Embryonic and posthatching treatments with sex steroids demasculinize the motivational aspects of crowing behavior in male Japanese quail

Demasculinizing action of embryonic estrogen on crowing behavior in male Japanese quails was examined. Eggs were treated with either 20 μg of estradiol benzoate (EB) or vehicle on the 10th day of incubation. Chicks hatched from both groups of eggs were injected daily with either testosterone propionate (TP; 10 μg/g b.w.), 5α-dihydrotestosterone (DHT, a non-aromatizable androgen; 10 μg/g b.w.), or vehicle from 11 to 50 days after hatching, and during this period their calling behaviors were observed. Irrespective of embryonic treatments, all birds received posthatching treatment with either TP or DHT, but not with vehicle, emitted crows in place of distress calls in a stress (non-sexual) context of being isolated in a recording chamber. The posthatching TP, but not posthatching DHT, induced crowing in a sexual context (crowing in their home-cages) from much earlier age than posthatching vehicle in the birds received control embryonic treatment with vehicle. The same TP treatment, however, completely eliminated the crowing in a sexual context in the birds received EB during their embryonic life. In the birds treated with either posthatching DHT or posthatching vehicle, the crowing in a sexual context was only slightly decreased by embryonic EB treatment. These data suggest that posthatching estrogen, derived from testosterone aromatization, enhances the demasculinizing action of embryonic estrogen, and thus strongly reduces the sexual motivation for crowing behavior. This demasculinizing action, however, would not influence vocal control system which generates acoustic pattern of crowing in the presence of androgens allowing the birds to crow in a non-sexual context.     

Effects of estradiol benzoate, estrone, and propionates of testosterone or dihydrotestosterone on sexual and related behaviors of ovariectomized rhesus monkeys

Ovariectomized adult rhesus monkeys were injected daily for 10 days with either 1 mg of dihydrotestosterone propionate (DHTP), 1 mg of testosterone propionate (TP), 10 μg of estradiol benzoate (EB), or 500 μg of estrone (El). On the 5th and 10th days of treatment, females received two 24-min behavioral tests with each of two adult males. All females received every hormonal treatment during the course of the study, with the order of treatments counterbalanced. Prior to the initiation of an hormonal treatment, each subject received two tests with no hormone treatment (NORX). Three behaviors related to female proceptivity were recorded. Treatment with DHTP had no influence on any aspect of proceptivity measured, in comparison to the NORX condition, whereas El or TP treatment augmented the frequencies of two of the proceptive behaviors and EB increased all three. The response of the male toward the female was influenced by the female's hormonal condition. Treatment with TP or DHTP did not increase the frequency of male contact or the mount rate in comparison to the NORX condition, whereas EB or El treatment did. In addition, DHTP was the only steroid which failed to increase the percentage of tests with intromission or ejaculation when compared to NORX. Female receptivity, as measured by acceptance or rejection of male contacts, was not different for the NORX-, TP-, EB-, or El-treated conditions. DHTP treatment, however, reduced female receptivity in comparison to all other conditions. Treatment with DHTP or TP resulted in an increase in the frequency of female yawning behavior, whereas neither estrogen treatment showed any effect on this behavior. The influences of TP on female proceptive and male sexual behavior were never duplicated or even approximated by treatment of females with the nonaromatizable DHTP. Nor was there any evidence that TP inhibited female receptivity below the level characteristic of NORX females, as was true for DHTP.     

Effects of testosterone, estrogen, and dihydrotestosterone upon aggressive and sexual behavior of female rats

Groups of female TMD rats were treated either with estradiol benzoate (EB), dihydrotestosterone propionate (DHTP), testosterone propionate (TP), EB + DHTP (EB/DHTP), or with oil. These groups of females were tested for social aggression and for masculine and feminine sexual behavior. In addition, patterns of masculine and feminine sexual responses during the aggressive encounters, were investigated. TP-treated females of the same strain were used as opponents in the tests for aggression. In accordance with previous results, EB did not activate aggression whereas TP treatment resulted in a significant increase in aggression in females. Aggressive responses were activated by adding DHTP to EB, up to levels equal to those activated by TP. Sexual responses were observed in the tests for aggression as well as in tests for sexual behavior. The results indicated that feminine and masculine sexual responses were affected significantly by hormonal treatment. Mounting behavior in the test for aggression was activated by TP and by EB/DHTP. Lordosis and proceptive responses were inhibited in these groups as compared to EB-treated females, both in tests for aggression and in tests for sexual behavior. The results are consistent with the idea that dihydrotestosterone inhibits feminine and activates masculine sexual activity. The results also indicate that EB and DHTP synergistically activate aggression.     

