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PubMed2001

Testosterone restoration of copulatory behavior correlates with medial preoptic dopamine release in castrated male rats.

The medial preoptic area (MPOA) is an important integrative site for male sexual behavior. We have reported an increase in dopamine (DA) release in the MPOA of male rats shortly before and during copulation. Postcastration loss of copulatory ability mirrored the loss of the precopulatory DA response to an estrous female. The present study investigated the time courses of restoration, rather than loss, of the MPOA DA response to a receptive female and of copulation in long-term castrates. Male rats were castrated and tested for loss of copulatory ability 21 days later. They then received 2, 5, or 10 daily subcutaneous injections of testosterone propionate (TP, 500 microg) or oil. Microdialysate samples were collected from the MPOA during baseline, exposure to a female behind a barrier, and copulation. Extracellular DA was measured using HPLC-EC. None of the six 2-day-TP-treated animals copulated, nor did they show elevated DA release in the MPOA in the presence of a receptive female. Five of the nine 5-day-TP-treated animals ejaculated; three intromitted without ejaculating; and one failed to copulate, with all but the noncopulating animal showing elevated DA release. All of the six 10-day-TP-treated animals copulated and also demonstrated an increase in MPOA DA. None of the oil controls copulated or showed an increase in DA release. Therefore, a consistent relationship between MPOA DA release during exposure to a receptive female and the subsequent ability of the male to copulate was observed.

Analysis of Variance↗
PubMed1992

Copulatory analgesia in male rats ensues from arousal, motor activity, and genital stimulation: blockage by manipulation and restraint.

The effect of copulation on the vocalization threshold to tail shock (VTTS) was assessed in freely-moving, sexually experienced, Wistar male rats. Mean VTTS during the first copulation was 40% above the baseline values and slightly decreased during the first postejaculatory interval (PEI; 28% above baseline). VTTS mean values further increased during the second copulatory series (93% above baseline and 63% during the PEI). Testing at the same intervals in noncopulating rats did not induce analgesia. VTTS values also increased after the display of either one ejaculation (E), five intromissions (I), or five mounts (M), the analgesia persisting for at least 20 min. Analgesia following M was smaller and shorter than that observed after either I or E. Analgesia developed gradually during copulation since VTTS values after five I were significantly higher than those after one I. The incidence of vocalizations to suprathreshold shocks (STS, 20% above the VTTS) occurring during various phases of copulation was also studied to determine the onset and short-term fluctuations of copulatory analgesia. The proportion of STS inducing vocalizations decreased from 91% (mean of individual proportions) before copulation, to 24% during copulation, and to 25% during the PEI. A maximal reduction in the proportion of vocalizations to STS was found during the last third of copulation. Nearly all vocalizations to STS during copulation occurred when subjects were quiescent, while few or no vocalizations occurred when rats were engaged in sexual activity, i.e., during M, I, or penile grooming. In contrast to the above-mentioned data, no significant analgesia was observed using the tail-flick latency test (TFL) following either E, five I, or five M.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗
PubMed1999

Prolonged mating in prairie voles (Microtus ochrogaster) increases likelihood of ovulation and embryo number.

Prairie voles are induced ovulators that mate frequently in brief bouts over a period of approximately 24 h. We examined 1) impact of mating duration on ovulation and embryo number, 2) incidence of fertilization, 3) temporal pattern of embryo development, 4) embryo progression through the reproductive tract over time, and 5) embryo development in culture. Mating was videotaped to determine first copulation, and the ovaries were examined and the reproductive tracts flushed at 6, 8, 10, 12, 16, 20, and 24 h and 2, 3, and 4 days after first copulation. The number of mature follicles and fresh corpora lutea and the number and developmental stage of embryos were quantified. One, two-, and four-cell embryos were cultured in Whitten's medium. Mature follicles were present at the earliest time examined (6 h). Thirty-eight percent of females that had been paired for < 12 h after the first copulation ovulated, whereas all females paired >/= 12 h after the first copulation ovulated. Virtually all (> 99%) oocytes recovered from females paired for >/= 12 h after first copulation were fertilized. Pairing time after first copulation and mean copulation-bout duration were significant (p < 0.05) determinants of embryo number. Embryos entered the uterine horns and implanted on Days 3 and 4, respectively, after first copulation (Day 0). Embryos cultured in vitro underwent approximately one cell division per day, a rate similar to that in vivo. We conclude that prairie voles ovulate reliably after pairing for >/= 12 h, although some females showed exceptional sensitivity not predicted by the variables quantified. Prolonged mating for longer than 12 h increased the total embryos produced. This mechanism likely has adaptive significance for increasing offspring number.

