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Brain localization of cholinergic influence on male sex behavior in rats: antagonists.

The muscarinic receptor antagonist scopolamine was microinjected into either the preoptic area or the lateral ventricle, preceding sexual behavior tests. In Experiment 1 unilateral ventricular injections of scopolamine delayed the initiation of copulation, while unilateral preoptic injections had no effect. In Experiment 2 bilateral injections into the preoptic area produced dose-related decreases in the percentages of animals intromitting and ejaculating. In Experiment 3 scopolamine, injected alone into the preoptic area, again decreased the percentages of animals mounting, intromitting, and ejaculating. The muscarinic agonist oxotremorine, injected alone into the preoptic area, decreased ejaculatory threshold (i.e., decreased the number of intromissions preceding ejaculation) as previously reported. Concurrent oxotremorine and scopolamine injections into the preoptic area were not different from vehicle; thus, scopolamine blocked oxotremorine's effect. These data suggest that some cholinergic activation of the preoptic area is critical for normal copulation, since bilateral blockade of muscarinic receptors there dramatically decreased the number of animals copulating. However, increased cholinergic activity there only reduced ejaculation threshold.

Animals↗

Contingent tolerance to the disruptive effects of alcohol on the copulatory behavior of male rats.

Sexually active male rats received five 30-min copulation tests with sexually receptive females, one every 4 days. One group of rats received alcohol (1 g/kg, IP) 45 min before, and an equivalent volume of saline 45 min after, each test; a second group received saline before and alcohol after each test; and a third, control group received saline both before and after. Four days after the last of the five tolerance-development trials, each rat received an injection of alcohol (1 g/kg, IP) 45 min before a copulation test so that the development of tolerance in the three groups could be compared. Tolerance to the disruptive effects of alcohol on mount, intromission, and ejaculation latencies, and on the duration of the postejaculatory interval was found to be significantly greater in the rats injected with alcohol before each copulation test than it was in the rats in the other two groups. These results constitute the first experimental evidence that tolerance develops to the disruptive effects of alcohol on male sexual behavior, and they support the theory that tolerance is an adaptive response to the disruptive effects of drugs on concurrent patterns of neural activity, rather than to drug exposure per se.

Animals↗

Light experience during development affects asymmetry of forebrain function in chickens.

The chicken brain has asymmetry of forebrain function of attack and copulation behaviors. This study reports that this asymmetry, revealed by intracranial injections of glutamate, is affected by light exposure. Eggs were incubated in 3 forced-draught incubators matched for noise, humidity and temperature control, so that light and laboratory noise were the only systematic variables. All eggs were incubated in darkness for the first 17 days of incubation after which they were allocated to one of 3 incubating conditions, L+S+, L-S+, L+S-, where + or - indicates the presence or absence of light (L) or laboratory noise (S). Light exposure was necessary to establish lateralization of attack and copulation, sound was not since only the L+S+ and L+S- groups were lateralized. The lateralization of auditory habituation was not affected by these incubating conditions. Further experiments established that on day 19 of incubation exposure for 4 h to a light of constant intensity produced lateralization. 1 h exposure to a light of constant intensity or to a flickering light (1.7 Hz) failed to produce lateralization. Examination of the data for individuals suggests that light exposure synchronizes the direction of lateralization rather than generates its presence. Environmental and genetic factors interact to produce the lateralization of attack and copulation.

Acoustic Stimulation↗

An animal model of fetishism.

An animal model of sexual fetishism was developed with male Japanese quail based on persistence of conditioned sexual responding during extinction to an inanimate object made of terrycloth (Experiments 1 and 3). This persistent responding occurred only in subjects that came to copulate with the terrycloth object, suggesting that the copulatory behavior served to maintain the fetishistic behavior. Sexual conditioning was carried out by pairing a conditioned stimulus (CS) with the opportunity to copulate with a female (the unconditioned stimulus or US). Copulation with the CS object and persistent responding did not develop if the CS was a light (Experiment 1) or if conditioning was carried out with a food US (Experiment 2). In addition, subjects that showed persistence in responding to the terrycloth CS did not persist in their responding to a light CS (Experiment 3). The results are consistent with the hypothesis that conditioned copulatory behavior creates a form of self-maintenance that leads to persistent responding to an inanimate object. The development of an animal model of such fetishistic behavior should facilitate experimental analysis of the phenomenon.