Sexual differentiation of behavior in the zebra finch: effect of early gonadectomy or androgen treatment

Treatment of nestling zebra finches with estradiol benzoate (EB) has been shown to masculinize singing in females and demasculinize copulatory behavior in males, suggesting that sexual differentiation of these behaviors is under hormonal control such that testicular hormones induce the capacity for song and ovarian hormones suppress the capacity for mounting. Two experiments were carried out to obtain a more complete picture of sexual differentiation in this species. In Experiment 1, nestlings were injected daily for the first 2 weeks after hatching with testosterone propionate (TP), dihydrotestosterone propionate (DHTP), or a combination of DHTP and EB. As adults, birds were gonadectomized and implanted with TP prior to testing, then tested again after implantation with EB. Singing was not increased in females by any of the treatments. The only effect of either TP or DHTP given alone was defeminization of female proceptive behavior by DHTP. Thus androgens appear to have less influence than estrogens on sexual differentiation of behavior in this species. The combination of DHTP and EB demasculinized mounting in males. In Experiment 2, nestlings were gonadectomized at 7-9 days of age and implanted with TP prior to testing in adulthood. Early gonadectomy had little effect on later behavior; early castrated males sang, danced, and copulated normally and early ovariectomized females neither sang nor mounted.     

Effect of implanting bull calves with testosterone propionate, dihydrotestosterone propionate or oestradiol-17 beta prepubertally on the pituitary-testicular axis and on postpubertal social and sexual behaviour.

Twelve non-implanted crossbred bull calves served as controls and 30 crossbred bull calves (10/treatment) were implanted for 82 days, beginning at 34 days of age, to determine the influence of testosterone propionate (TP), dihydrotestosterone propionate (DHTP) and oestradiol-17 beta (E2) on prepubertal and pubertal pituitary-testicular function and on postpubertal social and sexual behaviour. Compared with control bulls, concentrations of serum luteinizing hormone (LH), follicle-stimulating hormone (FSH) and inhibin concentrations were suppressed (P less than 0.01) in all implanted bulls. Testosterone (T) concentration increased (P less than 0.001) in TP-implanted, but decreased (P less than 0.01) in DHTP and E2 bulls during the implant period. LH response to gonadotrophin-releasing hormone (GnRH) challenge during the implant period (2.5 months of age) was less (P less than 0.01) in TP, E2 and DHTP bulls than in controls. A small but significant T response to GnRH occurred in control bulls at 2.5 months of age. LH and T responses to GnRH challenge at 7 months of age (100 days after implant removal) was similar (P greater than 0.20) in control and implanted bulls. Steroid implants administered prepubertally had no effect (P greater than 0.10) on postpubertal social and sexual behaviours, including number of flehmen responses, abortive mounts, services and competitive order score. Body weight did not differ (P greater than 0.10) between treatment groups, but testis size was reduced (P less than 0.01) during the implant period and up to 10 months of age in treated bulls compared with controls. Testes remained smaller in E2-treated bulls up to the end of the study (23 months of age), but daily sperm production and epididymal weight did not differ (P greater than 0.10) between treatment groups at slaughter. Control bulls reached puberty earlier (P less than 0.01; 270 +/- 11 days of age) than did TP (302 +/- 11 days), DHTP (309 +/- 11 days) or E2 (327 +/- 11 days) bulls. Although puberty was delayed in all implant groups, there was no difference in scrotal circumference at puberty (average 28.4 +/- 0.4 cm) between treatment groups. Our findings indicate that TP, DHTP and E2 implants administered prepubertally result in acute suppression of serum LH, FSH and inhibin during the implant period and in post-implant suppression of testis size and delayed puberty in bulls. The lack of treatment effect on behaviour suggests that steroidal programming of sexual behaviour occurs before 1 month of age in bulls.     

The influence of androgenic and estrogenic hormones on sexual behavior in castrated adult male pigs

The objectives of these studies were to evaluate the influence of testosterone propionate (TP), estradiol cypionate (EC), dihydrotestosterone propionate (DHTP), EC + TP, EC + DHTP, and TP + DHTP on traits of masculine sexual behavior in castrated adult male pigs of different breeds. Masculine sexual behavior was restored and maintained by TP, whereas EC initially activated sexual behavior, including copulation and ejaculation, but was unable to sustain copulatory behavior for the 8- to 18-week periods that were evaluated. Treatment with DHTP was ineffective for stimulation of sexual behavior; thus, it is suggested that testosterone promotes some aspects of masculine sexual behavior in male pigs via aromatization to estrogen, but both androgen and estrogen are required for maintenance of the full complement of masculine sexual behavior traits.     

