity along the accessory olfactory pathway that accompanied the decrease in preference for female odors in CMV-AR males, compared to both Nestin-AR and Wt male siblings. Together, results indicate that androgens act via non-neural AR to mediate olfactory preference and neural responses to olfactory stimuli, and further suggest that AR in non-neural tissues can promote androphilic odor preferences in male mice.     

Self-grooming induced by sexual chemical signals in male root voles (Microtus oeconomus Pallas)

Sniffing is one-way animals collect chemical signals, and many males self-groom when they encounter the odor of opposite-sex conspecifics. We tested the hypothesis that sexual chemical signals from females can induce self-grooming behavior in male root voles (Microtus oeconomus Pallas). Specifically, we investigated the sniffing pattern of male root voles in response to odors from the head, trunk, and tail areas of lactating and non-lactating females. The self-grooming behavior of males in response to female individual odorant stimuli was documented, and the relationship between self-grooming and sniffing of odors from the head, trunk, and tails areas were analyzed. Sniffing pattern results showed that males are most interested in odors from the head area, and more interested in odors from the tail as compared to the trunk area. Males displayed different sniffing and self-grooming behaviors when they were exposed to odors from lactating females as compared to non-lactating females. Males also spent more time sniffing and engaged in more sniffing behaviors in response to odors from the lactating females’ tail area as compared to the same odors from non-lactating females. Similarly, males spent more time self-grooming and engaged in more self-grooming behaviors in the presence of individual odors from lactating females as compared to individual odors from non-lactating females. Partial correlation analyses revealed that the frequency of self-grooming was significantly correlated with the frequency of tail area sniffs. Results from this experiment suggest that sexual attractiveness of lactating females is stronger than that of non-lactating females. Furthermore, the partial correlation analysis demonstrated that self-grooming in males is induced by odors from the tail area of females. Collectively, these results support the hypothesis that sexual chemical signals from females can induce self-grooming behavior in male root voles. Self-grooming may also reflect the groomer's sexual motivation and facilitate sexual interactions.     

Sexual partner preference requires a functional aromatase (cyp19) gene in male mice

Sexual motivation, sexual partner preference, and sexual performance represent three different aspects of sexual behavior that are critical in determining the reproductive success of a species. Although the display of sexual behavior is under strict hormonal control in both sexes, the relative roles of androgen and estrogen receptors in activating the various components of male sexual behavior are still largely unknown. A recently developed mouse model that is deficient in estradiol due to targeted disruption of exons 1 and 2 of the Cyp19 gene (aromatase knockout (ArKO) mice) was used here to analyze the role of estradiol in the control of all three aspects of male sexual behavior. When tested in a Y-maze providing volatile olfactory cues, male ArKO mice did not show a preference for the odors from an estrous female over those from an intact male, whereas wild-type (WT) and heterozygous (HET) males clearly preferred to sniff estrous odors. When provided with visual and olfactory cues, male ArKO mice also failed to show a preference for an estrous female when given a choice between an estrous female and an empty arm. However, sexual partner preferences of male ArKO mice were not sex-reversed: they did not prefer to investigate an intact male over an estrous female or empty arm. Thus, male ArKO mice seemed to have general deficits in discriminating between conspecifics by using olfactory and visual cues. Male coital behavior was also severely impaired in male ArKO mice: they displayed significantly fewer mounts, intromissions, and ejaculations than WT and HET males. Latencies to first mount or intromission were also significantly longer in ArKO males compared to WT and HET males, in addition to them showing less interest in investigating olfactory and visual cues in a Y-maze, suggesting that they were sexually less motivated. However, three out of seven male ArKO mice were capable of siring litters provided they were housed with a female for a prolonged period of time. In conclusion, aromatization of testosterone to estradiol appears to be essential for sexual motivation and sexual partner preference. By contrast, estradiol may play only a limited role in the expression of male coital behaviors.     

