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Lesions to the medial preoptic area affect singing in the male European starling (Sturnus vulgaris).

The aromatization of testosterone (T) in the medial preoptic nucleus (POM) is known to regulate male courtship and sexual behaviors expressed prior to, and in anticipation of, copulation. Singing in male European starlings is used to attract mates prior to physical sexual contact, suggesting that the POM might be involved. The present study was performed to examine the effects of lesions targeting the POM on singing and courtship behavior in reproductively active male starlings. A significant decrease in song output and the gathering of green nest materials was observed in males with lesions to the POM compared to males with damage to brain areas outside of the POM. Lesions did not affect a male's tendency to remain near a female or to occupy a nestbox, suggesting that the effects of POM lesions were specific to courtship behaviors. Behavioral differences were not related to testis mass or volume, and GnRH immunoreactivity was observed within the hypothalamus and median eminence for each male, suggesting that the effects of POM lesions were related specifically to POM involvement in song expression rather than to a disruption of the GnRH axis. These results suggest a general role for the POM in the expression of behaviors related to sexual arousal or anticipation, including song.

Analysis of Variance↗

Sex and context: hormones and primate sexual motivation.

Gonadal hormones regulate the ability to copulate in most mammalian species, but not in primates because copulatory ability has been emancipated from hormonal control. Instead, gonadal hormones primarily influence sexual motivation. This separation of mating ability from hormonally modulated mating interest allows social experience and context to powerfully influence the expression of sexual behavior in nonhuman primates, both developmentally and in adulthood. For example, male rhesus monkeys mount males and females equally as juveniles, but mount females almost exclusively as adults. Having ejaculated with a female better predicted this transition to female mounting partners than did increased pubertal testosterone (T). It is proposed that increased pubertal T stimulates male sexual motivation, increasing the male's probability of sexual experience with females, ultimately producing a sexual preference for females. Eliminating T in adulthood reduces male sexual motivation in both humans and rhesus monkeys, but does not eliminate the capacity to engage in sex. In male rhesus monkeys the effects of reduced androgens on sexual behavior vary with social status and sexual experience. Human sexual behavior also varies with hormonal state, social context, and cultural conventions. Ovarian hormones influence female sexual desire, but the specific sexual behaviors engaged in are affected by perceived pregnancy risk, suggesting that cognition plays an important role in human sexual behavior. How the physical capacity to mate became emancipated from hormonal regulation in primates is not understood. This emancipation, however, increases the importance of motivational systems and results in primate sexual behavior being strongly influenced by social context.

Animals↗

Seasonal changes of reproductive behavior and fecal steroid concentrations in Père David's deer.

From July 1997 to September 1997 and from March 1998 to July 1998, we studied reproductive behaviors of Père David's deer in Dafeng, China. During the field behavioral observations, we collected fresh voided fecal samples from the Père David's stags and hinds periodically and kept those samples under -20 degrees C until laboratory analysis. We analyzed the fecal testosterone, estradiol, and progesterone concentrations in those samples using radioimmunoassay. During this study, we also recorded 17 types of male reproductive behaviors and nine types of female reproductive behaviors. Reproductive behaviors and the fecal steroid concentrations showed overt seasonal fluctuations. There were statistically significant correlations between some male reproductive behaviors, such as anogenital sniffing, urine sniffing, urine spraying, wallowing, bellowing, antler adorning, Antler swags mud, chasing, herding hinds, chin resting, mounting and copulating, with the fecal testosterone concentrations. These results suggested that seasonal reproductive behaviors in stags are strongly associated with circulating testosterone. We also found that some female reproductive behaviors fluctuate corresponding with changes in fecal estrogen concentrations. Although there was no direct evidence to confirm the correlations between female reproductive behaviors and fecal estrogen in our experiment, we could not rule out that reproductive activities of hinds were largely related to ovarian estrogen secretion, and estrogen is necessary for inducing female reproductive behaviors.

Acclimatization↗

Only self-paced mating is rewarding in rats of both sexes.

When rats are mated in a traditional mating chamber (with one male and one female) in which the male dictates the pace of the copulatory sequence, males develop a reward state as evaluated by conditioned place preference (CPP). In this mating situation no reward state is induced in females. However, when female rats are able to control (pace) the rate of sexual stimulation, thereby reducing the aversive consequences associated with mating, a clear CPP is observed. In the present study the CPP paradigm was used to determine whether if the reinforced state induced by coital interactions in male rats can be maintained when females pace the sexual interaction. Adult male and female rats were mated in one of two different conditions: (1) where subjects were able to pace their coital interactions or (2) where subjects were not able to pace their sexual contacts. The results showed that when males had control over the sexual interaction they developed a clear place preference while males that mated with females that paced their coital contacts did not develop CPP. Similarly, only females that were able to pace their sexual contacts developed place preference. These results suggest that coital interactions in males, as well as in females, can induce a reward state only when they are able to control the sexual interaction. Under seminatural conditions sexual behavior in rats is highly promiscuous, they mate in groups and repeatedly change partners in the middle of copulation. This behavioral sequence allows both, male and female to control the rate of sexual interaction, assuring the induction of a reward state outlasting the actual performance of coital responses.

Animals↗

Activity of peptidergic neurons in the amygdala during sexual behavior in the male rat.