Sexual behavior and penile reflexes of neonatally castrated male rats treated in infancy with estrogen and dihydrotestosterone

Male rats castrated neonatally and treated with a combination of 0.5 μg estradiol benzoate (EB) plus 50μg dihydrotestosterone propionate (DHTP) for the next 14 days displayed normal sexual behavior when injected with testosterone propionate (TP) in adulthood. Neither EB nor DHTP alone had this developmental effect inasmuch as only 20–25% of the neonatal castrates treated with just 0.1, 0.5, or 10 μg EB, or 50 μg DHTP, displayed ejaculatory responses. The periodic application of mildly painful electric shock, which has been previously shown to markedly facilitate ejaculatory responding in normal male rats, failed to improve sexual performance in these latter subjects. This was true even of the castrates treated neonatally with DHTP which frequently intromitted. Castrates treated with EB or DHTP alone neonatally were subjected to spinal transection (after testing of sexual behavior) for examination of penile reflexes. Those treated with DHTP showed normal reflexes, characterized by numerous erections and flips, indicating the presumably nonaromatizable DHTP has developmental effects on penile reflexes similar to those of testosterone. Subjects treated with EB, including four animals that had ejaculated at least once, displayed very few, if any, erections on reflex tests and no flips. These results show that sometimes intromissive and ejaculatory patterns can occur even though the animal appears to have little or no capacity for penile reflexes.     

Organizational effects of testosterone,estradiol, and dihydrotestosterone on vasopressin mRNA expression in the bed nucleus of the stria terminalis

In adulthood, male rats express higher levels of arginine vasopressin (AVP) mRNA in the bed nucleus of the stria terminalis (BST) than do female rats. We tested whether this sex difference is primarily due to differences in neonatal levels of testosterone. Male and female rats were gonadectomized on the day of birth and treated with testosterone propionate (TP) or vehicle on postnatal days 1, 3, and 5 (P1, P3, and P5). Three months later, all rats were implanted with testosterone-filled capsules. Two weeks later, brains were processed for in situ hybridization to detect AVP mRNA. We found that neonatal TP treatment significantly increased the number of vasopressinergic cells in the BST over control injections. We then sought to determine the effects of testosterone metabolites, estradiol and dihydrotestosterone, given alone or in combination, on AVP expression in the BST. Rat pups were treated as described above, except that instead of testosterone, estradiol benzoate (EB), dihydrotestosterone propionate (DHTP), a combination of EB and DHTP (EB+DHTP), or vehicle was injected neonatally. Neonatal treatment with either EB or EB+DHTP increased the number of vasopressinergic cells in the BST over that of DHTP or oil treatment. However, treatment with DHTP also significantly increased the number of vasopressinergic cells over that of oil treatment. Hence, in addition to bolstering evidence that estradiol is the more potent metabolite of testosterone in causing sexual differentiation of the brain, these data provide the first example of a masculinizing effect of a nonaromatizable androgen on a sexually dimorphic neuropeptide system.     

Androgens and the social behavior of male and female lizards (Anolis carolinensis)

This study investigated the androgen specificity of aggressive and sexual behavior in the lizard Anolis carolinensis and the capacity of females of this species to exhibit male-typical copulation. Gonadectomized males and females were injected with testosterone propionate (TP) or dihydrotestosterone propionate (DHTP) or were implanted with Silastic tubing containing TP or DHTP. Either TP or DHTP activated male-typical sexual behavior in both males and females and activated aggressive behavior in males; DHTP activated aggressive behavior in females. Thus conversion of androgen to estrogen is not essential for these behavior patterns, and endogenous dihydrotestosterone may be important. TP but not DHTP stimulated receptivity in females, suggesting that conversion of testosterone to estrogen may underlie TP-stimulated receptivity. Females treated with TP did not differ from males in their display of male-typical courtship, neck-clasping, and intromission.     

