Is sexual motivational state linked to dopamine release in the medial preoptic area?

The medial preoptic area (mPOA) is a key site for the dopaminergic enhancement of male sexual behavior. Dopamine release increases in the rat mPOA with mating, supporting the critical stimulatory role played by preoptic dopamine on male sexual behavior. However, it has been questioned whether dopamine is specifically related to the occurrence of male sexual behavior and not simply involved in general arousal. To address this question, we asked whether dopamine release in the mPOA is linked to the production of male sexual behavior in Japanese quail (Coturnix japonica), a species that exhibits a much shorter temporal pattern of copulation than rats and does not have an intromittent organ, resulting in a very different topography of their sexual response. Extracellular samples from the mPOA of adult sexually experienced male quail were collected every 6 min before, during, and after exposure to a female using in vivo microdialysis and analyzed using high-performance liquid chromatography with electrochemical detection. Extracellular dopamine significantly increased in the presence of a female and returned to baseline after removal of the female. However, quail that failed to copulate did not display this increased release. These findings indicate that it is not solely the presence of a female that drives dopamine release in males, but how a male responds to her. Furthermore, in quail that copulated, dopamine release did not change in samples collected during periods of no copulation. Together, these findings support the hypothesis that dopamine action in the mPOA is specifically linked to sexual motivation and not only to copulatory behavior or physical arousal. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Methamphetamine impairs sexual motivation but not sexual performance in male Japanese quail.

The present study investigated the effects of chronic pre-exposure to methamphetamine on sexual motivation and performance in male Japanese quail. Quail were pre-exposed to methamphetamine (1.0 or 3.0 mg/kg ip) or saline (ip) once daily for 10 days and locomotor activity was measured. After a 10 day washout period, sexual motivation was measured in a straight-arm runway with visual access to a female at one end. Three to 5 hr after sexual motivation tests, males were allowed to copulate with a receptive female quail and copulatory behavior was assessed. Tests were conducted once per day for 10 days. Results showed that males pre-exposed to methamphetamine had decreased locomotor activity compared to saline controls. Males pre-exposed to METH later ran slower toward a female in the runway and spent less time near her. In contrast, methamphetamine pre-exposed males showed similar copulatory behavior as saline pre-exposed males. The findings suggest that chronic pre-exposure to methamphetamine may impair sexual motivation but not sexual performance. The findings are discussed from a comparative perspective and with regard to their clinical relevance. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Relative Contributions of the Male and the Female to Sexual Behavior and Reproductive Success in the Japanese Quail (Coturnix japonica).

Two contrasting investigative techniques provided evidence consistent with the interpretation that female quail (Coturnix japonica) regulate male copulatory behavior by the duration of their immobility and through this behavioral mechanism they also control the rate of fertilization of their eggs. In Experiment 1, copulation tests with males and females from different genetic lines showed that the type of female that participated in a copulatory test significantly influenced the latency of the male's grab, mount, and cloacal contact responses and also determined the efficiency of the male's copulatory behavior. These measures of male performance were correlated with female immobility in Experiment 2, which used a more homogeneous population of quail. Furthermore, 2 of these measures (copulatory efficiency and the latency to make cloacal contact) were correlated with fertilization rate. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Sexual fetishism in a quail (Coturnix japonica) model system: Test of reproductive success.

In the present study, the authors explored the reproductive consequences of fetishistic behavior in a previously developed animal model of sexual fetishism (F. K?ksal et al., 2004). Male domesticated quail (Coturnix japonica) received sexual conditioning trials in which a terrycloth object (the conditioned stimulus [CS]) was paired with the opportunity to copulate with a female quail (the unconditioned stimulus). Approximately half of the male quail came to copulate with the CS object and were considered to have developed fetishistic behavior. Each of the male quail was then tested with a female quail, whose eggs were incubated to determine rates of fertilization. The CS object was present for 30 s before and during the copulation test. Fetishistic male quail were slower to achieve cloacal contact with the female quail and showed less efficient copulatory behavior. However, they fertilized a greater proportion of eggs than nonfetishistic male quail. These results are unexpected from previous studies of the relationship between reproductive success and copulatory behavior and are discussed in terms of how fetishistic behavior directed toward an inanimate object may modify male-female interactions. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Effects of enclosure size on sexual behavior of Japanese quail (Coturnix japonica).

