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Prolonged copulatory behavior facilitates pregnancy success in the musk shrew.

Female musk shrews (Suncus murinus) are sexually receptive and actively seek contact with males during the first ten days of pregnancy. In the series of experiments reported here we examined the effects of prolonged copulatory behavior on pregnancy success. Females received multiple ejaculations either on the same day or over the course of up to six days. When ejaculations were distributed over an interval of several days, the number of females delivering litters was significantly increased. The same outcome was found regardless of whether the first fertile mating was followed by subsequent copulations with normal or vasectomized males. One ejaculation followed by daily contact with soiled bedding taken from a male's cage did not increase pregnancy rates. Three ejaculations received over a three-day interval caused significantly more females to ovulate than three ejaculations given on the same day. The results demonstrate that prolonged copulatory behavior, which includes vaginocervical stimulation, facilitates ovulation and pregnancy in this species.

Animals↗

Androgen and estrogen action in the preoptic area and activation of copulatory behavior in quail.

The sites of androgen and estrogen action on sexual behavior were studied in the preoptic area of castrated male Japanese quail by stereotaxic implantation of hormones, antihormones and metabolism inhibitors. The first experiment demonstrated that bilateral implantation of the aromatase inhibitor, androstatrienedione (ATD), in the sexually dimorphic nucleus (POM) of the preoptic area can completely suppress the behavioral activation produced by a systemic treatment with testosterone. The effects of ATD were only observed if the implants were located in the POM. In the second experiment, implants in the POM of the synthetic estrogen, diethylstilbestrol, restored copulatory behavior in castrated males while implants of the synthetic nonaromatizable androgen, methyltrienolone, were almost ineffective. During the third experiment, the activating effects of a systemic treatment with testosterone were blocked by stereotaxic implants in the POM of the antiestrogen, tamoxifen, or the antiandrogen, flutamide. The effects of tamoxifen were more pronounced than those of flutamide. In addition, tamoxifen was active in all parts of the POM while a behavioral inhibition was observed only for flutamide implants which were located in the caudal part of the nucleus. Taken together, these results demonstrate that the sexually dimorphic POM is the area where the behaviorally active estrogenic metabolites of T have to be produced. The estradiol derived from T aromatization presumably acts within the POM to activate copulation as demonstrated by the effectiveness of DES implanted in this region. Androgens also have a direct action on sexual behavior as suggested by the partial inhibition observed in flutamide-treated birds. It is, however, suggested that androgens and estrogens do not act in the same brain area to activate behavior.

Androgens↗

Influence of daylength on male hamster sexual behavior: masking effects of testosterone.

Exposure of male hamsters to short photoperiods for 6-8 weeks cause deficits in sexual behavior with receptive females. The present experiment tested the hypothesis that short photoperiodic effects on behavior could be masked in the presence of chronic and stable levels of testosterone. Males were castrated and administered Silastic capsules of testosterone while housed in long (16L:8D) or short (8L:16D) photoperiodic conditions for 7 weeks. Sexual behavior tests at this time indicated that the short photoperiod males copulated less well, but group differences were not robust. Testosterone capsules were then removed and half the animals in both 16L:8D and 8L:16D were transferred to the opposite photoperiod. Sexual behavior was tested 18 days later as the effects of this functional castration developed. These tests indicate that photoperiodic effects were much more obvious in the absence of testosterone than they were during week 7 tests when testosterone was still present. The behavior of the males that were transferred from one photoperiod to the other demonstrated that exposure to the short photoperiod for only 18 days was not sufficient to generate short photoperiod-like sexual behavior deficits. In contrast, exposure to the long photoperiod for 18 days was sufficient to reverse short photoperiodic effects that had already developed.

Animals↗

Penile spines affect copulatory behaviour in a primate (Callithrix jacchus).

