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The preoptic area (POA) has long been recognized as a sleep center, first proposed by von Economo. The POA, especially the medial POA (MPOA), is also involved in the regulation of various innate functions such as sexual and parental behaviors. Consistent with its many roles, the MPOA is composed of subregions that are identified by different gene and protein expressions. This review addresses the current understanding of the molecular and cellular architecture of POA neurons in relation to sleep and reproductive behavior. Optogenetic and pharmacogenetic studies have revealed a diverse group of neurons within the POA that exhibit different neural activity patterns depending on vigilance states and whose activity can enhance or suppress wake, non-rapid eye movement (NREM) sleep, or rapid eye movement (REM) sleep. These sleep-regulating neurons are not restricted to the ventrolateral POA (VLPO) region but are widespread in the lateral MPOA and LPOA as well. Neurons expressing galanin also express gonadal steroid receptors and regulate motivational aspects of reproductive behaviors. Moxd1, a novel marker of sexually dimorphic nuclei (SDN), visualizes the SDN of the POA (SDN-POA). The role of the POA in sleep and other innate behaviors has been addressed separately; more integrated observation will be necessary to obtain physiologically relevant insight that penetrates the different dimensions of animal behavior. [ABSTRACT FROM AUTHOR]

Brain serotonin (5-HT) neurotransmission plays an important role in male sexual behavior and it is well established that activating 5-HT1A receptors in rats facilitate ejaculatory behavior. However, the relative contribution of 5-HT1A somatodendritic autoreceptors and heteroreceptors in this pro-sexual behavior is unclear. Moreover, it is unclear whether the contribution of somatodendritic 5-HT1A autoreceptors and postsynaptic 5-HT1A heteroreceptors alter when extracellular 5-HT levels are chronically increased. Serotonin transporter knockout (SERT–/–) rats exhibit enhanced extracellular 5-HT levels and desensitized 5-HT1A receptors. These rats model neurochemical changes underlying chronic SSRI-induced sexual dysfunction. We want to determine the role of presynaptic versus postsynaptic 5-HT1A receptors in the pro-sexual effects of 5-HT1A receptor agonists in SERT+/+ and in SERT–/– rats. Therefore, acute effects of the biased 5-HT1A receptor agonists F-13714, a preferential 5-HT1A autoreceptor agonist, or F-15599, a preferential 5-HT1A heteroreceptor agonist, and S15535 a mixed 5-HT1A autoreceptor agonist/heteroreceptor antagonist, on male sexual behavior were assessed. A clear and stable genotype effect was found after training where SERT+/+ performed sexual behavior at a higher level than SERT–/– rats. Both F-15599 and F-13714 induced pro-sexual activity in SERT+/+ and SERT–/– animals. Compared to SERT+/+, the F13714-dose-response curve in SERT–/– rats was shifted to the right. SERT+/+ and SERT–/– rats responded similar to F15599. Within both SERT+/+ and SERT–/– rats the potency of F-13714 was much stronger compared to F-15599. S15535 had no effect on sexual behavior in either genotype. In SERT+/+ and SERT–/– rats that were selected on comparable low sexual activity (SERT+/+ 3 or less ejaculations and SERT–/– 5 or less ejaculations in 10 weeks) S15535 also did not influence sexual behavior. The two biased compounds with differential effects on 5-HT1A auto- and hetero-receptors, exerted pro-sexual activity in both SERT+/+ and SERT–/– rats. Applying these specific pharmacological tools has not solved whether pre- or post-synaptic 5-HT1A receptors are involved in pro-sexual activity. Moreover, the inactivity of S15535 in male sexual behavior in either genotype was unexpected. The question is whether the in vivo pharmacological profile of the different 5-HT1A receptor ligands used, is sufficient to differentiate pre- and/or post-synaptic 5-HT1A receptor contributions in male rat sexual behavior. [ABSTRACT FROM AUTHOR]

