References of "Schumacher, M"
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See detailSexual Differentiation in Quail: Critical Period and Hormonal Specificity
Schumacher, M.; Hendrick, J. C.; Balthazart, Jacques ULg

in Hormones & Behavior (1989), 23(1), 130-49

There is a discrepancy between results showing that male quail are demasculinized by exogenous estrogens only if the treatment is given before Day 12 of egg incubation and results showing that ovariectomy ... [more ▼]

There is a discrepancy between results showing that male quail are demasculinized by exogenous estrogens only if the treatment is given before Day 12 of egg incubation and results showing that ovariectomy of females after hatching still affects their sexual differentiation which leads to the conclusion that female demasculinization by ovarian estrogens is a continuing process extending into posthatching life. The first experiment was performed to test different models which have been proposed to reconcile these apparently contradictory results. Male and female quail were treated with 0, 5, or 25 micrograms of estradiol benzoate (EB) on either Day 9 or Day 14 of embryonic life. Birds were castrated at the age of 4 days to avoid the confounding effects of postnatal gonadal hormones and were treated as adults with testosterone (T). Whereas EB-treatment demasculizined sexual behavior and cloacal gland growth of males when administered on Day 9, it was without effect on Day 14. This result confirms the presence of a "critical period" for sexual differentiation of behavior in embryonic life. However, the time course of sexual differentiation and the sensitivity to the demasculinizing actions of estrogens were not the same for different behavioral and morphological characteristics. Some dependent variables such as plasma levels of luteinizing hormone and crowing were still affected by the EB treatment on Day 14. These results show that the whole process of demasculinization is not retricted to the "critical period" ending on Day 12 of incubation. A second experiment was performed to determine if 5 beta-dihydrotestosterone (5 beta-DHT), a metabolite of testosterone, also exerts demasculinizing effects during embryonic life. A large dose of 5 beta-DHT (2 mg/egg) had no effects on behavior and morphology in males if administered on Day 9 of egg incubation. This suggests that 5 beta-DHT, which is a steroid devoid of behavioral effects in the adult bird, is also an inactive compound as far as sexual differentiation of the quail is concerned. The high 5 beta-reductase activity which was previously identified in the hypothalamus of the embryonic quail thus probably plays a protective role. By transforming testosterone into inactive nonaromatizable androgens, it prevents male embryos from being demasculinized by their endogenous testosterone acting through aromatization. [less ▲]

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See detailChanges in Progesterone Metabolism in the Chicken Hypothalamus During Induced Egg Laying Stop and Molting
Balthazart, Jacques ULg; Verheyen, G.; Schumacher, M. et al

in General and Comparative Endocrinology (1988), 72(2), 282-95

In the present study, we have established and validated a radioenzyme assay which permits us to quantify progesterone metabolism in the chicken brain. Progesterone metabolism was then studied in five ... [more ▼]

In the present study, we have established and validated a radioenzyme assay which permits us to quantify progesterone metabolism in the chicken brain. Progesterone metabolism was then studied in five brain areas obtained by microdissection from the telencephalon (part of the lobus paraolfactorius immediately rostral to the preoptic area), the preoptic area, and the hypothalamus. Three metabolites of progesterone were produced in large amounts in these brain regions and were quantified in this study: 5 beta-pregnane-3,20-dione (5 beta-DHP) as well as its metabolite 3 alpha-hydroxy-5 beta-pregnane-20-one (5 beta,3 alpha-ol) and 5 alpha-pregnane-3,20-dione (5 alpha-DHP). The unmetabolized progesterone was also recovered and quantified. The 5 beta-reduction of progesterone (production of 5 beta-DHP and 5 beta,3 alpha-ol) was very active but its 5 alpha-reduction (production of 5 alpha-DHP) was almost absent in the lobus paraolfactorius. An opposite pattern of metabolism was found in the preoptic area and the hypothalamus (higher 5 alpha- but lower 5 beta-reductase activity). The changes in progesterone metabolism in these brain areas were then studied in groups of hens submitted to induced egg laying stop and molting. A significant decrease in progesterone 5 alpha-reduction was found in the median hypothalamus of hens during the period of molt. Simultaneously, the experimental procedures induced significant decreases in the production of 5 beta-DHP by the lobus paraolfactorius, anterior, and medial hypothalamus but induced a significant increase in the production of this metabolite in the preoptic area. These changes are likely to be involved in the control of reproductive functions including sexual behavior and secretion of luteinizing hormone-releasing hormone, and a number of possible causal mechanisms are presented. These should now be tested experimentally especially in view of the very limited information which is now available on the biological effects of the metabolites of progesterone. [less ▲]

