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See detailTopography in the preoptic region: Differential regulation of appetitive and consummatory male sexual behaviors
Balthazart, Jacques ULg; Ball, Gregory F.

in Frontiers in Neuroendocrinology (2007), 28(4), 161-178

Several studies have suggested dissociations between neural circuits underlying the expression of appetitive (e.g., courtship behavior) and consummatory components (i.e., copulatory behavior) of ... [more ▼]

Several studies have suggested dissociations between neural circuits underlying the expression of appetitive (e.g., courtship behavior) and consummatory components (i.e., copulatory behavior) of vertebrate male sexual behavior. The medial preoptic area (mPOA) clearly controls the expression of male copulation but, according to a number of experiments, is not necessarily implicated in the expression of appetitive sexual behavior. In rats for example, lesions to the mPOA eliminate male-typical copulatory behavior but have more subtle or no obvious effects on measures of sexual motivation. Rats with such lesions still pursue and attempt to mount females. They also acquire and perform learned instrumental responses to gain access to females. However, recent lesions studies and measures of the expression of the immediate early gene c-fos demonstrate that, in quail, sub-regions of the mPOA, in particular of its sexually dimorphic component the medial preoptic nucleus, can be specifically linked with either the expression of appetitive or consummatory sexual behavior. In particular more rostral regions can be linked to appetitive components while more caudal regions are involved in consummatory behavior. This functional sub-region variation is associated with neurochemical and hodological specializations (i.e., differences in chemical phenotype of the cells or in their connectivity), especially those related to the actions of androgens in relation to the activation of male sexual behavior, that are also present in rodents and other species. It could thus reflect general principles about POA organization and function in the vertebrate brain. (C) 2007 Elsevier Inc. All rights reserved. [less ▲]

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See detailEstradiol, a key endocrine signal in the sexual differentiation and activation of reproductive behavior in quail
Balthazart, Jacques ULg

in Comparative Biochemistry & Physiology Part A : Molecular & Integrative Physiology (2007, August), 148(Suppl. 1), 27

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See detailSexual Behavior activity tracks rapid changes in brain estrogen concentrations
Taziaux, Mélanie ULg; Keller, Matthieu ULg; Bakker, Julie ULg et al

in Journal of Neuroscience (2007), 27(24), 6563-6572

Estrogens are classically viewed as hormones that bind to intracellular receptors, which then act as transcription factors to modulate gene expression; however, they also affect many aspects of neuronal ... [more ▼]

Estrogens are classically viewed as hormones that bind to intracellular receptors, which then act as transcription factors to modulate gene expression; however, they also affect many aspects of neuronal functioning by rapid nongenomic actions. Brain estrogen production can be regulated within minutes by changes in aromatase (estrogen synthase) activity as a result of calcium-dependent phosphorylations of the enzyme. To determine the effects of rapid changes in estrogen availability on male copulatory behavior, we mimicked in male mice the rapid upregulation and downregulation of brain estrogen concentration that should occur after inactivation or activation of aromatase activity. A single injection of different aromatase inhibitors [Vorozole, 1,4,6-androstatrien-3,17-dione (ATD), or its metabolite 17-OH-ATD (1,4,6-androstatrien-17beta-ol-3-one)] almost completely suppressed male sexual behavior (mounts and intromissions) expressed 10-20 min later by C57BL/6J mice but did not affect behavior in aromatase knock-out (ArKO) mice, activated by daily injections of estradiol benzoate, thereby confirming the specificity of the behavioral inhibition observed in wild-type mice. The rapid ATD-induced inhibition was reversed by the simultaneous injection of a large dose of estradiol. A single injection of estradiol to ArKO mice also activated male sexual behavior within 15 min. Thus, rapid increases or decreases in brain estrogen concentrations are followed within minutes by corresponding changes in male sexual behavior. Sexual behavior can thus be used to monitor changes in local estrogen concentrations and analyze the mechanisms mediating the rapid decline in estrogen signaling that takes place after inhibition of estrogen synthesis. [less ▲]

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See detailExposure to oestrogen prenatally does not interfere with the normal female-typical development of odour preferences
Bakker, Julie ULg; De Mees, C.; Szpirer, J. et al

in Journal of Neuroendocrinology (2007), 19(5), 329-334

The neural mechanisms controlling mate recognition and heterosexual partner preference are sexually differentiated by perinatal actions of sex steroid hormones. We previously showed that the most ... [more ▼]

