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See detailVasotocinergic Innervation of Areas Containing Aromatase-Immunoreactive Cells in the Quail Forebrain
Balthazart, Jacques ULg; Absil, Philippe ULg; Viglietti-Panzica, C. et al

in Journal of Neurobiology (1997), 33(1), 45-60

In the male quail forebrain, aromatase-immunoreactive (ARO-ir) elements are clustered within the sexually dimorphic medial preoptic nucleus (POM), nucleus striae terminalis (nST), nucleus accumbens (nAc ... [more ▼]

In the male quail forebrain, aromatase-immunoreactive (ARO-ir) elements are clustered within the sexually dimorphic medial preoptic nucleus (POM), nucleus striae terminalis (nST), nucleus accumbens (nAc), and ventromedial and tuberal hypothalamus. These ARO-ir cells are sensitive to testosterone and its metabolites: Their number and size increase after exposure to these steroids. The POM and lateral septum are also characterized by a dense vasotocinergic innervation that is also sensitive to testosterone. We analyzed here the anatomical relationships between ARO-ir elements and VT-ir fibers in the quail prosencephalon. Sequential staining for vasotocin, aromatase, or vasotocin plus aromatase was performed on adjacent 30-microm-thick cryostat sections. High concentrations of thin VT-ir fibers were observed within the POM, nST, lateral septum, periventricular mesencephalic central gray, and ventromedial and tuberal hypothalamus. There was a close correspondence between the extension of the ARO-ir cells and of VT-ir fibers. In double-labeled sections, all clusters of ARO-ir cells with the exception of those located in the nAc were embedded in a dense network of VT-ir fibers. Many of the VT-ir terminals appeared to end in the neuropile surrounding ARO-ir elements rather than directly on their cell bodies. This study supports the idea that the testosterone-dependent aromatase system is directly innervated by a testosterone-dependent peptidergic system. Aromatase-containing cells could therefore be modulated by steroids both directly and indirectly through the vasotocin system. Alternatively, this neuroanatomical arrangement may mediate the control of vasotocin synthesis or release by steroids. Functional studies demonstrate that both aromatase and vasotocin affect reproductive behavior in quail, and the present data provide anatomical support for the integration of these effects. [less ▲]

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See detailIdentification of Catecholaminergic Inputs to and Outputs from Aromatase-Containing Brain Areas of the Japanese Quail by Tract Tracing Combined with Tyrosine Hydroxylase Immunocytochemistry
Balthazart, Jacques ULg; Absil, Philippe ULg

in Journal of Comparative Neurology (The) (1997), 382(3), 401-28

In the quail brain, aromatase-immunoreactive (ARO-ir) neurons located in the medial preoptic nucleus (POM) and caudal paleostriatum ventrale/nucleus accumbens/nucleus striae terminalis complex (PVT/nAc ... [more ▼]

In the quail brain, aromatase-immunoreactive (ARO-ir) neurons located in the medial preoptic nucleus (POM) and caudal paleostriatum ventrale/nucleus accumbens/nucleus striae terminalis complex (PVT/nAc/nST) receive catecholaminergic inputs identified by the presence of tyrosine hydroxylase-immunoreactive (TH-ir) fibers and punctate structures. The origin of these inputs was analyzed by retrograde tracing with cholera toxin B subunit (CTB) or red latex fluospheres (RLF) combined with TH immunocytochemistry. CTB and RLF injected in the POM or PVT/nAc/nST were found in cells located in anatomically discrete areas in the telencephalon (hippocampus, septum, archistriatum), hypothalamus (many areas in periventricular position), thalamus, mesencephalon, and pons. In these last two regions, many retrogradely labeled cells were located in dopaminergic areas such as the retroruberal field (RRF), substantia nigra (SN), and area ventralis of Tsai (AVT) but also in noradrenergic cell groups such as the locus ceruleus and subceruleus. CTB tracing showed that most of these connections are bidirectional. Many retrogradely labeled cells contained TH-ir material. As a mean, 10-20% and 40-60% of the RLF-containing cells in the dopaminergic areas were TH-ir when RLF had been injected in the POM or PVT/nAc/nST, respectively. TH-ir cells projecting to the POM appeared to be mostly located in the periventricular hypothalamus and in AVT, whereas projections to the PVT/nAc/nST originated mainly in the SN (with significant contributions from the RRF and AVT). These data support the existence of functional relationships between aromatase and catecholamines. [less ▲]