Highlights • Reviews the current status of the behavior system for sexual learning. • In addition to reviewing research with quail (which was the focus of prior paper), in this treatment research with rats is also incorporated into the characterization of the behavior system. • Unlike prior papers which dealt primarily with sexual learning in males, in this review we also discuss in detail sexual learning in females. • For the first time the characterization of the sexual learning system includes detailed information on how learning facilitates fertilization and reproductive success. • Although much progress has been made in characterizing the sexual learning system, we include an extensive section discussing areas for future research. • A new perspective on how sexual learning can contribute to evolutionary change is included. Abstract In this paper we review and update evidence relevant to formulating a behavior system for sexual learning. We emphasize behavioral rather than neurobiological evidence and mechanisms. Our analysis focuses on three types of responses or response modes: general search, focal search, and consummatory or copulatory behavior. We consider how these response modes are influenced by three categories of stimuli: spatially distributed contextual cues, arbitrary localized stimuli, and species-typical cues provided by the sexual partner. We characterize behavior control by these types of stimuli before and after various Pavlovian conditioning procedures in which the unconditioned stimulus is provided by copulation with a sexual partner. The results document extensive Pavlovian modifications of sexual behavior. These conditioning effects reflect new conditioned responses that come to be elicited by various categories of stimuli. In addition, the conditioning of contextual cues and localized stimuli facilitate sexual responding to species-typical cues. Thus, learning experiences enhance how the species-typical cues of a sexual partner stimulate sexual behavior. These modulatory conditioning effects not only produce significant behavioral changes but also increase rates of fertilization of eggs and numbers of offspring produced. These latter findings suggest that sexual learning can lead to differential reproductive success, which in turn can contribute to evolutionary change. [ABSTRACT FROM AUTHOR]

Abstract Ethnopharmacological relevance Piper auritum Kunth is employed as an aphrodisiac in the traditional medicine, but corroborative evidence for such effect is scarce. Aim of the study The pro-sexual effect of an aqueous extract of P. auritum and its possible mechanisms were analyzed in two paradigms of male sexual function. Material and methods Effects of an aqueous extract of P. auritum (PA, single administration) were investigated in the fictive ejaculation, and copulatory behavior paradigms in sexually sluggish male rats. WAY 100635 (antagonist of 5-HT 1A receptors), atosiban (antagonist of oxytocinergic receptors), L-NAME (inhibitor of the nitric oxide synthase) and baclofen (antagonist of GABA B receptors) were used as pre-treatments in order to investigate the role of different neurotransmitter systems in PA actions. Chemical profile of PA was determined by Gases Chromatography and Ultra Performance Chromatography-Electrospray Ionization-Masses Spectrometry (UPLC-ESI-MS). Results In males with retarded ejaculation, PA stimulated ejaculatory behavior and recovered electromyographic activity of pelvic musculature participating in seminal emission and ejaculation. All pre-treatments blocked stimulating effects of PA on the fictive ejaculation; additionally WAY 100635 interfered with PA actions on ejaculatory behavior. Safrol, apigenin dimethylether, myristicin, vaccihein A, sakuranin and sakuranetin flavonoids, were main constituents of PA, with possible participation in its pro-sexual effects. Conclusions Pro-sexual effects of P. auritum elicited at level of ejaculation were mediated by several neurotransmitter systems, among which serotonin and its 5-HT 1A receptors play an important role. Present findings support P. auritum reputation as an aphrodisiac, with potential use in delayed ejaculation disorder. Graphical abstract fx1 [ABSTRACT FROM AUTHOR]

GABAergic transmission in the ventral tegmental area (VTA) exerts a tonic inhibitory influence on mesolimbic dopaminergic neurons’ activity. Blockade of VTA GABA A receptors increases dopamine release in the nucleus accumbens (NAcc). Increases in NAcc dopamine levels typically accompany sexual behavior display. Copulation to satiety is characterized by the instatement of a long lasting (72 h) sexual behavior inhibition and the mesolimbic system appears to be involved in this phenomenon. GABAergic transmission in the VTA might play a role in the maintenance of this long lasting sexual inhibitory state. To test this hypothesis, in the present work we investigated the effect of GABA A receptor blockade in sexually exhausted males 24 h after copulation to satiety, once the sexual inhibitory state is established, and compared it with its effect in sexually experienced rats. Results showed that low doses of systemically administered bicuculline induced sexual behavior expression in sexually exhausted rats, but lacked an effect on copulation of sexually experienced animals. Intra-VTA bilateral infusion of bicuculline did not modify sexual behavior of sexually experienced rats, but induced sexual behavior expression in all the sexually exhausted males. Hence, GABA plays a role in the control of sexual behavior expression at the VTA. The role played by GABAergic transmission in male sexual behavior expression of animals with distinct sexual behavior conditions is discussed. [ABSTRACT FROM AUTHOR]