The neural mechanisms controlling mate recognition and heterosexual partner preference are sexually differentiated by perinatal actions of sex steroid hormones. We previously showed that the most important action of oestrogen during prenatal development is to defeminise and, to some extent, masculinise brain and behaviour in mice. Female mice deficient in alpha-foetoprotein (AFP) due to a targeted mutation in the Afp gene (AFP-KO) do not show any female sexual behaviour when paired with an active male because they lack the protective action of AFP against maternal oestrogens. In the present study, we investigated whether odour preferences, another sexually differentiated trait in mice, are also defeminised and/or masculinised in AFP-KO females due to their prenatal exposure to oestrogens. AFP-KO females of two background strains (CD1 and C57Bl/6j) preferred to investigate male over female odours when given the choice between these two odour stimuli in a Y-maze, and thus remained very female-like in this regard. Thus, the absence of lordosis behaviour in these females cannot be explained by a reduced motivation of AFP-KO females to investigate male-derived odours. Furthermore, the presence of a strong male-directed odour preference in AFP-KO females suggests a postnatal contribution of oestrogens to the development of preferences to investigate opposite-sex odours. [less ▲]

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See detailThe microtubule-associated protein doublecortin is broadly expressed in the telencephalon of adult canaries
Boseret, Géraldine ULg; Ball, G. F.; Balthazart, Jacques ULg

in Journal of Chemical Neuroanatomy (2007), 33(3), 140-154

The protein doublecortin (DCX) is expressed in post-mitotic migrating and differentiating neurons in the developing vertebrate brain and, as a part of the microtubule machinery, is required for neuronal ... [more ▼]

The protein doublecortin (DCX) is expressed in post-mitotic migrating and differentiating neurons in the developing vertebrate brain and, as a part of the microtubule machinery, is required for neuronal migration. DCX expression is generally maximal during embryonic and early post-natal life but decreases markedly and almost disappears in older animals in parallel with the major decrease or cessation of neurogenesis. In several seasonally breeding songbird species such as canaries, the volume of several song control nuclei in the brain varies seasonally such that the largest nuclei are observed in the late spring and early summer. This variation is based on changes in cell size, dendritic branching, and, in nucleus HVC, on the incorporation of neurons newly born in adulthood. Because songbirds maintain an active neurogenesis and neuronal incorporation in their telencephalon throughout their lives, we investigated here the distribution of DCX-immunoreactive (ir) structures in the brain of adult male canaries. Densely stained DCX-ir cells were found exclusively in parts of the telencephalon that are known to incorporate new neurons in adulthood, in particular the nidopallium. Within this brain region, the boundaries of the song control nucleus HVC could be clearly distinguished from surrounding structures by a higher density of DCX-ir structures. In most telencephalic areas, about two thirds of these cells displayed a uni- or bipolar fusiform morphology suggesting they were migrating neurons. The rest of the DCX-ir cells in the telencephalon were larger and had a round multipolar morphology. No such staining was found in the rest of the brain. The broad expression of DCX specifically in adult brain structures that exhibit the characteristic of active incorporation of new neurons suggests that DCX plays a key role in the migration of new neurons in the brain of adult songbirds as it presumably does during ontogeny. (c) 2007 Elsevier B.V. All rights reserved. [less ▲]

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See detailNon-photoperiodic factors and timing of breeding in blue tits: Impact of environmental and social influences in semi-natural conditions
Caro, S. P.; Lambrechts, M. M.; Balthazart, Jacques ULg et al

in Behavioural Processes (2007), 75(1), 1-7

The seasonal development of life-history traits is influenced by many environmental factors. The impact of photoperiodic and non-photoperiodic factors on nest building and egg laying has been rarely ... [more ▼]

The seasonal development of life-history traits is influenced by many environmental factors. The impact of photoperiodic and non-photoperiodic factors on nest building and egg laying has been rarely investigated in non-domesticated avian species for which long term field data sets are available. Former investigations showed that blue tits originating from geographically close populations in the Mediterranean region do not respond in the same way to photoperiodic factors in semi-natural outdoor conditions. Here we show experimentally that nest building and onset of egg laying in captive blue tits is also proximately influenced by non-photoperiodic factors, including aspects related to aviary characteristics and social interactions between birds of the two sexes originating from different local Mediterranean study populations. In two successive experiments, we show that (1) increasing the volume of the aviary advanced the egg laying period of one specific population by almost I month, and (2) crossing pairs of birds from different origins strongly reduced the nest building and egg laying behaviours. These results indicate that obtaining biologically relevant breeding results in captivity with wild birds requires the control and experimental manipulation of a wide array of complex environmental cues. (c) 2007 Elsevier B.V All rights reserved. [less ▲]

