Androgen regulation of the motor copulatory pattern in the male New Zealand white rabbit

The effect of castration and testosterone propionate (TP) treatment on the motor copulatory pattern was studied by an accelerometric technique in five sexually experienced New Zealand white male rabbits. This technique permits the oscillographic recording of thrusting frequency, rhythmicity, and amplitude of the pelvic movements occurring during copulation. Castration resulted in a marked decrease in sexual activity in all rabbits. Mounting, including occasional intromissions, was retained by four of the rabbits for periods ranging from 2 to 15 weeks. Castration did not affect mount duration, but decreased strength and frequency of pelvic thrusting. Diminution in the frequency of pelvic thrusting was mainly due to intercalation of pauses within the mounting trains. The change in the rhythm of pelvic thrusting was related to the failure of most mounts performed by castrated rabbits to stimulate lordosis in the female. Testosterone propionate (TP, 10 mg daily for 15 days) restored mounting activity and increased strength and frequency of pelvic thrusting in all rabbits. It is concluded that TP, besides stimulating sexual motivation, regulates the vigor and rhythm of pelvic movements during copulation in the rabbit.     

Characteristics of the motor and genital copulatory responses of the male hamster

The temporal and dynamic characteristics (duration, frequency, rhythmicity, and vigor) of pelvic thrusting displayed by 17 male golden hamsters during their copulatory behavioral responses: mounts, intromissions, ejaculations, and long intromissions, as well as their temporal correlation with the genital contacts established by intravaginal penile insertion, were studied by an accelerometric and polygraphic technique. Pelvic thrusting in all copulatory behavioral responses appeared as series or trains of rhythmical, synchronic, vigorous movements. The various pelvic thrusting trains lasted around 1 s on average, but those of mounts were significantly longer than the trains displayed in the other behavioral responses. The frequencies of pelvic thrusting were similar in all responses (around 15 thrusts per s) excepting mounts which had lower values. In intromission, ejaculation, and long intromission responses, when penile insertion occurred, pelvic thrusting either was interrupted or showed changes in its characteristics: penile insertion was related to a period without thrusting in intromissions, to a series of intravaginal thrusting of higher frequency (16.4 thrusts per s) and lower vigor in ejaculations, and to a prolonged period of 6 to 25 s of slow intravaginal pelvic thrusting (1–2 thrusts per s) in long intromissions. Penile insertion lasted longer in ejaculations than in intromissions and it was significantly shorter in both of these responses than in long intromissions. These results provide information about some dynamic aspects of sexual behavior in hamsters, as well as a temporal correlation between the motor and genital components of this behavior.     

Crowding pregnant mice affects attack and threat behavior of male offspring

Attack and threat behavior of adult male offspring of female mice crowded during the final third of pregnancy was investigated. In 5-min test pairings with an anosmic "standard opponent" which had 50 microliter of male mouse urine applied to its fur, the prenatally stressed group of males showed significantly less attack behavior; attack latency was longer and number of attacks, bites, amount of time spent attacking, and composite aggression scores were all lower, compared with the control group. Similarly, less threat behavior was observed in offspring from crowded dams; there were lower frequencies of tail rattles, rough grooms, and upright threats. Additionally, proportionally fewer males in the prenatally stressed group attacked or displayed threats. A second experiment was designed to investigate the effects of exogenous androgen on the aggressiveness of males from crowded mice: testosterone propionate administration (500 micrograms/animal/day, for 5 days prior to testing) abolished differences both in the proportion of males from crowded mice that fought and also apparently abolished differences in intensities of attack and threat behavior between groups. However, trends toward reduced aggression in prenatally crowded males remained. More detailed analysis of these responses, based only on animals that displayed aggression, revealed significantly reduced intensity of aggression in offspring from females crowded during pregnancy, indicating that testosterone propionate therapy did not completely restore this behavior. In order to reduce postnatal effects due to possible differences in mothering, all offspring were fostered to untreated mothers at birth. The results are discussed in terms of in utero exposure of male fetuses of crowded dams to stress-liberated adrenal steroids of maternal origin, and the possible consequences for the endocrine integrity of these offspring.     

Estrogen-activated sexual behavior in male rats

Daily injections of 100 μg estradiol benzoate activated the whole pattern of sexual behavior in castrated sexually experienced male rats. If compared to rats treated daily with 100 μg testosterone propionate, the estrogen-treated males tended to have longer latencies and more mounts and intromissions prior to ejaculation. Fifty micrograms of estradiol benzoate stimulated the display of mounts and intromissions in prepuberally castrated male rats. No peripheral effects of the estrogen treatment were noted. These results suggest that estrogen has central “androgen-like” effects, but no such effects in the periphery. Estrogen treatment (5, 50, and 200 μg/kg for 3 weeks) of intact sexually experienced male rats resulted in testicular atrophy and loss of body weight, but had no significant effects on the sexual behavior.     

