Copulatory behavior of sexually inexperienced male hamsters paired with natural proestrous females

The copulatory behavior of sexually inexperienced male hamsters paired with natural proestrous females was observed between 1900 and 2000 hours under dim red light illumination. Each male was allowed 30 min to mate with a receptive female. Five out of 7 males showed at least one ejaculation within 30 min. Although the copulatory behavior of the above males was observed again, the activities of copulatory behavior did not facilitate.     

Endogenous oxytocin is necessary for preferential Fos expression to male odors in the bed nucleus of the stria terminalis in female Syrian hamsters

Successful reproduction in mammals depends on proceptive or solicitational behaviors that enhance the probability of encountering potential mates. In female Syrian hamsters, one such behavior is vaginal scent marking. Recent evidence suggests that the neuropeptide oxytocin (OT) may be critical for regulating this behavior. Blockade of OT receptors in the bed nucleus of the stria terminalis (BNST) or the medial preoptic area (MPOA) decreases vaginal marking responses to male odors; lesion data suggest that BNST, rather than MPOA, mediates this effect. However, how OT interacts with sexual odor processing to drive preferential solicitation is not known. To address this issue, intact female Syrian hamsters were exposed to male or female odors and their brains processed for immunohistochemistry for Fos, a marker of recent neuronal activation, and OT. Additional females were injected intracerebroventricularly (ICV) with an oxytocin receptor antagonist (OTA) or vehicle, and then tested for vaginal marking and Fos responses to sexual odors. Colocalization of OT and Fos in the paraventricular nucleus of the hypothalamus was unchanged following exposure to male odors, but decreased following exposure to female odors. Following injections of OTA, Fos expression to male odors was decreased in BNST, but not in MPOA or the medial amygdala (MA). Fos expression in BNST may be functionally relevant for vaginal marking, given that there was a positive correlation between Fos expression and vaginal marking for BNST, but not MPOA or MA. Together, these data suggest that OT facilitation of neuronal activity in BNST underlies the facilitative effects of OT on solicitational responses to male odors.     

Olfactory bulb cells generated in adult male golden hamsters are specifically activated by exposure to estrous females

Two experiments were carried out to test whether cells which are born in adulthood and migrate to the olfactory bulb of adult male golden hamsters are activated during sexual behaviors, to determine the time course over which such responsiveness appears, and to ask whether activation is specific to sexual cues. In the first experiment, adult male hamsters were injected with 5'-bromodeoxyuridine (BrdU, 50mg/kg b.w.) 3 times over the course of one week in order to mark dividing cells. Ten days, three weeks, or seven weeks after the first BrdU injection, the animals were allowed to mate with an estrous female for half an hour before being sacrificed. Confocal analysis of fluorescent immunostaining of BrdU and c-Fos first revealed dual labeled cells in the olfactory bulb 3 weeks after injection of the thymidine analog. In order to determine whether the activation of these newly generated cells is specific to sexual cues, we next compared the incidence of c-Fos expression in newborn (BrdU positive) cells among male hamsters exposed to an estrous female, an aggressive male, a cotton swab containing vaginal secretion from an estrous female hamster (FHVS), a cotton swab containing peppermint, or a cotton swab containing distilled water. In the mitral and glomerular layers of the accessory olfactory bulb, animals exposed to an estrous female had significantly more double labeled cells than did those given other treatments (p < 0.01). In the mitral layer of the main bulb, animals exposed to an estrous female had a significantly higher percentage of double labeled cells than those of other groups, except those exposed to an aggressive male (p < 0.05). No double labeled cells were seen in medial preoptic area (MPOA), medial nucleus of the amygdala (Me), the bed nucleus of the stria terminalis (BNST), or the hypothalamus. Our results indicate that cells born in adulthood are more responsive to cues arising from estrous females than other stimuli, and thus may participate in sociosexual behaviors.     

Conversion of testosterone to estradiol may not be necessary for the expression of mating behavior in male Syrian hamsters (Mesocricetus auratus)

Male sexual behavior is mediated in part by androgens, but in several species, mating is also influenced by estradiol formed locally in the brain by the aromatization of testosterone. The role of testosterone aromatization in the copulatory behavior of male Syrian hamsters is unclear because prior studies are equivocal. Therefore, the present study tested whether blocking the conversion of testosterone to estradiol would inhibit male hamster sexual behavior. Chronic systemic administration of the nonsteroidal aromatase inhibitor Fadrozole (2.0 mg/kg/day) for 5 or 8 weeks did not significantly increase mount latency or reduce mount frequency, intromission frequency, ejaculation frequency, or anogenital investigation relative to levels shown by surgical controls. However, Fadrozole effectively inhibited aromatase activity, as evidenced by the suppression of estrogen-dependent progesterone receptor immunoreactivity in the male hamster brain. The JZB39 anti-progesterone receptor antibody labeled significantly more neurons in brains of sham-treated hamsters than in brains of Fadrozole-treated hamsters. These data suggest that aromatization of testosterone to estradiol is not necessary for normal mating behavior in Syrian hamsters.     

