Differential effect of testosterone and dihydrotestosterone on the sexual behavior of prepuberally castrated male rabbits

The effect of testosterone propionate (Tp) and dihydrotestosterone propionate (DHTp) at doses of 1, 3 and 9 mg daily for 30 days on the copulatory behavior of prepuberally castrated male New Zealand white rabbits was studied. Tp was significantly more effective than DHTp in eliciting copulatory behavior at each dose level tested. Three milligrams Tp was the minimal dose required to elicit mounting consistently. DHTp at the high dose level (9 mg) only elicited sexual activity comparable to that observed with the low dose of Tp (1 mg). The results suggest that T does not need to be reduced to DHT to stimulate sexual behavior in the male rabbit.     

Effect of small doses of naloxone on sexual exhaustion in White New Zealand male rabbits

The objective of the present study was to determine the effect of small doses of naloxone on sexual exhaustion in White New Zealand male rabbits. Twelve young and 12 adult male rabbits 6–12 months old and 14–20 months of age, respectively, were selected from a commercial farm. Each male rabbit was housed individually in galvanized cages (90 cm × 60 cm × 40 cm). The rabbits were housed in an open shed exposed to natural photoperiod (12 L 12 D, 19°N). Daily temperature fluctuated through the year from 28 to 16 °C. Humidity was 45 ± 5%. Water and food (rabbit chow PMI) was supplied ad libitum. After sexual behaviour for each studied group was established, the males were given a 6-day rest, and 3 days before next trial, six males of each group (treated) received a subcutaneous implant of 8 mg of naloxone in a crystalline nitrocellulose pellet formulated to be completely absorbed in 15 days. The remaining six males were sham-treated (control). At the end of the resting period as previously described, the sexual behavior of each group was studied and compared using a Mann–Whitney statistical U-test. The effect of naloxone on sexual behavior was analyzed with a Wilcoxon test for correlated samples. With regard to sexual activity between young and adult rabbits, it was observed that there was a significant difference between groups (P = 0.00275, Z = 2.8823, adjusted Z = 2.99.43) showing that younger rabbits mounted/ejaculated from 9 to 10 females compared with 6 to 8 mounted/ejaculated by older rabbits. When naloxone was administered to both groups, there was a significant difference when comparing sexual behavior before and after administration of naloxone (table first and second trial). Young rabbits treated with naloxone mounted/ejaculated 11–12 females while older rabbits mounted nine females before reaching sexual exhaustion. A significant difference was observed when comparing the number of estrous females that were mounted/ejaculated between groups. Environmental photoperiod and temperature changes were not considered. It was concluded that endogenous opioids are important modulators of behavioral and hormonal interactions related to sexual behavior.     

Synergistic actions of estrogen and androgen on the sexual behavior of the castrated male rabbit.

Daily injections of 2.5 mg dihydrotestosterone (DHT) for 30 days induced sexual behavior in 19% of piepuberally and 62% of postpuberally castrated New Zealand white male rabbits. Combined treatment of 2.5 mgm DHT plus 5 μgm of estradiol benzoate (EB) activated sexual behavior in 100 and 85% of prepuberally and postpuberally castrated rabbits respectively. Moreover, subjects (Ss) receiving DHT + EB displayed sexual activity in a significantly higher percentage of tests and presented a higher frequency of mounts and intromissions than those Ss receiving only DHT. The results demonstrate that estrogen synergizes with androgen (DHT) to stimulate sexual behavior in the male rabbit.     

Effects of ATD on male sexual behavior and androgen receptor binding: a reexamination of the aromatization hypothesis