Animals↗

Sperm allocation and cost of mating in a tropical tephritid fruit fly.

Males that copulate repeatedly may suffer from reduced sperm stores. However, few studies have addressed sperm depletion from both the female and male perspective. Here, we show that male Anastrepha obliqua (Diptera: Tephritidae) do not ejaculate all available sperm and are left with mature sperm in the seminal vesicles even after copulating as often as three times in half a day. Ejaculate size was not related to male mating history; time elapsed since the last mating, copulation duration, female thorax length or head width. Larval host origin did not affect the number of sperm stored by females. More sperm was found in the ventral receptacle compared to sperm stored in the three spermathecae. Males apparently do not suffer a cost of mating in terms of longevity, although we cannot rule out other fitness costs. Sperm production in this species may not be as costly as it is for other species. Results suggest that males strategically allocate similar numbers of sperm among successive mates without exhausting sperm reserves for future encounters. We discuss the role that differential sperm storage may have in mediating sperm competition and tie our results to the unique natural history of A. obliqua.

Animals↗

Seminal fluids mediate sexual inhibition and short copula duration in mated female Queensland fruit flies.

Molecules in male seminal fluid transferred to female insects during mating can have potent effects on their subsequent sexual and reproductive behaviour. Like many other tephritids, female Queensland fruit flies (Bactrocera tryoni) typically have diminished sexual receptivity after their first mating. Also, copulations of females that do remate tend to be shorter than those of virgins. We here find that virgin females injected with small doses (0.1, 0.2 or 0.5 male equivalents) of extracts from the male reproductive tract accessory tissues, which consist of male accessory glands, ejaculatory apodeme and ejaculatory duct (AG/EA/ED), have diminished receptivity and short copula duration very similar to naturally mated females. In contrast, virgin females injected with saline or with high doses of AG/EA/ED (1 or 2 male equivalents) that likely exceed the range of natural variation retain the higher levels of sexual receptivity and longer copulations of un-injected virgins. We conclude that reduced sexual receptivity and shorter copulations of mated female Q-flies are mediated by products in the male seminal fluid derived from the male reproductive tract accessory tissues.

Animals↗

The perirhinal-entorhinal cortex, but not the hippocampus, is critical for expression of individual recognition in the context of the Coolidge effect.

The Coolidge effect is a phenomenon in which males show renewed sexual interest in a novel female following copulation to satiety with another female. In golden hamsters, this phenomenon depends on the ability to recognize conspecifics using chemosensory cues processed through the main olfactory system. Here we tested whether olfactory targets in the hippocampal system support this natural form of recognition memory. Male hamsters received ibotenic acid lesions of the perirhinal-entorhinal cortex (PR-ENT) or hippocampus (H) and were allowed to copulate to satiety with a female conspecific, then were presented with two anesthetized females, the familiar mate and an unfamiliar female that copulated with another male. Sham-operated and H-lesioned subjects preferentially investigated the novel female, indicating intact recognition of individual identity. By contrast, PR-ENT-lesioned males failed to discriminate familiar and novel females, and this deficit could not be attributed to abnormal copulatory behavior during mating. All subjects were able to detect and discriminate between female odors when presented in isolation during a habituation-discrimination test, indicating that behavioral deficits shown by PR-ENT males were not due to anosmia or a general investigatory deficit. Thus, the perirhinal-entorhinal cortex, but not the hippocampus, is critical for the recognition of familiar conspecifics in this naturalistic situation. This study reveals an essential role for the perirhinal-entorhinal cortex, but not the hippocampus, in a natural form of recognition memory within the social behavior of hamsters. The findings show a strikingly similar pattern to the effects of selective damage to the same brain regions on performance in standard recognition memory tasks by rats and monkeys. Therefore, the present data extend our understanding of the differential role of structures of the hippocampal memory system, showing continuity across species and between formal laboratory tests and the function of memory in natural social behavior.