Gonadal hormones have a developmental role in organization of the nervous system that regulates sexually dimorphic behavior. It is well known that androgen secreted from testes in the perinatal period is converted to estrogen by aromatase in rodent brain, and that estrogen and its receptor play a pivotal role in masculinization of brain structure and function. Treatment with flutamide, an androgen receptor (AR) antagonist, during the perinatal period inhibits development of malespecific brain structure and function, suggesting that androgen signaling via AR also influences brain masculinization. In this study, we investigated which stage during the postnatal period is critical for androgen signaling in brain masculinization. The postnatal period was designated as postnatal days (PD) 0–22, and divided into stages I (PD 0–7), II (PD 8–14), and III (PD 15–22). Newborn male rats were given flutamide subcutaneously in each stage. After adulthood, the effects of postnatal flutamide treatment on brain masculinization were evaluated byanalysis of male sexual behavior. Continuous inhibition of AR throughout stages I and II caused a robust reduction of the intromission ratio and ejaculation frequency compared with other groups. AR inhibition in stage I, II, or III did not cause any change. AR inhibition had no effect onmount behavior. These results show that stage-specific AR activation in the first two postnatal weeks may contribute to brain masculinization mediating male sexual behavior in adulthood. [ABSTRACT FROM AUTHOR]

Male sexual satiety has been associated with a decrease in dopamine levels. Spontaneous recovery of copulatory behavior begins at least 72 h after sexual satiety is reached or in the condition in which a sexually-satiated male is exposed to a new receptive female distinct from the one with which sexual satiety was reached. The aim of the present study was to explore whether dopaminergic activation by bromocriptine (BrCr) can reactivate copulatory behavior with the same sexual mate immediately after sexual satiety is reached. Male rats were divided into three groups exposed to one of the following three conditions: 1) administration of 2 mg/kg s.c. of BrCr and exposure to the same female with whom sexual satiety was previously reached; 2) administration of 0.3 mL s.c. of the vehicle solution with exposure to the same female with whom sexual satiety was reached; and, 3) exposure to a new receptive female after sexual satiety was reached. Results showed that BrCr significantly reactivated copulatory capability in sexually-satiated males with the same receptive female. In contrast, no males in the vehicle group ejaculated with the same female after reaching sexual exhaustion. Copulation was reactivated by BrCr in a way similar to that observed in untreated males exposed to a new receptive female ( i.e. , the Coolidge effect). The reversal of sexual satiety in the males treated with BrCr could be explained by its action on D2 family receptors, which promotes a reactivation of sexual motivation at a level sufficient to allow renewed copulation with the same female mate. [ABSTRACT FROM AUTHOR]

Highlights: [•] SCH 23390 (D1 antagonist) blocks the conditioned place preference induced by ejaculation. [•] SCH 23390 does not modify locomotion and have a transient effect on sexual behavior. [•] SCH 23390 affects only the first coital interaction. [•] SKF 38393 (D1 agonist) induces a conditioned place preference and it is blocked by SCH 23390. [•] Our results suggest that dopamine is involved in the sexual reward induced by ejaculation. [ABSTRACT FROM AUTHOR]

