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See detailEstradiol rapidly activates male sexual behavior and affects brain monoamine levels in the quail brain
Cornil, Charlotte ULg; Dalla, C.; Papadopoulou-Daifoti, Z. et al

in Behavioural Brain Research (2006), 166(1), 110-123

Steroids are generally viewed as transcription factors binding to intracellular receptors and activating gene transcription. Rapid cellular effects mediated via non-genomic mechanisms have however been ... [more ▼]

Steroids are generally viewed as transcription factors binding to intracellular receptors and activating gene transcription. Rapid cellular effects mediated via non-genomic mechanisms have however been identified and one report showed that injections of estradiol rapidly stimulate chemoinvestigation and mounting behavior in castrated male rats. It is not known whether such effects take place in other species and what are the cellular underlying mechanisms. We show here that a single injection of estradiol (500 wg/kg) rapidly and transiently activates copulatory behavior in castrated male quail pre-treated with a dose of testosterone behaviorally ineffective by itself. The maximal behavioral effect was observed after 15 min. In a second experiment, the brain of all subjects was immediately collected after behavioral tests performed 15 min after injection. The preoptic area-hypothalamus (HPOA), hindbrain, telencephalon and cerebellum were isolated and monoamines measured by HPLC-ED. Estradiol increased levels of the serotonin metabolite 5-hydroxyindoleacetic acid (5-HIAA) and 5-HIAA/serotonin ratios in the telencephalon and hindbrain independently of whether animals had mated or not. Estradiol also affected these measures in HPOA and cerebellum but this effect was correlated with the level of sexual activity so that significant effects of the treatment only appeared when sexual activity was used as a covariate. Interactions between estradiol effects and sexual activity were also observed for dopamine in the HPOA and for serotonin in the hindbrain and cerebellum. Together, these data demonstrate that a single estradiol injection rapidly activates male sexual behavior in quail and that this behavioral effect is correlated with changes in monoaminergic activity. (c) 2005 Elsevier B.V. All rights reserved. [less ▲]

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See detailRapid effects of aromatase inhibition on male reproductive behaviors in Japanese quail
Cornil, Charlotte ULg; Taziaux, Mélanie ULg; Baillien, M. et al

in Hormones & Behavior (2006), 49(1), 45-67

Non-genomic effects of steroid hormones on cell physiology have been reported in the brain. However, relatively little is known about the behavioral significance of these actions. Male sexual behavior is ... [more ▼]

Non-genomic effects of steroid hormones on cell physiology have been reported in the brain. However, relatively little is known about the behavioral significance of these actions. Male sexual behavior is activated by testosterone partly through its conversion to estradiol via the enzyme aromatase in the preoptic area (POA). Brain aromatase activity (AA) changes rapidly which might in turn be important for the rapid regulation of behavior. Here, acute effects of Vorozole (TM), an aromatase inhibitor, injected IP at different doses and times before testing (between 15 and 60 min), were assessed on male sexual behavior in quail. To limit the risk of committing both types of statistical errors (I and II), data of all experiments were entered into a meta-analysis. Vorozole (TM) significantly inhibited mount attempts (P < 0.05, size effect [g] = 0.527) and increased the latency to first copulation (P < 0.05, g = 0.251). The treatment had no effect on the other measures of copulatory behavior. Vorozole (TM) also inhibited appetitive sexual behavior measured by the social proximity response (P < 0.05, g = 0.534) or rhythmic cloacal sphincter movements (P < 0.001, g = 0.408). Behavioral inhibitions always reached a maximum at 30 min. Another aromatase inhibitor, androstatrienedione, induced a similar rapid inhibition of sphincter movements. Radioenzyme assays demonstrated that within 30 min Vorozole (TM) had reached the POA and completely blocked AA measured in homogenates. When added to the extracellular milieu, Vorozole (TM) also blocked within 5 min the AA in POA explants maintained in vitro. Together, these data demonstrate that aromatase inhibition rapidly decreases both consummatory and appetitive aspects of male sexual behavior. (c) 2005 Elsevier Inc. All rights reserved. [less ▲]