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See detailSteroid Control and Sexual Differentiation of Brain Aromatase
Balthazart, Jacques ULg

in Journal of Steroid Biochemistry & Molecular Biology (1997), 61(3-6), 323-39

Brain aromatase (ARO) activity in the quail is markedly enhanced by testosterone (T). This effect only becomes detectable after several hours and reaches its maximum within a few days, which suggests ... [more ▼]

Brain aromatase (ARO) activity in the quail is markedly enhanced by testosterone (T). This effect only becomes detectable after several hours and reaches its maximum within a few days, which suggests enzymatic induction at the genomic level. This idea is reinforced by the fact that T also increases the ARO protein, as observed by immunocytochemistry (ICC) and the ARO mRNA, as measured by reverse transcriptase-polymerase chain reaction (RT-PCR). These changes can be mimicked by the administration of estrogens and therefore presumably require T aromatization. In our first test, injection of the non-steroidal ARO inhibitor, R76713 (racemic vorozole), unexpectedly revealed an increase in ARO immunoreactivity in the preoptic area (POA) of treated birds. This property of R76713 was shared by another non-steroidal inhibitor, fadrozole, but not by two steroidal inhibitors, androstatrienedione (ATD) and 4-hydroxy-androstenedione (OHA). These last two compounds markedly decreased the concentration of brain ARO as estimated by ICC. In parallel, ATD and OHA decreased ARO mRNA concentration measured by RT-PCR but vorozole and fadrozole had no effect on these concentrations in the POA, and only caused them to decrease slightly in the posterior hypothalamus. Together, these data indicate that the removal of estrogens caused by steroidal inhibitors decreases the synthesis of ARO, presumably at the transcriptional level. Additional regulatory mechanisms apparently take place after the injection of non-steroidal inhibitors and probably include increased half-life of the protein. The induction of ARO activity by steroids appears to be greater in males than in females, but this difference has been difficult to localize and confirm by assay methods. We therefore analysed by ICC the tridimensional distribution of ARO-ir neurons in the POA of males and females that were sexually mature or gonadectomized and treated with T-filled or control empty implants. Localized sex differences and effects of T were detected in this way. In particular, males had more ARO-ir cells than females in the lateral POA but a difference in the opposite direction was evident in the medial part of this area. These sex differences are largely activational (i.e. caused by the higher T levels in males) but they may also reflect organizational effects of neonatal steroids. Castration decreased ARO-ir cell numbers in the lateral POA, but increased it in the periventricular region. This anatomically specialized control by T may be mediated by three potential mechanisms that are discussed and comparatively evaluated: a migration of ARO neurons towards the ventricle after castration; a differential colocalization of ARO with estrogen receptors or a differential modulation of ARO neurons by catecholaminergic inputs. [less ▲]

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See detailAromatase Inhibition Blocks the Activation and Sexual Differentiation of Appetitive Male Sexual Behavior in Japanese Quail
Balthazart, Jacques ULg; Castagna, C.; Ball, G. F.

in Behavioral Neuroscience (1997), 111(2), 381-97

Two experiments investigated the role of estrogens in the activation and sexual differentiation of appetitive sexual behavior (ASB) in Japanese quail (Coturnix japonica) as measured by a learned social ... [more ▼]

Two experiments investigated the role of estrogens in the activation and sexual differentiation of appetitive sexual behavior (ASB) in Japanese quail (Coturnix japonica) as measured by a learned social proximity response. Injection of the aromatase inhibitor R767 13 in castrated, testosterone (T)-treated male quail completely suppressed ASB, confirming that, like consummatory sexual behavior, ASB is mediated by T aromatization. ASB is not observed in female quail, even if they are treated with T as adults. The role of embryonic estrogens in the sexual differentiation of ASB was tested by blocking estrogen synthesis in ovo. Control male and T-treated female quail deprived of estrogens during embryonic life learned the social proximity response used to assess ASB, whereas control female quail did not, despite the presence of high T. Thus, ASB is demasculinized by the action of embryonic estrogens during ontogeny as is consummatory behavior. [less ▲]

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See detailA Direct Dopaminergic Control of Aromatase Activity in the Quail Preoptic Area
Baillien, M.; Balthazart, Jacques ULg

in Journal of Steroid Biochemistry & Molecular Biology (1997), 63(1-3, Sep-Oct), 99-113

In the quail preoptic area (POA) anatomical and pharmacological data suggest that catecholamines may be implicated in the control of testosterone (T) aromatization into estrogens. The biochemical ... [more ▼]