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See detailChanges in Serum Concentrations of Steroids During Embryonic and Post-Hatching Development of Male and Female Japanese Quail (Coturnix Coturnix Japonica)
Schumacher, M.; Sulon, Joseph ULg; Balthazart, Jacques ULg

in Journal of Endocrinology (1988), 118(1), 127-34

Serum concentrations of testosterone, 5 alpha-dihydrotestosterone, oestradiol and progesterone were measured by radioimmunoassay combined with Celite chromatography in male and female Japanese quail ... [more ▼]

Serum concentrations of testosterone, 5 alpha-dihydrotestosterone, oestradiol and progesterone were measured by radioimmunoassay combined with Celite chromatography in male and female Japanese quail (Coturnix coturnix japonica) during the second half of embryonic life (days 9-17 of incubation) and during the first 5 weeks after hatching. The mean level of each of the four steroids was significantly affected by the age of the birds. An overall effect of sex was detected by analysis of variance only on oestradiol concentrations, with females having higher serum concentrations than males during most of the age range studied. Significant peaks of testosterone and progesterone were also detected around hatching time. These results are consistent with the view that oestradiol is the major hormone implicated in the sexual differentiation of reproductive behaviour in the quail. The relationships between the circulating concentrations of oestradiol during ontogeny and the critical period of differentiation as postulated by currently accepted models is also discussed. [less ▲]

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See detailNeuroanatomical Distribution of Testosterone-Metabolizing Enzymes in the Japanese Quail
Schumacher, M.; Balthazart, Jacques ULg

in Brain Research (1987), 422(1), 137-48

We describe a very sensitive and precise assay which allows one to study the metabolism of testosterone (T) in small brain nuclei dissected out according to the method of Palkovits and Brownstein. With ... [more ▼]

We describe a very sensitive and precise assay which allows one to study the metabolism of testosterone (T) in small brain nuclei dissected out according to the method of Palkovits and Brownstein. With this method, the neuroanatomical distributions of aromatase, and 5 alpha- and 5 beta-reductase activities were studied in adult male quail (Coturnix coturnix japonica). The different enzymes show different neuroanatomical distributions. Production of estradiol-17 beta (E2) was highest in the sexually dimorphic nucleus preopticus medialis (POM). We showed previously that the preoptic aromatase activity is higher in male than in female quail. As the POM is a central and very large structure within the preoptic area, the present results suggest a relationship between the neuroanatomical and the biochemical sex differences. By contrast, the production of 5 alpha-DHT was highest in the lateral hypothalamic area (LHY), the bed nucleus of the pallial commissure (BPC) and the lateral septum (SL). The 5 beta-reductase activity was highest in the lateral septum and in the ventral part of the archistriatum (AV). Moreover, there was a rostral to caudal decrease in 5 beta-reductase activity in the hypothalamus. [less ▲]

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See detailSteroid Hormones, Behavior and Sexual Dimorphism in Animals and Men: The Nature-Nurture Controversy
Schumacher, M.; Legros, Jean-Jacques ULg; Balthazart, Jacques ULg

in Experimental & Clinical Endocrinology (1987), 90(2), 129-56

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See detailInteractions des androgenes et des oestrogenes dans le controle de la reproduction
Schumacher, M.; Alexandre, Corine; Balthazart, Jacques ULg

in Comptes Rendus de l'Académie des Sciences. Série III, Sciences de la Vie (1987), 305(14), 569-74