The authors determined whether results of experiments on copulatory and affiliative behavior of pairs of Japanese quail (Coturnix japonica) conducted in a closely confining apparatus would predict behavior in a large enclosure in which female quail could avoid contact with male quail. As found previously in studies of closely confined quail, in a large enclosure containing numerous barriers, both unmated female quail and mated female quail laying unfertilized eggs were more likely to remain near a confined male quail than were mated female quail laying fertilized eggs. Furthermore, the number of copulations that a pair engaged in when closely confined predicted the number of copulations that they engaged in when they were in the large enclosure. Patterns of affiliation and of mating in a confining laboratory apparatus thus predicted behavior in a larger enclosure that provided female quail with opportunity to avoid contact with male quail. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Hormonal control of behavior in the Japanese quail.

Adult male and female Japanese quail were functionally castrated by exposure to a short photoperiod and then injected with either estradiole benzoate (EB) or testosterone propionate (TP). Females injected with EB (n = 24) exhibited normal sexual receptivity again but displayed no male courtship behavior. Males injected with EB (n = 10) showed both male and female copulatory patterns. Females injected with TP (n = 14) showed little sexual behavior of any sort, while males injected with TP (n = 10) displayed male sexual behavior. The male quail is thus more bisexual than the female. In contrast with mammalian species, the female is more bisexual. The difference is discussed in relation to the differences in the embryology and cytogenetics of birds and mammals. (29 ref.) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

A comparative approach to the study of dopamine and male sexual behavior: What can Japanese quail teach us? A reply to Pfaus (2010).

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 (see record 2010-24688-009) 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. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Sexual-discrimination learning in male Japanese quail (Coturnix coturnix japonica).

We investigated how male Japanese quail (Coturnix coturnix japonica) learn through extensive social and sexual experience to discriminate between male and female conspecifics. Opportunity for heterosexual copulation was important for this learning, but even extensive copulatory opportunity was not sufficient to produce a sexual discrimination; subjects also required exposure to other males. Exposure to females after copulatory opportunity did not produce a sexual discrimination but facilitated its acquisition. Time or exposure to only the visual features of male birds (provided by taxidermic models) after copulatory opportunity did not result in differential responding to male and female conspecifics. Finally, presenting stimulus birds one at a time proved to be a more sensitive test of sexual-discrimination learning than presenting two stimulus birds at the same time. The results indicate that sexual-discrimination learning is similar to conventional associative learning. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Stimulus control of copulatory behavior in sexually naive male Japanese quail (Coturnix japonica): Effects of test context and stimulus movement.

Sexually experienced male quail (Coturnix japonica) are more likely to engage in copulatory behavior than sexually naive ones. These experiments suggest that sexual experience in a particular place may facilitate later copulatory responding because of increased familiarity with the contextual cues of the environment. Male quail in Exp 1 did not copulate reliably with taxidermic models of females in a novel context, even though some of the subjects were allowed to copulate with female quail in their home cages. In contrast, sexually naive males in Exps 2 and 3 copulated vigorously with taxidermic models of females in a familiar context. In Exp 4, sexually naive males tested in an unfamiliar context were more likely to copulate with a moving than with a static model. The stimulus control of copulatory behavior in sexually naive male quail was similar to that in sexually experienced ones but only in familiar contexts. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Contextual conditioning and the control of copulatory behavior by species-specific sign stimuli in male Japanese quail.

Three experiments were conducted to identify species-specific sign stimuli sufficient to elicit copulatory behavior in male Japanese quail and to determine how learning is involved in the control of behavior by these sign stimuli. In Exp 1, sexually experienced Ss were tested for copulatory behavior with a live female quail and with a model consisting of a female quail's head and neck mounted in front of a foam pad. Comparable levels of copulatory behavior were observed in the two tests, indicating that static visual cues provided by a female quail's head and neck are sufficient to elicit copulatory behavior in this species. Exp 2 showed that male birds that previously received numerous opportunities to copulate with a live female quail in the test situation were significantly more likely to copulate with the head?+?neck model than were sexually inexperienced Ss. Exp 3 showed that prior sexual experience with live quail facilitated responding to the head?+?neck model only if the sexual experience was provided in the same place where Ss were later tested with the model. This finding suggests that sexual experience facilitates control of copulatory behavior by species-specific sign stimuli through contextual conditioning. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Dopamine: Helping males copulate for at least 200 million years: Theoretical comment on Kleitz-Nelson et al. (2010).