In the quail preoptic area (POA) anatomical and pharmacological data suggest that catecholamines may be implicated in the control of testosterone (T) aromatization into estrogens. The biochemical mechanism(s) mediating this control of the enzyme activity is (are) however unexplored. The present studies were carried out to investigate whether the catecholamines, dopamine (DA) and norepinephrine (NE) are able to directly affect aromatase activity (AA) measured during in vitro incubations of POA homogenates. AA was quantified in the POA-hypothalamus of adult male Japanese quail by measuring the tritiated water production from [1beta-3H]-androstenedione. Enzyme activity was linear as a function of the incubation time and of the protein content of homogenates. It exhibited a typical Michaelis-Menten kinetics, with an apparent Km of 2.8 nM and a Vmax of 266.6 fmol h(-1) mg wet weight(-1). AA was then measured at a substrate concentration of 25 nM in the presence of catecholamines and some of their receptor agonists or antagonists, at two concentrations, 10(-3) and 10(-6) M. Norepinephrine and prazosin (alpha1-adrenergic antagonist) had no or very limited effects on AA at both concentrations. In contrast, DA and some D1 and/or D2 receptor agonists (apomorphine[D1/D2], SKF-38393 [D1] and RU-24213 [D2]) depressed AA by 40 to 70% at the 10(-3) M concentration. One D2 receptor antagonist also produced a major inhibition of AA (sulpiride) while other antagonists either had no significant effect or only produced moderate decreases in enzyme activity (SCH-23390 [D1], spiperone [D2], pimozide [D2]) as did two DA indirect agonists, amfonelic acid and nomifensine. The inhibitory effect of the agonists was not antagonized by the less active antagonists, SCH-23390 [D1] or spiperone [D2]. Taken together these results suggest that the inhibitory effects do not involve specific binding of DA or its agonists/antagonists to dopaminergic receptors mediating changes in cAMP concentration. This conclusion is also supported by the observation that addition of dibutyryl cAMP did not change brain AA. It appears more likely that DA and dopaminergic drugs inhibit AA by a direct effect on the enzyme, as suggested by the competitive nature of DA and SKF-38393 inhibition of AA (Ki's of 59 and 84 microM, respectively). The functional significance of this effect should still be demonstrated but this mechanism may represent an important physiological pathway through which neurotransmitters could rapidly affect steroid-dependent processes such as the neural synthesis of estrogens. This would provide a mean by which environmental stimuli could affect reproductive behavior and physiology. [less ▲]

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See detailDo Sex Differences in the Brain Explain Sex Differences in the Hormonal Induction of Reproductive Behavior? What 25 Years of Research on the Japanese Quail Tells Us
Balthazart, Jacques ULg; Tlemcani, O.; Ball, G. F.

in Hormones & Behavior (1996), 30(4), 627-61

Early workers interested in the mechanisms mediating sex differences in morphology and behavior assumed that differences in behavior that are commonly observed between males and females result from the ... [more ▼]

Early workers interested in the mechanisms mediating sex differences in morphology and behavior assumed that differences in behavior that are commonly observed between males and females result from the sex specificity of androgens and estrogens. [less ▲]

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See detailDistribution of Aromatase-Immunoreactive Cells in the Forebrain of Zebra Finches (Taeniopygia Guttata): Implications for the Neural Action of Steroids and Nuclear Definition in the Avian Hypothalamus
Balthazart, Jacques ULg; Absil, Philippe ULg; Foidart, Agnès ULg et al

in Journal of Neurobiology (1996), 31(2), 129-48

Cells immunoreactive for the enzyme aromatase were localized in the forebrain of male zebra finches with the use of an immunocytochemistry procedure. Two polyclonal antibodies, one directed against human ... [more ▼]

