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Hyperactivity in hyposexual male rats.

It is widely accepted that some male rats fail to copulate because of a decrease in arousability, measured as decreased general locomotor activity (hypoactivity). This relationship, however, failed to explain an observation made in our laboratory that rats that failed to copulate exhibited increased general locomotor behavior. To directly address this issue, we quantified open-field and male sexual behaviors in 360 rats from two different strains. Twenty-two out of 49 hyposexual males were also hyperactive; this was a significantly greater number than would be expected by chance (p < 0.002, binomial test). Interestingly, only 6 of the 49 hyposexual males were hypoactive; this number was actually significantly smaller than would be expected by chance (p < 0.02). There was no correlation between behavioral measures and plasma levels of testosterone or progesterone. A decrease in selective attention and a failure to be stimulated by amphetamine was apparent in all hyperactive rats--those normal-sexual as well as hyposexual. The hyperactive rats were not hypertensive. We conclude that a significant percentage of hyposexual rats are hyperactive, and that hypoactive rats generally exhibit normal levels of sexual behavior. Decreased selective attention and decreased responsiveness to amphetamine do not explain this result, which is also not related to blood pressure or androgen levels.

Amphetamine↗

Sensitivity to testosterone varies with strain, sex, and site of action in chickens.

Day-old chicks (cockerels and pullets) of two strains of chicken (a commercial breed and strain of feral fowl) were treated intramuscularly with 25 mg/0.1 ml of testosterone oenanthate on day 1 posthatching. Controls received 0.1 ml of the vehicle. Attack and copulatory behavior were scored from days 7 to 14 using standard hand-thrust tests, which rank the responses from 1 to 10. The sizes of the comb and testes were also measured. All measures showed strain and sex differences. Copulation and attack scores were highest in males of the feral strain. Even untreated males of the feral strain had high scores, and these were further elevated by the testosterone treatment. In fact, the attack scores of feral males were marginally higher than those of the males of the commercial strain treated with testosterone. Testosterone treatment of the feral females also elevated attack and copulation to a much greater extent than in the commercial strain. Opposite effects occurred for the development of the comb. Comb volume, absolute and adjusted for body weight, was much greater in treated chicks of the commercial strain than in those of the feral strain. These results indicate that chicks of the feral strain may have more central and fewer peripheral receptors for testosterone or that their receptors for testosterone are more sensitive than those of the commercial strain. Alternatively, there may be strain, as well as sex, differences in the metabolism of testosterone.

Aggression↗

Mating-induced alterations in the regulatory mechanisms of male rats.

The effect of mating behavior on the regulatory mechanisms of male rats was investigated. Moderate levels of copulatory behavior (two ejaculations/week for 6 weeks) reduced body weight gain, and decreased food efficiency. Mating also affected carcass composition: it caused a decrease in fat content and a rise in nitrogen and water levels. The altered regulatory functions were observed in two age groups: 14-17-week- old male rats and 27-week- old male rats. Controls for the non-copulatory components of mating demonstrated the specificity of the copulation effects. The regulatory functions of the control groups (Social-Interaction, Voluntary-Exercise, and Novel-Cage) were similar to those of the sexually rested, Home-Cage group. These results demonstrate that a naturally occurring behavior, mating, significantly affects regulatory mechanisms in the male rat. Since growth and longevity have been shown to be interrelated, these findings support the hypothesis that copulation can affect aging.

Adipose Tissue↗

Opioids and sexual behavior in the male rat.

Naloxone in the doses of 4 or 16 mg/kg failed to affect copulatory behavior of testosterone-treated castrated male rats. Morphine 10 mg/kg, administered 60 min before behavioral observation, reduced the proportion of animals displaying sexual behavior. Doses of 2.5 or 5 mg/kg reduced the latency to the second ejaculation, whereas the few animals still copulating after morphine 10 mg/kg showed a reduced latency to the first ejaculation. The same doses of morphine administered 5 min before behavioral observation produced a dose-dependent reduction of mount, intromission and ejaculation percentages. However, those animals that did copulate showed a normal copulatory behavior. D-Ala2-Met5 enkephalinamide (DALA) infused into the left cerebral ventricle in a dose of 5 micrograms 5 or 60 min before tests had no effect. When the peptide was infused 30 sec after the first intromission, the number of intromissions as well as the latency to ejaculation were reduced. Opioids may facilitate ejaculatory mechanisms, perhaps as a consequence of their rewarding properties. Moreover, in animals treated with DALA after the first intromission, the number of intromissions and the latency to ejaculation were similar for the first and second copulatory series, while these parameters were much reduced upon the second ejaculation for control animals. It is possible that liberation of endogenous opioids is the cause of ejaculation-induced facilitation of subsequent sexual behavior.