Castrated male quail were injected with the synthetic oestrogen, diethylstylbestrol (DES) or the synthetic androgen, methyltrienolone (R 1881) or both compounds simultaneously. Both R 1881 and DES ... [more ▼]

Castrated male quail were injected with the synthetic oestrogen, diethylstylbestrol (DES) or the synthetic androgen, methyltrienolone (R 1881) or both compounds simultaneously. Both R 1881 and DES activated male sexual behaviour, inhibited LH and FSH secretion and increased hypothalamic aromatase activity. Additive effects between R 1881 and DES were observed for the induction of brain aromatase and for the inhibition of FSH secretion. As a consequence, mechanisms mediated by androgen and estrogen receptors must be involved in the control of these reproductive characteristics. [less ▲]

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See detailSexual differentiation and hormonal control of the sexually dimorphic medial preoptic nucleus in the quail.
Panzica, G. C.; Viglietti-Panzica, C.; Calacagni, M. et al

in Brain Research (1987), 416(1), 59-68

We recently identified a sexually dimorphic nucleus in the preoptic region of the Japanese quail, the medial preoptic nucleus (POM), which is significantly larger in males than in females. In the present ... [more ▼]

We recently identified a sexually dimorphic nucleus in the preoptic region of the Japanese quail, the medial preoptic nucleus (POM), which is significantly larger in males than in females. In the present study, we investigated the hormonal control of this morphological neuroanatomical difference and the possible relationships between the sexual dimorphism in POM volume and in copulatory behavior. Treatments which are known to affect sexual behavior were thus applied to different groups of birds and the POM volume was then measured. In one experiment, male and female quails were either gonadectomized, gonadectomized and treated with testosterone or left intact. The larger size of the POM in males was confirmed and treatments significantly affected the nucleus size which was decreased by gonadectomy and restored by testosterone treatment in both sexes to a level similar to that seen in intact males. In two other experiments, eggs were injected with estradiol benzoate on day 9 of incubation and the POM volume was measured in adulthood either in intact birds or in gonadectomized birds receiving a replacement therapy with testosterone. Despite the fact that estradiol benzoate treatment completely suppressed copulatory behavior, it did not affect the volume of the POM or slightly increased it. These data thus show that the POM volume is controlled by testosterone levels in adulthood and could thus be an interesting model for the study of the effects of steroids on the brain. [less ▲]

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See detailA two-step model for sexual differentiation.
Balthazart, Jacques ULg; Schumacher, M.

in Annals of the New York Academy of Sciences (1986), 474

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See detailTestosterone-induced brain aromatase is sexually dimorphic.
Schumacher, M.; Balthazart, Jacques ULg

in Brain Research (1986), 370(2), 285-93

Female quail are less sensitive than males to the activating effects of testosterone (T) on behavior. Testosterone induces hypothalamic aromatase activity more efficiently in males than in females (higher ... [more ▼]

Female quail are less sensitive than males to the activating effects of testosterone (T) on behavior. Testosterone induces hypothalamic aromatase activity more efficiently in males than in females (higher Vmax but similar Km). As the conversion of T to estradiol (E2) is important for the activation of male sexual behavior, we propose that the incapacity of the female hypothalamus to produce behaviorally active E2 contributes to their insensitivity to T. The 5 alpha-reductase, which converts T into 5 alpha-dihydrotestosterone (5 alpha-DHT), is induced by T specifically in the anterior hypothalamus of both sexes. [less ▲]

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See detailComparison of monoamine concentrations in the brains of adult male and female Japanese quail.
Ottinger, M. A.; Schumacher, M.; Clarke, R. N. et al

in Poultry Science (1986), 65(7), 1413-20

A fluorometric assay measuring brain tissue concentrations of norepinephrine, dopamine, and serotonin has been validated for Japanese quail. Accuracy, precision, specificity, and parallelism were ... [more ▼]