Cells immunoreactive for the enzyme aromatase were localized in the forebrain of male zebra finches with the use of an immunocytochemistry procedure. Two polyclonal antibodies, one directed against human placental aromatase and the other directed against quail recombinant aromatase, revealed a heterogeneous distribution of the enzyme in the telencephalon, diencephalon, and mesencephalon. Staining was enhanced in some birds by the administration of the nonsteroidal aromatase inhibitor, R76713 racemic Vorozole) prior to the perfusion of the birds as previously described in Japanese quail. Large numbers of cells immunoreactive for aromatase were found in nuclei in the preoptic region and in the tuberal hypothalamus. A nucleus was identified in the preoptic region based on the high density of aromatase immunoreactive cells within its boundaries that appears to be homologous to the preoptic medial nucleus (POM) described previously in Japanese quail. In several birds alternate sections were stained for immunoreactive vasotocin, a marker of the paraventricular nucleus (PVN). This information facilitated the clear separation of the POM in zebra finches from nuclei that are adjacent to the POM in the preoptic area-hypothalamus, such as the PVN and the ventromedial nucleus of the hypothalamus. Positively staining cells were also detected widely throughout the telencephalon. Cells were discerned in the medial parts of the ventral hyperstriatum and neostriatum near the lateral ventricle and in dorsal and medial parts of the hippocampus. They were most abundant in the caudal neostriatum where they clustered in the dorsomedial neostriatum, and as a band of cells coursing along the dorsal edge of the lamina archistriatalis dorsalis. They were also present in high numbers in the ventrolateral aspect of the neostriatum and in the nucleus taeniae. None of the telencephalic vocal control nuclei had appreciable numbers of cells immunoreactive for aromatase within their boundaries, with the possible exception of a group of cells that may correspond to the medial part of the magnocellular nucleus of the neostriatum. The distribution of immunoreactive aromatase cells in the zebra finch brain is in excellent agreement with the distribution of cells expressing the mRNA for aromatase recently described in the finch telencephalon. This widespread telencephalic distribution of cells immunoreactive for aromatase has not been described in non-songbird species such as the Japanese quail, the ring dove, and the domestic fowl. [less ▲]

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See detailLocalization of Testosterone-Sensitive and Sexually Dimorphic Aromatase-Immunoreactive Cells in the Quail Preoptic Area
Balthazart, Jacques ULg; Tlemcani, O.; Harada, N.

in Journal of Chemical Neuroanatomy (1996), 11(3), 147-71

The distribution of aromatase-immunoreactive cells was studied in the medial preoptic nucleus of male and female quail that were sexually mature and gonadally intact, or gonadectomized, or gonadectomized ... [more ▼]

The distribution of aromatase-immunoreactive cells was studied in the medial preoptic nucleus of male and female quail that were sexually mature and gonadally intact, or gonadectomized, or gonadectomized and treated with testosterone. The study first confirmed the existence of a significant difference in the number of aromatase-immunoreactive cells between males and females (males > females) and the marked effect of castration and testosterone treatment which, respectively, decrease and restore the number of these cells. An analysis of the distribution in space of this neurochemically defined cell population was also carried out. This study revealed that castration does not uniformly decrease the density of aromatase-immunoreactive cells, but local increases are observed in an area directly adjacent to the third ventricle. A number of new sex differences in the organization of the medial preoptic nucleus and its population of aromatase cells have, in addition, been identified. The density of aromatase-immunoreactive cells is not higher in males than in females throughout the nucleus, but a higher density of immunoreactive cells is present in the ventromedial part of the nucleus in females as compared to males. In addition, the cross-sectional area of the nucleus as defined by the population of aromatase-immunoreactive cells is larger in males than in females in its rostral part and its shape is more elongated in the dorso-ventral direction in females than in males. Some of these differences (e.g. higher density of ARC-ir cells in the ventromedial part of the female POM, shape of the nucleus) appear to be organizational in nature, because they are still present in birds exposed to the same endocrine conditions during adult life (e.g. gonadectomized and treated with a same dose of testosterone). This conclusion should now be tested by experiments manipulating the endocrine environment of quail embryos. The anatomical heterogeneity of the medial preoptic nucleus revealed by this study also suggests a functional heterogeneity and the specific roles of the medial and lateral parts of the nucleus should also be investigated. [less ▲]

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See detailEffects of Testosterone and Its Metabolites on Aromatase-Immunoreactive Cells in the Quail Brain: Relationship with the Activation of Male Reproductive Behavior
Balthazart, Jacques ULg; Foidart, Agnès ULg; Absil, Philippe ULg et al

in Journal of Steroid Biochemistry & Molecular Biology (1996), 56(1-6 Spec No), 185-200

The enzyme aromatase converts testosterone (T) into 17 beta-estradiol and plays a pivotal role in the control of reproduction. In particular, the aromatase activity (AA) located in the preoptic area (POA ... [more ▼]