The aromatization hypothesis asserts that testosterone (T) must be aromatized to estradiol (E2) to activate copulatory behavior in the male rat. In support of this hypothesis, the aromatization inhibitor, ATD, has been found to suppress male sexual behavior in T-treated rats. In our experiment, we first replicated this finding by peripherally injecting ATD (15 mg/day) or propylene glycol into T-treated (two 10-mm Silastic capsules) or control castrated male rats. In a second experiment, we bilaterally implanted either ATD-filled or blank cannulae into the medial preoptic area (MPOA) of either T-treated or control castrated male rats. With this more local distribution of ATD, a lesser decline in sexual behavior was found, suggesting that other brain areas are involved in the neurohormonal activation of copulatory behavior in the male rat. To determine whether in vivo ATD interacts with androgen or estrogen receptors, we conducted cell nuclear androgen and estrogen receptor binding assays of hypothalamus, preoptic area, amygdala, and septum following treatment with the combinations of systemic T alone. ATD plus T, ATD alone, and blank control. In all four brain areas binding of T to androgen receptors was significantly decreased in the presence of ATD, suggesting that ATD may act both as an androgen receptor blocker and as an aromatization inhibitor. Competitive binding studies indicated that ATD competes in vitro for cytosol androgen receptors, thus substantiating the in vivo antiandrogenic effects of ATD. Cell nuclear estrogen receptor binding was not significantly increased by exposure to T in the physiological range. No agonistic properties of ATD were observed either behaviorally or biochemically. Thus, an alternative explanation for the inhibitory effects of ATD on male sexual behavior is that ATD prevents T from binding to androgen receptors.     

Effect of some antiestrogens and aromatase inhibitors on androgen induced sexual behavior in castrated male rats

The effect of antiestrogens (MER-25, ICI-46474, and cis-clomiphene) and aromatase inhibitors (5-α-androstanedione, metopirone, and aminoglutethimide) on androgen induced copulatory behavior was tested in sexually inexperienced castrated male tats. Daily injections of 1 mg testosterone (T) for 21 days induced sexual activity in most subjects (61% mounting). Daily pretreatment with MER-25 or cis-clomiphene at three dose levels did not block the behavioral response to T. ICI-46474 at the high dose level (1 mg/kg) elicited a significant depressory effect on the sexual behavior of the T treated castrated rats. A single injection of 6 mg testosterone propionate (TP) induced mounting behavior in 56% of the tested rats within 120 hr. Treatment with metopirone or 5 α-androstanedione (injections every 12 hr for 96 hr) did not inhibit the response to TP. By contrast, aminoglutethimide (5 or 15 mg every 12 hr for 96 hr) abolished the behavioral response to androgen.     

Male sexual behavior in deer mice (Peromyscus maniculatus) following castration and hormone replacement

Sexually experienced male deer mice (Peromyscus maniculatus bairdi) were castrated and tested for male sexual behavior. In the weeks following castration male sexual behavior decreased. Ejaculation disappeared first, followed by intromission and, finally, mounting. Castrated males failing to copulate were assigned to one of four treatment groups: 200 μg testosterone propionate (TP); 200 μg dihydrotestosterone propionate (DHTP); 2 μg estradiol benzoate (EB); or sesame oil (OIL). TP and DHTP were equally effective in restoring the complete male sexual behavior pattern. In contrast, EB was effective in stimulating mounting and minimally effective in stimulating intromissions (vaginal penetration), but did not stimulate ejaculatory responses. These data indicate that in deer mice testosterone may mediate male sexual behavior through reduction to dihydrotestosterone rather than through aromatization to estradiol.     

The reversible inhibition of steroid-induced sexual behavior by intracranial cycloheximide

Cycloheximide(Cyclo), an inhibitor of protein synthesis by a direct action on protein synthesis at the ribosomal level, was used to reversibly inhibit estrogen-induced sexual receptivity. Cyclo (100 μg per rat) was infused into the preoptic area(POA) of ovariectomized rats at varying times before, simultaneously with, and after 3 μg of subcutaneous estradiol benzoate (EB). All animals received 0.5 mg progesterone (P) 36 hr after EB, and were tested for sexual receptivity 4–6 hr after P. The females were placed with stud males and a lordosis quotient was computed for each female (lordosis quotient = number of lordosis responses/20 mounts by the male × 100). Females receiving Cyclo 6 hr before, simultaneously with, or 12 hr after EB showed significantly lower levels of sexual receptivity when compared to females receiving Cyclo 36 hr before and 18 and 24 hr after EB. When those animals that showed low levels of sexual behavior after Cyclo infusion were reprimed with EB and P 7 days later and presented with a male they showed high levels of sexual receptivity. Thus, the effect of Cyclo was reversible. Only Cyclo infusions into the POA (bilateral) and third ventricle were effective in suppressing sexual behavior. Caudate nucleus, lateral ventricle, and unilateral POA infusions were without effect.The data presented are in agreement with earlier work that utilized actinomycin D to inhibit steroid-induced sexual behavior. Cyclo was found to be less toxic than actinomycin D. All of the available evidence is consistent with the hypothesis that estrogen stimulates RNA and/or protein synthesis in its facilitation of sexual behavior in the female rat.     