A fluorometric assay measuring brain tissue concentrations of norepinephrine, dopamine, and serotonin has been validated for Japanese quail. Accuracy, precision, specificity, and parallelism were determined. The sensitivity of the assays was 6 ng/tube, which allowed individual assay of 1 to 2 mg hypothalamic tissue. In Experiment 1, relatively large areas of brain from adult, reproductively active males and females were found to differ significantly in norepinephrine content in optic lobes and for dopamine in right telencephalon. A microdissection technique was used in Experiment 2 to sample small portions of hypothalamic tissue. Sex differences were observed for norepinephrine in the sections containing the lobus paraolfactorius and the preoptic, anterior, and medial hypothalamus. Differences in monoamine content were most apparent when smaller areas dissected by microdissection were analyzed. These results give evidence for sex differences in the monoamine content in specific areas of the brain. [less ▲]

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See detailBrain testosterone metabolism during ontogeny in the zebra finch.
Balthazart, Jacques ULg; Schumacher, M.; Prove, E.

in Brain Research (1986), 378(2), 240-50

The testosterone metabolism in different parts of the brain and in the syrinx was studied by an in vitro radioenzyme assay in male and female zebra finch (Taeniopygia guttata castanotis) at various ages ... [more ▼]

The testosterone metabolism in different parts of the brain and in the syrinx was studied by an in vitro radioenzyme assay in male and female zebra finch (Taeniopygia guttata castanotis) at various ages between 1 day post-hatch and adulthood. The 5 beta-reductase appeared in all experiments as the main enzyme involved in testosterone metabolism. Two experiments allowed to determine the kinetic parameters of the 5 beta-reductase. It was shown that the Vmax of the 5 beta-reduction is much higher in very young animals than in adults but that the Km of the enzyme is smaller (higher affinity) in the latter than in the former. A major decrease of 5 beta-reductase activity is observed during the ontogeny of both males and females in all the tissue samples which were studied including the hypothalamus and brain regions containing nuclei of the song system. This high reductase activity which produces behaviorally inactive compounds is likely to play a protective role during the sexual differentiation of the birds and in this way prevent the masculinization in females of the song system by the relatively high levels of circulating androgens. [less ▲]

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See detailInteraction of androgens and estrogens in the control of sexual behavior in male Japanese quail.
Balthazart, Jacques ULg; Schumacher, M.; Malacarne, G.

in Physiology & Behavior (1985), 35(2), 157-66

A series of 4 experiments was performed to study the relative contribution of androgens and estrogens in the activation of sexual behavior in castrated male quail. The synthetic androgen methyltrienolone ... [more ▼]

A series of 4 experiments was performed to study the relative contribution of androgens and estrogens in the activation of sexual behavior in castrated male quail. The synthetic androgen methyltrienolone (R 1881) which is not metabolized in androgen target tissues activated sexual behavior in castrated birds and at the dose level of 0.5-1 mg/day/animal had the same potency as testosterone (T). However R 1881 was much more active than T in the induction of cloacal gland growth and activation of crowing, two typically androgen-dependent responses. This suggests that sexual behavior is not controlled by exactly the same mechanism as crowing or cloacal gland growth. In another experiment, estradiol (E2) alone activated sexual behavior but it is only at very high doses which had clear toxic effects that a significant behavioral activation could be observed. This questions the role of E2 as the physiological agent stimulating copulation in intact birds unless it is assumed that centrally administered E2 would be much more active compared to peripheral E2 which is exposed to a very intense peripheral catabolism. In the last two experiments, a clear synergism could be detected between 5 alpha-dihydrotestosterone (5 alpha-DHT) and E2 in the activation of sexual activity and doses of hormones could be defined which had almost no activity by themselves but significantly stimulated sexual behavior when given simultaneously. It was however impossible to define a hormonal treatment with T metabolites which restored behavior to its precastration level, a result very easily achieved with T treatments. Taken together, these data suggest that activation of sexual behavior in quail does not depend only on E2, nor 5 alpha-DHT nor even on their combined action. Considering that specific T receptors which probably do not bind 5 alpha-DHT are present in the brain, it would seem justified to reconsider the possible role played by T itself in the activation of behavior. [less ▲]