The enzyme aromatase converts testosterone (T) into 17 beta-estradiol and plays a pivotal role in the control of reproduction. In particular, the aromatase activity (AA) located in the preoptic area (POA) of male Japanese quail is a limiting step in the activation by T of copulatory behavior. Aromatase-immunoreactive (ARO-ir) cells of the POA are specifically localized within the cytoarchitectonic boundaries of the medial preoptic nucleus(POM), a sexually dimorphic and steroid-sensitive structure that is a necessary and sufficient site of steroid action in the activation of behavior. Stereotaxic implantation of aromatase inhibitors in but not around the POM strongly decreases the behavioral effects of a systemic treatment with T of castrated males. AA is decreased by castration and increased by aromatizable androgens and by estrogens. These changes have been independently documented at three levels of analysis: the enzymatic activity measured by radioenzymatic assays in vitro, the enzyme concentration evaluated semi-quantitatively by immunocytochemistry and the concentration of its messenger RNA quantified by reverse transcription-polymerase chain reaction (RT-PCR). These studies demonstrate that T acting mostly through its estrogenic metabolites regulates brain aromatase by acting essentially at the transcriptional level. Estrogens produced by central aromatization of T therefore have two independent roles: they activate male copulatory behavior and they regulate the synthesis of aromatase. Double label immunocytochemical studies demonstrate that estrogen receptors(ER) are found in all brain areas containing ARO-ir cells but the extent to which these markers are colocalized varies from one brain region to the other. More than 70% of ARO-ir cells contain detectable ER in the tuberal hypothalamus but less than 20% of the cells display this colocalization in the POA. This absence of ER in ARO-ir cells is also observed in the POA of the rat brain. This suggests that locally formed estrogens cannot control the behavior and the aromatase synthesis in an autocrine fashion in the cells where they were formed. Multi-neuronal networks need therefore to be considered. The behavioral activation could result from the action of estrogens in ER-positive cells located in the vicinity of the ARO-ir cells where they were produced (paracrine action). Alternatively, actions that do not involve the nuclear ER could be important. Immunocytochemical studies at the electron microscope level and biochemical assays of AA in purified synaptosomes indicate the presence of aromatase in presynaptic boutons. Estrogens formed at this level could directly affect the pre-and post-synaptic membrane or could directly modulate neurotransmission namely through their metabolization into catecholestrogens (CE) which are known to be powerful inhibitors of the catechol- omicron - methyl transferase (COMT). The inhibition of COMT should increase the catecholaminergic transmission. It is significant to note, in this respect, that high levels of 2-hydroxylase activity, the enzyme that catalyzes the transformation of estrogens in CE, are found in all brain areas that contain aromatase. On the other hand, the synthesis of aromatase should also be controlled by estrogens in an indirect, transynaptic manner very reminiscent of the way in which steroids indirectly control the production of LHRH. Fibers that are immunoreactive for tyrosine hydroxylase (synthesis of dopamine), dopamine beta-hydroxylase (synthesis of norepinephrine) or vasotocine have been identified in the close vicinity of ARO-ir cells in the POM and retrograde tracing has identified the origin of the dopaminergic and noradrenergic innervation of these areas. A few preliminary physiological experiments suggest that these catecholaminergic inputs regulate AA and presumably synthesis. [less ▲]

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See detailPre- and Post-Translational Regulation of Aromatase by Steroidal and Non-Steroidal Aromatase Inhibitors
Foidart, Agnès ULg; Tlemcani, O.; Harada, N. et al

in Brain Research (1995), 701(1-2), 267-78

Treatment of castrated quail with testosterone (T) reliably activates male copulatory behavior and, at the same time, increases the aromatase activity (AA), the number of aromatase-immunoreactive (ARO-ir ... [more ▼]

Treatment of castrated quail with testosterone (T) reliably activates male copulatory behavior and, at the same time, increases the aromatase activity (AA), the number of aromatase-immunoreactive (ARO-ir) cells and the concentration of aromatase mRNA as measured by RT-PCR in the brain. All these effects can be mimicked by estrogens. The behavioral effects of T can be blocked by a variety of aromatase inhibitors and, in parallel, the AA is strongly inhibited in the preoptic area (POA). We showed recently that the steroidal inhibitor, 4-OH-androstenedione (OHA) markedly decreases the immunostaining density of brain ARO-ir cells while the non-steroidal inhibitor, R76713 (racemic Vorozole; VOR) unexpectedly increased the density of this staining, despite the fact that the enzyme activity was completely inhibited. To generalize these findings and try to identify the underlying mechanism, we compared here the effects of two steroidal (OHA and androstatrienedione [ATD]) and two non-steroidal (VOR and Fadrozole [FAD]) aromatase inhibitors on the aromatase immunostaining and aromatase mRNA concentration in the brain of castrated quail concurrently treated with T. The 4 inhibitors significantly blocked the activation by T of male copulation. The two steroidal inhibitors decreased the immunostaining of brain ARO-ir cells but both VOR and FAD markedly enhanced the density of this staining. In parallel, OHA and ATD completely blocked the T-induced increase in aromatase mRNA concentration, while VOR and FAD had no effect on these RNA concentrations in the POA-anterior hypothalamus and they decreased them only slightly in the posterior hypothalamus. Taken together these results suggest that the inhibition of AA by ATD or OHA and the subsequent removal of locally produced estrogens blocks the synthesis of aromatase presumably at the transcriptional level. By contrast, the two non-steroidal inhibitors tested here block AA but in parallel increase the aromatase immunostaining. This effect does not result from an enhanced transcription and it is therefore speculated that these compounds increase either the translation of the aromatase mRNA or the half-life of the protein itself. [less ▲]