Short-latency effect of testosterone on copulatory behaviour and ejaculation in sexually experienced intact male rats

The effect of a single injection of testosterone on the number of stimuli required to trigger ejaculation and on copulatory behaviour was investigated in sexually experienced intact male rats. An intraperitoneal injection of 100 microgram free testosterone 1 h before testing facilitated the copulatory behaviour and decreased the number of mounts and intromissions needed for ejaculation; the latter effect was obtained in a test in which a constant interval was kept between each intromission.     

Aromatase inhibition depresses ultrasound production and copulation in male hamsters

Rates of ultrasound production and copulatory behavior were observed in castrated male hamsters maintained on 100 micrograms/day of injected testosterone propionate (TP). Groups matched on their initial levels of behavior received either continued treatment with TP alone, or TP together with 6 mg/day injections of the aromatase inhibitor 1,4,6-androstatriene-3,17-dione (ATD). Testing at 11-15 days after the start of these treatments revealed deficits in the sexual behaviors of the subjects in the latter group. Specifically, these males showed lower rates of ultrasound production and intromission during, as opposed to before, treatment with ATD. These results support previous work suggesting that aromatization plays significant roles in the mediation of androgenic effects on both the courtship and copulatory behaviors of male hamsters.     

Erectile function and bulbospongiosus EMG activity in estrogen-maintained castrated rats vary with behavioral context.

Electromyographic (EMG) activity in the bulbospongiosus muscles (BS) was recorded to monitor potential castration-induced alterations in muscle activity during copulation and reflexive erections. EMG recordings were made from intact male rats and from castrated rats maintained from 7 to 50 days on estradiol benzoate (300 micrograms/day) or testosterone (200 micrograms/day). Despite a 40-50% postcastration reduction in the weight of the BS and accessory sexual glands in estrogen-treated rats, the pattern of EMG activity during copulation was similar across groups. In estradiol-treated males, the EMG burst frequency during mounts and burst duration during intromissions exceeded the parameters of intact males and of castrated males maintained on testosterone. Between intromissions, and following ejaculatory patterns, estrogen-treated males displayed spontaneous muscle bursts accompanied by visually confirmed erection of the glans penis, but these males quickly lost the capacity for reflexive erections. These data demonstrate that despite castration-induced atrophy of the penile muscles and, presumably, their spinal motor nuclei, the motor output to these muscles is maintained following androgen removal. The capacity for substantial penile erection is retained during copulation long after reflexive erections have diminished.     

Female-typical sexual behavior of rhesus and defeminization by androgens given prenatally

Adult rhesus monkeys were observed in standardized tests for female-typical sexual and related social responses. In the first experiment reported, 7 castrated males and 5 spayed females were paired with each of 4 intact males on two occasions following intramuscular injection with estradiol benzoate (EB) (6 micrograms/kg X 14 days) and on two other occasions without such treatment. In tests without EB, males and females did not behave differently toward the intact male partners, and all responses were displayed at low frequencies. In tests with EB, females showed reliably higher frequencies than males of approaching, sitting close to, grooming, and soliciting, and they presented to a higher proportion of the male partner's sexual contacts. EB reliably increased the frequency of display of all of these same five responses in females but not in castrated males. The intact male partners displayed reliably fewer approaches, sexual contacts, mounts, intromissions, and ejaculations to castrated males than to spayed females regardless of estrogenization. In a second experiment 10 intact adult pseudohermaphroditic females and 6 intact control females were tested following EB injections with each of the same 4 intact males. Pseudohermaphrodites were experimentally produced by injecting pregnant females with either testosterone propionate (TP) or dihydrotestosterone propionate (DHTP). Pseudohermaphrodites, regardless of type of androgen used in their production, showed reliably fewer solicits than controls to male partners. Moreover, they displayed most of the other responses at lower average frequencies than controls. Frequencies of intromission and ejaculation by intact male partners were reliably lower with pseudohermaphrodites than with control females, but frequencies of approach, sexual contact, and mount were not reliably different. We conclude that in this testing and measurement situation male and female rhesus monkeys differ markedly in the degree of expression of female-typical sexual behaviors, and genotypic males are behaviorally less responsive to estrogens than females. Exposing genotypic females to androgens during fetal life decreases the expression of female-typical, estrogen-influenced responses, and the effect is most pronounced on those soliciting responses that subserve proceptivity.     