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See detailSexual dimorphism in the hypothalamic metabolism of testosterone in the Japanese Quail (Coturnix coturnix japonica).
Schumacher, M.; Balthazart, Jacques ULg

in Progress in Brain Research (1984), 61

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See detailChanges in testosterone metabolism by the brain and cloacal gland during sexual maturation in the Japanese quail (Coturnix coturnix japonica).
Balthazart, Jacques ULg; Schumacher, M.

in Journal of Endocrinology (1984), 100(1), 13-8

Testosterone metabolism in the brain and pituitary and cloacal glands of male and female Japanese quail was studied in vitro during sexual maturation (from 1 day to 5 weeks after hatching). The production ... [more ▼]

Testosterone metabolism in the brain and pituitary and cloacal glands of male and female Japanese quail was studied in vitro during sexual maturation (from 1 day to 5 weeks after hatching). The production of 5 alpha-dihydrotestosterone in the hyperstriatum and cloacal gland and that of androstenedione in the cloacal gland of males was highest at 1 day after hatching, which could be related to the peak of plasma androgens previously demonstrated in neonatal quail. 5 beta-Reductase activity was very high in the brain, but not the pituitary or cloacal glands of young chicks and decreased markedly, especially in the hypothalamus, during sexual maturation. As 5 beta-reduced metabolites of testosterone are inactive androgens, it is suggested that the decrease of 5 beta-reductase activity with age corresponds to a potentiation of the effects of testosterone at the level of the brain. [less ▲]

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See detailRelative potencies of testosterone and 5 alpha-dihydrotestosterone on crowing and cloacal gland growth in the Japanese quail (Coturnix coturnix japonica).
Balthazart, Jacques ULg; Schumacher, M.; Malacarne, G.

in Journal of Endocrinology (1984), 100(1), 19-23

It has been suggested that testosterone is less effective at inducing crowing behaviour in young birds than in adults because of the presence of higher levels of steroid 5 beta-reductase in the young ... [more ▼]

It has been suggested that testosterone is less effective at inducing crowing behaviour in young birds than in adults because of the presence of higher levels of steroid 5 beta-reductase in the young brain, which converts testosterone to inactive 5 beta-reduced metabolites. This hypothesis was tested indirectly by comparing the relative potencies of 5 alpha-dihydrotestosterone (5 alpha-DHT), which cannot be converted to 5 beta-metabolites, and testosterone at inducing crowing in young gonadectomized male and female quail. The promotion of cloacal gland growth by these treatments was also assessed since there are no age-related changes in 5 beta-reductase in this organ. Silicone elastomer implants (2 X 5, 5 and 10 mm) containing 5 alpha-DHT were more effective at stimulating crowing than similar implants of testosterone whilst there was little difference in their potency at inducing cloacal gland growth. These results are consistent with the hypothesis that brain steroid 5 beta-reductase regulates the behavioural activity of testosterone in the brain of young birds. [less ▲]

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See detailPartial characterization of testosterone-metabolizing enzymes in the quail brain.
Schumacher, M.; Contenti, E.; Balthazart, Jacques ULg

in Brain Research (1984), 305(1), 51-9

The properties of 5 beta-reductase, 5 alpha-reductase and aromatase, 3 testosterone metabolizing enzymes, were studied in the quail brain by an in vitro incubation technique. The results describe the ... [more ▼]

The properties of 5 beta-reductase, 5 alpha-reductase and aromatase, 3 testosterone metabolizing enzymes, were studied in the quail brain by an in vitro incubation technique. The results describe the changes in time of metabolite production and the effects of temperature, enzyme and cofactor concentrations. The apparent Km and Vmax were evaluated for the 3 enzymes. Aromatase and 5 alpha-reductase have a higher affinity but a lower capacity than 5 beta-reductase. The kinetics of the latter enzyme are complex and suggest the presence of two types of enzymes. These characteristics fit in well with the role probably played by the enzymes in vivo. [less ▲]

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See detailOrganization and activation of behavior in quail: role of testosterone metabolism.
Balthazart, Jacques ULg; Schumacher, M.

in Journal of Experimental Zoology (The) (1984), 232(3), 595-604

In quail, the hypothalamus enzymatically transforms testosterone (T) into estradiol (E2), 5 alpha-dihydrotestosterone (5 alpha-DHT), and 5 beta-dihydrotestosterone (5 beta-DHT). During the embryonic life ... [more ▼]