Animals↗

Repeated interactions with females elevate metabolic capacity in the limbic system of male rats.

The effect of heterosexual social experience on brain metabolic capacity was investigated by measuring the activity of cytochrome oxidase, a rate-limiting enzyme in oxidative metabolism. Male Sprague-Dawley rats were kept naïve or allowed to copulate with receptive females three (3 F males) or 16 times (16 F males). Throughout the vomeronasal system and other limbic areas, 16 F males had elevated metabolic capacity relative to naïve and 3 F males, whereas no significant differences in brain metabolism were found between 3 F and naïve males. Behavioral differences were also found between 3 F and 16 F males. In a second experiment, we assessed differences in brain metabolism between sexually active and inactive males given only one opportunity to copulate and found no significant difference in neural metabolism between these males. This suggests that the differences found in the first experiment were primarily driven by differences in repeated experience rather than by sexual performance between 16 F and 3 F males. We speculate that these changes in brain metabolic capacity could be related to immediate early gene expression during copulation and could underlie the long-term behavioral changes accompanying heterosexual social experience.

Animals↗

Regulation of sexual receptivity of female Mediterranean fruit flies: old hypotheses revisited and a new synthesis proposed.

The objective of this study was to examine the relative contributions of copula duration and sperm transfer to the inhibition of sexual receptivity of female Mediterranean fruit flies (Ceratitis capitata, Diptera: Tephritidae). Females choosing to remate had significantly fewer sperm in their spermathecae than females who chose not to remate. Duration of a female's first copulation did not affect her subsequent receptivity. Furthermore, on the first day following copulation significantly more females whose first mate was sterile and from a laboratory strain (sterile males transfer fewer sperm than wild males) chose to copulate than did females whose mate was fertile and recently derived from wild stock. Finally, we offer a synthesis of the available information on remating in this species, and suggest that while females are facultatively polyandrous, copula duration, sperm transfer and male accessory gland secretions act in succession to inhibit female receptivity.

Animals↗

The development of olfactory conditioned ejaculatory preferences in the male rat. I. Nature of the unconditioned stimulus.

We have demonstrated previously that repeated pairing of a neutral odor with copulation produces a subsequent conditioned ejaculatory preference (CEP) for a female bearing that odor. Here we examine the copulatory components that comprise the unconditioned stimulus (UCS). In Experiment 1, male Long-Evans rats were allowed to copulate with scented females for nine sessions in which they achieved two ejaculations, one ejaculation plus the first intromission following the postejaculatory interval (PEI), one ejaculation without a PEI, or five intromissions without ejaculation. Only the males that achieved two ejaculations or one ejaculation plus the PEI displayed significant CEP. In Experiment 2, males were allowed to remain in the presence of the scented female without access to her after different amounts of copulatory stimulation. Under these conditions, both one and two ejaculations, but not five intromissions, supported the development of CEPs. In Experiment 3, males were allowed to copulate to ejaculation with an unscented female followed by exposure without access to a scented female. This treatment also supported the development of CEP. These results indicate that ejaculation plus a PEI are necessary for the development of CEPs and that the female must be present during the PEI for this to occur. These findings indicate that events during the PEI are the critical components of the UCS for CEP development.

Animals↗

Hypothalamic and olfactory control of sexual behavior and partner preference in male rats.

Sexually active male rats prefer a sexually receptive female to a nonreceptive female, and partner-preference tests provide one way of studying sexual motivation. Surgical deafferentation of the olfactory bulbs from all the known chemosensory systems of the nasal septum renders rats anosmic. In Experiment 1, we show that, although bulb deafferentation of male rats decreases some aspects of sexual performance, most deafferented males copulate and partner preference is not affected. In Experiment 2, we show that large excitotoxin lesions of the preoptic hypothalamus eliminate copulation, an effect that is correlated with damage to the anterior portions of the medial and lateral preoptic area. Lesions also decrease partner preference, an effect that is correlated (r = 0.82) with damage to a small part of the bed nucleus of the stria terminalis. Most males who do not copulate after hypothalamic lesions show a persistent, albeit reduced, preference for receptive females over nonreceptive females. This preference appears to depend on olfactory ability because bulb deafferentation of lesioned, noncopulating males virtually eliminates partner preference.