Influence of environmental events immediately after birth on postnatal testosterone secretion and adult sexual behavior in the male rat

A rise in plasma testosterone (T) levels occurs in male rats during the first 2 hr after birth which is of importance for the process of sexual differentiation. To study the influence of environmental factors on the postnatal T surge and sexual development, newborn male rats were subjected to various treatments immediately after cesarean delivery including cooling, ether anesthesia, and mother-infant separation. In adulthood, the animals were observed for masculine and feminine sexual behavior. Males anesthetized at 0 hr showed elevated levels of feminine sexual behavior and impaired masculine sexual behavior. Pups subjected to cooling or mother-infant separation showed slightly prolonged intromission latencies, but otherwise normal levels of feminine sexual behavior. Significantly elevated plasma T levels were found in intact pups 2 hr after birth but not in pups subjected to cooling or ether anesthesia. Significantly higher levels of T were observed in pups subjected to cooling 4 hr after birth, suggesting a delay of the T surge. The most pronounced impairing effects were seen in the defeminization process, but the masculinization process also is affected by ether anesthesia. It was concluded that ether anesthesia immediately after birth may permanently interfere with the sexual development by suppressing the neonatal T surge.     

The relationship between circulating testosterone levels and male sexual behavior in rats

The relationships between plasma testosterone (T) and various parameters of male sexual behavior were examined in intact and castrated T-treated male rats. Repeated blood sampling and behavioral testing revealed no correlation between any measure of sexual behavior and plasma T in normal untreated sexually active males. T-Filled Silastic capsules, implanted subcutaneously at the time of castration, were found to produce plasma T levels proportional to capsule size. Plasma T titers less than 10% of normal (0.2 ng/ml) maintained ejaculatory behavior near normal levels for the 58 days of the experiment. Measures of sexual behavior which showed androgen dependence were intromission latency, postejaculatory interval, and intromission frequency. The plasma T concentration required to maintain these parameters within the intact range was 0.7 ng/ml, which is less than one-third of the mean intact level (2.6 ng/ml). No significant improvement in the sex behavior measures was seen with plasma T levels between 0.7 and 3.1 ng/ml. It was concluded that the absence of relationships between circulating T and sexual behavior in the normal rat population is due to the androgen requirement for this behavior being less than the amount normally present. Findings on T levels and T treatment in noncopulator males are also presented.     

Inhibition of testosterone-induced sexual behavior in the castrated male rat by aromatase blockers

The effect of some aromatase inhibitors (aminoglutethimide, 1,4,6-androstatrien-3, 17-dione, and 4-hydroxy-androstenedione) on testosterone propionate (TP)-induced copulatory behavior was tested in sexually inexperienced castrated male rats. A single injection of 6 mg of TP induced mounting in 48% and ejaculatory pattern in 19% of the rats within 120 hr. Treatment with the aromatase inhibitors (injections every 12 hr for 108 hr) suppressed ejaculation in all but one rat and significantly reduced the number of rats mounting and intromitting. Concurrent administration of estradiol benzoate (EB, 1 or 3 μg every 12 hr) prevented the inhibitory effect of aromatase blockers. No inhibitory effect of the aromatization blockers was observed in rats in which sexual behavior was induced by dihydrotestosterone (1 mg/day) and EB (2.5 μg/day) for 20 days. The results support the concept that aromatization is an essential step for the induction of male sexual behavior by androgen in the rat.     