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See detailDifferential c-fos expression in the brain of male Japanese quail following exposure to stimuli that predict or do not predict the arrival of a female
Taziaux, Mélanie ULg; Lopez, J.; Cornil, Charlotte ULg et al

in European Journal of Neuroscience (2007), 25(9), 2835-2846

We investigated the effects of presenting a sexual conditioned stimulus on the expression of c-fos in male Japanese quail. Eight brain sites were selected for analysis based on previous reports of c-fos ... [more ▼]

We investigated the effects of presenting a sexual conditioned stimulus on the expression of c-fos in male Japanese quail. Eight brain sites were selected for analysis based on previous reports of c-fos expression in these areas correlated with sexual behaviour or learning. Males received either paired or explicitly unpaired presentations of an arbitrary stimulus and visual access to a female. Nine conditioning trials were conducted, one per day, for each subject. On the day following the ninth trial, subjects were exposed to the conditional stimulus (CS) for 5 min. Conditioning was confirmed by analysis of rhythmic cloacal sphincter movements (RCSM), an appetitive sexual behaviour, made in response to the CS presentation. Subjects in the paired condition performed significantly more RCSM than subjects in the unpaired group. Brains were collected 90 min following the stimulus exposure and stained by immunolhistochemistry for the FOS protein. Significant group differences in the number of FOS-immunoreactive (FOS-ir) cells were found in two brain regions, the nucleus taeniae of the amygdala (TnA) and the hippocampus (Hp). Subjects in the paired condition had fewer FOS-ir cells in both areas than subjects in the unpaired condition. These data provide additional support to the hypothesis that TnA is implicated in the expression of appetitive sexual behaviours in male quail and corroborate numerous previous reports of the involvement of the hippocampus in conditioning. Further, these data suggest that conditioned and unconditioned sexual stimuli activate different brain regions but have similar behavioural consequences. [less ▲]

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See detailSexual versus individual differentiation: the controversial role of avian maternal hormones
Carere, C.; Balthazart, Jacques ULg

in Trends in Endocrinology and Metabolism (2007), 18(2), 73-80

Avian embryos are exposed not only to endogenous sex steroids, which are produced by their gonads and have a key role in sexual differentiation, but also to maternal steroids transferred into the egg yolk ... [more ▼]

Avian embryos are exposed not only to endogenous sex steroids, which are produced by their gonads and have a key role in sexual differentiation, but also to maternal steroids transferred into the egg yolk, which can modulate the development of individual differences in behavior. Studies of maternal hormones have primarily focused on ultimate questions (evolutionary trade-offs, functional significance), whereas proximate mechanistic questions have been largely ignored. A central problem that must be addressed is how exposure to maternal hormones affects the individual phenotype without interfering with sexual differentiation. Separate effects could result from the action of different hormones, at different doses or at different times, on different targets. [less ▲]

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See detailNeuroanatomical specificity of sex differences in expression of aromatase mRNA in the quail brain
Voigt, C.; Ball, G. F.; Balthazart, Jacques ULg

in Journal of Chemical Neuroanatomy (2007), 33(2), 75-86

In birds and mammals, aromatase activity in the preoptic-hypothalamic region (HPOA) is usually higher in males than in females. It is, however, not known whether the enzymatic sex difference reflects the ... [more ▼]