Animals↗
PubMed1997

Behavioral and endocrine characteristics of the reproductive cycle in wild muriqui monkeys, Brachyteles arachnoides.

The analysis of fecal ovarian steroids provides a powerful noninvasive method to obtain insights into ovulatory cycles, gestation length, and the timing of sexual interactions relative to the periovulatory period in wild primates. Techniques developed to collect and assay feces from free-ranging muriqui monkeys (Brachyteles arachnoides) for estradiol and progesterone yield the first explicit reproductive data on this species, and provide the first opportunity to evaluate the timing of observed copulations with muriqui ovarian cycles. Hormonal profiles from seven females indicate average cycle lengths of 21.0 +/- 5.4 days (n = 20). Females conceived after 3-6 ovulatory cycles. Gestation length averaged 216.4 +/- 1.5 days for the five females for which conception cycles were sampled. Discrete copulation periods spanned an average of 2.1 +/- 1.2 days (n = 29), with intervals between these concentrated periods of copulations averaging 15.6 +/- 6.7 days (n = 20). There were no significant differences among females in cycle lengths, copulation period lengths, or copulation interval lengths. Ejaculation was visible following 71.8 +/- 26.7% of copulations during the females preovulatory periods. All females copulated outside the periovulatory period. The proportion of copulation days outside the periovulatory period was slightly greater (p = 0.08) for primiparous females (64.8 +/- 28.3%) than for multiparous females (28.7 +/- 19.7%).

Animals↗
PubMed2006

Mating behaviors of the proboscis monkey (Nasalis larvatus).

The mating behaviors of the proboscis monkey were observed in a riverine forest along a tributary of the Kinabatangan River, Sabah, Malaysia, for a period of 30 months. Solicitation for copulation was initiated frequently by males and occasionally by females. Most copulations involved only one mount; however, some multiple-mount copulations were observed and a maximum of six mounts per copulation were recorded. The mean duration of mounts was about 27 sec. Nonsexual mounts (female-female, female-juvenile/infant, juvenile-juvenile, and juvenile-infant) were also observed. Female-female mounts occurred shortly after failed solicitations toward males were observed. Harassment by juveniles and/or infants was observed during copulation; however, these harassments apparently did not interfere with copulation. Sexual swelling was evident in 77.4% of copulating females, with copulating subadult females showing the most distinct swelling.

Animals↗
PubMed1990

Medial preoptic area kindling induces sexual behavior in sexually inactive male rats.

In the present experiments, sexually experienced and non-copulating male rats were kindled in the medial preoptic area (MPOA) or the amygdala (AMG) with the purpose to investigate if the widespread modification of brain function produced by kindling induces sexual behavior in non-copulating rats and if kindling facilitates sexual behavior in copulating rats. Since kindling in the MPOA has been little studied, a description of this phenomenon is also presented. The animals were stimulated 4 times daily on odd days and sexual behavior monitored on even days. The results showed that the MPOA required higher stimulus intensity to elicit an afterdischarge (AD) than the AMG. The AD duration was significantly longer in animals kindled in the MPOA than in animals kindled in the AMG. No difference was found in the development of kindling between these brain structures. Kindling had no effect upon sexual behavior in copulating rats. In contrast, MPOA kindling induced copulation in non-copulating male rats. Seven out of 9 animals displayed sexual behavior. No significant facilitation was observed after AMG kindling in non-copulating male rats. Furthermore, the sexual behavior displayed by the previously non-copulating MPOA kindled rats was similar to the sexual behavior displayed by sexually experienced animals. It is proposed that sexual behavior is facilitated in these male rats by local neural changes produced by kindling.