Animals↗

Mounting and brief noncontact exposure of males to receptive females facilitate reflexive erection in rats, even after hypogastric nerve section.

In three experiments, reflexive erection in male rats was facilitated by housing the males for 2 min with inaccessible sexually receptive females. In Experiment 1, males were sexually naive or experienced and received two reflexive erection tests, 1 week apart, immediately after the males were exposed to receptive females, to unreceptive females, or to no females (n = 8 per group). In both tests, experienced males exposed to estrous females had the shortest reflexive erection latencies; in Test 1 the differences among groups were of borderline significance (p = 0.057), but in Test 2 the differences among groups were highly reliable (p<0.01). Further analysis indicated that only experienced males exposed to receptive females were significantly different from other groups. In Experiment 2, sexually experienced males (n = 11) received four reflexive erection tests: after being with no female, and 0, 5, or 10 min after exposure to estrous females. As the interval between exposure and test increased, the males had progressively shorter erection latencies (p<0.01) and more intense glans erections (p<0.03). Experiments 1 and 2 may be viewed as demonstrating the psychogenic facilitation of reflexive erections. In Experiment 3, males underwent sham surgery (sham, n = 10) or bilateral transection of the hypogastric nerves (HgNx, n = 10), which are conventionally viewed as mediating psychogenic erection. After males mounted a receptive female for 5 min without intromission or had 2 min of non-contact exposure to receptive females, the males had shorter erection latencies (p<0.001) and more erections (p<0.02). These facilitative effects of pretest stimulation were unaffected by HgN transection. During copulation tests, HgNx males had longer ejaculation latencies (p<0.05) and lower intromission ratios (p<0.05), possible signs of impaired erectile function. However, in Experiment 4, other males were tested twice for reflexive erection and copulation after sham (n = 8) or HgNx (n = 9) surgery, and there were no significant effects of surgery on reflexive erection or copulatory behavior. Collectively, these experiments indicate (a) that brief noncontact exposure of sexually experienced males to estrous females facilitates reflexive erection, (b) that this facilitation increases for at least 10 min after the females are removed, and (c) that the hypogastric nerves do not mediate these facilitative effects. The evidence for a role for the HgN in copulation was inconclusive.

Animals↗

Peripheral nerves mediating penile erection in the rat.

We examined the effects in male rats of bilateral transection of two nerves previously implicated in erectile function, the viscerocutaneous branch of the pelvic nerve (Vc) and the hypogastric nerve (HgN). In Experiment 1 (conducted in Storrs), males underwent simultaneous or successive section of Vc and HgN and were tested for copulation, reflexive erection, and noncontact erection (NCE), i.e. in response to remote cues from estrous females. NCE is considered to be analogous to 'psychogenic' erection in humans, for which the HgN has been ascribed a significant role. In all three types of test, males had a moderate to severe deficit in erectile function after Vc transection. Section of HgN alone had no apparent pro- or anti-erectile effect in any context, nor did it affect the decrement resulting from Vc surgery. Regardless of treatment, all groups retained some erectile potential in each type of test. The loss of bladder function after Vc surgery and of seminal plug deposition after HgN section gave evidence that the targeted nerves were in fact severed. In Experiment 2 (conducted in Xalapa), males were tested only for NCE, but (a) they were tested every 3 days beginning 3 days after each surgery, (b) the interval between the two surgeries was more than 2 weeks, rather than 1 week as in Experiment 1, to allow more time for recovery from general effects of surgery and for hypothetical plasticity of neural function. In the first test after the first surgery, all groups had a modest reduction in the proportion of males displaying NCE, relative to sham-operated males. However, this deficit did not extend to measures of NCE latency or number, and was absent after the second test. After the second surgery, when all males except those with sham operations had both nerves cut, none of the groups exhibited a significant deficit in NCE, and all groups had at least one test in which at least half the males responded. Thus, (a) HgN section did not significantly impair NCE, reflexive erection, or copulation; (b) Vc section impaired, but did not eliminate, erection in all three contexts, but even those effects may be transient; and (c) transection of both nerves, simultaneously or successively, did not cause a greater impairment in erection than did cutting just the Vc. We infer that the HgN may have no pro-erectile role in erection in rats, even in a model analogous to psychogenic erection. The Vc is probably the most important nerve mediating pro-erectile function in NCE, as in reflexive erection and copulation, but this nerve may not be essential for erection in rats in any context, at least in some males.