雄性布氏田鼠对不同熟悉程度和动情状态下雌鼠气味的辨别

通过雄性布氏田鼠（Microtus brandti)对配偶和陌生雌鼠气味的辨别实验发现：（1）雌鼠动情状况对雄鼠气味行为反应的影响,当配偶雌鼠处于动情期时,被试雄鼠对动情陌生雌鼠巢垫物的溴闻和舔舐的持续时间都明显多于对非动情陌生雌鼠巢垫物的嗅闻和舔舐时间,而其他行为没有明显判别当配偶雌鼠处于非动情期时,雄鼠对动情的和非动情的陌生雌鼠的气味行为反应都没有明显判别当陌生雌鼠处于动情期时,被试雄鼠对动情配偶气味的嗅闻时间明显多于对非动情配偶的嗅闻时间,而其他行为反应没有明显差异;当陌生雌鼠处于非动情期时,被试雄鼠对动情配偶雌鼠巢垫物的嗅闻时间,挖掘频次,舔舐频次和时间都明显多于对非动情配偶气味的嗅闻,而且雄鼠在动情配偶气味周围搔扒体侧行为的发生频次和持续时间也多于在非动情配偶气味周围的搔扒行为;（2）熟悉性对雄鼠气味行为反应的影响,当配偶雌鼠和陌生雌鼠都处于动情期或都处于非动情期时,被试雄鼠对陌生雌鼠气味的探究行为（嗅问、挖掘和舔噬）和搔扒体侧行为明显多于对配偶雌鼠气味的探究;雄鼠访问陌生雌鼠气味源箱的频次和在其中的停留时间也都多于访问配偶雌鼠气味源箱的频次和停留时间;当配偶雌鼠处于动情期而陌生雌鼠处于非动情期时,被试雄鼠对配偶气味的嗅闻,挖掘和自身修饰行为明显多于对陌生雌鼠气味的行为反应,进入配偶气味源箱中的访问频次也多于进入陌生雌鼠气味源箱的访问频次。结果说明：雌鼠与被试雄鼠的熟悉程度和雌鼠的动情状况直接影响雄性布氏田鼠的社会探究和气味选择行为,即雄鼠偏好并选择动情雌鼠和陌生雌鼠,雄性布氏田鼠对雌鼠气味的行为反应也表现出田鼠属动物典型的多配制种类特征。     

Divergent roles for estrogens and androgens in the expression of female goat sexual behavior

We tested the hypothesis that the activation of the androgen receptor (AR) is required for full expression of female goat sexual behavior. Once a week for 6 weeks, ovariectomized (OVX) females were given priming doses of progesterone 72 and 48 h before behavioral observation. Estradiol (E(2); 100 microg), testosterone (T; 100 mg), or sesame oil was supplied 14 h before behavioral testing. Six goats received the AR antagonist flutamide (9 mg/kg sc) 8 h before and 4 h after steroid injection. Six goats received the carrier only. After 3 weeks, flutamide and carrier treatments were switched so that all females received all treatments. Treatments with E(2) and T were equally effective in eliciting estrus-typical behaviors (sniffing, courting, leg kicks, mount attempts by males, bouts of thrusting by males, ejaculations, and flehman responses) compared to treatment with oil. Flutamide treatment enhanced proceptive behaviors in E(2)-treated females compared to other treatment groups; this was most likely via enhanced tail wagging. Moreover, compared to goats given T + carrier, T + flutamide significantly reduced receptivity in females. The results of this experiment implicate the AR as an important facilitator of some aspects of the female goat sexual behavior. However, the results of this experiment do not show whether androgens influence estrous behaviors alone or in some combination with estrogen.     

The bed nucleus of the stria terminalis is critical for sexual solicitation, but not for opposite-sex odor preference, in female Syrian hamsters

Successful reproduction in vertebrates depends critically upon a suite of precopulatory behaviors that occur prior to mating. In Syrian hamsters (Mesocricetus auratus), these behaviors include vaginal scent marking and preferential investigation of male odors. The neural regulation of vaginal marking and opposite-sex odor preference likely involves an interconnected set of steroid-sensitive nuclei that includes the medial amygdala (MA), the bed nucleus of the stria terminalis (BNST), and the medial preoptic area (MPOA). For example, lesions of MA eliminate opposite-sex odor preference and reduce overall levels of vaginal marking, whereas lesions of MPOA decrease vaginal marking in response to male odors. Although BNST is densely interconnected with both MA and MPOA, little is known about the role of BNST in female precopulatory behaviors. To address this question, females received either bilateral, excitotoxic lesions of BNST (BNST-X) or sham lesions (SHAM), and were tested for scent marking and for investigatory responses to male and female odors. Whereas SHAM females vaginal marked more to male odors than female odors on two days of the estrous cycle, BNST-X females marked at equivalent levels to both odors. This deficit is not due to alterations in social odor investigation, as both BNST-X and SHAM females investigated male odors more than female odors. Finally, BNST lesions did not generally disrupt the cyclic changes in reproductive behaviors that occur across the estrous cycle. Taken together, these results demonstrate that BNST is critical for the normal expression of solicitational behaviors by females in response to male odor stimuli.     