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See detailRapid changes in production and behavioral action of estrogens.
Balthazart, Jacques ULg; Cornil, Charlotte ULg; Taziaux, Mélanie ULg et al

in Neuroscience (2006), 138(3), 783-91

It is well established that sex steroid hormones bind to nuclear receptors, which then act as transcription factors to control brain sexual differentiation and the activation of sexual behaviors ... [more ▼]

It is well established that sex steroid hormones bind to nuclear receptors, which then act as transcription factors to control brain sexual differentiation and the activation of sexual behaviors. Estrogens locally produced in the brain exert their behavioral effects in this way but mounting evidence indicates that estrogens also can influence brain functioning more rapidly via non-genomic mechanisms. We recently reported that, in Japanese quail, the activity of preoptic estrogen synthase (aromatase) can be modulated quite rapidly (within minutes) by non-genomic mechanisms, including calcium-dependent phosphorylations. Behavioral studies further demonstrated that rapid changes in estrogen bioavailability, resulting either from a single injection of a high dose of estradiol or from the acute inhibition of aromatase activity, significantly affect the expression of both appetitive and consummatory aspects of male sexual behavior with latencies ranging between 15 and 30 min. Together these data indicate that the bioavailability of estrogens in the brain can change on different time-scales (long- and short-term) that match well with the genomic and non-genomic actions of this steroid and underlie two complementary mechanisms through which estrogens modulate behavior. Estrogens produced locally in the brain should therefore be considered not only as neuroactive steroids but they also display many (if not all) functional characteristics of neuromodulators and perhaps neurotransmitters. [less ▲]

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See detailRapid decreases in preoptic aromatase activity and brain monoamine concentrations after engaging in male sexual behavior
Cornil, Charlotte ULg; Dalla, C.; Papadopoulou-Daifoti, Z. et al

in Endocrinology (2005), 146(9), 3809-3820

In Japanese quail, as in rats, the expression of male sexual behavior over relatively long time periods (days to weeks) is dependent on the local production of estradiol in the preoptic area via the ... [more ▼]

In Japanese quail, as in rats, the expression of male sexual behavior over relatively long time periods (days to weeks) is dependent on the local production of estradiol in the preoptic area via the aromatization of testosterone. On a short-term basis (minutes to hours), central actions of dopamine as well as locally produced estrogens modulate behavioral expression. In rats, a view of and sexual interaction with a female increase dopamine release in the preoptic area. In quail, in vitro brain aromatase activity (AA) is rapidly modulated by calcium-dependent phosphorylations that are likely to occur in vivo as a result of changes in neurotransmitter activity. Furthermore, an acute estradiol injection rapidly stimulates copulation in quail, whereas a single injection of the aromatase inhibitor vorozole rapidly inhibits this behavior. We hypothesized that brain aromatase and dopaminergic activities are regulated in quail in association with the expression of male sexual behavior. Visual access as well as sexual interactions with a female produced a significant decrease in brain AA, which was maximal after 5 min. This expression of sexual behavior also resulted in a significant decrease in dopaminergic as well as serotonergic activity after 1 min, which returned to basal levels after 5 min. These results demonstrate for the first time that AA is rapidly modulated in vivo in parallel with changes in dopamine activity. Sexual interactions with the female decreased aromatase and dopamine activities. These data challenge established views about the causal relationships among dopamine, estrogen action, and male sexual behavior. [less ▲]

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See detailDopamine modulates male sexual behavior in Japanese quail in part via actions on noradrenergic receptors
Cornil, Charlotte ULg; Dejace, C.; Ball, G. F. et al

in Behavioural Brain Research (2005), 163(1), 42-57

In rats, dopamine (DA) facilitates male sexual behavior through its combined action on D1- and D2-like receptors, in the medial preoptic area (MPOA) as well as other brain areas. In Japanese quail ... [more ▼]