Androgen-dependent, keratinized "spines" occur on the glans penis in many rodents, primates and other mammals. Since penile spines overlie dermal tactile receptors, they may play a role in copulatory behaviour. An experiment was conducted to test this hypothesis. Sixteen sexually experienced adult male marmosets were paired with ovariectomized females before, and after, removal of penile spines (using thioglycollate cream applied to the glans under anaesthesia) or a sham operation. Spine removal resulted in an increased duration of preintromission pelvic thrusting (mean +/- s.e.m. from 6.87 +/- 1.09 to 14.94 +/- 3.32 s, p = 0.05) and of intromitted thrusting (from 1.73 +/- 0.11 to 2.0 +/- 0.11 s, p less than 0.05). Three males exhibited partial intromissions during some postspinectomy tests, an effect which had not been observed prior to the operation. Sham operations had no behavioural effects. Results indicate that penile spines play a significant (but not indispensible) role in sensory feedback during copulation in this primate species.

Androgens↗

Lesions of the medial amygdala produce severe impairment of copulatory behavior in sexually inexperienced male rats.

The effects of amygdaloid lesions on masculine copulatory behavior were examined in male rats. Sexually inexperienced male rats were castrated and subjected to bilateral lesions in one of the following areas: the medial amygdala, the cortical amygdala, or the basolateral amygdala. Three weeks later, all rats received implantation of silastic capsules containing testosterone. Then, four observations of copulatory behavior were carried out every 5 days following the implantation of testosterone. Rats with medial amygdala lesions showed a severe deficit of copulatory behavior, whereas rats with basolateral amygdala lesions showed no change in the performance of copulation. As for rats with cortical amygdala lesions, although their copulatory behavior was impaired, the effect was confined to a deficit in intromission and ejaculation responses. These findings suggest that the medial amygdala plays a critical role in regulating masculine sexual behavior in the rat.

Amygdala↗

Copulatory behavior and reflexive penile erection in rats after section of the pudendal and genitofemoral nerves.

Two experiments probed the roles of the pudendal and genitofemoral nerves in sexual behavior. Male rats were tested for copulatory behavior and reflexive erections after transection of the sensory (SP) or motor (MP) branches of the pudendal nerves and, in Experiment 2, section of the genitofemoral (GF) nerve alone or in combination with the SP nerves. Damage to the GF nerve had no apparent effects. Division of the SP nerve severely impaired the ability of males to achieve intromission, and hence ejaculation, and reflexive erections were drastically reduced. However, this treatment caused impairments less complete than those previously described for more distal deafferentation by section of the dorsal penile nerves or by application of topical anesthetics to the glans penis. Penile autotomy following SP section was delayed but not avoided by daily treatment with amitriptyline. Transection of the MP nerves had the most drastic effects, preventing reflexive erections, although some intromissions (but no ejaculations) occurred during copulation. Although urinary function was also disrupted by MP transection, the impairment in sexual function is tentatively ascribed to chronic hyperinvolution of the penile corpora following loss of phasic vasoconstrictive stimulation normally supplied via the MP nerves. The pattern of results suggests that the pudendal nerves make different contributions to penile erection in different contexts.

Animals↗

Sexual and aggressive behaviour of adult male marmosets (Callithrix jacchus) castrated neonatally, prepubertally, or in adulthood.

During early postnatal development, males of some primate species exhibit higher levels of plasma testosterone than females. The possible significance of this for behavioural development was examined in male marmosets castrated neonatally (NC), prepubertally (PC), or in adulthood (AC) and reared in their natal groups. Age-matched intact (AI) adult males served as controls. All males were removed from their natal groups as adults for pair testing with unfamiliar conspecific males and females. NC males did not mount, but were frequently aggressive during tests with females. NC males also elicited aggression from females but rarely from intact males. PC and AC males attempted copulation during the majority of tests with females; less aggression occurred but more so in tests with PC males than between females and AC males. Aggression between PC or AC males and intact males was much more frequent than when NC males met intacts in paired encounters. Results show that secretion of testicular hormones during infancy in male marmosets has important effects upon the development of sexual and aggressive behaviour. Castration in infancy has measurably different effects upon behavioural development than castration performed later in life (prepubertally or in adulthood).