Pre-exposure to female chemosignals or intracerebral GnRH restores mating behavior in naive male hamsters with vomeronasal organ lesions

Chemosensory input is essential for mating in male hamsters and the vomeronasal organ is critical to mating in naive males. In studies to investigate the convergence of vomeronasal chemosensory input and the neurohormone gonadotrophin releasing hormone (GnRH), we have unexpectedly found that pre-exposure to pheromone-containing chemosignals from female hamsters will also eliminate mating deficits normally seen in naive male hamsters with vomeronasal organs removed (VNX). In the present studies, naive-intact and naive-VNX male hamsters were given intracerebroventricular injections of GnRH or saline and exposed to female pheromones found in hamster vaginal fluid (HVF) or to water 40 min prior to a 5 min mating test. VNX males given saline injections and exposed to water had severe mating deficits, but VNX males given saline injections and exposed to HVF mated normally. As shown previously, males given GnRH injections and exposed to water also mated normally. HVF exposure prior to a mating test apparently acted to compensate for the lack of vomeronasal input in these males.     

Acoustic orientation of male Aedes diantaeus during mating

The reaction of males of Aedes diantaeus to different auditory stimuli was studied in natural environment during swarming and under laboratory conditions. The males were attracted by 250-420 Hz that corresponds to the main frequencies of flight tones of sympatric females and the range of sensitivity of male Johnston's organs. Behavioural data show that in natural environment swarming males are attracted by flying conspecific and heterospecific females rather than by dead or immobilized females. Contact with heterospecific females always ended in parting of mosquitoes while contact with cospecific females resulted in pairing. Thus, swarming males of Aedes diantaeus localize females by their flight tones but recognize the females of their own species in contact.     

Vomeronasal damage, not nasopalatine duct damage, produces mating behavior deficits in male hamsters

The experiments reported here show that the deficits in malehamster mating behavior that follow removal of the vomeronasalorgan (Meredith, 1986) cannot be mimicked by palatal damageor suture of the nasopalatine ducts (which connect the nasaland oral cavities). A recent report (Mackay-Sim and Rose, 1986)concluded that deficits in mating behavior in female hamsterscould be caused either by removal of the vomeronasal organ orby suture of the nasopalatine ducts. In the experiments reportedhere, removal of the vomeronasal organs produced severe deficitsin about 40% of sexually naive male hamsters, supporting previousconclusions (Meredith, 1986) that vomeronasal organ damage beforesexual experience is particularly debilitating for male hamsters.Mating behavior as measured here was not impaired in animalswith nasopalatine ducts sutured bilaterally, or in animals withpalatal surgery alone (sham vomeronasal organ removal).     

Testosterone, androstenedione, and androstenediol: effects on the initiation of mating behavior of inexperienced castrated male rats

The effectiveness of testosterone, androstenediol, and androstenedione in initiating sexual behavior in inexperienced castrated male rats was studied. Androgens were injected daily for 32 days at three dose levels (0.3, 1, and 3 mg) for each androgen. Among the three effective androgens tested, testosterone was found to be the most potent one in initiating copulatory behavior while androstenedione was the least potent one.     

Lifespan in male ants linked to mating syndrome

Male ants have long been thought to leave the nest, swarm, mate and die in quick succession (male aggregation syndrome). However, the ancestors of the ants likely used female calling, where females advertise with pheromones for longer lived and presumably free living males. In this study, male lifespan was compared in four species from a Panama rain forest. Males of two species with aggregation syndrome (Atta colombica and Azteca sp.) lived only days after collection at a light trap, and their lifespan failed to increase when supplied sugar water ad libitum. In contrast, two species with female calling syndrome (Ectatomma ruidum and Paraponera clavata) lived up to 116 days when fed. These results link male lifespan to mating systems, and provide a framework to examine variation in how ant colonies invest in males.     