Animals↗
PubMed1995

Sperm competition: evolutionary causes and consequences.

The interaction between functional and mechanistic approaches to sperm competition and between male and female perspectives are described and illustrated by a study of the zebra finch, Taeniopygia guttata. Sperm competition experiments in the laboratory show that last male sperm precedence occurs (as it does in many other taxa) although the mechanism is unknown (as in most other taxa). Empirically-derived values were used to construct a mathematical model of sperm competition in the zebra finch. The model indicates that precedence occurs as a consequence of: (i) the temporal pattern of pair copulations; (ii) the rate at which sperm are lost from the female tract; and (iii) more sperm being transferred during extra-pair copulations than during pair copulations. The latter effect is a consequence of males seeking extra-pair copulations after their own pair copulation period has ended. The effect of sperm numbers on the pattern of sperm precedence may be further increased by: (i) extra-pair males increasing ejaculate size (sperm numbers) (for which there is no evidence); (ii) extra-pair males being of a better quality and transferring more sperm or better quality sperm (for which there is some evidence); and (iii) cryptic female choice. Females eject over 99% of sperm following insemination; if they eject fewer sperm from males chosen as extra-pair copulation partners, the potential for cryptic female choice is considerable. However, this is still being investigated. The model also predicts the optimal time for an extra-pair copulation to occur (from either a male or female perspective). A comparison between the predicted and observed pattern suggests that the optimal timing of extra-pair copulations is constrained in both sexes.

Animals↗
PubMed2005

Novel male trait prolongs survival in suicidal mating.

Male redback spiders (Latrodectus hasselti) maximize paternity if they copulate twice with their cannibalistic mate. Facilitating cannibalistic attack during their first copulation yields paternity benefits. However, females have paired sperm-storage organs inseminated during two separate copulations, so males that succumb to partial cannibalism during the first copulation lose at least 50% of their paternity to rivals. In this paper, we describe a novel male trait--an abdominal constriction that appears during courtship--that allows males to survive and mate with females for a second time, despite the substantial cannibalistic damage inflicted during the first copulation. Constricted males that were wounded to simulate early cannibalism had higher endurance, greater survivorship, longer subsequent courtship and higher mating success than wounded males that were not constricted. Constriction was not found in a non-sacrificial congener that rarely survived simulated cannibalism, and the protective effect of constriction in redbacks was specific to the type of damage inflicted by females during the first copulation. Thus, the abdominal constriction allows males to overcome the potential fitness limit imposed by their own suicidal strategy-paradoxically, by prolonging survival across two cannibalistic copulations.

Animals↗
PubMed1998

Sperm protection and mate assessment in the bushcricket Coptaspis sp. 2.

Several adaptive explanations regarding the function of lengthy copulations in insects have been proposed. They may represent a form of mate guarding, where the male physically prevents the female from copulating with rival males. Alternatively, they may function to ensure full insemination of the male's sperm when copulation duration covaries with the amount of sperm transferred and male fertilization success. Finally, lengthy copulations may serve to allow males to assess female quality in terms of mating status and body weight. In this study I examine these hypotheses for the function of lengthy copulations in the Australian bushcricket Coptaspis sp. 2 (Orthoptera: Tettigoniidae). Unlike most other bushcrickets, males of this species do not produce a large spermatophylax that the female feeds on during insemination, but remain attached to the female's genitals up to 6 h after spermatophore attachment. Experimental manipulation of the duration of spermatophore attachment showed it to be related to the amount of sperm transferred. This suggests that the main function of copulation duration is to ensure complete transfer of the male's ejaculate. Males also discriminated between females, and provided mated females with more sperm which resulted in longer copulations than with virgin females. It is possible that possession of a large spermatophylax has been lost evolutionarily in this species, with males themselves acting as a sperm protection device during insemination.Copyright 1998 The Association for the Study of Animal Behaviour

Journal Article↗
PubMed1995

Sexual activity increases dopamine transmission in the nucleus accumbens and striatum of female rats.