Organizational effects of testosterone,estradiol, and dihydrotestosterone on vasopressin mRNA expression in the bed nucleus of the stria terminalis

In adulthood, male rats express higher levels of arginine vasopressin (AVP) mRNA in the bed nucleus of the stria terminalis (BST) than do female rats. We tested whether this sex difference is primarily due to differences in neonatal levels of testosterone. Male and female rats were gonadectomized on the day of birth and treated with testosterone propionate (TP) or vehicle on postnatal days 1, 3, and 5 (P1, P3, and P5). Three months later, all rats were implanted with testosterone‐filled capsules. Two weeks later, brains were processed for in situ hybridization to detect AVP mRNA. We found that neonatal TP treatment significantly increased the number of vasopressinergic cells in the BST over control injections. We then sought to determine the effects of testosterone metabolites, estradiol and dihydrotestosterone, given alone or in combination, on AVP expression in the BST. Rat pups were treated as described above, except that instead of testosterone, estradiol benzoate (EB), dihydrotestosterone propionate (DHTP), a combination of EB and DHTP (EB+DHTP), or vehicle was injected neonatally. Neonatal treatment with either EB or EB+DHTP increased the number of vasopressinergic cells in the BST over that of DHTP or oil treatment. However, treatment with DHTP also significantly increased the number of vasopressinergic cells over that of oil treatment. Hence, in addition to bolstering evidence that estradiol is the more potent metabolite of testosterone in causing sexual differentiation of the brain, these data provide the first example of a masculinizing effect of a nonaromatizable androgen on a sexually dimorphic neuropeptide system. © 2003 Wiley Periodicals, Inc. J Neurobiol 54: 502–510, 2003     

Circadian rhythms of testosterone-dependent behaviors,crowing and locomotor activity,in male Japanese quail

Summary An apparatus was devised to record crowing (mate calling by males) together with locomotor activity and recorded data was analyzed by several methods for rhythm analysis. Crowing and locomotor activity of Japanese quail held on long days were recorded during sexual development as estimated from circulating gonadotropins and testosterone. Both behaviors were testosterone-dependent but commencement of crowing preceded the increase in locomotor activity. When the two behaviors attained their maximum levels, crowing showed consistent daily rhythms in which two peaks were apparent, a major one at the onset of light and a broader one 8 hours later. Locomotor activity also showed a clear daily rhythm with a peak between the two peaks of crowing rhythm suggesting a fixed phase relationship between the two rhythms.Both rhythms free-ran under constant dim light with periods shorter than 24 h. They persisted in birds which had been castrated and then supplied with exogenous testosterone via implanted Silastic capsules. The durations of both rhythms were quite comparable to each other and they maintained a fixed phase relationship similar to that found under LD cycles.The results indicate that testosterone is essential for the induction of crowing and for the enhancement of locomotor activity but the formation of the rhythms in behavior was strictly dependent on a circadian oscillatory mechanism.Abbreviations LH luteinizing hormone - FHS follicle-stimulating hormone - LD light-dark - LDim light-dim light     

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.     

Restoration of male sexual behavior by adult exogenous estrogens in male aromatase knockout mice

We previously found that male aromatase knockout (ArKO) mice that carry a targeted mutation in exons 1 and 2 of the CYP19 gene and as a result cannot aromatize androgen to estrogen show impaired sexual behavior in adulthood. To determine whether this impairment was due to a lack of activation of sexual behavior by estradiol, we studied here male coital behavior as well as olfactory investigation of sexually relevant odors in male ArKO mice following adult treatment with estradiol benzoate (EB) or dihydrotestosterone propionate (DHTP). Again, we found that gonadally intact ArKO males show pronounced behavioral deficits affecting their male coital behavior as well as their olfactory investigation of volatile body odors but not that of soiled bedding. Deficits in male coital behavior were largely corrected following adult treatment with EB and the androgen DHTP, suggesting that estradiol has prominent activational effects on this behavior. By contrast, adult treatment with EB to either castrated or gonadally intact ArKO males did not stimulate olfactory investigation of volatile body odors, suggesting that this impairment may result from a lack of proper organization of this behavior during ontogeny due to the chronic lack of estrogens. In conclusion, the present studies suggest that the behavioral deficits in sexual behavior in male ArKO mice result predominantly from a lack of activation of the behavior by estrogens. This is in contrast with earlier pharmacological studies performed on rats and ferrets that have suggested strong organizational effects of estradiol on male sexual behavior.     