Male copulatory patterns in the lesser bushbaby (Galago moholi) in captivity

I observed seven pairs of sexually mature, captiveGalago moholi during the females’ ovarian cycles to determine the pattern of male copulatory behavior and its relationship to female receptivity. In this species, a copulatory lock was not apparent, a variety of thrusting patterns was displayed, and the male was capable of ejaculating following a single mount and intromission and may have ejaculated more than once during the 30-min test. Male mating indices did not differ significantly on consecutive days of the mating period, though declines in certain mating latencies on the second and third days of mating suggested increased female receptivity. Mating indices did not differ notably between the first and the second copulation within a test, though both showed some differences compared to the third copulation. InG. moholi, male copulatory behavior is typical of a prosimian species with a dispersed — nongregarious — mating system, in which copulation has a single prolonged intromission that extends into the postejaculatory period. This is characteristic of species in which more than one male has access to an estrous female.     

Reversal of reproductive deficiency in the hpg male mouse by neonatal androgenization.

Some aspects of reproductive function in the GnRH-deficient hypogonadal (hpg) mutant mouse can be restored by transplanting normal fetal brain tissue containing GnRH cells into the central nervous system of adult hpg mice. However, hpg males showing physiological response to the graft fail to display sexual behavior and are infertile. We hypothesized that the reproductive deficit of these males is due to insufficient perinatal exposure to testicular androgens as a consequence of the GnRH deficiency. To test this hypothesis we androgenized hpg males by giving them neonatal injections of testosterone propionate (TP). Controls consisted of hpg males not androgenized neonatally and of normal males. All three groups received a TP implant in adulthood, and their copulatory behavior and reproductive capability were recorded. In addition, other hpg males, not androgenized neonatally, received fetal brain transplants containing GnRH neurons and were also tested for copulatory behavior and reproductive capability before and after receiving a TP implant. Three of 8 neonatally androgenized hpg males expressed the full repertoire of male sexual behavior, including intromission and ejaculation, and sired several litters. Three of 7 control hpg males that were not androgenized neonatally but received TP implants in adulthood also displayed mounting and intromission, but there was no evidence of ejaculation, and these males failed to impregnate normal females. Of the 8 hpg males that responded to a fetal transplant with testicular growth, only 1 displayed mounting behavior. However, when given a TP implant, 4 of 8 hpg males with grafts displayed mounting and intromissions.(ABSTRACT TRUNCATED AT 250 WORDS)     

Study of pharmacological activation of the male sexual behavior

The model of sexual exhaustion of male rats was used in the study. The specific aim was to compare the effects of cocaine on the mount latency, number of mounts, intromissions and ejaculations during the entire copulatory cycle with the same indices during the first 30 min of observation (the latter is most frequently used in laboratory settings). Experiments consisted of four 3-day test periods. On days 1 and 2, male rats were allowed to interact with receptive females until the satiation criterion was reached (30 min without mounts). On day 3, male rats were injected with cocaine (5, 10, or 30 mg/kg, i.p.) or its vehicle 15 min prior to testing. Cocaine administration in a dose-dependent way increased the total number of mounts and ejaculations during the entire observation session but not during the initial 30-min period. The mount latency was unaffected by cocaine treatment. The results suggest that the expression of the stimulating effects of cocaine on sexual behavior depends on the duration of the observation period.     

Sexual and aggessive behaviour of castrated male sheep after injection of gonadal steroids and implantation of androgens in the hypothalamus: A preliminary study

The combined effects of hypothalamic steroid implants and subcutaneous hormone injections on the courtship, copulatory and aggressive behaviour of five castrated male sheep (wethers) were assessed in thrice weekly tests with sexually receptive ewes. The animals were prepared with bilateral guide tubes for the insertion of removable hormone-containing cannulae aimed at the preoptic/anterior hypothalamic region of the brain. The sheep were treated as follows: weeks 1-4, cholesterol implants + oestradiol dipropionate injections; weeks 5-7, dihydrotestosterone propionate implants + oestradiol dipropionate injections; weeks 10-12, testosterone terone propionate implants + oil injections; weeks 13-15, testosterone propionate implants + dihydrotestosterone propionate injections. During peripheral treatment with oestradiol dipropionate (weeks 1-4), two of five sheep displayed ejaculatory reflexes in the absence of erection and intromission. Moreover, no obvious behavioural effects could be attributed to the additional presence of central dihydrotestosterone propionate implants (weeks 5-7). By contrast, testosterone propionate implants at the same central sites (weeks 10-12) maintained sexual behaviour in four of five sheep, induced mounting in the remaining animal and stimulated aggressive behaviour in all five sheep. Subsequently, additional peripheral treatment with dihydrotestosterone propionate (weeks 13-15) also stimulated three of the animals to exhibit penile erections.     