Abstract: Sexual behavior of the male rat can be described as a ‘sequence’: a series of behavioral transitions eventually leading to a consummatory act: ejaculation. A ‘funnel-model’ is presented to describe the behavioral progression during the sexual sequence. The ejaculation itself is extensively controlled by the ‘spinal ejaculation generator’, consisting of several elements with afferent sources of genitosensory information, with ascending projection fibers to inform the brainstem and forebrain as well as with descending afferent fibers providing the supraspinal control mechanisms with the opportunity to restrict ejaculations to the optimal moments and circumstances. The messages ascending from the spinal cord reach several interconnected thalamic, hypothalamic and limbic brain areas and are integrated with olfactory information. These brain areas play a role in mechanisms like ‘sexual satiety’ or a temporary interruption of sexual activities (post-ejaculatory interval), but the exact facilitatory and inhibitory mechanisms involved have not been elucidated yet. In the ‘downward’ mechanisms controlling the spinal ‘release’ of an ejaculation, the medial preoptic nucleus plays an important role in cooperation with a number of brainstem areas. This nucleus is also explicitly involved in the rewarding experiences coming with an ejaculation. Finally, the role of several neurotransmitters and-peptides on male sexual behavior are discussed shortly, because sometimes they show remarkable effects on specific aspects of the behavioral sequence. We conclude that, despite our increased knowledge about the brain mechanisms involved in the control of ejaculation, we are still far away from a complete understanding and quite a few questions remain to be resolved. [Copyright &y& Elsevier]

Abstract: Several neurotransmitters, among them neuropeptides, have been implicated in the control of male sexual behavior. Opioids are involved in mediating the positive affective states generated by the execution of sexual behavior in both males and females. We have previously shown that intracerebroventricular administration of endomorphin-1 (EM-1), the specific ligand for the μ opioid receptor, increased the ejaculation latency and interintromission interval and reduced the number of ejaculations during the test. In the present study we evaluated the effect of EM-1 upon male sexual behavior and socio-sexual interactions when infused in the medial preoptic area (MPOA) or the medial amygdala (Me). The results indicate that the administration of EM-1 in the MPOA increased mount and intromission latencies while infusion in the Me increased the number of mounts before ejaculation as well as the ejaculation latency. With respect to socio-sexual interactions, the duration of pursuit was significantly increased after administration of EM-1 in the MPOA. The effects upon sexual behavior and socio-sexual interactions were blocked by administration of the opioid antagonist naloxone. These results indicate that EM-1 modulates the appetitive aspect of sexual behavior in the MPOA and the consummatory phase in the Me. [Copyright &y& Elsevier]

Abstract: Discoveries in the past two decades have continued to improve our understanding of the pathophysiology of erectile dysfunction disease and animal models have played a significant role to define the basic mechanisms of erectile dysfunction treatment. Both in vitro and in vivo models have been developed in the past years to study the aphrodisiac agents. Methods that are used in aphrodisiac study can be categorized into physical methods including male sexual behavior (mount frequency, mount latency, intromission frequency, intromission latency, ejaculation frequency, post-ejaculatory interval, couplatory rate, index of libido, computed male sexual behavior parameter), pendiculation study, orientation behavior, determination of hesitation time & attraction towards female, test of potency, test for libido, penile microcirculation study, Intracavernous pressure study and biochemical methods, histopathology, sperm count, Fructose content in seminal vesicles, sperm preservation, organ weight, hormonal determination, assay of nitric oxide synthase, In vitro nitric oxide release & androgen receptor protein. This review aims to highlight some of the new and currently used experimental models that are used for the evaluation of aphrodisiac agents. [Copyright &y& Elsevier]

The degree to which male sexual behavior and territorial aggression are regulated by gonadal steroid hormones depends strongly on species and experience. While castration abolishes male sexual behavior in most laboratory rodents, approximately one third of B6D2F1 mice retain the full repertoire of male sexual behaviors long term ("maters"). It is not yet known whether maters retain other behaviors that typically rely on gonadal steroids to a greater extent than non-maters. In this study, we tested aggressive behavior in B6D2F1 males and males of each parental strain (C57BL/6J and DBA/2J) in the resident intruder paradigm before and after castration, as well as male sexual behavior after castration. Before castration, B6D2F1 residents displayed more attacks compared to DBA/2J males (p < 0.05). There was no difference in attack frequency between B6D2F1 and C57BL/6J males nor between DBA/2J and C57BL/6J males (p > 0.2). A greater proportion of hybrid males demonstrated intromissions and the ejaculatory reflex compared to males of either parental strain (p < 0.01). After castration, B6D2F1 residents attacked more than C57BL/6J males, but not DBA/2 J males (p < 0.05; p > 0.2). There was no difference in post-castration attack frequency between maters and non-maters (p > 0.7). Finally, residents that attacked during all 3 pre-castration resident intruder tests displayed more attacks post-castration than animals that attacked during 1 pre-castration test (p < 0.05). These data suggest that strain and experience influence the expression of aggressive behavior after castration and warrant future study in experience-induced transient increases in extragonadal testosterone. • Intact B6D2F1 males attacked more than DBA/2J males in the resident intruder test. • Persistent maters and non-maters did not differ in post-castration aggression. • B6D2F1 and DBA/2J, but not C57BL/6J residents, continued to attack post-castration. • Residents that attacked during 3 pre-castration tests attacked more post-castration. [ABSTRACT FROM AUTHOR]