In vivo microdialysis was used to monitor extracellular concentrations of dopamine (DA), and its metabolites dihydroxyphenylacetic acid (DOPAC) and homovanillic acid (HVA) in the nucleus accumbens and dorsal striatum of sexually active female rats during tests of locomotor activity, exposure to a novel chamber, exposure to sex odors, the presentation of a sexually active male rat, and copulation. DA increased slightly but significantly in the nucleus accumbens when a sexually active male was placed behind a wire-mesh screen, and further during copulation. DA also increased significantly in the dorsal striatum during copulation; however, the magnitude of this effect was significantly lower than that observed in the nucleus accumbens. The metabolites DOPAC and HVA generally followed DA with a delay, and increased significantly during copulation in both regions. In contrast, forced locomotion on a rotating drum, exposure to a novel testing chamber, and exposure to sex odors did not increase DA significantly in either region, although forced locomotion increased DOPAC significantly in both regions, and HVA significantly in the nucleus accumbens. The magnitude of DA release in the nucleus accumbens was significantly greater during copulation than running, whereas no significant difference was detected for striatal DA release between these two behavioral conditions. These results indicate that novelty or locomotor activity alone do not account for the increase in DA observed in the nucleus accumbens of female rats during copulation, and suggest that DA transmission in the nucleus accumbens is associated with anticipatory and consummatory aspects of sexual activity, as it is in male rats. In the dorsal striatum, however, DA release during copulation may reflect an increase in locomotor activity associated with active pacing of the male.

3,4-Dihydroxyphenylacetic Acid↗
PubMed1999

Hormone-neurotransmitter interactions in the control of sexual behavior.

The stimuli from a receptive female and/or copulation itself leads to the release of dopamine (DA) in at least three integrative hubs. The nigrostriatal system promotes somatomotor activity; the mesolimbic system subserves numerous types of motivation; and the medial preoptic area (MPOA) focuses the motivation onto specifically sexual targets, increases copulatory rate and efficiency, and coordinates genital reflexes. The previous (but not necessarily concurrent) presence of testosterone is permissive for DA release in the MPOA, both during basal conditions and in response to a female. One means by which testosterone may increase DA release is by upregulating nitric oxide synthase, which produces nitric oxide, which in turn increases DA release. Hormonal priming in females may also increase DA release in the MPOA, and copulatory activity may further increase DA levels in females. One of the intracellular effects of stimulation of DA D1 receptors in the MPOA of male rats may be increased expression of the immediate-early gene c-fos, which may mediate longer term responses to copulation. Furthermore, increased sexual experience led to increased immunoreactivity to Fos, the protein product of c-fos, following copulation to one ejaculation. Another intracellular mediator of DA's effects, particularly in castrates, may be the phosphorylation of steroid receptors. Finally, while DA is facilitative to copulation, 5-HT is generally inhibitory. 5-HT is released in the LHA, but not in the MPOA, at the time of ejaculation. Increasing 5-HT in the LHA by microinjection of a selective serotonin reuptake inhibitor (SSRI) increased the latency to begin copulating and also the latency to the first ejaculation, measured from the time the male first intromitted. These data may at least partially explain the decrease in libido and the anorgasmia of people taking SSRI antidepressants. One means by which LHA 5-HT decreases sexual motivation (i.e. increases the latency to begin copulating) may be by decreasing DA release in the NAcc, a major terminal of the mesolimbic system. Thus, reciprocal changes in DA and 5-HT release in different areas of the brain may promote copulation and sexual satiety, respectively.

Animals↗
PubMed2004

Extreme repeated mating as a counter-adaptation to sexual conflict?

The Australian scaly cricket, Ornebius aperta, can copulate over 50 times with the same partner; the benefits of such extreme repeated copulation are unclear. We support the hypothesis that repeated copulation increases insemination success, as the number of sperm transferred increases with each spermatophore. This probably increases paternity for males, as on average a female mates with over 40 males. Despite intense sperm competition each ejaculate has only a few hundred sperm, orders of magnitude less than in related crickets. We show that all sperm are transferred from each spermatophore in the few seconds before a female removes and eats it. Repeated copulation increases effective copulation duration while a small ejaculate ensures that this strategy is not excessively costly. Thus repeated copulation in these crickets may have arisen as a counter-adaptation to female-imposed limits on copulation.