Characteristic frequency bands of the cortico-frontal EEG during the sexual interaction of the male rat as a result of factorial analysis

The electrocorticogram (ECoG) from the prefrontal cortex was simultaneously recorded with the accelerometric signals of pelvic thrusting performed by male rats during sexual behavior. The changes in the prefrontal ECoG were precisely correlated in time with well defined elements of male rat copulation. Principal component analysis allowed to identify three distinct bands of frequencies in the frontal ECoG: the absolute power (AP) of the 4–16 Hz band was increased in the 500-ms periods before, during, and after the execution of pelvic thrusting in mount, intromission and ejaculation responses; the AP of the 18–24 Hz band was selectively increased during the execution of pelvic thrusting at the three copulatory responses, whereas the AP of the 26–32 Hz band was increased only during the pelvic movements of mount and intromission responses. These results show that the electroencephalographic activity of the prefrontal cortex of the male rat is related to the performance of sexual behavior, supporting the concept that this cortical area is involved in the organization of sequential behaviors, as sexual behavior.     

Effects of testosterone metabolites on copulation and medial preoptic dopamine release in castrated male rats

The medial preoptic area (MPOA) is an important integrative site for male sexual behavior. Dopamine (DA) is released in the MPOA of male rats shortly before and during copulation. The recent presence of testosterone (T) may be necessary for this precopulatory increase in release. Previously, the postcastration loss of copulatory ability mirrored the loss of the DA response to an estrous female, and the restoration of copulation with exogenous T was concurrent with the reemergence of this DA response. The present study investigated the effectiveness of the two major metabolites of T in maintaining copulation and basal and female-stimulated DA levels. Adult male rats were castrated and received daily injections of estradiol benzoate (EB), dihydrotestosterone benzoate (DHTB), EB + DHTB, testosterone propionate (TP), or oil vehicle for 3 weeks. Microdialysis samples were collected from the MPOA during baseline conditions, exposure to an estrous female behind a barrier, and copulation testing. EB + DHTB- and TP-treated animals had normal basal DA levels and showed a precopulatory DA response, and most copulated normally. EB-treated castrates had high basal DA levels, but failed to show a female-stimulated increase; most intromitted, but none ejaculated. DHTB- and oil-treated groups had low basal levels of extracellular DA that did not increase during copulation testing; most failed to mount and none ejaculated. These results suggest that E maintains normal basal levels of extracellular DA in the MPOA, which are sufficient for suboptimal copulation, but that androgen is required for the female-stimulated increase in DA release and for facilitation of ejaculation.     

Deterioration of male sexual behavior in rats by the new prolactin-secreting tumor 7315b

The effects of hyperprolactinemia on male copulatory behavior in adult male and female rats were studied. Hyperprolactinemia was induced by the transplantable purely prolactin-secreting tumor 7315b. Male rats were castrated and received testosterone-filled capsules of different sizes which induced normal and subnormal testosterone levels. After sexual training the rats of the experimental groups were inoculated with tumor 7315b. Three weeks after tumor-inoculation high prolactin levels (2000-30000 ng/ml) were found. During this hyperprolactinemia ejaculation latency increased significantly, while the mount frequency and intromission frequency remained unchanged. Only 9 out of 22 rats ejaculated 19 days after inoculation. Moreover, it appeared that the inhibitory effect of the tumor was as strong in the presence of normal (2.33 +/- 0.07 ng/ml) as in the presence of low (0.35 +/- 0.01 ng/ml) testosterone levels. The inhibitory effect of tumor 7315b on copulatory behavior was not influenced by adrenalectomy. In gonadectomized female rats bearing testosterone-filled capsules tumor 7315b induced prolactin levels of about 2000 ng/ml and an almost complete cessation of mounts and intromission patterns 4 weeks after tumor-inoculation. It was concluded that tumor 7315b causes a strong inhibitory effect on male copulatory behavior in male and female rats and that this effect is not influenced by the presence of normal or low testosterone levels or removal of the adrenals, suggesting a direct effect of prolactin on brain functions.     

Effect of pineal indoles and testosterone on sex behavior

The possible action of melatonin and 5-methoxytryptophol, two pineal indoles, upon castrated male rat sexual behaviour with three testosterone dose levels was studied. Both indoles reduced the number of mounts and intromissions. 5-Methoxytryptophol, in addition, decreased the number of ejaculations. The sexual actions of the pineal indoles are prevented by the administration of high testosterone dose levels (500 micrograms/day).     