Brain dopamine (DA) systems are implicated in a variety of behavioral responses and clinical syndromes, including sex, drug addiction, feeding, satiety, sleep, wakefulness, arousal, attention, reward, decision-making, depression, anxiety, psychosis, and movement disorders. The paper in this issue (see record 2010-24688-004) by Kleitz-Nelson, Dominguez, and Ball (2010) shows how DA release in the medial preoptic area of male quail are activated in an androgen-dependent manner during appetitive and consummatory phases of sexual behavior, similar to that reported previously in male rats. Those data suggest that the steroid-dependent role of hypothalamic DA in male sexual behavior has been conserved through evolutionary time. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Aggression in male mice lacking functional estrogen receptor α.

Estrogen receptor alpha knockout (ERαKO) male mice fail to display sexual behavior. The authors hypothesized that ERαKOs require higher testosterone (T) concentrations than wild-type (WT) males to exhibit copulatory behavior. Increasing T stimulated sexual behavior and preference for females in WT males but failed to do so in ERαKOs. However, T did induce female-directed aggression in ERαKOs. In aggression tests, WT residents selectively attacked T-treated male intruders. ERαKO residents attacked female, T-treated male, and estrogen-treated male intruders equally. Increased access to olfactory cues prior to direct contact reduced overall aggression in ERαKO versus WT males but did not cause ERαKOs to differentially attack male and female opponents. Results suggest that ERα is essential for normal social behavior, perhaps via processing of chemoinvestigatory cues, which are required to discriminate males from females. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Sexual approach conditioning: Unconditioned stimulus factors.

Unconditioned stimulus (US) factors were investigated in a Pavlovian sexual conditioning paradigm with male Japanese quail (Coturnix coturnix japonica). Copulation with a female quail was more effective than exposure to a female without copulation, but the latter also produced conditioned responding (Exp 1). The greater effectiveness of copulatory opportunity as a US was probably not due to nonassociative effects of copulation (Exp 2). Visual cues of the female, as presented on a taxidermic model, were not effective unless the Ss had prior sexual experience that increased their response to the model (Exps 3a and 3b). Successful conditioning with noncopulatory female exposure is significant because it allows for sexual learning to occur in a broader range of circumstances than does conditioning with copulation and because it permits conducting multiple trials per day because males do not become satiated as rapidly. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

6-Hydroxydopamine Lesions in Female Hamsters (Mesocricetus auratus) Abolish the Sensitized Effects of Sexual Experience on Copulatory Interactions With Males.

This study examined the effects of sexual experience in female hamsters (Mesocricetus auratus) on copulatory interactions with male hamsters. Female sexual experience improved the copulatory efficiency of sexually naive males, an effect that persisted for at least 6 weeks without further sexual behavior testing. In a 2nd study, dopamine lesions made in the region of the nucleus accumbens prior to sexual experience specifically blocked the effects of the female's sexual experience on the hit rate of naive males. These results suggest that sexual experience in female hamsters increases the efficiency of copulatory interactions with males, that these effects persist in the absence of further sexual experience, and that dopamine neurotransmission in the basal forebrain underlies this effect of sexual experience. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Electrical stimulation of dorsal and ventral striatum differentially alters the copulatory behavior of male rats.

Sexual behavior is a natural reward that activates striatal dopaminergic (DA) circuits, and dopamine exerts a facilitative influence on copulation. Electrical stimulation of the striatum has been shown to be rewarding, but its effect on male sexual behavior display has not been established. The objective of the present work was to assess the effects of low- and high-frequency electrical stimulation of the dorsal and ventral striatum on male rat sexual behavior expression. To this aim, copulatory activity of sexually experienced male rats was recorded during electrical stimulation of the nucleus accumbens (NAcc) or caudate-putamen (CP), at each stimulation frequency, before and after sexual exhaustion. Results showed that electrical stimulation of the NAcc at both frequencies increased the number of ejaculations that male rats were able to show in a 30-min period. By contrast, stimulation delivered to the CP inhibited sexual behavior by slowing its display. Each effect was more pronounced at low than at high stimulation frequencies. In the same rats, once sexually exhausted, electrical stimulation of these brain areas did not reverse the sexual behavior inhibition that characterizes the sexual exhaustion state. It is concluded that dorsal and ventral striatal DA brain regions exert opposite influences on copulatory behavior expression of sexually experienced male rats. Also, that the facilitative effect of NAcc electrical stimulation on sexual activity, with the stimulation parameters used, cannot surmount the sexual behavior inhibition resulting from copulation to satiation. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Effects of novelty on the reproductive behavior of male Japanese quail (Coturnix coturnix japonica).