In rats, dopamine (DA) facilitates male sexual behavior through its combined action on D1- and D2-like receptors, in the medial preoptic area (MPOA) as well as other brain areas. In Japanese quail, systemic injections of dopaminergic drugs suggested a similar pharmacology but central injections have never been performed. Recent electrophysiological experiments demonstrated that DA effects in the MPOA of quail are mediated mainly through the activation of alpha(2)-noradrenergic receptors. Previous studies of DA action on behavior used specific dopaminergic agonists/antagonists and therefore unintentionally avoided the potential cross-reaction with a-receptors. The present study was thus designed to investigate directly the effects of DA on male sexual behavior and to test whether the interaction of DA with heterologous receptors affects this behavior. Intracerebroventricular (i.c.v.) injection of DA or NE inhibited copulation in a dose-dependent manner. Systemic injections of yohimbine, an alpha(2)-noradrenergic antagonist, modulated copulation in a bimodal manner depending on the dose injected. Interestingly, a behaviorally ineffective dose of yohimbine markedly reduced the inhibitory effects of DA when injected 15 min before. Together, these results show for the first time that i.c.v. injections of DA itself inhibit male sexual behavior in quail and suggest that the interaction of DA with alpha(2)-receptors has behavioral significance. (C) 2005 Elsevier B.V. All rights reserved. [less ▲]

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See detailDifferences in neural activation following expression of appetitive and consummatory male sexual behavior in the quail brain
Taziaux, Mélanie ULg; Cornil, Charlotte ULg; Dejace, C. et al

in Hormones & Behavior (2005, June), 48(1), 130

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See detailAndrogen mediation of conditioned rhythmic cloacal sphincter movements in Japanese quail (Coturnix Japonica)
Holloway, Kevin S.; Balthazart, Jacques ULg; Cornil, Charlotte ULg

in Journal of Comparative Psychology (2005), 119(1), 49-57

Demonstrations of increased reproductive success due to sexual conditioning in Japanese quail (Coturnix japonica) have been reported, although the mechanisms that underlie these effects have remained ... [more ▼]

Demonstrations of increased reproductive success due to sexual conditioning in Japanese quail (Coturnix japonica) have been reported, although the mechanisms that underlie these effects have remained elusive. One possible mechanism is conditioned rhythmic cloacal sphincter movements (RCSM). Two experiments were conducted with male quail to determine whether associations between a conditioned stimulus (CS) and a hen would result in the ability of the CS to elicit RCSM, and to explore the androgen mediation of conditioned RCSM. The results suggest that a focal CS paired with visual access to a female will elicit RCSM via a representation of the hen activated by the CS. Further, the available evidence indicates that conditioned RCSM is androgen mediated and that this learning may transfer across breeding seasons. [less ▲]

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See detailRapid changes in production and behavioral action of estrogens
Balthazart, Jacques ULg; Baillien, Michelle; Charlier, Thierry ULg et al

in Trabajos del Instituto Cajal (2005), 80

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See detailAromatase inhibition blocks the expression of sexually-motivated cloacal gland movements in male quail
Taziaux, Mélanie ULg; Cornil, Charlotte ULg; Balthazart, Jacques ULg

in Behavioural Processes (2004), 67(3), 461-469

In Japanese quail (Coturnix japonica), activation of appetitive and consummatory aspects of male sexual behavior requires aromatization of testosterone (T) into estrogens. Appetitive male sexual behavior ... [more ▼]

In Japanese quail (Coturnix japonica), activation of appetitive and consummatory aspects of male sexual behavior requires aromatization of testosterone (T) into estrogens. Appetitive male sexual behavior (ASB) is usually assessed with the use of a learned social proximity procedure. In the present experiment, we investigated the role of estrogens in the activation of an another index of ASB. the female-induced activation of rhythmic cloacal sphincter movements (RCSMs) that are produced in reaction to the visual presentation of a female. Consummatory sexual behavior (CSB) was also assessed by the frequency and latency of copulatory behaviors. Castrated male quail were treated with Silastic implants filled with T in association with chronic injections of the aromatase inhibitor Vorozole(TM) (R83842; 1 mg/kg twice a day; CX + T + VOR group). Control birds were implanted with T capsules only (CX + T group). CSB was almost completely blocked by injections of the aromatase inhibitor. The RCSM frequency decreased progressively in the CX + T + VOR group by comparison with the CX + T group and was therefore significantly reduced at the end of the experiment. These results demonstrate that the frequency of RCSM, a second measure of ASB is, like the social proximity response and CSB, blocked by inhibition of estrogen production. It was shown previously that lesions of the preoptic area inhibit both aspects of the appetitive sexual behavior (proximity response and RCSM). It is therefore, likely that both responses are controlled, like copulation, by aromatase-containing neurons of the preoptic area. (C) 2004 Elsevier B.V. All rights reserved. [less ▲]