In birds and mammals, aromatase activity in the preoptic-hypothalamic region (HPOA) is usually higher in males than in females. It is, however, not known whether the enzymatic sex difference reflects the differential activation of aromatase transcription or some other control mechanism. Although sex differences in aromatase activity are clearly documented in the HPOA of Japanese quail (Coturnix japonica), only minimal or even no differences at all were observed in the number of aromatase-immunoreactive (ARO-ir) cells in the medial preoptic nucleus (POM) and in the medial part of the bed nucleus striae terminalis (BSTM). We investigated by in situ hybridization the distribution and possible sex differences in aromatase mRNA expression in the brain of sexually active adult quail. The distribution of aromatase mRNA matched very closely the results of previous immunocytochemical studies with the densest signal being observed in the POM, BSTM and in the mediobasal hypothalamus (MBH). Additional weaker signals were detected in the rostral forebrain, arcopallium and mesencephalic regions. No sex difference in the optical density of the hybridization signal could be found in the POM and MBH but the area covered by mRNA was larger in males than in females, indicating a higher overall expression in males. In contrast, in the BSTM, similar areas were covered by the aromatase expression in both sexes but the density of the signal was higher in females than in males. The physiological control of aromatase is thus neuroanatomically specific and with regard to sex differences, these controls are at least partially different if one compares the level of transcription, translation and activity of the enzyme. These results also indirectly suggest that the sex difference in aromatase enzyme activity that is present in the quail HPOA largely results from differentiated controls of enzymatic activity rather than differences in enzyme concentration. (c) 2007 Elsevier B.V. All rights reserved. [less ▲]

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See detailSex differences in projections from preoptic area aromatase cells to the periaqueductal gray in Japanese quail
Carere, C.; Ball, G. F.; Balthazart, Jacques ULg

in Journal of Comparative Neurology (2007), 500(5), 894-907

In many vertebrate species the medial preoptic area projects to a premotor nucleus, the periaqueductal central gray (PAG). This connection plays an important role in the control of reproductive behavior ... [more ▼]

In many vertebrate species the medial preoptic area projects to a premotor nucleus, the periaqueductal central gray (PAG). This connection plays an important role in the control of reproductive behavior. In male Japanese quail (Coturnix japonica) specifically, the medial preoptic nucleus (POM), where various types of sensory inputs converge, is a critical site for the activational action of testosterone on male sexual behavior. To activate male copulatory behavior, testosterone must be aromatized to estradiol within the POM and aromatase-immunoreactive cells in the POM are the main source of projections to the PAG. The POM-PAG connection is thus an important functional circuit integrating the sensory with premotor components of sexual behavior. Contrary to what is observed in males, testosterone does not activate male-typical copulatory behavior in females and we investigated here via retrograde tracing methods whether this behavioral sexual difference is associated with a sex difference in connectivity between POM and PAG. Fluorescent microspheres were injected in the PAG of male and female quail and retrogradely labeled fluorescent cells counted in four fields of the POM in sections that had been immunolabeled for aromatase. Males had more aromatase-immunoreactive neurons projecting to the PAG than females and this difference was most prominent in the caudolateral part of the nucleus that has been specifically implicated in the control of male copulatory behavior. These data therefore support the hypothesis that sex differences in POM-PAG connectivity are causally linked to the sex difference in the behavioral response to testosterone. [less ▲]

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See detailFunctional significance of the rapid regulation of brain estrogen action: Where do the estrogens come from?
Cornil, Charlotte ULg; Ball, G. F.; Balthazart, Jacques ULg

in Brain Research (2006), 1126

Estrogens exert a wide variety of actions on reproductive and non-reproductive functions. These effects are mediated by slow and long lasting genomic as well as rapid and transient non-genomic mechanisms ... [more ▼]

Estrogens exert a wide variety of actions on reproductive and non-reproductive functions. These effects are mediated by slow and long lasting genomic as well as rapid and transient non-genomic mechanisms. Besides the host of studies demonstrating the role of genomic actions at the physiological and behavioral level, mounting evidence highlights the functional significance of non-genomic effects. However, the source of the rapid changes in estrogen availability that are necessary to sustain their fast actions is rarely questioned. For example, the rise of plasma estrogens at pro-estrus that represents one of the fastest documented changes in plasma estrogen concentration appears too slow to explain these actions. Alternatively, estrogen can be synthesized in the brain by the enzyme aromatase providing a source of locally high concentrations of the steroid. Furthermore, recent studies demonstrate that brain aromatase can be rapidly modulated by afferent inputs, including glutamatergic afferents. A role for rapid changes in estrogen production in the central nervous system is supported by experiments showing that acute aromatase inhibition affects nociception as well as male sexual behavior and that preoptic aromatase activity is rapidly (within min) modulated following mating. Such mechanisms thus fulfill the gap existing between the fast actions of estrogen and their mode of production and open new avenues for the understanding of estrogenic effects on the brain. (c) 2006 Elsevier B.V. All rights reserved. [less ▲]