The bed nucleus of the stria terminalis is critical for sexual solicitation,but not for opposite-sex odor preference,in female Syrian hamsters

Successful reproduction in vertebrates depends critically upon a suite of precopulatory behaviors that occur prior to mating. In Syrian hamsters (Mesocricetus auratus), these behaviors include vaginal scent marking and preferential investigation of male odors. The neural regulation of vaginal marking and opposite-sex odor preference likely involves an interconnected set of steroid-sensitive nuclei that includes the medial amygdala (MA), the bed nucleus of the stria terminalis (BNST), and the medial preoptic area (MPOA). For example, lesions of MA eliminate opposite-sex odor preference and reduce overall levels of vaginal marking, whereas lesions of MPOA decrease vaginal marking in response to male odors. Although BNST is densely interconnected with both MA and MPOA, little is known about the role of BNST in female precopulatory behaviors. To address this question, females received either bilateral, excitotoxic lesions of BNST (BNST-X) or sham lesions (SHAM), and were tested for scent marking and for investigatory responses to male and female odors. Whereas SHAM females vaginal marked more to male odors than female odors on two days of the estrous cycle, BNST-X females marked at equivalent levels to both odors. This deficit is not due to alterations in social odor investigation, as both BNST-X and SHAM females investigated male odors more than female odors. Finally, BNST lesions did not generally disrupt the cyclic changes in reproductive behaviors that occur across the estrous cycle. Taken together, these results demonstrate that BNST is critical for the normal expression of solicitational behaviors by females in response to male odor stimuli.     

Preferences for salivary odor cues by female hamsters

There is a growing body of data which indicates that saliva may function as a chemosignal in mammals. The salience of this cue in hamsters is investigated. Twelve females were tested in a four-choice olfactorium to determine whether the subjects could differentiate and demonstrate a preference for salivary cues as a function of the dominance status of the stimulus donors and the subjects estrous state. Clear differences in responding were found for estrous as opposed to diestrous females. Estrous females detected and exhibited a preference for male stimuli on the basis of entry and approach measures. Subjects were also found to exhibit sniffing preferences for male odors and possible an avoidance of female odors relative to the saline control. Estrous females appeared to be unable to distinguish between dominant and subordinate males.     

Dendritic spines of the medial amygdala: plasticity, density, shape, and subcellular modulation by sex steroids

The medial nucleus of the amygdala (MeA) is a complex component of the "extended amygdala" in rats. Its posterodorsal subnucleus (MePD) has a remarkable expression of gonadal hormone receptors, is sexually dimorphic or affected by sex steroids, and modulates various social behaviors. Dendritic spines show remarkable changes relevant for synaptic strength and plasticity. Adult males have more spines than females, the density of dendritic spines changes in the course of hours to a few days and is lower in proestrous and estrous phases of the ovarian cycle, or is affected by both sex steroid withdrawal and hormonal replacement therapy in the MePD. Males also have more thin spines than mushroom-like or stubby/wide ones. The presence of dendritic fillopodia and axonal protrusions in the MePD neuropil of adult animals reinforces the evidence for local plasticity. Estrogen affects synaptic and cellular growth and neuroprotection in the MeA by regulating the activity of the cyclic AMP response element-binding protein (CREB)-related gene products, brain-derived neurotrophic factor (BDNF), the anti-apoptotic protein B-cell lymphoma-2 (Bcl-2) and the activity-regulated cytoskeleton-related protein (Arc). These effects on signal transduction cascades can also lead to local protein synthesis and/or rearrangement of the cytoskeleton and subsequent numerical/morphological alterations in dendritic spines. Various working hypotheses are raised from these experimental data and reveal the MePD as a relevant region to study the effects of sex steroids in the rat brain.     

Ability of mice to detect estrous odor in bovine urine: roles of hormones and behavior in odor discrimination

This study was designed to evaluate the ability of mice to discriminate cow urinary odor from different reproductive phases, with a view toward detecting the estrous phase. Experiments were also carried out to establish the relationship between androgen and mouse behaviors during estrous detection. Further, the study was also intended to establish the relationship between androgen and behaviors in estrous detection. Bovine urine was collected during estrus and non-estrous periods, i.e., prepubertal, preovulatory, ovulatory, postovulatory, pregnancy, and lactation. Behavioral analyses were carried out in a Y-maze apparatus, in which the mice were acclimatized in before odor-preference tests. The number and duration of visits, and grooming behavior by male responders towards the urine samples, were recorded. Intact male mice showed a higher response towards estrus urine samples than towards non-estrous urine. By contrast, orchidectomized mice failed to discriminate estrous urine, whereas castrated mice treated with testosterone regained the ability to discriminate estrus odor. A higher level of grooming behavior was found in males exposed to estrous urine than to urine of other phases. These results suggest that normal mice have the ability to detect estrus, and that this discriminating ability is androgen dependent. The grooming behavior shown by males in response to estrous urine may be taken as a key parameters in estrous detection. The results further suggest that bovine estrous urine produces specific odors that probably involve both intraspecific and interspecific communication.