Sex-specific, behavior-specific actions of dihydrotestosterone: Activation of aggression, but not mounting in ovariectomized guinea pigs

Adult ovariectomized guinea pigs were tested for aggressive behavior during treatments with estradiol benzoate (EB), testosterone propionate (TP), dihydrotestosterone propionate (DHTP), or with DHTP + EB. Aggression was not influenced by EB, but was augmented by all other steroid treatments. DHTP given by itself was not as effective as TP, but was significantly potentiated by the concurrent administration of EB. When tested for mounting behavior, ovariectomized guinea pigs were refractory to DHTP and to DHTP + EB, whereas they mounted when given TP. The findings suggest that the hormone-sensitive neural systems which mediate aggression in female guinea pigs have in part different steroid requirements from those subserving the activation of mounting. In addition, the findings emphasize that DHTP + EB administration does not always duplicate the effects of TP for behavioral endpoints, since DHTP + EB and TP had similar effects on aggression, but quite different effects on mounting in female guinea pigs. These results stand in contrast to those obtained with male guinea pigs, in which DHTP has been reported to be as effective as TP for the activation of mounting. It is hypothesized that both sex-specific and hormone-specific activational phenomena may be genetically regulated by factors separate from those responsible for the establishment of prenatal hormonal conditions.     

Neonatal dihydrotestosterone and estrogen stimulation: Effects on sexual behavior of male rats

Male rats castrated on the second day after birth (Day 2) were, for the next 10 days, given daily injections of one of five steroids or steroid combinations: 200 μg of testosterone propionate (TP); 200 μg of dihydrotestosterone propionate (DHTP); 5 μg of estradiol benzoate (EB); 5 μg of estradiol benzoate plus 200 μg of dihydrotestosterone propionate; oil vehicle (VH). Control male rats castrated on Day 90 received a sham castration and oil vehicle in the neonatal period. All animals were given TP in adulthood and tested for male sexual behavior. There was no difference in mounting activity among the subjects. Day 2 DHTP subjects displayed intromissions but were incapable of ejaculating. The more frequent display of intromissions by Day 2 DHTP animals in comparison to Day 2 VH animals could be solely due to their larger and more highly developed penes. On the other hand, the ejaculatory failure of the Day 2 DHTP subjects was attributed to some deficiency in central neural processes controlling ejaculatory mechanisms rather than inadequate penile development. Equivocal results were obtained with the Day 2 EB and Day 2 EB-DHTP animals in that only a few animals in both groups showed an ejaculatory pattern.     

Mounting behavior of female guinea pigs after prenatal and adult administration of the propionates of testosterone, dihydrotestosterone, and androstanediol.

Female guinea pigs were exposed prenatally (Day 28–58) to the propionates of testosterone (TP), dihydrotestosterone (DHTP), or androstanediol (ADP). Only TP females failed to display lordosis in adulthood after estrogen and progesterone treatment. When given TP in adulthood, females in all groups mounted, but TP and DHTP females showed augmented intromission frequencies and higher percentages of correctly oriented mounts relative to controls. Moreover, TP females responded more quickly to TP injections in adulthood and had higher over-all mounting frequencies than other groups, while DHTP females displayed mounting frequencies intermediate to controls and TP females. ADP females were not different from controls for any measure of mounting behavior.No female in any group mounted when given DHTP in adulthood, even after 7 wk of daily injections. Since male guinea pigs do mount in response to DHTP given in adulthood, the results raise the possibility that mechanisms determining sensitivity to specific steroids may not be mediated exclusively by steroids during critical periods of embryological differentiation.     

Implantation of dihydrotestosterone propionate into the lateral septum or medial amygdala facilitates copulation in castrated male rats given estradiol systemically

Two experiments were conducted to determine whether unilateral implantation of dihydrotestosterone propionate (DHTP) into different brain regions of castrated rats, bearing silastic capsules containing estradiol, could augment sexual behavior without appreciable leakage of androgen into the peripheral circulation. In Experiment 1 implanation of pulverized crystalline DHTP (via 25-gauge, 1-mm-long pellets) facilitated mating significantly without stimulating penile spine growth, provided the pellets were positioned in the lateral septum or medial amygdala. Insertion of DHTP pellets into the preoptic area-anterior hypothalamus, caudate-putamen, or the border of the substantia nigra and ventral tegmental nucleus or of cholesterol pellets into lateral septum or medial amygdala had no behavioral effects. Implanation of DHTP into the lateral septum also failed to activate penile erections in rats restrained in a supine position. In Experiment 2 implantation of different bone wax dilutions of DHTP (via 25-gauge, 1-mm-long pellets) into the preoptic area-anterior hypothalamus augmented males' sexual performance only in that group in which penile spine growth was also significantly stimulated. The results sugggst that 5α-reduced androgen is capable of activating mating in the male rat by acting locally in the lateral septum and/ or medial amygdala.     