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See detailSocial context affects testosterone-induced singing and the volume of song control nuclei in male canaries (Serinus canaria)
Boseret, Géraldine ULg; Carere, C.; Ball, G. F. et al

in Journal of Neurobiology (2006), 66(10), 1044-1060

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See detailSimultaneous pituitary-gonadal recrudescence in two Corsican populations of male blue tits with asynchronous breeding dates
Caro, S. P.; Lambrechts, M. M.; Chastel, O. et al

in Hormones & Behavior (2006), 50(3), 347-360

Animal populations living in geographically variable environments respond to different selection pressures. The adaptive character of the responses to environmental information determines the degree of ... [more ▼]

Animal populations living in geographically variable environments respond to different selection pressures. The adaptive character of the responses to environmental information determines the degree of synchrony of the breeding period with local optimal conditions. An example is provided by two populations of Mediterranean blue tits (Parus caeruleus) in Corsica, breeding in different habitats, with a 1-month difference in the onset of egg laying. This difference in the onset of lay is supposed to be adaptive because, although chicks from both populations are raised mostly on caterpillars, the timing of the appearance of caterpillars is earlier for populations of tits associated with deciduous oak trees than those associated with evergreen oak trees. Here, we show that, despite the difference in the timing of egg laying, males from these two populations start seasonal hypothalamo-hypophysial-testicular development at approximately the same time, in late winter. Specifically, the vernal recrudescence of brain GnRH-I perikarya and fibers, testes volume and song activity began around the same dates and proceeded at the same pace in late winter in both populations. Plasma testosterone and LH levels displayed seasonal variations that were shifted by less than 2 weeks compared to the 1-month difference in egg laying periods. We hypothesize that the strong selection pressures on these two populations to adapt the timing of their breeding seasons to their local environment may have acted mostly on the female egg laying dates, and not so much on the initiation and rate of seasonal recrudescence of the hypothalamo-hypophysial-testicular activity in males. (c) 2006 Elsevier Inc. All rights reserved. [less ▲]

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See detailThe proximate basis of adaptive micro-geographic variation in reproductive phenology in male and female Blue Tits
Caro, S.; Balthazart, Jacques ULg; Lambrechts, M.

in Journal of Ornithology (2006, August), 147(5, Suppl. 1), 47-48

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See detailRapid effects of estrogens on sexual appetite and consummation in quail
Balthazart, Jacques ULg

in Journal of Ornithology (2006, August), 147(5, Suppl. 1), 56

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See detailIn vivo MR imaging of the seasonal volumetric and functional plasticity of song control nuclei in relation to song output in a female songbird
Van Meir, V.; Pavlova, D.; Verhoye, M. et al

in Neuroimage (2006), 31(3), 981-992

In temperate zone songbird species, seasonal plasticity in the morphological and functional state of brain regions involved in song production occurs in association with seasonal changes in song output ... [more ▼]

In temperate zone songbird species, seasonal plasticity in the morphological and functional state of brain regions involved in song production occurs in association with seasonal changes in song output. Following MHCl2-injections in HVC (used as proper name) of female starlings, in vivo tract-tracing by Manganese Enhanced-Magnetic Resonance Imaging (ME-MRI) provided repeated measures of the volume of two HVC targets, the nucleus robustus arcopallii (RA) and area X, along with measures of the activity of the caudal motor pathway and rostral basal-ganglia pathway that control singing. Mn2+- labeling (volume labeled and signal intensity) of both nuclei was dramatically reduced in July (post-breeding season) when birds did not sing, compared to March (breeding season) when birds produced song. Seasonal changes in telencephalon volume did not exceed 4% and were not significant but were surprisingly correlated with individual measures of song rate and song bout length. Although individual song rates were variable in March, all MnCl2-injections led to a reliable labeling of area X and RA. In July, delineation of area X was only possible in two birds and RA could be delineated in 50% of the population; its volume had decreased by 46% as compared to March. The birds in which RA could be delineated in July had in March a higher activity of the HVC to area X projection as reflected by the total amount of Mn2+ accumulated in area X, which suggests unexpected relationships between the two types of HVC projection neurons. (c) 2006 Elsevier Inc. All rights reserved. [less ▲]