Hemolymph loss during nuptial feeding constrains male mating success in sagebrush crickets

Although costs of mating have been widely documented in females,intrinsic costs of copulation have been poorly documented inmales, and there is little evidence that such costs constrainmale mating success under natural conditions. Male sagebrushcrickets, Cyphoderris strepitans, offer females an unusual somaticfood gift at copulation that may constitute a significant costof copulation: females chew on the ends of the males' fleshyhind wings and ingest hemolymph seeping from the wounds theyinflict. Previous studies have shown that once a male has mated,his probability of obtaining an additional copulation is reducedrelative to that of a virgin male seeking to secure his firstmating. If the future mating prospects of nonvirgin males arediminished because of the costs of copulation, this could stemeither from the resources required to manufacture a new spermatophoreor through the energy needed to replenish hemolymph lost throughfemale wing-feeding. To distinguish between these two alternatives,we experimentally depleted virgin males of varying amounts hemolymphin a way that mimicked hemolymph loss of nonvirgin males, withoutthe attendant costs of spermatophore production. After theyhad been treated, males were released in the field and recapturedover the course of the breeding season to monitor their matingsuccess. Control males mated significantly sooner than did malesdepleted of hemolymph. We conclude, therefore, that the depletionof hemolymph that occurs through female wing feeding is sufficientby itself to diminish a nonvirgin male's ability to secure anothermating.     

Pheromone-induced anorexia in male Syrian hamsters

Transition from long days (LDs) to short days (SDs) triggers seasonal obesity in Syrian hamsters. We report here that SD-exposed males housed near females exhibit obesity resistance, episodic weight loss, and reduced adiposity. Negative energy balance is achieved by reduced eating, elevated motor activity, and increased caloric efficiency without metabolic compensation. Circulating leptin, insulin, testosterone, corticosterone, and cortisol are normal or reduced in obesity-resistant hamsters. When males are housed in chambers that block physical, visual, and auditory, but not pheromonal, signals from females, resistance to seasonal obesity persists. Moreover, inhalation of extracts from pheromone-releasing flank glands of females suppresses eating and weight gain in SD-exposed males. This novel phenomenon, pheromone-induced anorexia, shows that female pheromones play a critical role in the seasonal energy balance of male hamsters. These findings provide a model to study neural and endocrine mechanisms that underlie eating disorders.     

Cell proliferation and survival in the mating circuit of adult male hamsters: effects of testosterone and sexual behavior

The transient actions of gonadal steroids on the adult brain facilitate social behaviors, including reproduction. In male rodents, testosterone acts in the posterior medial amygdala (MeP) and medial preoptic area (MPOA) to promote mating. Adult neurogenesis occurs in both regions. The current study determined if testosterone and/or sexual behavior promote cell proliferation and survival in MeP and MPOA. Two experiments were conducted using the thymidine analog BrdU. First, gonad-intact and castrated male hamsters (n = 6/group) were compared 24 h or 7 weeks after BrdU. In MeP, testosterone-stimulated cell proliferation 24 h after BrdU (intact: 22.8 ± 3.9 cells/mm², castrate: 13.2 ± 1.4 cells/mm²). Testosterone did not promote cell proliferation in MPOA. Seven weeks after BrdU, cell survival was sparse in both regions (MeP: 2.5 ± 0.6 and MPOA: 1.7 ± 0.2 cells/mm²), and was not enhanced by testosterone. In Experiment 2, gonad-intact sexually-experienced animals were mated weekly to determine if regular neural activation enhances cell survival 7 weeks after BrdU in MeP and MPOA. Weekly mating failed to increase cell survival in MeP (8.1 ± 1.6 vs. 9.9 ± 3.2 cells/mm²) or MPOA (3.9 ± 0.7 vs. 3.4 ± 0.3 cells/mm²). Furthermore, mating at the time of BrdU injection did not stimulate cell proliferation in MeP (8.9 ± 1.7 vs. 8.1 ± 1.6 cells/mm²) or MPOA (3.6 ± 0.5 vs. 3.9 ± 0.7 cells/mm²). Taken together, our results demonstrate a limited capacity for neurogenesis in the mating circuitry. Specifically, cell proliferation in MeP and MPOA are differentially influenced by testosterone, and the birth and survival of new cells in either region are not enhanced by reproductive activity.     

Olfactory sensory input increases gill ventilation in male round gobies (Neogobius melanostomus) during exposure to steroids

In teleostean fish, ventilation increases have been observed in response to low dissolved oxygen levels, visual stimuli, and gustatory cues. However, olfactory sensory input may also stimulate gill ventilation rate. We investigated whether olfactory sensory input mediates gill ventilation responses, as suggested by the observation that steroidal compounds detected by the olfactory system elicited increases in opercular activity in the perciform teleost, the round goby (Neogobius melanostomus). Close parallels between gill ventilation and olfactory responses, led us to conduct an empirical study that used two different olfactory sensory deprivation techniques to seek a causal relationship between olfactory epithelial activity and hyperventilation. Chemical lesion of olfactory sensory neurons or mechanical occlusion of the nasal cavities inhibited gill ventilation responses of reproductive male round gobies to estrone (1,3,5(10)-estratrien-3-ol-17-one) and to ovarian extracts. This direct evidence demonstrates the role of olfactory sensory input for the gill ventilation response to putative reproductive pheromones and may represent an important regulatory mechanism for odorant sampling during pheromone communication.     