Animals↗

Relation of spontaneous wet dog shakes and copulatory behavior in male rats.

Pharmacological manipulation of 5-HT2 activity has yielded equivocal effects on male rat sexual behavior. Both facilitation and inhibition of copulation have been reported following treatment with 5-HT2 antagonists. "Wet dog shake" (WDS), a component of the Serotonin Behavioral Syndrome, is largely mediated by 5-HT2 receptors. The present series of experiments were aimed at determining whether WDS might yield a spontaneous behavioral measure of 5-HT2 activity that converges on the pharmacological data. In Experiment 1, spontaneous WDS was recorded in 58 male rats paired with receptive females. Normal copulators (Studs) exhibited significantly less WDS than did noncopulators (Duds). In Experiment 2, the males were additionally paired with males and nonreceptive females. In these situations, WDS did not discriminate Duds from Studs, but Studs scores differed from each other across the three different partner conditions. Lastly, in Experiment 3, treatment with the selective 5-HT2 agonist DOI potently inhibited copulatory responses and increased WDS. Overall, the data from the three experiments suggest that 5-HT2 activity mediates an inhibition of male rat sexual behavior.

Amphetamines↗

Dopaminergic drugs in the medial preoptic area and nucleus accumbens: effects on motor activity, sexual motivation, and sexual performance.

In two experiments, dopamine agonists and/or antagonists were injected into the medial preoptic area (MPOA) or the nucleus accumbens (NAcc) of male rats. The animals were then tested in an X-mase with four goal boxes, which contained a receptive female, a male, or were empty. In Experiment 1, the D1 antagonist SCH-23390 and the D2 antagonist raclopride in the MPOA decreased the percentage of trials on which the female's chamber was chosen, a measure of sexual motivation. Raclopride also decreased the number of animals that copulated after choosing the female's chamber. The 10-micrograms dose of the D3/D2 agonist quinelorane increased the latency to reach the female's chamber, slowed the onset of copulation, and decreased the number of intromissions preceding an ejaculation. In Experiment 2, 1- and 5-micrograms doses of quinelorane and of the mixed D1/D2 agonist apomorphine were injected bilaterally into the NAcc. Both doses of quinelorane increased the number of times that the subject did not select a chamber within 60 s. No drug in the NAcc affected specifically sexual motivation or performance. The results are consistent with differential influence of the MPOA and the NAcc on motor activity, sexual motivation, and sexual performance.

Animals↗

Control of ovulation, vaginal estrus, and behavioral receptivity in voles (Microtus).

Rodents of the genus Microtus provide an excellent focus for the study of the effects of environmental and behavioral influences on reproductive physiology. Despite some early reports, there is little definitive evidence of truly spontaneous ovulation in Microtus. Stimuli from copulation with males appear capable of triggering ovulation in all species: the requisite amount of stimulation for ovulation and a functional luteal phase varies with the species. In addition, however, a certain percentage of the females of some, but not all, species ovulate when placed in proximity to males when copulation is presented. There are few reports of regular cycles of cell types in the vaginal smears of Microtus females; the relative preponderance of cornified, nucleated, and leukocytic cells varies with species and conditions. Behavioral receptivity is not invariably correlated with smear type. The complexity of environment-behavior relationships in different species suggests the need for a more comprehensive classification schema for systems of female reproductive physiology and a study of their adaptive significance.

Animals↗

"Central arousal" and sexual responsiveness in the snail, Helix aspersa.