Animals↗

The nature of the conditioned response mediating olfactory conditioned ejaculatory preference in the male rat.

We have developed a model to study the influence of conditioning on sexual partner preference in the rat. In this model, pairing a neutral odor (almond) with copulation to ejaculation produces a subsequent preference to ejaculate with females bearing that odor. We refer to this phenomenon as a conditioned ejaculatory preference (CEP). The present study investigates the nature of the conditioned response that mediates CEP. Given equal mount and intromission distributions, but an unequal ejaculation distribution, we hypothesized that two mechanisms could account for CEP: facilitated ejaculation or selective ejaculation. To test these hypotheses, we examined the effect of omitting the olfactory conditioned stimulus (CS) or applying it to a nonreceptive female. Males trained with the CS paired with copulation (paired-trained males) failed to display evidence of delayed ejaculation when copulating with unscented females. Conversely, paired-trained males displayed CS-elicited copulatory behavior with CS-bearing nonreceptive females. In addition, we re-analyzed the data from the copulatory preference tests of previous experiments for CEP-displaying males. Again, we failed to find evidence of facilitated ejaculation with the scented female. However, the time between the last mount or intromission and ejaculation was increased if either occurred with the scented female. Furthermore, more mounts were directed towards the scented female near the end, but not at other points, of an ejaculatory series. These findings suggest that the paired-trained males attend to the scented female near the point of ejaculation, and are consistent with the hypothesis that CEP is mediated by selective ejaculation.

Animals↗

Differential contributions of the two visual pathways to functional lateralization in chicks.

The contribution of the two visual pathways to lateralization of visual behaviour in chicks was assessed using unilateral injections of 0.5 microliters of 100 mM monosodium glutamate into localized regions of the forebrain. Chicks treated with glutamate in the left visual hyperstriatum made more errors in a visual discrimination task (pebble-floor test) than did chicks treated in the right visual hyperstriatum. Glutamate injection into the left visual hyperstriatum also elevated attack and copulation scores, but this did not occur following injection of the right visual hyperstriatum. The performance of chicks treated in the right visual hyperstriatum did not differ from that of sham-operated controls. Thus, only the left visual hyperstriatum is involved in the control of these three visually guided behaviours. By contrast, glutamate injections of the left ectostriatum affected only the attack behavior and not performance in the pebble-floor test or copulation responses. Glutamate treatment of the right ectostriatum had no affect on any of the behaviours tested and this was also the case for glutamate treatment of both the left and right neostriata. Although injecting glutamate in a larger volume that allows glutamate to spread over a wide area of the left hemisphere is known to retard auditory habituation, localized injection of glutamate in the areas chosen for this study had no effect on auditory habituation. The results suggest that the tectofugal and thalamofugal pathways have different roles in the lateralization of visual functions. The forebrain region which receives input from the thalamofugal visual system has a lateralized role in categorising pebbles as different from food grains, and also a role in controlling attack and copulation responses. The forebrain region which receives input from the tectofugal visual system is involved in the control of attack responses only.

Agonistic Behavior↗

Semen collection, cryopreservation and artificial insemination in the dromedary camel.