Sex and context: hormones and primate sexual motivation

Gonadal hormones regulate the ability to copulate in most mammalian species, but not in primates because copulatory ability has been emancipated from hormonal control. Instead, gonadal hormones primarily influence sexual motivation. This separation of mating ability from hormonally modulated mating interest allows social experience and context to powerfully influence the expression of sexual behavior in nonhuman primates, both developmentally and in adulthood. For example, male rhesus monkeys mount males and females equally as juveniles, but mount females almost exclusively as adults. Having ejaculated with a female better predicted this transition to female mounting partners than did increased pubertal testosterone (T). It is proposed that increased pubertal T stimulates male sexual motivation, increasing the male's probability of sexual experience with females, ultimately producing a sexual preference for females. Eliminating T in adulthood reduces male sexual motivation in both humans and rhesus monkeys, but does not eliminate the capacity to engage in sex. In male rhesus monkeys the effects of reduced androgens on sexual behavior vary with social status and sexual experience. Human sexual behavior also varies with hormonal state, social context, and cultural conventions. Ovarian hormones influence female sexual desire, but the specific sexual behaviors engaged in are affected by perceived pregnancy risk, suggesting that cognition plays an important role in human sexual behavior. How the physical capacity to mate became emancipated from hormonal regulation in primates is not understood. This emancipation, however, increases the importance of motivational systems and results in primate sexual behavior being strongly influenced by social context.     

Action of androgen and estrone implants on sexual behavior and reproductive organs of castrated male rabbits

Sexually mature but inexperienced male rabbits were castrated, immediately implanted with either testosterone (T), estrone (E₁), dihydrotestosterone (DHT), T + E₁, or DHT + E₁, and tested for male sexual behavior. Other castrates were not implanted, and testing was either begun immediately (Ca-I) or delayed for 4 weeks (Ca-D). Intact males served as controls (C). Latency to mount a teaser female and to ejaculate into an artificial vagina was tested twice in a morning three times per week for 8 weeks. Then, animals were sacrificed, and reproductive organs were measured. The Ca-I group responded slowly to sexual training and ceased nearly all sexual activity by 8 weeks, whereas the Ca-D males seldom displayed interest in the teaser female. Intact controls and the T and T + E₁, groups all responded to the teaser and mounted and ejaculated within a few seconds. DHT and E₁, individually maintained the sexual activity of castrates equivalent to that of C for 4–5 weeks, but the time required to mount and, particularly, to ejaculate increased thereafter. The results with DHT + E₁ were equivocal in that castrates with this hormone combination sustained sexual activity equivalent to that of the controls for 7 weeks, but one animal in particular became sexually inactive the last week of the experiment. Penis weight was at least partially maintained in all implanted castrates. Accessory sex gland weight was smallest in the DHT group and was significantly increased in the T + E₁ and DHT + E₁ groups. The largest ejaculates of fluid were obtained in the group receiving E₁ alone. These results may be interpreted to indicate a role of both androgen and estrogen centrally and peripherally in the rabbit.     

Aging and primate male sexual behavior

Old male rhesus macaques display less sexual behavior than young and middle-aged males. The decrease in sexual activity occurs without a statistically significant decline in gonadal hormones or change in diurnal patterns of serum T, DHT, or LH. Levels of sexual activity are not increased by administering T to old intact males. However, the hormone is effective in increasing sexual behavior in old long-term-castrated males. Performance can be increased to levels observed in equally old untreated intact males. Readily detectable physical disabilities of old age have been observed to impair sexual performance, but the observed general decline in sexual activity cannot be accounted for by known physical disabilities. Novelty, as represented by a change in female partner or by a change in environment, has not increased sexual performance in old rhesus males. Only when old males were paired with empirically selected preferred females is their sexual behavior increased to levels displayed by young males. Drugs reported to increase levels of sexual behavior in rats have thus far been less effective in old rhesus males than powdered rhinoceros horn has been in man. The probable absence of a placebo effect in rhesus males should increase their usefulness as an animal model for the study of sexual behavior in aging men.     

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.     

Adrenal contribution to the induction of sexual behavior in the female musk shrew.