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See detailSexual Differentiation of Brain and Behavior in Quail and Zebra Finches: Studies with a New Aromatase Inhibitor, R76713
Foidart, Agnès ULg; Balthazart, Jacques ULg

in Journal of Steroid Biochemistry & Molecular Biology (1995), 53(1-6), 267-75

In many species of vertebrates, major sex differences affect reproductive behavior and endocrinology. Most of these differences do not result from a direct genomic action but develop following early ... [more ▼]

In many species of vertebrates, major sex differences affect reproductive behavior and endocrinology. Most of these differences do not result from a direct genomic action but develop following early exposure to a sexually differentiated endocrine milieu. In rodents, the female reproductive phenotype mostly develops in the absence of early steroid influence and male differentiation is imposed by the early action of testosterone, acting at least in part through its central conversion into estrogens or aromatization. This pattern of differentiation does not seem to be applicable to avian species. In Japanese quail (Coturnix japonica), injection of estrogens into male embryos causes a permanent loss of the capacity to display male-type copulatory behavior when exposed to testosterone in adulthood. Based on this experimental result, it was proposed that the male reproductive phenotype is "neutral" in birds (i.e. develops in the absence of endocrine influence) and that endogenous estradiol secreted by the ovary of the female embryo is responsible for the physiological demasculinization of females. This model could be recently confirmed. Females indeed display a higher level of circulating estrogens that males during the second part of their embyronic life. In addition, treatment of female embryos with the potent aromatase inhibitor, R76713 or racemic vorozole which suppresses the endogenous secretion of estrogens maintains in females the capacity to display the full range of male copulatory behaviors. The brain mechanisms that control this sexually differentiated behavior have not been identified so far but recent data suggest that they should primarily concern a sub-population of aromatase-immunoreactive neurons located in the lateral parts of the sexually dimorphic preoptic nucleus. The zebra finch (Taeniopygia guttata) exhibits a more complex, still partly unexplained, differentiation pattern. In this species, early treatment with exogenous estrogens produces a masculinization of singing behavior in females and a demasculinization of copulatory behavior in males.(ABSTRACT TRUNCATED AT 400 WORDS) [less ▲]

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See detailAromatase-Immunoreactive Cells Are Present in Mouse Brain Areas That Are Known to Express High Levels of Aromatase Activity
Foidart, Agnès ULg; Harada, N.; Balthazart, Jacques ULg

in Cell & Tissue Research (1995), 280(3), 561-74

The transformation of testosterone into estradiol in the brain plays a key role in several behavioral and physiological processes, but it has been so far impossible to localize precisely the cells of the ... [more ▼]

The transformation of testosterone into estradiol in the brain plays a key role in several behavioral and physiological processes, but it has been so far impossible to localize precisely the cells of the mammalian brain containing the relevant enzyme, viz., aromatase. We have recently established an immunohistochemical technique that allows the visualization of aromatase-immunoreactive cells in the quail brain. In this species, a marked increase in the optical density of aromatase-immunoreactive cells is observed in subjects that have been treated with the aromatase inhibitor, R76713 or racemic Vorozole. This increased immunoreactivity, associated with a total blockade of aromatase activity, has been used as a tool in the present study in which the distribution of aromatase-immunoreactive material has been reassessed in the brain of mice pretreated with R76713. As expected, the aromatase inhibitor increases the density of the immunoreactive signal in mice. Strongly immunoreactive cells are found in the lateral septal region, the bed nucleus of the stria terminalis, the central amygdala, and the dorso-lateral hypothalamus. A less dense signal is also present in the medial preoptic area, the nucleus accumbens, several hypothalamic nuclei (e.g., paraventricular and ventromedial nuclei), all divisions of the amygdala, and several regions of the cortex, especially the cortex piriformis. These data demonstrate that, contrary to previous claims, aromatase-immunoreactive cells are present in all brain regions that have been shown previously to contain high aromatase activity. [less ▲]