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See detailPreoptic aromatase modulates male sexual behavior: slow and fast mechanisms of action
Balthazart, Jacques ULg; Baillien, M.; Cornil, Charlotte ULg et al

in Physiology & Behavior (2004), 83(2), 247-270

In many species, copulatory behavior and appetitive (anticipatory/motivational) aspects of male sexual behavior are activated by the action in the preoptic area of estrogens locally produced by ... [more ▼]

In many species, copulatory behavior and appetitive (anticipatory/motivational) aspects of male sexual behavior are activated by the action in the preoptic area of estrogens locally produced by testosterone aromatization. Estrogens bind to intracellular receptors, which then act as transcription factors to activate the behavior. Accordingly, changes in aromatase activity (AA) result from slow steroid-induced modifications of enzyme transcription. More recently, rapid nongenomic effects of estrogens have been described and evidence has accumulated indicating that AA can be modulated by rapid (minutes to hour) nongenomic mechanisms in addition to the slower transcriptional changes. Hypothalamic AA is rapidly down-regulated in conditions that enhance protein phosphorylation, in particular, increases in the intracellular calcium concentration, such as those triggered by neurotransmitter (e.g., glutamate) activity. Fast changes in brain estrogens can thus be caused by aromatase phosphorylation as a result of changes in neurotransmission. In parallel, recent studies demonstrate that the pharmacological blockade of AA by specific inhibitors rapidly (within 15-45 min) down-regulates motivational and consummatory aspects of male sexual behavior in quail while injections of estradiol can rapidly increase the expression of copulatory behavior. These data collectively support an emerging concept in neuroendocrinology, namely that estrogen, locally produced in the brain, regulates male sexual behavior via a combination of genomic and nongenomic mechanisms. Rapid and slower changes of brain AA match well with these two modes of estrogen action and provide temporal variations in the estrogen's bioavailability that can support the entire range of established effects for this steroid. (C) 2004 Elsevier Inc. All rights reserved. [less ▲]

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See detailThe effects of aromatase inhibition on testosterone-dependent conditioned rhythmic cloacal sphincter movements in male Japanese quail
Cornil, Charlotte ULg; Holloway, K. S.; Taziaux, Mélanie ULg et al

in Physiology & Behavior (2004), 83(1), 99-105

Male Japanese quail produce a foam that, along with semen, is transferred to the quail hen during copulation. This foam has been reported to increase fertility, prolong sperm motility, and enhance sperm ... [more ▼]

Male Japanese quail produce a foam that, along with semen, is transferred to the quail hen during copulation. This foam has been reported to increase fertility, prolong sperm motility, and enhance sperm competition. Action of the cloacal sphincter muscles in response to visual exposure to a female produces the foam. The rhythmic cloacal sphincter movements (RCSM) responsible for foam production in male quail is elicited by a conditioned stimulus (CS) previously paired with access to a quail hen. These conditioned RCSM are testosterone-dependent. The present experiment was conducted to explore whether, as is the case with most other testosterone-dependent male sexual behaviors in the quail, conditioned RCSM are mediated by the aromatization of testosterone. Castrated, testosterone-treated male quail were presented with paired presentations of an arbitrary focal CS and visual access to a female. Once conditioned RCSM had developed, subjects received twice daily injections of the aromatase inhibitor Vorozole(TM) (R083842) during a series of extinction test presentations of the CS. Injections of Vorozole(TM) significantly decreased the number of RCSM elicited by a sexual CS. This decrease was specific to sexual RCSM; cloacal sphincter movements that occurred following defecation were not affected by Vorozole. Conditioned sexual RCSM are therefore mediated by the aromatization of testosterone, most likely due to effects on central aromatase activity related to sexual motivation. (C) 2004 Elsevier Inc. All rights reserved. [less ▲]