Female and male contribution to egg size in salmonids

Egg size contributes to other life history traits of an individual. It is traditionally considered as a maternally determined characteristic to which the male does not have any direct contribution. However, a recent finding in insects suggests that males can affect egg size also directly. In fish, the male effect could take place only during egg swelling, as the final egg size is reached after that. We studied egg size in four freshwater salmonid species (the land-locked Atlantic salmon, the brown trout, the Arctic charr and the lake trout) right after fertilisation (initial egg size) and after the swelling phase (final egg size). The results showed that the final egg size is affected not only by the initial egg size but also by both the female and the male through the process of egg swelling. This study suggests that paternal contribution may form a previously largely ignored source of variation in early life history traits in salmonid fish.     

Testosterone and photoperiod interact to regulate locomotor activity in male hamsters

Testicular size, plasma testosterone levels, copulatory behavior, and daily locomotor activity are reduced in male hamsters after 10 weeks of exposure to short days. The role of testosterone in the short day-induced decline in locomotor activity was investigated, determining whether or not photoperiod could alter the effect of testosterone on activity. Castrated adult hamsters were allowed to acclimate to running wheels (wired to digital counters) and then were kept on either long (L:D 14:10) or short (L:D 6:18) days for 60 days. On Day 60, half of the animals on each light cycle were implanted with 12-mm-long testosterone-filled Silastic capsules; half received empty capsules. Digital counting of wheel-running activity continued for another 140 days. Blood samples taken on Day 200 confirmed L:D 14:10 and L:D 6:18 testosterone-treated hamsters had equivalent plasma testosterone levels. After an initial decline in activity, L:D 14:10 animals exhibited a progressive rise in mean running activity (from ~2000 to ~5000 wheel revolutions per day) through 100 days after the initiation of testosterone treatment. In contrast, activity levels in testosterone-treated L:D 6:18 animals remained uniform (~2000 wheel revolutions per day) during this time, indicating exposure to short days rendered the hamsters less sensitive to the stimulatory effect of testosterone on activity. Of further interest was a marked increase in activity after 160–200 short days in animals treated with either testosterone-filled or empty capsules. It appears the total amount of daily locomotor activity in the hamster is modulated by circulating testosterone levels in a manner which is dependent upon the environmental photoperiod.     

Neuroendocrine changes in male hamsters following photostimulation

Transfer of gonadally regressed male golden hamsters from a short (5 L:19 D) to a stimulatory (14 L:10 D) photoperiod elicits, within 24 hr, significant changes in hypothalamic dopamine, serotonin, and possibly norepinephrine metabolism. Hypothalamic LHRH content was significantly elevated in short-photoperiod animals, but within 24 hr of transfer to a 14:10 photoperiod, LHRH declined to levels not different from those in hamsters maintained continuously in a long photoperiod. Plasma FSH levels were also significantly elevated within 24 hr of transfer, but increases in plasma LH were somewhat slower. Chronic treatment with the tyrosine hydroxylase inhibitor, alpha-methyl tyrosine (alpha MPT), which inhibits catecholamine synthesis, blocked the effect of a stimulatory photoperiod on plasma FSH levels, while treatment of 5:19 hamsters with the catecholamine precursor, L-dopa, mimicked the effects of photostimulation on plasma FSH levels. Testicular weights were not affected by alpha MPT or L-dopa treatment for 1 week. From these data, it appears that endocrine events associated with photoperiod-induced testicular recrudescence are under the control of hypothalamic neurotransmitters.     

Strategic male mating effort and cryptic male choice in a scorpionfly

In animal species with high male mating effort, males often find themselves in a dilemma: by increasing their mating effort, the gain from each copulation increases but simultaneously reduces available resources and, thus, the opportunity for future copulations. Therefore, we expect males to spend less reproductive resources on matings that provide low reproductive potential, thereby saving resources for future copulations, possibly with high-quality females, a sort of cryptic male choice. However, the strength of the trade-off between investment in a current mating and resources available for future matings must not be the same for all males. Males with relatively high mating costs should allocate their limited resources more cautiously than males with more plentiful resources. Here, we examine this prediction in the scorpionfly Panorpa cognata. Prior to copulation, males produce a large salivary mass on which females feed during copulation. We show that the production of larger salivary masses leads to longer copulations. Moreover, the size of the salivary gland and salivary mass increases with increasing male condition. However, males in poor condition make a relatively higher mating investment than males in good condition. We therefore expect male condition to influence cryptic male choice. In accordance with our hypothesis, only males in poor condition choose cryptically, producing larger salivary masses in copulations with females of high fecundity.