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See detailAppetitive as Well as Consummatory Aspects of Male Sexual Behavior in Quail Are Activated by Androgens and Estrogens
Balthazart, Jacques ULg; Reid, J.; Absil, Philippe ULg et al

in Behavioral Neuroscience (1995), 109(3), 485-501

Appetitive male sexual behavior was measured in male quail with the use of a learned social proximity procedure that quantified the time spent by a male in front of a window providing a view of a female ... [more ▼]

Appetitive male sexual behavior was measured in male quail with the use of a learned social proximity procedure that quantified the time spent by a male in front of a window providing a view of a female that was subsequently released into the cage, providing an opportunity for copulation. The learned response is not acquired by castrated males but can be acquired when castrates are treated with testosterone (T) or with the synthetic estrogen diethylstilbestrol or with the endogenous estrogen 17 beta-estradiol. Only birds that become sexually active acquire the response. Conversely, birds in which the consummatory copulatory behavior is disrupted by treatment with the antiestrogen tamoxifen lose the anticipatory response. These results demonstrate that appetitive sexual behavior is, like copulation, activated by T and by estrogens. This suggests that intracerebral aromatization of T also plays a critical role in the activation of this behavior. [less ▲]

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See detailCritical Re-Examination of the Distribution of Aromatase-Immunoreactive Cells in the Quail Forebrain Using Antibodies Raised against Human Placental Aromatase and against the Recombinant Quail, Mouse or Human Enzyme
Foidart, Agnès ULg; Reid, J.; Absil, Philippe ULg et al

in Journal of Chemical Neuroanatomy (1995), 8(4), 267-82

Mouse and quail aromatase cDNAs were isolated from libraries of mouse ovary and quail brain by using a human aromatase cDNA fragment (hA-24) as a probe. These three cDNAs were inserted into plasmid ... [more ▼]

Mouse and quail aromatase cDNAs were isolated from libraries of mouse ovary and quail brain by using a human aromatase cDNA fragment (hA-24) as a probe. These three cDNAs were inserted into plasmid vectors and expressed in Escherichia coli. Antisera against these purified recombinant proteins were raised in rabbit and purified by ammonium sulfate fractionation and affinity chromatography. The three antibodies directed against recombinant human, mouse and quail proteins were used to visualize aromatase-immunoreactive cells in the quail brain. They were compared with the antibody raised against human placental aromatase used in previous experiments and with another antibody recently developed by similar methods. The signal obtained with all antibodies was completely abolished by preadsorption with the homologous recombinant antigens and the signal produced by the two antibodies raised against placental aromatase was similarly abolished by a preadsorption with recombinant quail aromatase. The antibodies raised against recombinant proteins identified the major groups of aromatase cells previously described in the quail brain. The antibodies directed against the mouse and quail antigen identified more positive cells and stained them more densely than the antibodies raised against human recombinant antigen or purified placental aromatase. The new cell groups identified by the antibody raised against quail recombinant aromatase were located in an area ventral to the fasciculus prosencephali lateralis, the nucleus accumbens, the paleostriatum ventrale, the nucleus taeniae, the area around the nucleus ovoidalis, the caudal tuber and the mesencephalic central gray. A critical re-examination of the distribution and nomenclature of the aromatase-positive cells is proposed based on these new findings. [less ▲]

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See detailLes phéromones humaines: vestige animal ou réalité non reconnue
Foidart, Agnès ULg; Legros, Jean-Jacques ULg; Balthazart, Jacques ULg

in Revue Médicale de Liège (1994), 49(12), 662-80

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See detailEffects of Steroidal and Non Steroidal Aromatase Inhibitors on Sexual Behavior and Aromatase-Immunoreactive Cells and Fibers in the Quail Brain
Foidart, Agnès ULg; Harada, N.; Balthazart, Jacques ULg

in Brain Research (1994), 657(1-2), 105-23

Castrated quail were treated with Silastic implants filled with testosterone (T) in association with injections of the aromatase inhibitors, R76713 (racemic vorozole; 1 mg/kg twice a day) or 4 ... [more ▼]

Castrated quail were treated with Silastic implants filled with testosterone (T) in association with injections of the aromatase inhibitors, R76713 (racemic vorozole; 1 mg/kg twice a day) or 4-hydroxyandrostenedione (OHA; 5 mg/bird twice a day). [less ▲]