In quail, the hypothalamus enzymatically transforms testosterone (T) into estradiol (E2), 5 alpha-dihydrotestosterone (5 alpha-DHT), and 5 beta-dihydrotestosterone (5 beta-DHT). During the embryonic life, the 5 beta-reductase activity is very high, which probably protects the brain of males from being behaviorally demasculinized by their endogenous T. 5 beta androstanes are inactive androgens. The decrease of 5 beta reductase with age during sexual maturation corresponds to a potentiation of the effects of T as shown by experiments that compared the effects of T and 5 alpha-DHT in adult and young quail. T metabolism is also involved in the activation of male behavior in the adult. T aromatization is probably essential for behavioral activation, but nonaromatizable androgens such as methyltrienolone, and to some extent 5 alpha-DHT, can also stimulate sexual behavior in castrates. These enzymatic activities show a clear neuroanatomical localization and are sexually dimorphic. Males produce more active metabolites (E2, 5 alpha-DHT) than females, which could explain the male's greater sensitivity to T treatments. It thus appears that T metabolism is involved in the differentiation and activation of behavior in quail. [less ▲]

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See detailThe postnatal demasculinization of sexual behavior in the Japanese quail (Coturnix coturnix japonica).
Schumacher, M.; Balthazart, Jacques ULg

in Hormones and Behavior (1984), 18(3), 298-312

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 ... [more ▼]

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. [less ▲]

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See detailEstradiol contributes to the postnatal demasculinization of female Japanese quail (Coturnix coturnix japonica).
Balthazart, Jacques ULg; Schumacher, M.

in Hormones and Behavior (1984), 18(3), 287-97

Two experiments were performed to characterize the process of postnatal demasculinization in Japanese quail. In the first experiment, it was shown that estradiol (E2) can complete female demasculinization ... [more ▼]

Two experiments were performed to characterize the process of postnatal demasculinization in Japanese quail. In the first experiment, it was shown that estradiol (E2) can complete female demasculinization during the first 4 weeks of life. By contrast, E2 did not demasculinize sexual behavior and cloacal gland in neonatally castrated males. Neonatally gonadectomized females preferentially performed mount attempts when tested in their home cage by comparison to a test arena. In Experiment 2, E2 Silastic implants (40-mm) maintained full copulatory behavior in castrated males but not in females. This large dose of E2 did not demasculinize adult sexually active birds (males or females) even if treatment lasted for 1 month. It is concluded that E2 can demasculinize sexual behavior only in females and only if treatment is performed in very young birds. [less ▲]

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See detailThe effects of testosterone and its metabolites on sexual behavior and morphology in male and female Japanese quail.
Schumacher, M.; Balthazart, Jacques ULg

in Physiology & Behavior (1983), 30(3), 335-9

Adult Japanese quail are sexually dimorphic. Even when implanted with testosterone (T), ovariectomized females fail to copulate and their cloacal glands are smaller than those of males. This may be due to ... [more ▼]

Adult Japanese quail are sexually dimorphic. Even when implanted with testosterone (T), ovariectomized females fail to copulate and their cloacal glands are smaller than those of males. This may be due to a reduced capacity of the females to transform testosterone into active metabolites (estradiol-17 beta and 5 alpha-dihydrotestosterone). Indeed, in the male quail, estradiol-17 beta (E2) activates copulation whereas 5 alpha-dihydrotestosterone (5 alpha-DHT) activates crowing, strutting and the development of the cloacal gland. To test this hypothesis, we studied the effects of in vivo treatments of male and female quail with the different T-metabolites. Forty-one castrated male and female quail were implanted with subcutaneous silastic implants of T, 5 alpha-DHT, E2 and E2 in combination with 5 alpha-DHT. When implanted with these metabolites, females failed to copulate and their cloacal glands remained less developed than those of males. Sexual differences in behavior and morphology thus cannot be entirely explained by sexual dimorphism of the metabolism. [less ▲]

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