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See detailIs brain estradiol a hormone or a neurotransmitter?
Balthazart, Jacques ULg; Ball, G. F.

in Trends in Neurosciences (2006), 29(5), 241-249

Mounting evidence indicates that, besides their well-known hormonal mode of action at the genetic level, estrogens such as 17 beta-estradiol also influence brain function by direct effects on neuronal ... [more ▼]

Mounting evidence indicates that, besides their well-known hormonal mode of action at the genetic level, estrogens such as 17 beta-estradiol also influence brain function by direct effects on neuronal membranes. Experimentally induced rapid changes in estradiol bioavailability in the brain have been shown to alter the expression of male sexual behavior significantly within minutes - probably too quickly to be accounted for by conventional genetic mechanisms. In parallel, recent studies indicate that aromatase, the enzyme that converts testosterone to estradiol in the brain, is expressed in presynaptic terminals and modulated within minutes by Ca2+-dependent phosphorylation. In this article, we develop the hypothesis that brain estrogens display many, if not all, functional characteristics of neuromodulators or even neurotransmitters. [less ▲]

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See detailNeuroanatomical specificity in the expression of the immediate early gene c-fos following expression of appetitive and consummatory male sexual behaviour in Japanese quail
Taziaux, Mélanie ULg; Cornil, Charlotte ULg; Dejace, C. et al

in European Journal of Neuroscience (2006), 23(7), 1869-1887

We investigated the neural sites related to the occurrence of appetitive (ASB) and consummatory (CSB) aspects of male sexual behaviour in Japanese quail. Castrated males treated with testosterone were ... [more ▼]

We investigated the neural sites related to the occurrence of appetitive (ASB) and consummatory (CSB) aspects of male sexual behaviour in Japanese quail. Castrated males treated with testosterone were exposed for 5 min to one of four experimental conditions: (i) free interaction with a female (CSB group); (ii) expression of rhythmic cloacal sphincter movements in response to the visual presentation of a female (ASB-F group); (iii) or a male (ASB-M group), and (iv) handling as a control manipulation. Brains were collected 90 min after the start of behavioural tests and stained by immunocytochemistry for the FOS protein. An increase in FOS expression was observed throughout the rostro-caudal extent of the medial preoptic nucleus (POM) in CSB males, whereas the view of a female (ASB-F) induced an increased FOS expression in the rostral POM only. In the CSB group, there was also an increase in FOS expression in the bed nucleus striae terminalis, and both the CSB and ASB-F groups exhibited increased FOS expression in aspects of the ventro-lateral thalamus (VLT) related to visual processing. Moreover, both the CSB and ASB-M groups showed increased FOS expression in the lateral septum. These data provide additional support to the idea that there is a partial anatomical dissociation between structures involved in the control of both aspects of male sexual behaviour and independently provide data consistent with a previous lesion study that indicated that the rostral and caudal POM differentially control the expression of ASB and CSB in quail. [less ▲]

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See detailAlpha-fetoprotein protects the developing female mouse brain from masculinization and defeminization by estrogens
Bakker, Julie ULg; De Mees, C.; Douhard, Quentin ULg et al

in Nature Neuroscience (2006), 9(2), 220-226

Two clearly opposing views exist on the function of alpha-fetoprotein (AFP), a fetal plasma protein that binds estrogens with high affinity, in the sexual differentiation of the rodent brain. AFP has been ... [more ▼]

Two clearly opposing views exist on the function of alpha-fetoprotein (AFP), a fetal plasma protein that binds estrogens with high affinity, in the sexual differentiation of the rodent brain. AFP has been proposed to either prevent the entry of estrogens or to actively transport estrogens into the developing female brain. The availability of Afp mutant mice (Afp(-/-)) now finally allows us to resolve this longstanding controversy concerning the role of AFP in brain sexual differentiation, and thus to determine whether prenatal estrogens contribute to the development of the female brain. Here we show that the brain and behavior of female Afp(-/-) mice were masculinized and defeminized. However, when estrogen production was blocked by embryonic treatment with the aromatase inhibitor 1,4,6-androstatriene-3,17-dione, the feminine phenotype of these mice was rescued. These results clearly demonstrate that prenatal estrogens masculinize and defeminize the brain and that AFP protects the female brain from these effects of estrogens. [less ▲]

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