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See detailSynergistic Control by Androgens and Estrogens of Aromatase in the Quail Brain
Balthazart, Jacques ULg; Stoop, R.; Foidart, Agnès ULg et al

in Neuroreport (1994), 5(14), 1729-32

Castrated quail were injected with testosterone or with the synthetic hormones diethylstilbestrol (DES) or methyltrienolone (R1881) to analyse the steroid specificity in the induction of brain aromatase ... [more ▼]

Castrated quail were injected with testosterone or with the synthetic hormones diethylstilbestrol (DES) or methyltrienolone (R1881) to analyse the steroid specificity in the induction of brain aromatase. R1881 produced a moderate (generally non-significant) increase in the number of aromatase-immunoreactive cells. DES significantly increased the number of positive cells in most brain areas. A clear synergism between DES and R1881 was observed in all brain regions: more immunoreactive cells were found in birds receiving both compounds than in those injected with DES or R1881 alone. DES and R1881 are highly specific ligands for oestrogen and androgen receptors respectively. It appears likely that both androgens and oestrogens directly modulate brain aromatase, presumably at the transcription level. [less ▲]

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See detailAfferent and Efferent Connections of the Sexually Dimorphic Medial Preoptic Nucleus of the Male Quail Revealed by in Vitro Transport of Dii
Balthazart, Jacques ULg; Dupiereux, V.; Aste, N. et al

in Cell & Tissue Research (1994), 276(3), 455-75

The medial preoptic nucleus of the Japanese quail is a testosterone-sensitive structure that is involved in the control of male copulatory behavior. The full understanding of the role played by this ... [more ▼]

The medial preoptic nucleus of the Japanese quail is a testosterone-sensitive structure that is involved in the control of male copulatory behavior. The full understanding of the role played by this nucleus in the control of reproduction requires the identification of its afferent and efferent connections. In order to identify neural circuits involved in the control of the medial preoptic nucleus, we used the lipophilic fluorescent tracer DiI implanted in aldheyde-fixed tissue. Different strategies of brain dissection and different implantation sites were used to establish and confirm afferent and efferent connections of the nucleus. Anterograde projections reached the tuberal hypothalamus, the area ventralis of Tsai, and the substantia grisea centralis. Dense networks of fluorescent fibers were also seen in several hypothalamic nuclei, such as the anterior medialis hypothalami, the paraventricularis magnocellularis, and the ventromedialis hypothalami. A major projection in the dorsal direction was also observed from the medial preoptic nucleus toward the nucleus septalis lateralis and medialis. Afferents to the nucleus were seen from all these regions. Implantation of DiI into the substantia grisea centralis also revealed massive bidirectional connections with a large number of more caudal mesencephalic and pontine structures. The substantia grisea centralis therefore appears to be an important center connecting anterior levels of the brain to brain-stem nuclei that may be involved in the control of male copulatory behavior. [less ▲]

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See detailTestosterone Effects on Neurotensin-Immunoreactive Cells in the Quail Preoptic Area
Absil, Philippe ULg; Balthazart, Jacques ULg

in Neuroreport (1994), 5(9), 1129-32

In the quail preoptic area (POA), neurotensin-immunoreactive (NT-ir) cells are mostly located in the sexually dimorphic medial preoptic nucleus (POM). They are more numerous in females than in males and ... [more ▼]

In the quail preoptic area (POA), neurotensin-immunoreactive (NT-ir) cells are mostly located in the sexually dimorphic medial preoptic nucleus (POM). They are more numerous in females than in males and, in females, their number varies during the ovulatory cycle. This suggests a control by steroids of NT expression in the POA. This idea was tested by comparing the NT-ir populations in gonadectomized males and females treated or not with testosterone. No sex differences in the NT-ir cell number was present in gonadectomized birds. Testosterone treatment increased the number of immunoreactive perikarya. This effect was anatomically specific and the increase was observed in the area caudal to the anterior commissure but not in POM itself. [less ▲]

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See detailSex Difference in the Neurotensin-Immunoreactive Cell Populations of the Preoptic Area in Quail (Coturnix Japonica)
Absil, Philippe ULg; Balthazart, Jacques ULg

in Cell & Tissue Research (1994), 276(1), 99-116

The distribution of neurotensin-immunoreactive cells and fibers was analyzed by immunocytochemistry in the forebrain of male and female Japanese quail (Coturnix japonica) by using an antibody directed ... [more ▼]

The distribution of neurotensin-immunoreactive cells and fibers was analyzed by immunocytochemistry in the forebrain of male and female Japanese quail (Coturnix japonica) by using an antibody directed against the C-terminal part of the molecule. [less ▲]

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