The relationship between visual exploratory behavior and spontaneous activity in rhesus monkeys (Macaca mulatta)

Ten rhesus monkeys underwent a visual exploration test with steady or flickering light as a stimulus. The frequency and cumulative time of peeping or observing responses was measured for each stimulus. The results showed that the relative effectiveness of steady or flickering light as an incentive could not be found differentially; the visual stimulus, however, whether steady or flickering light, elicited more peeping responses than the slit without any light, i.e., the operant condition. These peeping responses were rapidly satiated not only within a session, but also between-sessions as well as between-days. Independent measures of spontaneous activity obtained concurrently with visual exploration showed that spontaneous activity is relatively stable regardless of ongoing peeping behavior. It was found that two measures of frequency and duration on peeping behavior were highly correlated.     

Effects of exogenous steroids on androgen receptors in fetal guinea pig brain

We treated pregnant guinea pigs on Day 50 of gestation with 10 mg testosterone propionate (TP), obtaining fetuses 2, 4, 8, or 18 h later as well as after 5 days of treatment. In a second group of pregnant guinea pigs, dihydrotestosterone propionate (DHTP), estradiol benzoate (E2B), progesterone (P), or cortisol was given 2 h before obtaining fetuses. Although TP treatment elevated fetal serum T (p less than 0.05), brain cytosolic androgen receptor (ARc) content was unchanged in fetuses of either sex. In female fetuses, nuclear androgen receptors (ARn) increased 10-fold in medial-basal hypothalamus (MBH) and preoptic area (POA) at 2 and 4 h (respectively) after treatment, while fetal male ARn content was unchanged. Maternal injection of other steroids (E2B, P, or cortisol, but not DHTP) significantly increased these hormones in the fetus 2 h later (p less than 0.05). Only androgens affected fetal androgen receptor (AR) content. While TP increased ARn in female MBH, DHTP decreased ARc in fetal anterior pituitary of both sexes. In this latter case, a metabolite of DHT may mediate the effects. We conclude that T crosses the guinea pig placenta and activates ARn in POA and MBH of female fetuses; male ARn appear to be maximally occupied by endogenous T. Steroids of other classes do not induce AR responses in fetal guinea pig brain. These AR changes may represent an initial cellular mechanism in brain sexual differentiation.     

Sexual differentiation in quail: Conversion of androgen to estrogen mediates testosterone-induced demasculinization of copulation but not other male characteristics

Previous research has shown that administration of either testosterone or estradiol to male quail embryos will demasculinize behavior and morphology. Six experiments in which embryos were treated were conducted to test the hypothesis that this testosterone-induced demasculinization is due to conversion of testosterone to estrogen (aromatization). In Experiment 1, dihydrotestosterone propionate, a nonaromatizable androgen, failed to demasculinize copulatory behavior, but did demasculinize crowing, strutting, and proctodeal glands. In Experiment 2, injection of the aromatizable androgens testosterone propionate (TP), testosterone, or androstenedione demasculinized copulatory behavior, the nonaromatizable androgen androsterone failed to have such an effect, and all androgens demasculinized proctodeal glands. In Experiment 3, Silastic implants of testosterone demasculinized all male characteristics, whereas implants of androsterone demasculinized only proctodeal glands. In Experiment 4, the antiestrogen tamoxifen prevented TP from demasculinizing copulatory behavior, but had no such effect with respect to crowing and strutting. In Experiments 5 and 6, the aromatization inhibitor 1,4,6-androstatrien-3,17-dione (ATD) prevented TP but not estradiol benzoate from demasculinizing copulatory behavior. Thus (1) in quail, testosterone-induced demasculinization of copulatory behavior is due to androgen aromatization, whereas testosterone-induced demasculinization of crowing, strutting, and proctodeal glands is not; (2) the distinct components of normal male reproductive behavior exhibit different patterns of steroid specificity during the organizational period, as was previously shown for the activational period; (3) the steroid specificity of crowing, strutting, and proctodeal glands changes between the organizational and activational periods. During organization, there is little specificity, whereas during activation, these characteristics respond only to androgens, never to estrogens. This difference suggests that developmental changes have occurred in the underlying biochemical substrates.