Aggression↗

Progesterone modulation of androgen-dependent sexual behavior in male rats.

The present study examines the effects of physiological levels of progesterone (P) on copulatory behavior in sexually naive male rats. Two weeks after gonadectomy males were implanted with either empty Silastic capsules (BL) or Silastic capsules containing testosterone (T), P, or both (P+T). When tested with an estrous female, all of the gonadally intact males (intact) and none of the BL controls exhibited mounting/intromission behaviors. Mounting was observed in 75% of the T-alone males. More than half (64%) of the P-alone males and 100% P+T males exhibited mounting. In most cases, mounting was followed by intromission responses. Subsequently, intact and gonadectomized males received daily injections of the P antagonist RU486 along with hormone treatment. After receiving RU486, only 63% of the intact males and 71% of the T-alone males mounted successfully. The facilitatory effects of P on copulatory behavior were completely abolished by RU486 treatment. The present studies provide the first evidence in mammals suggesting that P-dependent mechanisms influence neurochemical pathways involved in copulation.

Androgens↗

Physiology and mechanics of rat levator ani muscle: evidence for a sexual function.

The levator ani (LA) of male rodents is a classic model tissue for the study of hormone-muscle interactions, although its functions remain unknown. Recordings during copulation from chronic electromyographic (EMG) electrodes in the LA and bulbospongiosus (BS) revealed that EMG activity in the LA and BS was tightly coordinated. The LA was not active during noncopulatory behaviors, including the 1-min interval surrounding defecation. Electrical stimulation of the LA motor nerves increased penile bulb pressure. Increases in penile bulb pressure following BS nerve stimulation were markedly attenuated after LA denervation and were reduced further by LA removal. Stimulation of the LA nerve yielded insignificant changes in rectal pressure. Perineal motion analysis demonstrated that the LA acts upon the penile bulb and the surrounding BS exclusively. Apparently the rodent LA muscle is an active component in a highly coordinated neuromuscular system augmenting penile erection and, contrary to its name, is most unlikely to participate in alimentary function.

Animals↗

Male reproductive systems under chronic fluoxetine or trimipramine treatment.

Adult male Long-Evans rats (n = 9 per group) received daily exposure for 4 weeks to fluoxetine (0.75 mg FLUOX/kg body weight) or trimipramine (1.6 mg TRIMI/kg body weight). Separate tests of copulation, sexual motivation, and intermale aggressive behaviors were used to evaluate functional changes during chronic exposure to either typical or atypical antidepressant drugs with more or less serotonin specificity. Circulating hormones, primary and secondary sex structures, and concentrations of dopamine (DA) and serotonin (5-HT) from mesolimbic tissue were assessed at necropsy. Results of tests with estrous females and untreated males revealed progressive disruption to sexual performance and aggressive responsiveness over time of treatment with TRIMI and, to a lesser extent, with FLUOX. By contrast, motivation, testosterone, and all measures of reproductive physiology were indistinguishable from controls. Ratios of transmitter metabolites relative to the parent compounds indicated similar reductions of 5-HT turnover with FLUOX and TRIMI. However, influences on DA turnover were significantly less with FLUOX than with TRIMI. Conclusions are that long-term intervention with antidepressant drugs may disrupt sociosexual exchanges without compromising male rats' interest in sexual contact or integrity of their reproductive physiology. Lessened disruption of sociosexual behaviors with this regimen of chronic FLUOX treatment may be related to the greater selectivity on serotonin relative to dopamine turnover.

Adrenal Cortex Hormones↗

Scanning electron microscopic study of hedgehog uteri.