Sexually experienced female musk shrews (Suncus murinus) lack an ovarian, vaginal, and behavioral estrous cycle. Females, once induced by their initial contact with a male, are able to display copulatory behavior whenever a male is present (Rissman, Silveira, and Bronson, 1988). Based on plasma levels of steroids, and on hormone replacement studies conducted after ovariectomy (OVX), we have shown that testosterone (T) plays an essential role in the regulation of female sexual behavior (Rissman and Crews, 1988; Rissman, Clendenon, and Krohmer, 1990a; Rissman, 1991). To date we have only examined the potential contribution of adrenal steroids to female sexual behavior in a preliminary manner. After adrenalectomy, gonadally intact females display significantly lower levels of sexual behavior than controls (Rissman and Bronson, 1987). The following experiments were conducted to examine the role the adrenal steroids (in contrast to the medullary hormones) play in the induction of female sexual behavior in the musk shrew. In the first experiment gonadally intact females were treated with dexamethasone (DEX) to reduce the secretion of adrenal steroids. Significantly fewer females receiving DEX demonstrated sexual behavior as compared with controls. In the second study, OVX females received T-filled Silastic implants. When DEX was administered to OVX + T females at a dose that dropped circulating T levels to those found in ovary and adrenal intact females, no effect on sexual behavior was noted. The data show that the adrenals are a behaviorally important source of T and contribute toward the hormonal control of sexual behavior in these female mammals.     

Effects of sexual stimulation on the sexual performance of rams

The sexual performance of bulls and male goats is improved if they are allowed to view the hetero-sexual behavior of other males as a prelude to mating. The purpose of the following study was to determine whether sexual stimulation enhances the sexual performance of rams. In Experiment 1, 11 sexually experienced ram lambs ( 9 months of age) and 18 sexually experienced yearling and 2-year-old rams were individually exposed to 4 unrestrained, hormone-induced estrous ewes for 60 min after viewing the courtship and mounting behaviors of a male conspecific for 20 min (two tests) and in the absence of stimulator animals (two tests). In contrast to the results with bulls and bucks, the rams were hardly influenced by the sexual stimulation treatment. Latencies for first mount and first ejaculation were shorter for sexually stimulated ram lambs; otherwise, treatment differences were negligible.

A similar follow-up experiment was administered to 12 mature rams using restrained females in the sexual performance tests. Again, treatment differences were minor.

It was concluded that sexual stimulation does not functionally enhance the sexual performance of rams. Species differences in response to sexual stimulation are discussed in terms of female sexual behaviors that may result in a selective (competitive) advantage to males that are stimulated to locate and mate with females early in the estrous period.     

锰胁迫和性别竞争交互处理下沙棘雌雄幼苗生理响应特征

沙棘(Hippophae rhamnoides)是重要的雌雄异株人工林防护树种,但对其环境胁迫的性别响应差异研究不足,性别竞争与胁迫因子的交互效应响应特征尚不清楚。为了探讨锰胁迫和性别竞争交互处理下沙棘雌雄植株的生理响应特征和耐受能力,旨在为沙棘修复土壤重金属污染提供实践指导,该文研究了锰胁迫(4 000 mg·kg^-1)和3种不同性别组合模式(雌雄、雌雌、雄雄)处理下沙棘的生理响应,分别测定雌雄沙棘叶片中叶绿素、过氧化物酶(peroxidase,POD)、超氧化物歧化酶(superoxide dismutase,SOD)、丙二醛(malondialdehyde,MDA)、总酚(total phenols, TP)、游离脯氨酸(free proline,Pro)、可溶性糖(soluble sugar,SS)、甜菜碱(glycine betaine, GB)以及锰含量。结果表明:(1)锰胁迫下,在所有竞争组合中,性间竞争的雄株(M/FM)SOD活性最高,而MDA含量与对照相比未有明显升高,表明雄株的抗氧化能力更好,膜氧化损伤程度更小。(2)锰胁迫时M/FM积累了更多的游离脯氨酸,表现出更好的渗透调节能力和耐受能力。(3)交互效应分析显示性别互作和锰胁迫交互处理显著影响了沙棘雌雄叶片的光合色素、抗氧化酶活性和渗透调节能力; 主成分分析显示SOD、POD、MDA、叶绿素b(chlorophyll b, Chlb)、SS、Pro可作为重要的生理响应指示参数。该研究结果对于利用沙棘修复土壤重金属污染可提供一定的参考。     