Collection of semen with a bovine artificial vagina (AV) was attempted with each of 14 camels over a period of 2 years. Semen samples were evaluated, extended and cryopreserved. Frozen thawed semen, diluted cooled semen or whole semen was used to inseminate some female camels which were induced to ovulate with hCG. Males ejaculated semen into the AV in 74.6% collection attempts. The male copulated for at least 200s in 62.9% attempts. The remaining copulations were of shorter duration. Similarly, 49.3% ejaculates were at least 3ml of semen. Libido and donation of semen improved from December onwards and reached a peak after mid January with peak performance persisting until April. It declined during May. The majority of camels had lost libido and refuse to donate semen by the end of May. Camel semen is in gel form. While 35.9% of 203 semen samples exhibited no individual sperm motility, 28.5% exhibited low to fair grade individual sperm motility and only 35.4% exhibited >50% sperm motility. Differences existed between animals (P<0.01) and months (P<0.05) of collection, while effect of copulation time was not significant. Mass motility was not observed in camel semen. Individual sperm motility develops after liquefaction of semen. Addition of caffeine but not chymotrypsin improved the individual motility. The mean live percent sperm count and normal acrosome were 73.3+/-1.0 and 92.0+/-0.5, respectively. Only 51.1% of 45 semen samples with pre-freeze motility of >50% and 25% of 16 semen samples from low pre-freeze motility group with an overall success of 44.2% of 61 semen samples were successfully preserved. Wide variation was observed in the freezability of semen from different males. Attempts to impregnate female camels with liquid semen, frozen thawed semen and whole semen after hCG induced ovulation resulted in 0/10, 1/13 and 4/10 pregnancies.

Acrosome↗

Sexual behavior of mares.

Behavior during the estrous phase of the ovulatory cycle of the mare is analogous in most ways to that of estrous females of other species. Proceptive behaviors bring the mare into the proximity of the male and attract his attention. Positioning facilitates mounting, intromission, and ejaculation. Estrous signs appear to be more intense in the few days prior to ovulation than during the transition periods that separate the recurring estrous and diestrous phases. Sexual behavior is absent during diestrus. Detection of estrus in mares is problematic in that it requires the presence (or at least facsimile acoustic or tactile stimuli) or a stallion. Unexplained conditions such as silent or subestrus make reliable determinations even more difficult. The choice of methods for distinguishing estrous from nonestrous mares depends, among other factors, upon characteristics of individual mares, as well as the number of mares to be evaluated. Detection of true estrus--that is, sexual behavior that is associated with follicular growth and ovulation--is confounded by the occurrence of estrus-like behavior independent of gonadal stimulation. Seasonally anovulatory and ovariectomized mares may regularly solicit and accept copulation despite low or even no circulating levels of ovarian steroids. Instead, endocrine support of sexual behavior in these mares appears to be provided by adrenal cortical androgens and/or estrogens. A smaller percentage of pregnant mares may also show some signs of estrus but are not likely to be receptive to mating. The ability of the mare to dissociate sexual behavior from ovulation is unique among infraprimate species studied. The display of estrous behavior and willingness to copulate, even at times when conception is impossible, may have functional significance in promoting band cohesiveness in wild and feral populations. Even without copulation, continued motivation to remain in proximity to the band stallion may contribute to the stability of the social unit outside the ovulatory season.

Animals↗

The effect of feeding status on sexual attractiveness of Ixodes ricinus (Acari: Ixodidae) females.

Sexual attractiveness of unengorged, semiengorged and fully engorged females of both field and laboratory populations of Ixodes ricinus was studied under laboratory conditions by means of a computerised video tracking system. A male and a female were allowed to walk freely in a glass arena during 1 h and their behavioural interactions were observed. Obtained results revealed that the feeding status of L. ricinus females affects their sexual attractiveness. The highest attractiveness was observed in engorged females, the lowest in unengorged females of the field population. Copulation occurred in all experimental groups with the highest frequency observed in the unengorged virgin laboratory females. Despite engorged females being highly attractive for males on distance, the frequency of copulation was low. Differences between field and laboratory populations were found in both behavioural parameters and frequency of copulation.

Animals↗