Testosterone implanted in the preoptic area of male Japanese quail must be aromatized to activate copulation

Intracranial implantation of minute pellets of gonadal steroids was combined with aromatase inhibitor treatment to determine if aromatization within the preoptic area (POA) is necessary for androgens to activate sexual behavior in the Japanese quail (Coturnix japonica). In this species, implantation of pellets of testosterone propionate (TP) or estradiol benzoate (EB) in the POA of castrated males restores male-typical copulatory behavior. In Experiment 1, adult male castrated quail were implanted intracranially with 200-micrograms pellets of equimolar mixtures of crystalline TP + cholesterol (CHOL), TP + 1,4,6-androstatriene-3,17-dione (ATD, an aromatase inhibitor), EB + ATD, or CHOL and behavior-tested with intact males and females. Copulation was stimulated by POA implants containing TP or EB (three of six CHOL + TP males and two of seven ATD + EB males copulated vs zero of four CHOL males), but copulation was not inhibited by combining ATD with TP (three of four ATD + TP males copulated). In Experiment 2, adult male castrated quail were injected systemically with ATD or oil for 6 days prior to and 14 days after intracranial implantation of 200-micrograms pellets containing the same amounts of TP or EB as in Experiment 1. The ATD injections completely blocked copulatory behavior in males with TP implants in the POA such that ATD/TP and Oil/TP mount frequencies differed significantly, but failed to block copulation in males with EB implants in the POA (proportions of males copulating were ATD/EB, 6/8; ATD/TP, 0/6; Oil/TP, 4/7). The cloacal foam gland, an androgen-sensitive secondary sex character, was unaffected by the dose of ATD used. We conclude that activation of copulatory behavior by TP implants in the POA is not due to nonspecific effects of high local testosterone concentrations but rather to aromatization. These results support the hypothesis that cells within the POA aromatize testosterone to estrogens, which directly stimulate the cellular processes leading to activation of male-typical copulatory behavior.     

Group mating among Norway rats II. The social dynamics of copulation: Competition,cooperation, and mate choice

The social interactions within groups of Norway rats (Rattus norvegicus) had a strong impact on the individual pattern of copulation which, in turn, affects sperm precedence and the probability of implantation in this species. Males alternated uninterrupted ejaculatory series, augmenting each others' copulatory investment. Females took turns mating after receiving an intromission, collectively potentiating the males' copulatory behaviour; increasing the number of oestrous females increased the number of intromissions and ejaculations achieved by each male but did not affect the amount of copulation experienced by each female. These turn-taking patterns within each sex provided the opportunity to change partners and permitted the emergence of different sex-typical patterns of copulation. Furthermore, the dominant male contributed more intromissions and tended to give each female more ejaculations than the subordinates did. Dominant males were also more likely to inhibit the subordinates' sperm transport. Females competed among themselves for the opportunity to mate with a male as he approached ejaculation and were likely to protect more of the dominant male's sperm transport than the subordinate male's.     

Actions of esters of testosterone, dihydrotestosterone, or estradiol on sexual behavior in castrated male guinea pigs

Prepuberally castrated male guinea pigs were treated in adulthood with estradiol benzoate, testosterone propionate, dihydrotestosterone propionate or corn oil (vehicle control). Both corn oil and estradiol benzoate were ineffective in augmenting or inducing any aspect of adult male sexual behavior. Dihydrotestosterone propionate and testosterone propionate were both effective in establishing the complete male sexual behavior pattern, although they differed in the manner in which they affected specific components. For example, males treated with testosterone propionate showed more non-intromissive but not more intromissive mounts than males treated with dihydrotestosterone propionate. In addition, the average frequency of thrusts per intromission was greater for males treated with dihydrotestosterone propionate than for males treated with testosterone propionate.     

Indirect influence of s-triazines on rat gonadotropic mechanism at early postnatal period

Daily s.c. injections of atrazine and deethylatrazine to rat mothers during pregnancy only or during pregnancy and lactation influenced the pituitary-gonadal axis of male and female offsprings. In female and male offspring, slow maturation of gonadotropic system is evident and as a consequence modified male and female pituitary 5 alpha-R activity is present. The number of specific steroid-hormone receptor sites at the offsprings' gonads is unchanged if the mothers were treated only during pregnancy, but 5 alpha-DHT prostate receptors are strongly inhibited in offspring of mothers treated with atrazine and deethylatrazine during pregnancy and lactation.