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-OHATD (1,4,6-androstatrien-17β-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. [ABSTRACT FROM AUTHOR]

Abstract: The correlation between monoamine levels in the nucleus accumbens (NAcc) and male sexual behavior was studied in middle-aged rats. Male rats (18–19months) were assigned to three groups: (1) Group MIE consisted of rats showing mounts, intromissions, and ejaculations; (2) Group MI was composed of rats showing mounts and intromissions, but no ejaculation; and (3) Group NC were non-copulators showing no sexual behavior. Young adult rats (4–5months), displaying complete copulatory behavior, were used as the control group. Levels of dopamine (DA), serotonin, and norepinephrine and their metabolites in the NAcc were measured by high-pressure liquid chromatography with electrochemical detection. No difference was seen in DA levels between MIE rats and young controls, whereas DA levels in NC rats were significantly lower than those in both MIE and MI rats. Serotonin levels in NC rats were significantly higher than those in MIE and MI rats. Conversely, norepinephrine levels in NC rats were lower than those in MIE rats. These results suggest that monoamine levels in the NAcc correlate with sexual performance in male rats and that changes in NAcc monoamine levels might affect male sexual behavior in middle-aged rats. [Copyright &y& Elsevier]

Oxytocin (OT) modulates adult mammalian sexual behavior, sperm production and transport, and steroidogenesis; however, the consequences of developmental manipulations of oxytocin have received little attention. The purpose of this experiment was to determine whether neonatal exposure to OT, an oxytocin antagonist (OTA), saline (SAL), or handling (HAN)-only would have long-term effects on reproductive potential in male prairie voles (Microtus ochrogaster). Adult males were observed for 24 h with a sexually receptive female and sexual behavior was recorded. Females were subsequently lavaged and smears were examined for sperm. Reproductive parameters including motility of epididymal sperm, testis weight, and plasma androgen levels were in the normal range. OT-treated males that did not mate within the first 30 min did not mate at all, and in comparison to controls, a higher proportion of those OT-treated and OTA-treated males that did mate did not transfer sperm to the females. OTA-treated males also had significantly higher testicular sperm concentrations than HAN-only males, and significantly lower epididymal sperm concentrations. These differences suggest that in males, developmental manipulations of OT may have the potential to influence the subsequent expression of sexual behavior and sperm transport. [Copyright &y& Elsevier]

Adult male rats were exposed to inhale halothane in the following regime: 15 ppm/4 h/5 days/week/9 weeks. Sexual behavior observations and sperm motility test were made before halothane exposure (0 days) and at 15, 30, 45 and 60 days of exposure. Fifteen days after halothane exposure, this anesthetic inhibited the proportion of animals displaying ejaculation. In those animals ejaculating, halothane produced an inhibition of masculine sexual behavior reflected as an increase in the intromission latency, number of mounts and postejaculatory interval. At 30 days after exposure, only an increase in the intromission latency was observed. At 45 and 60 days, the inhibitory effect of halothane on sexual behavior disappeared. Similarly, at 15 and 30 days, but not at 45 or 60 days of halothane exposure, a reduced sperm motility was observed. Such transient effects of halothane suggest the development of tolerance to the inhibitory actions of this anesthetic on sexual behavior and sperm motility. These halothane effects are in line with an inhibition of masculine sexual behavior after stimulation of the GABAergic system. [Copyright &y& Elsevier]