Adaptation, Biological↗
PubMed2001

Strategic male mating effort and cryptic male choice in a scorpionfly.

In animal species with high male mating effort, males often find themselves in a dilemma: by increasing their mating effort, the gain from each copulation increases but simultaneously reduces available resources and, thus, the opportunity for future copulations. Therefore, we expect males to spend less reproductive resources on matings that provide low reproductive potential, thereby saving resources for future copulations, possibly with high-quality females, a sort of cryptic male choice. However, the strength of the trade-off between investment in a current mating and resources available for future matings must not be the same for all males. Males with relatively high mating costs should allocate their limited resources more cautiously than males with more plentiful resources. Here, we examine this prediction in the scorpionfly Panorpa cognata. Prior to copulation, males produce a large salivary mass on which females feed during copulation. We show that the production of larger salivary masses leads to longer copulations. Moreover, the size of the salivary gland and salivary mass increases with increasing male condition. However, males in poor condition make a relatively higher mating investment than males in good condition. We therefore expect male condition to influence cryptic male choice. In accordance with our hypothesis, only males in poor condition choose cryptically, producing larger salivary masses in copulations with females of high fecundity.

Animals↗
PubMed1967

Temporal patterns of sexual behavior in rabbits as determined by an automatic recording technique.

Sexual behavior between rabbits was automatically recorded by means of electrodes attached externally to the unrestrained animals. Current flowed only when the animals came into physical contact; the duration and amplitude of the current flow provided a means of automatic differentiation between sexual and other types of physical contact. Various temporal aspects of sexual behavior were revealed: (1) copulation was preceded by a series of thrusts that occurred at a constant frequency of 13 to 14 thrusts per sec; (2) the duration of a copulation was about 1.5 sec and could be as brief as 0.3 sec; (3) copulation was much more likely during the hours between dusk and dawn despite constant illumination; (4) copulations occurred in groups or "runs" characterized by many copulations within a few hours; (5) each run began with a high rate of copulation that often decreased over time; (6) between runs, several hours or days passed with little or no sexual activity; and (7) a run of copulations was noted at the end of every 16- to 20-day period (which corresponds to the known period of pseudopregnancy in rabbits) whether or not other runs occurred within this period.

Animals↗
PubMed1993

Effects of an oral contraceptive on sexual behavior of chimpanzees (Pan troglodytes)

Behavioral and physiological effects of a combined oral contraceptive (OC) were studied in chimpanzees for comparative purposes related to (i) the ambiguity surrounding the effects of OCs on the sexuality of humans, (ii) the close biological relationship between chimpanzees and humans, especially with respect to endogenous sex hormones and sexual behavior, and (iii) the relatively greater behavioral sensitivity of the chimpanzee to changes in sex hormone levels such as those that accompany the use of OCs. Two different types of pair tests and detailed behavioral assessments were used to differentiate the hormonal effects of female behavior from social effects imposed by the male. Anogenital swelling and copulation were reduced during OC cycles, but the effect of the OC on copulation was directly related to the social and sexual relationship of the pair during natural cycles. The more compatible and more frequently copulating pairs in the natural cycles continued to copulate during the OC cycles, albeit at reduced rates, whereas the less compatible and less frequently copulating pairs ceased copulating altogether when the female received the OC. There was no independent effect of the OC on ejaculation per se. Both male and female sexual initiative were reduced in the OC cycles, but female responsiveness to male sexual solicitations and direct copulatory attempts were not affected. Data indicate that despite generally adverse effects of the OC on certain physiological and behavioral dimensions of sexuality, social factors ultimately determined the degree of response to the OC.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗
PubMed1984

Periovulatory changes in female sexual behavior and patterns of ovarian steroid secretion in group-living rhesus monkeys.