In molluscs, a "central arousal" system is thought to positively modulate both an animal's level of activity and its behavioral responsiveness. This hypothesis is examined in Helix aspersa by testing the relationships between activity, feeding, and sexual behavior. Activity, feeding, and mating exhibit parallel daily rhythms. Snails are most active, and eat and mate most frequently, during scotophase and the first 3 h of photophase. Handling and injections of serotonin (5 x 10(-7) m/kg body wt) increase general activity. Inducing activity during late photophase increases food consumption, but it does not induce sexual activity. Moreover, serotonin and handling have no effect on sexual arousal; treated snails show no increase in genital eversion stage and require the same length of courtship as controls. If snails are sexually deprived, increasing activity in late photophase does increase copulation frequency, but snails do not copulate more frequently than do unhandled snails tested during early photophase. These results suggest that while manipulations thought to increase central arousal do increase activity and responsiveness to food stimuli, they do not directly affect sexual proclivity or sexual arousal. Moreover, the relationship between feeding and sexual behavior differs between Helix and Aplysia. This difference between the two species may be related to the differences in their reproductive strategies.

Animals↗

Brain monoaminergic control of male reproductive behavior. II. Dopamine and the post-ejaculatory refractory period.

This study was designed to examine the role of central dopaminergic mechanisms in the control of copulation and the subsequent post-ejaculatory refractory period in the male rat. Disruption of central dopaminergic pathways was achieved in two separate groups of animals by: (1) selective electrolytic lesions of the substantia nigra (the major locus for dopamine cell bodies in the brain); or (2) localized intracerebral injection of 6-hydroxydopamine, a specific neurotoxin for catecholaminergic pathways. A third group of animals was tested for sexual behavior following administration of the dopamine receptor blocker pimozide. Both electrolytic and neurochemical lesions localized in the substantia nigra produced a significant increase in the length of the post-ejaculatory refractory period. Dopamine receptor blockade following administration of pimozide also caused a significant increase in refractory period duration. These results support the hypothesis that central dopaminergic pathways are involved in the motivational or arousal component of copulation and may be integral to the maintenance of a normal post-ejaculatory refractory period.

Amygdala↗

Light input and the reversal of functional lateralization in the chicken brain.

During its later stages of development, the chicken embryo is oriented in the egg so that it occludes its left eye with its body and the right eye is positioned so that it can receive light input. This lateralized light input has been shown to play a decisive role in determining the direction of brain lateralization for two behavioural functions, copulation and performance of a visual discrimination task known as the 'pebble floor test', since the direction of lateralization for these functions can be reversed by occluding the right eye of the embryo on day 19/20 of incubation and at the same time exposing the left eye to light. The sensitive period during which this role of lateralized light input influences the lateralization for copulation extends to day 1 posthatching if the eggs are incubated and hatched in darkness, but it is over by day 3. For the pebble floor test the sensitive period is already over by day 1 posthatching. By exposing eggs to light for various times on day 19 of incubation, it was possible to determine that between 2.5 and 6 h of lateralized light exposure is necessary to stabilise the normal direction of lateralization so that it can no longer be reversed by occlusion of the right eye. Thus, in the developing chicken embryo substantial neural reorganization must occur in response to a brief period of lateralized light input.

Animals↗

Parental care and sexual interactions in Mongolian gerbils (Meriones unguiculatus) during the postpartum estrus.

The behavior of male and female Mongolian gerbils was continuously video-recorded from 24 h before the parturition to the end of postpartum estrus. We noted that the mean interval between the delivery of the last pup and the first mounting was 13 h and 32 min. Over the whole duration of the postpartum estrus (7 h and 41+/-57 min), females spent significantly more time in crouching over, pup licking, nest building and "digging" activities, and were more recipient of allogrooming than males. Maternal pup retrievals were not very frequent (14.0+/-5.6 episodes), but males never retrieved pups nor exhibited a full sequence of nest building activity. Males spent a longer mean time in bodily interactions with females, as compared to the time they spent with pups, and engaged in intense copulatory activity (592.8+/-40.5 mounts or 1 episode per 46.7s); even during the delivery process males attempted to force copulation, but females rejected mounting in all cases. We conclude that females exhibited higher levels of parental care than males, and our findings suggest that males compete with pups in attracting the female's attention, since they actively disrupt maternal care or persistently persecute the female in order to copulate.

Animals↗

Function of multiple sperm storage organs in female Mediterranean fruit flies (Ceratitis capitata, Diptera: Tephritidae).