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See detailAromatase activity modulates conditioned cloacal sphincter movements, an appetitive sexual behavior, in Japanese quail
Holloway, K. S.; Cornil, Charlotte ULg; Taziaux, Mélanie ULg et al

in Hormones & Behavior (2004, June), 46(1), 92

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See detailRapid regulation of brain aromatase activity by afferent inputs: Behavioral implications
Balthazart, Jacques ULg; Baillien, M.; Cornil, Charlotte ULg

in Hormones & Behavior (2004, June), 46(1), 127

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See detailEffects of central administration of Naloxone on the extinction of appetitive sexual responses
Holloway, Kevin; Cornil, Charlotte ULg; Balthazart, Jacques ULg

in Behavioural Brain Research (2004), 153(2), 567-572

Several studies indicate that opioids are involved in the control of consummatory sexual behavior in male Japanese quail. Naloxone has been reported to increase copulatory responses. In the current study ... [more ▼]

Several studies indicate that opioids are involved in the control of consummatory sexual behavior in male Japanese quail. Naloxone has been reported to increase copulatory responses. In the current study, the effect of naloxone on appetitive sexual behaviors was assessed during extinction test trials. Naloxone was found to substantially reduce appetitive responding, suggesting that opioids differentially affect anticipatory and contact components of sexual behavior. [less ▲]

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See detailBehavioral effects of rapid changes in aromatase activity in the central nervous system
Balthazart, Jacques ULg; Baillien, Michelle; Cornil, Charlotte ULg et al

in Kordon, C.; Gaillard, R. C.; Christen, Y. (Eds.) Research and perspectives in endocrine action (2004)

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See detailMultiple mechanisms control brain aromatase activity at the genomic and non-genomic level
Balthazart, Jacques ULg; Baillien, Michelle; Charlier, Thierry ULg et al

in Journal of Steroid Biochemistry & Molecular Biology (2003), 86

Evidence has recently accumulated indicating that aromatase activity in the preoptic area is modulated in parallel by both slow (hours to days) genomic and rapid (minutes to hours) non-genomic mechanisms ... [more ▼]

Evidence has recently accumulated indicating that aromatase activity in the preoptic area is modulated in parallel by both slow (hours to days) genomic and rapid (minutes to hours) non-genomic mechanisms. We review here these two types of control mechanisms and their potential contribution to various aspects of brain physiology in quail. High levels of aromatase mRNA, protein and activity (AA) are present in the preoptic area of this species where the transcription of aromatase is controlled mainly by steroids. Estrogens acting in synergy with androgens play a key role in this control and both androgen and estrogen receptors (ER; alpha and beta subtypes) are present in the preoptic area even if they are not necessarily co-localized in the same cells as aromatase. Steroids have more pronounced effects on aromatase transcription in males than in females and this sex difference could be caused, in part, by a sexually differentiated expression of the steroid receptor coactivator 1 in this area. The changes in aromatase concentration presumably control seasonal variations as well as sex differences in brain estrogen production. Aromatase activity in hypothalamic homogenates is also rapidly (within minutes) down-regulated by exposure to conditions that enhance protein phosphorylation such as the presence of high concentrations of calcium, magnesium and ATP. Similarly, pharmacological manipulations such as treatment with thapsigargin or stimulation of various neurotransmitter receptors (alpha-amino-3-hydroxy-methyl-4-isoxazole propionic acid (AMPA), kainate, and N-methyl-d-aspartate (NMDA)) leading to enhanced intracellular calcium concentrations depress within minutes the aromatase activity measured in quail preoptic explants. The effects of receptor stimulation are presumably direct: electrophysiological data confirm the presence of these receptors in the membrane of aromatase-expressing cells. Inhibitors of protein kinases interfere with these processes andWestern blotting experiments on brain aromatase purified by immunoprecipitation confirm that the phosphorylations regulating aromatase activity directly affect the enzyme rather than another regulatory protein. Accordingly, several phosphorylation consensus sites are present on the deduced amino acid sequence of the recently cloned quail aromatase. Fast changes in the local availability of estrogens in the brain can thus be caused by aromatase phosphorylation so that estrogen could rapidly regulate neuronal physiology and behavior. The rapid as well as slower processes of local estrogen production in the brain thus match well with the genomic and non-genomic actions of steroids in the brain. These two processes potentially provide sufficient temporal variation in the bio-availability of estrogens to support the entire range of established effects for this steroid. [less ▲]

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