In Exp I, 8 sexually deprived male Japanese quail were exposed to 1 of 7 female Japanese quail until they satisfied a 20-min sexual satiation criterion. The female was then replaced with a novel female or was allowed to remain in the test cage undisturbed. Introduction of the novel female did not reinvigorate sexual behavior above levels observed when the female was allowed to remain undisturbed. In Exp II, with 9 males and 15 females, males that were not sexually deprived were tested in the presence of 2 females. During experimental tests, 1 of the females was novel and the other was familiar. During control tests, both of the females were familiar. The presence of a novel female increased overall levels of male sexual behavior and also increased the number of males engaged in reproductive behavior. However, the enhanced sexual behavior was not differentially directed toward the novel or familiar female. Implications of these findings for the mating system of Japanese quail and for studies of the role of novelty in sexual behavior are discussed. (17 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Effects of neurochemical lesions of the preoptic area on male sexual behavior in the Japanese quail

Two experiments were carried out during which the noradrenergic neurotoxin, 5-amino-2,4-dihydroxy-alpha-methylphenylethylamine (5-ADMP) was applied to the brain of quail in order to evaluate the role of the noradrenergic system in the control of male copulatory behavior. In the first experiment, the ICV injection of 5-ADMP slightly enhanced the sexual behavior observed in testosterone (T)-treated castrated male quail. This brings additional support to the notion that norepinephrine tonically inhibits male copulatory behavior in quail. In the second experiment, 5-ADMP implanted directly into the preoptic area disrupted the restoration by T of copulatory behavior in castrated quail and, at the same time, produced a brain lesion that partly destroyed the sexually dimorphic medial preoptic nucleus, a previously established site of T action on behavior. These lesions produced by a high (presumably too high) concentration of neurotoxin provided an independent confirmation of effects previously observed after electrolytic lesions. Correlation analyses also confirmed that the medial part of the POM just rostral to the anterior commissure is more closely associated with copulatory behavior and may, therefore, represent a key center for steroid action on this behavior.     

Appetitive as well as consummatory aspects of male sexual behavior in quail are activated by androgens and estrogens.

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 17β-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. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Lesions of the medial preoptic area interfere with the display of a conditioned place preference for vaginocervical stimulation in rats.

The present study examined the effect of ibotenic acid lesions of the medial preoptic area (mPOA) on the expression of a conditioned place preference (CPP) for vaginocervical stimulation. Rats with bilateral lesions of the mPOA failed to display the CPP for vaginocervical stimulation shown by rats with sham or incomplete lesions. These findings provide additional support for the role of the mPOA in the neural circuitry underlying the reinforcing effects of female sexual behavior and raise the possibility that the altered pattern of approach and withdrawal behavior observed following lesions of the mPOA may be attributable in part to a diminution of the reinforcing effects of vaginocervical stimulation. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

An NMDA antagonist impairs copulation and the experience-induced enhancement of male sexual behavior in the rat.

Sexual experience facilitates subsequent male sexual behavior; activation of the N-methyl-D-aspartate (NMDA) glutamate receptor may play a role in this experience-induced enhancement. In this article, the authors report that systemic injections of MK-801, an NMDA receptor antagonist, impaired male sexual behavior in sexually naive and sexually experienced male rats. Furthermore, saline-treated rats that received 7 daily exposures to an inaccessible estrous female instead of sexual experience displayed enhancement of copulation on the following day. Injections of MK-801 before each of these exposures inhibited the experience-induced enhancement on the drug-free test on Day 8. These data suggest that stimulation of NMDA receptors enhances sexual performance immediately and mediates the experience-induced enhancement of subsequent copulatory behavior. (PsycINFO Database Record (c) 2010 APA, all rights reserved)