Role of aromatization in anticipatory and consummatory aspects of sexual behavior in male rats

The present study was designed to test the hypothesis that aromatization is involved in the maintenance by testosterone of the appetitive component of male sexual behavior. We measured appetitive sexual behavior by administering behavioral tests in bilevel chambers and quantifying anticipatory level changes during a 5-min period prior to introduction of a stimulus female. In addition, we recorded standard measures of consummatory male sexual behavior after the female was introduced. Following 3 weekly tests, level-changing behavior reached a plateau and remained stable for up to 10 weeks. After 10 bilevel tests, rats were given subcutaneous testosterone capsules to clamp circulating androgen at physiological levels. Rats were tested and divided into two groups that were matched for measures of sexual behavior. One group was then treated with the nonsteroidal aromatase inhibitor, Fadrozole (2.5 mg/kg/day), given subcutaneously in beta-cyclodextrin and the other group was treated with vehicle. Within 1 week of Fadrozole treatment, the number of anticipatory levels changes was significantly reduced, but not the latency to begin searching. Fadrozole treatment also significantly reduced all measures of copulatory behavior over the period of treatment and increased latencies to first mount, intromission, and ejaculation. After 8 weeks, both treatment groups were given an additional Silastic capsule filled with estradiol and tested for 4 additional weeks. Estrogen treatment partially restored level-changing behavior, mounts, and intromissions but had little effect on ejaculations. These results support the view that aromatization is important for maintaining both the appetitive and the consummatory aspects of sexual behavior in male rats.     

The postnatal demasculinization of sexual behavior in the Japanese quail (Coturnix coturnix japonica)

Three experiments were performed to analyze the time course of demasculinization in the Japanese quail and to test the activating and organizing effects of estradiol (E2) in adult sexually active birds. In Experiment 1, males and females were castrated at the age of 1 day or 1, 2, 4, and 6 weeks and treated as adults with testosterone (T). The age of castration had no effect on behavior and morphology in males. Plasma gonadotrophins (LH and FSH) were, however, higher in males castrated at or before than in those castrated after 2 weeks of age. This suggests that postnatal testicular secretions have organizing effects on the pituitary activity. Females which were castrated before 1 week of age were less sensitive to the activating effects of T than males, but were not fully demasculinized. The demasculinization of different reproductive characteristics such as male sexual behavior, cloacal gland size, and weight of the syringeal muscles is achieved in females at different times posthatching. In Experiment 2, castration of male and female quail at the ages of 4 days or 4 weeks confirmed that postnatal ovarian secretions contribute to the full behavioral and morphological demasculinization of females. It is easier to elicit mounting in T-treated females when they are tested in their home cage instead of a test arena. This difference was not observed in males. During Experiment 3, it was impossible to demasculinize sexually active adult males or females by treatment with Silastic implants of E2. E2 did not maintain sexual behavior in ovariectomized females showing male sexual behavior when treated with T but maintained the behavior in males.     

Theophylline, otentiation of testosterone propionate or estradiol bensoate in inducing male but not female sexual behavior in the female rat.

Two experiments were carried out to assess the possible involvement of 3′:5′cyclic adenosine monophosphate (cAMP) in the hormonally mediated activation of masculine and feminine sexual behavior in female rats. In Experiment I, theophylline, a compound shown to be effective in inhibiting the degradation of cAMP, was combined with estradiol benzoate (EB) in an attempt to potentiate the action of estradiol for inducing feminine or masculine sexual behavior. Theophylline, when administered in combination with EB to ovariectomized females, resulted in an increase in masculine sexual behavior but no potentiating action on female receptivity. In Experiment II, theophylline, when given to female rats, potentiated the action of testosterone propionate in stimulating male but not female sexual behavior. These data suggest that estradiol and testosterone may be activating masculine sexual behavior through similar biochemical mechanisms. Likewise, cAMP may be involved in the activation of masculine but not feminine sexual behavior by gonadal steroids.