Ultrastructural studies of hedgehog uteri (Erinaceus europaeus L.) have been made using animals in anestrus, in estrus and in estrus after sojourn of a week with a male. In estrus and anestrus the uterine epithelium is homogeneous, regularly interrupted by orifices of glands. It is composed of microvillous cells only. Microvilli decrease in number and length in anestrus. A new type of cell, a ciliated cell, appears after copulation. Probable correlation of ultrastructural aspects of endometrium with hormonal situation is discussed.

Animals↗

The male copulatory apparatus in an opisthobranch mollusc, Runcina.

The copulatory apparatus of a primitive opisthobranch, Runcina, is described. The apparatus is comprised of the following organs: the spermatic bulb, the prostrate, the penis and the penial sac. The spermatic bulb wall consists of cuboidal epithelium with forked microvilli and densely arranged cilia. Prior to copulation the interior is tightly packed with sperm. The prostate is lined with alternating glandular and supporting cells, the latter being compressed but with a mushroom-shaped apex bearing a few forked microvilli and many cilia. The glandular cells produce differing secretions, each cell producing a single type. Large paracrystalline structures enclosed in cells close to the penial area are particularly striking. Considerable amounts of the secretory products are accumulated in the protruding cell apices. One type of inclusion is found at a later stage, packed around the sperm mass within the spermatophore; its function, and the fate of the other secretions is not yet clear. The epithelium of the penis is of more columnar structure, covered with forked microvilli and extremely long cilia which are anchored by long rootlets in the cells. Some of the cells contain large electron-dense secretory granules, others hold accumulations of small secretory vesicles in their apices. It seems likely that these contribute towards the outer layer of spermatophore. The wall of the penial sac is lined by one to two rows of flat-cuboidal cells bearing sparse forked microvilli and cilia.

Animals↗

Brain localization of cholinergic influence on male sex behavior in rats: agonists.

Cholinergic agonists were microinjected into either the lateral ventricle or the preoptic area of sexually experienced male rats. In Experiment 1 carbachol, injected into the lateral ventricles, delayed the initiation of sexual behavior. When injected into the preoptic area, carbachol again delayed the onset of copulation, but these delays were shorter than after ventricular injections. In addition, preoptic injections reduced the number of intromissions preceding ejaculation. In Experiment 2 ventricular injections of the muscarinic agonist oxotremorine again delayed initiation of sexual behavior and also slowed its rate. However, oxotremorine injections into the preoptic area, through cannulae angled to miss all ventricles, only decreased the number of intromissions preceding ejaculation. These data suggest that cholinergic synapses in proximity to the ventricles may decrease sexual arousal, while cholinergic mechanisms in or near the preoptic area may reduce ejaculatory threshold.

Animals↗

Copulatory pelvic thrusting in the male rat is insensitive to the perispinal administration of glycine and GABA antagonists.

The role of the inhibitory neurotransmitters glycine and GABA in the pacing of pelvic thrusting during copulation was assessed in male rats by an accelerometric technique. Either strychnine, an antagonist of glycine (10 micrograms), bicuculline, an antagonist of GABA (1 microgram), or a combination of strychnine (5 micrograms) plus bicuculline (0.3 microgram), and saline as control, were administered intrathecally to sexually active males. Administration of the antagonists either alone or in combination, at these dose levels, produced sensory effects (skin hyperalgesia, scratching or biting the skin) in all rats. Generalized motor seizures occurred in only a few animals. The incidence of ejaculations, but not of mounts, tended to decrease after treatment with the amino acids antagonists. On the other hand, the values of the instantaneous frequency, duration, and rhythmicity of the copulatory thrusting movements performed during mounts, intromissions or ejaculations did not differ significantly from the values obtained under saline treatment. These findings indicate that the duration and rhythmicity of copulatory movements in the male rat are either controlled by synapses that are insensitive or inaccessible to strychnine and bicuculline, or these copulatory components are independent of glycinergic and GABAergic control and are under the control of other neurotransmitter systems.

Animals↗

Pharmacological manipulation of anxiety and male rat sexual behavior.