The behavior of nine intact group-living adult female rhesus was observed for 30 min daily with each of four adult male rhesus across a verified ovulatory menstrual cycle. Blood samples collected from females daily or on alternate days were analyzed for estradiol, testosterone, and progesterone. Female patterns of approach, follow, and initiate proximity increased several days prior to the estradiol peak, peaked on the day of the estradiol peak, then declined completely or to very low frequencies. Mounts, intromissions, and ejaculations increased significantly on the day of the estradiol peak, remained elevated for 2 more days, then declined completely by the fifth day after peak estradiol. Ejaculations never occurred outside of a 10-day period starting 4 days before the estradiol peak and ending 5 days after the estradiol peak. During this period females initiated over 90% of all approaches. Female hand slap, threaten away, and stand up increased significantly on the first day of increased copulation, remained elevated while copulation was significantly elevated, then decreased along with the decline in copulation. Ten of eleven patterns of female behavior correlated significantly with estradiol level prior to the estradiol peak. All were significantly inversely correlated with progesterone level after the estradiol peak. No pattern of female behavior correlated significantly with testosterone either before or after the estradiol peak. Similarly, male patterns of behavior correlated with female levels of estradiol and progesterone, but not testosterone. These results demonstrate a relationship between increased serum estradiol and increased female initiation of sexual behavior. The finding that some patterns of female behavior increase several days prior to copulation, whereas other behaviors increase coincident with increased copulation suggests that the behavior of group-living rhesus females serves two functions. The first is to communicate sexual interest and the second is to maintain the consort pair and increase the probability that ejaculation will occur. In addition, the strong correlation between preovulatory female behavior and estradiol level suggests that the female's behavior provides precise information about her reproductive state and could thus coordinate copulation with maximal fertility.

Animals↗
PubMed1994

Coital behavior of male Pteropus scapulatus (little red flying foxes) in captivity.

Copulation in Pteropus scapulatus, the little red flying fox (suborder Megachiroptera), is characterised by brief orogenital grooming for 20 s; a single mount lasting 175 s; a single intromission, which includes a mean of 20 s thrusting in most copulations; ejaculation, lasting only about 9 s, observed in about 20% of copulations; and lock et copula, for about 155 s. In 65% of copulations, mating between the same pair resumes, usually after about 35 s. Each pair typically undertakes a series of two-three copulations at a time. Copulation is the dorso-ventral position, while both animals hang by their feet in their normal head-down posture. An unusual feature of their coitus is that each animal supports its own weight, and the female appears undisturbed by the male's thrusting movements. Mating is essentially silent and still. Peak mating occurs in November and December (late spring/early summer), with fertile matings being rare at other times.

Animals↗
PubMed2004

Androgen receptor blockade in the MPOA or VMN: effects on male sociosexual behaviors.

Previously, our laboratory has shown that androgen receptors in the medial preoptic area (MPOA) and ventromedial nucleus (VMN) are necessary for copulation in male rats. The present study examined whether these receptors are required for other sociosexual behaviors. In Experiment 1, different regions of the VMN were implanted with the antiandrogen hydroxyflutamide (OHF). We found that implants located in anterodorsal portions of the VMN were more effective at inhibiting the restoration of copulation than implants in the posteroventral VMN. In Experiment 2, a second set of male rats was pretested for copulation and other sociosexual behaviors and was castrated. Experimental animals then received Silastic capsules filled with testosterone (T) plus intracranial (IC) implants filled with OHF to selectively block androgen receptors in either the MPOA or VMN. We found that androgen receptor blockade in the MPOA inhibited the restoration of copulation but had no effect on other sociosexual behaviors. OHF directed at the VMN inhibited the restoration of copulation and 50-kHz vocalizations but had no effect on scent marking. Two tests were used to assay sexual motivation: partner preference and conditioned place preference (CPP). Both methods revealed impairments in sexual motivation in the VMN group but not in animals receiving OHF in the MPOA. Taken together, these data suggest that androgen receptors in the MPOA are essential for copulatory performance, while androgen receptors in the VMN are important for copulation, sexual motivation, and androgen-dependent vocalizations.

Analysis of Variance↗