Sperm storage organs allow females to temporally separate insemination from fertilization, manipulate ejaculates and control fertilization. In the reproductive tract of female fruit flies (Diptera: Tephritidae), sperm are found in two different organs--a pair or triplet of spermathecae, and a "fertilization chamber". In order to understand the specific function of each of these organs, we tested the following hypotheses: (1) Sperm are distributed equally amongst the various sperm storage organs; (2) Both organ types maintain sperm viability; and (3) Sperm used in fertilization come from the fertilization chamber. We counted sperm in spermathecae and fertilization chamber of Mediterranean fruit flies (Ceratitis capitata) every 3 days for 18 days following insemination, and used a live/dead staining technique to determine the viability of sperm in these organs. Finally, by extirpating spermathecae from inseminated females and allowing them to oviposit, we were able to identify the fertilization chamber as the source of fertilizing sperm. Numbers of sperm in the spermathecae declined from an average of 3575 on the day of copulation to 649, 18 days later. Conversely, the fertilization chamber maintained a fairly constant level of sperms, ranging between an average of 207 cells on day 3 to 115 sperms on day 18. Throughout the period we monitored, we found high levels of sperm viability in both organs (> 80%). Sperm viability was similarly high in the fertilization chambers of females without spermathecae. However, fertility of eggs laid by these females declined rapidly, as did the number of sperm in the fertilization chamber. We conclude that both the spermathecae and the fertilization chamber are active sperm storage organs, with separate functions: the spermathecae for long-term storage and the fertilization chamber, periodically filled by the spermathecae, a staging point for fertilizing sperm. We suggest that the use of both organs by females results in sperm economy, which adaptively prolongs the intervals between copulations.

Animals↗

Remating inhibition in female Queensland fruit flies: effects and correlates of sperm storage.

Reproductive success of male insects commonly hinges both on their ability to secure copulations with many mates and also on their ability to inseminate and inhibit subsequent sexual receptivity of their mates to rival males. We here present the first investigation of sperm storage in Queensland fruit flies (Tephritidae: Bactrocera tryoni; a.k.a. 'Q-flies') and address the question of whether remating inhibition in females is directly influenced by or correlated with number of sperm stored from their first mates. We used irradiation to disrupt spermatogenesis and thereby experimentally reduce the number of sperm stored by some male's mates while leaving other aspects of male sexual performance (mating probability, latency until copulating, copula duration) unaffected. Females that mated with irradiated rather than normal males were less likely to store any sperm at all (50% vs. 89%) and, if some sperm were stored, the number was greatly reduced (median 11 vs. 120). Despite the considerable differences in sperm storage, females mated by normal males and irradiated males were similarly likely to remate at the next opportunity, indicating (1) number of sperm stored does not directly drive female remating inhibition and (2) factors actually responsible for remating inhibition are similarly expressed in normal and irradiated males. While overall levels of remating were similar for mates of normal and irradiated males, factors responsible for female remating inhibition were positively associated with presence and number of sperm stored by mates of normal but not irradiated males. We suggest seminal fluids as the most likely factor responsible for remating inhibition in female Q-flies, as these are likely to be transported in proportion to number of sperm in normal males, be uninfluenced by irradiation, and be transported without systematic relation to sperm number in irradiated males.

Animals↗

Endocrine and behavioral observations during transition of non-breeding into breeding season in female American bison (Bison bison).

This study provides endocrine data in relation to behavioral events during the transition of the non-breeding into the breeding season in American bison (Bison bison). Fecal progesterone metabolite patterns (20-oxo-P) were obtained in 13 adult female American bison and hormonal data were correlated with behavioral observations; i.e. copulation, male tending, female tail-up behavior and gestation length. Based on fecal progesterone metabolite patterns, the breeding season started between the middle of July and early August. Predictable short cycles reflected the transition from non-breeding to the breeding season; the luteal phase of these cycles was 4.10+/-0.86 days. Copulations and female tail-up behavior were reliably associated with the hormonally detected ovulation. Male tending behavior was more loosely associated with hormonally detected ovulation. The observed hormonal pattern in the study females indicated that 9 of 10 pregnant cows conceived during the second ovulatory period in the breeding season. One other cow conceived during her third ovulatory period, and one cow did not conceive until later in the breeding season by beginning of October. Gestation duration was on average 266.30+/-1.00 days. In summary, this study confirmed that the bison is a seasonally polyestrous species; the transition from the non-breeding into the breeding season was characterized by short cycles with low progesterone metabolite values.