Several anxiolytic/anxiogenic treatments were evaluated on male rat sexual behavior. The anxiolytic drug diazepam (1.0 mg/kg) inhibited copulatory behavior as indicated by an increase in the number of mounts preceding ejaculation, prolongation of the ejaculation latency and the postejaculatory interval. These changes were not accompanied by alterations in motor coordination as tested on a treadmill apparatus. A lower dose of diazepam (0.5 mg/kg) did not affect the sexual behavior. The anxiogenic drug Zk 39106 (2 and 4 mg/kg) facilitated the copulatory behavior by reducing the number of intromissions preceding ejaculation. A higher dose of Zk 39106 (8 mg/kg) inhibited sexual behavior. The selective benzodiazepine antagonist, Ro 15-1788 (5 and 10 mg/kg), did not modify male sexual behavior, but effectively antagonized the effects of Zk 39106 and diazepam. The administration of Zk 39106 (2 mg/kg) reversed the inhibitory action of diazepam (1.0 mg/kg) on copulation; however, diazepam did not prevent the facilitory effect of Zk 39106. The data are discussed in terms of the possible relationship existing between anxiety and masculine sexual behavior.

Animals↗

DOI-induced inhibition of copulatory behavior in male rats: reversal by 5-HT2 antagonists.

Relatively little is known regarding the role of 5-HT2 receptor activity in male rat sexual behavior. Previous work has yielded equivocal results, and both facilitation and inhibition of copulation have been reported to follow administration of selective 5-HT2 antagonists. In the present series of experiments, the ability of a variety of 5-HT2 antagonists to block inhibition induced by the 5-HT2/5-HT1C agonist DOI was examined. Systemic ritanserin, pirenperone and ketanserin all potently blocked DOI-induced (1.0 mg/kg SC) inhibition of mounts, intromissions and ejaculations. None of these drugs influenced the sexual behavior of the male rats when given alone in doses that effectively blocked DOI-induced inhibition. In addition, unlike ritanserin and ketanserin, pirenperone produced a biphasic effect on DOI-induced inhibition, exhibiting a diminished blockade at higher doses. This may be due to activity at receptors other than 5-HT2. Overall, the present data suggest that activity at 5-HT2 receptors mediates an inhibition of male rat sexual behavior.

Amphetamines↗

Increased sexual behavior in male Macaca arctoides monkeys produced by atipamezole, a selective alpha 2-adrenoceptor antagonist.

The effect of a highly selective and potent alpha 2-adrenoceptor antagonist, atipamezole, on sexual behavior was studied in three stumptail macaques (Macaca arctoides). Following IM administration of atipamezole or saline control, the behavior of the male monkey with a female monkey was observed for 30 min. Atipamezole dose dependently (0.01-0.15 or 0.30 mg/kg) produced a significant increase in the number of ejaculations in all three monkeys, including an old one with decreased sexual activity in control conditions. Both ejaculations obtained by copulation and masturbation were increased. It is concluded that atipamezole is effective in increasing sexual behavior in male stumptail monkeys.

Adrenergic alpha-Antagonists↗

Participation of opiatergic, GABAergic, and serotonergic systems in the expression of copulatory analgesia in male rats.

Copulation in the male rat provoked an abrupt and significant rise in the threshold to induce vocalization by electrical shock to the tail (copulatory analgesia, CA). The possible effect on CA of the intrathecal (IT) administration of receptor antagonists to neurotransmitters participating in nociception was ascertained in this study. CA was significantly reduced, though not abolished, by IT injections of either naloxone, picrotoxin, or methysergide, but not by strychnine or yohimbine. This analgesic effect was achieved without significantly altering copulatory behavior. Results suggest that both brain and spinal systems participate in the development of CA. Brain effects would be mediated by descending serotonergic fibers, although intrinsic spinal systems would involve both opiate and GABA interneurons.

Analgesia↗