In rodents, gonadal steroids play a critical yet variable role in behaviors such as social interaction and cognitive performance. Gonadal steroids organize sex differences observed in spatial working memory, while the absence of activational effects induced by castration generally impedes spatial learning and memory. Although male sexual behavior is typically inhibited following castration, a significant proportion of gonadectomized B6D2F1 hybrid males retains the complete repertoire of male reproductive behavior. In a prior study, amyloid precursor protein and tau, proteins involved in cognitive behavior, facilitated steroid-independent male sex behavior in B6D2F1 hybrid male mice. We used this strain to investigate the relationship between gonadal steroid-independent male sexual behavior and cognition. After identifying "maters" (animals retaining steroid-independent male sex behavior) and "non-maters," we tested spatial memory in an 8-arm radial arm maze. Although neither group demonstrated a decrease in errors as a function of time, maters committed fewer errors compared to non-maters overall (p < 0.05). Maters also completed the maze more quickly than non-maters (p < 0.05). We measured mRNA expression of APP and MAPT as well as LEPR and D2R to probe potential roles of metabolism and motivation. Uniquely among maters, increased relative expression of D2R and LEPR in the hippocampus was associated with a longer latency to complete the maze during the last 3 or across all trials, respectively. These data demonstrate that maters outperform non-maters in the radial arm maze, warranting further study of potential differences in acquisition of spatial memory tasks or learning strategy between these groups. • ~30% B6D2F1 male mice retain male sex behavior (MSB) long after castration (maters). • Maters commit fewer errors and complete the radial arm maze more quickly compared to non-maters. • Steroid-independent retention of male sexual behavior and spatial memory may be linked. • Motivation and metabolism may be involved in this relationship. [ABSTRACT FROM AUTHOR]

Enduring sex differences in the brain are established during a developmental process known as brain sexual differentiation and are mainly driven by estrogens during a critical period. In rodents, the masculinization of the preoptic area by estrogens derived from the central aromatization of testosterone depends in part on the interaction between microglia and prostaglandin E 2 (PGE 2), a pro-inflammatory hormone of the prostanoid subclass. In contrast, in birds, estrogens produced by females induce a demasculinization, but whether an interaction with the neuro-immune system is involved in this process is unknown. This study addressed this question by testing the effects of blockade of cyclo‑oxygenases (COX), the rate-limiting enzymes for prostanoid synthesis, on embryonic microglia and the sexual differentiation of brain and behavior using the Japanese quail as an animal model. The results show that COX inhibition does not affect the behavior of females, but impairs male sexual behavior and suppresses the sex difference in microglial profiles at embryonic day 12 (E12) in the medial preoptic nucleus by increasing the number of microglia in males only. However, neither prostanoid concentrations nor PGE 2 receptors differed between sexes in the hypothalamus and preoptic area (HPOA) during development. Overall, these results uncovered a potential role of prostanoids in the demasculinization of Japanese quail. Moreover, the parallel effect of COX inhibition on behavior and microglia suggests an interaction between prostanoids and microglia in brain demasculinization, thus fueling the hypothesis of a conserved role of the neuroimmune system in the organization of the brain by estrogens. • Inhibiting embryonic cyclo‑oxygenases (COX) impairs male sexual behavior in quail. • Early COX inhibition prevents the embryonic sex difference in preoptic microglia. • Brain prostanoid levels and PGE 2 receptors did however not differ between sexes. • These data suggest a role of prostanoids in the demasculinization of the avian brain. • This process may also involve an interaction between prostanoids and microglia. [ABSTRACT FROM AUTHOR]

One of the most robust findings about dopamine (DA) is that the stimulation of dopaminergic systems promotes the activation of male sexual behavior. The commentary by Pfaus (2010) included a thorough review of studies of DA and male sexual behavior. We agree with him that the release of DA in the preoptic region in male quail in response to females and in association with the exhibition of male sexual behavior appears to be highly conserved and that it seems to have evolved very early in the evolutionary history of the vertebrate brain. However, additional data have been collected indicating that there may be significant species differences in the dopaminergic regulation of male behavior in quail compared with rats. In this response, we take the opportunity to make a few broader points about DA and male sexual behavior in light of other studies that have been conducted in birds and introduce some interesting taxonomic variation that is still not well understood. [ABSTRACT FROM AUTHOR]