Animals↗

Rapid inhibition of female sexual behavior by gonadotropin-inhibitory hormone (GnIH).

Gonadotropin-releasing hormone (GnRH) is largely responsible for the initiation of sexual behaviors; one form of GnRH activates a physiological cascade causing gonadal growth and gonadal steroid feedback to the brain, and another form is thought to act as a neurotransmitter to enhance sexual receptivity. In contrast to GnRH, gonadotropin-inhibitory hormone (GnIH) inhibits gonadotropin release. The distribution of GnIH in the avian brain suggests that it has not only hypophysiotropic actions but also unknown behavioral actions. GnIH fibers are present in the median eminence (ME) and are in apparent contact with chicken GnRH (cGnRH)-I and -II neurons and fibers. In birds, cGnRH-I regulates pituitary gonadotropin release, whereas cGnRH-II enhances copulation solicitation in estradiol-primed females exposed to male song. In the present study, we determined the effects of GnIH administered centrally to female white-crowned sparrows. A physiological dose of GnIH reduced circulating LH and inhibited copulation solicitation, without affecting locomotor activity. Using rhodaminated GnIH, putative GnIH binding sites were seen in the ME close to GnRH-I fiber terminals and in the midbrain on or close to GnRH-II neurons. These data demonstrate direct effects of GnIH upon reproductive physiology and behavior, possibly via separate actions on two forms of GnRH.

Animals↗

Intersexuality and behavior in crayfish: the de-masculinization effects of androgenic gland ablation.

In crustaceans, male differentiation and primary and secondary characteristics are regulated by the androgenic gland (AG). In gonochoristic crustaceans, the AG is also linked to intersexuality. Whereas the co-occurrence of various male and female characteristics has been demonstrated in intersex crustaceans, little is known regarding sexually dimorphic behavior patterns in such individuals. In the present study, we used an intersex crayfish model to investigate--for the first time in crustaceans--the agonistic and mating behavior of intersex individuals, and to explore the effects of AG ablation on behavior, morphology and physiology. As was the case for their morphological and physiological reproductive traits, intersex individuals--despite being genotypically females--generally resembled males in terms of behavior: they engaged in fighting with males and copulated with receptive females. However, fighting durations of intersex animals were intermediate between those of males and females, and the durations of the copulations were remarkably short. Adult intersex individuals that had been AG ablated at the juvenile stage were unlikely to engage in fighting with males (similar behavior to females) and did not exhibit any mating behavior with receptive females. AG ablation resulted in feminine morphological and physiological shifts in the treated intersex individuals and enabled vitellogenin gene transcription and the onset of secondary vitellogenesis. It thus appears that an as-yet-unknown AG-secreted factor(s) regulating maleness also seems to regulate the organization of male behavior in crustaceans.

Agonistic Behavior↗

Post-conception mating in wild long-tailed macaques (Macaca fascicularis): characterization, endocrine correlates and functional significance.

In many anthropoid primates, mating activity is not restricted to the ovarian cycle but also occurs during pregnancy. Although it has been suggested that the main function of this post-conception mating is to confuse paternity, studies showing whether or not male primates can distinguish between the fertile phase of the conception cycle (FPCC) and the period of peak post-conception mating (peak PCM) are almost non-existent. Here, we examine whether the pattern of female sexual traits (specific sexual behaviors, sexual swelling) and female attractiveness to males differ between FPCC and peak PCM in 6 wild female long-tailed macaques. We also use fecal hormone analysis to investigate whether female traits during peak PCM are related to changes in female sex hormones. All females exhibited a distinct period of heightened mating activity around days 45-60 of gestation. During peak PCM, swelling size and frequency of female solicitations (but not reaching back) were significantly correlated with changes in the estrogen to progestogen ratio. Swelling size, frequency of female sexual behaviors and copulations and proportion of male-initiated copulations and ejaculations were not significantly different between FPCC and peak PCM. Although males spent significantly less time consorting females during peak PCM, all (particularly low-ranking and non-resident males) invested heavily in terms of reproductive costs associated with mate-guarding and mating during pregnancy. We conclude that post-conception mating in wild long-tailed macaques is not merely a by-product of endocrine changes and devoid of adaptive function. Our results more strongly support the hypothesis that it may form part of a female reproductive strategy to confuse paternity, which appears to apply particularly to low-ranking and extra-group males.

Animals↗