The medial amygdaloid nucleus (AME) occupies a central position in the circuitry that organizes sexual behavior in the male rat. It receives a projection from olfactory structures that are activated by pheromonal cues indicating receptivity in the female and projects in turn to limbic and hypothalamic structures that are thought to organize aspects of coitus. Electrical stimulation of the AME elicits a behavioral state that is indistinguishable by several measures from the post-ejaculatory interval. We used chronic single-unit recording techniques to determine the behavioral conditions in which the AME is normally active. We found that the cells indeed fired selectively during the presence of a receptive female, but that the discharge considerably anticipated copulation in time. We propose that sexual behavior in the male rat is a reaction chain of fixed action patterns, each one acting as a releaser for the next. The AME mediates an early event in the reaction chain, namely recognition of the receptive female, but electrical activation of the AME causes the reaction chain to proceed to its culminating behavior, the post-ejaculatory interval.

Amygdala↗

Sexual behavior in laboratory strains of Drosophila mercatorum that have spontaneously adopted parthenogenesis.

Three instances are described in which bisexual laboratory strains spontaneously adopted an exclusively parthenogenetic mode of reproduction, even in the presence of fertile, bisexual males. The few males produced by the parthenogenetic strains lack a Y chromosome and are sterile but, nevertheless, showed no correlated impairment of normal mating behavior. In contrast, females show a strong reluctance to accept copulation. This behavioral correlate of parthenogenesis also has been observed previously in experimentally produced parthenogenetic lines. We suggest that genetic breakdown in female mating behavior may contribute to an evolutionary stimulus that results in a selective increase in the frequency of diploidizing events in unfertilized eggs. This ultimately might lead to the origin of an exclusively parthenogenetic reproductive mode.

Animals↗

Influence of food on frequency-dependent sexual activity of Drosophila melanogaster.

When two types of Drosophila are in competition, the frequency dependence of mating successes frequently is measured by direct observation of copulating pairs in "Elens-Wattiaux" observation chambers, the relative frequency of both types being varied. The present experiments, concerning white-ebony mutants in competition with wild-type Canton-S flies, show that the presence of food in the mating chamber influences the sexual activity of flies, this influence differing in the two types when food is present. Thus, in order to have a realistic estimation of frequency dependence, it would seem prudent to conduct these experiments with food in the chamber. Three methods are used to analyze observation data: K coefficients of Petit and Ehrman, regression equations of Ayala and Campbell, and regression equations of Wattiaux and Lichtenberger. These three methods are compared and discussed.

Animals↗

The behavioral phenotype and mating behavior of two inbred strains of Drosophila melanogaster.

Time to copulation was measured in matings within and between two inbred Drosophila melanogaster strains, Edinburgh (E) and 6C/L, using groups of one or five flies of each sex. The E males, which mated faster, usually mate with E females, which are less likely to be fertilized and have fewer progeny whose adult viability is lower. Although generally E males mated far faster with E females than did 6C/L males, there was no difference between the males when with 6C/L females. This need not imply any mating discrimination by E females, but only differences between the strains in the use of preening as a general repulsion movement toward other flies. It is suggested that a greater variety of behavioral and other characteristics (the "behavioral phenotype") should be considered in studies of mating speed and assortative mating, especially where mating speed is being considered as a major component of fitness.

Animals↗

Role of presynaptic serotonergic receptors on the mechanism of action of 5-HT1A and 5-HT1B agonists on masculine sexual behaviour: physiological and pharmacological implications.

In order to establish whether the 5-HT1A or the 5HT1B agonists, 8-OH-DPAT or TFMPP, produce their facilitatory or inhibitory actions on masculine sexual behaviour via a mechanism involving: (a) the serotonin synthesis or release; (b) the stimulation of presynaptic receptors, or (c) the stimulation of somatodendritic receptors, three series of experiments were performed. The administration of the serotonin synthesis inhibitor, p-chlorophenylalanine (p-CPA, 300 mg/kg x 3 days), facilitated sexual behaviour but does not interfere neither with the inhibitory nor with the facilitatory effects of TFMPP (0.5 mg/kg) or 8-OH-DPAT (0.5 mg/kg), respectively. The icv or the intraraphé administration of the serotonergic neurotoxin, 5,7-dihydroxytryptamine (5,7-DHT), slightly stimulated masculine sexual behaviour and produced a decrease in serotonin and its metabolite levels. In lesioned animals TFMPP (0.5 mg/kg) resulted in an inhibitory effect reflected as a prolongation of the ejaculation latency. The inhibitory effect of this drug on mounting behaviour was not observed in 5,7-DHT treated rats. In lesioned animals 8-OH-DPAT (0.5 mg/kg) produced the same facilitatory effect. Present data indicate that serotonergic postsynaptic receptors mediate both the inhibitory and the facilitatory actions of TFMPP or 8-OH-DPAT in copulation. All data further support the idea that endogenous serotonin acts via the stimulation of 5-HT1B receptors to induce its inhibitory effects on masculine sexual behaviour.

5,7-Dihydroxytryptamine↗

Homosexual behavior in a laboratory group of stumptail monkeys (Macaca arctoides): forms, contexts, and possible social functions.

Observations of stumptail monkeys revealed frequent prolonged, intensive genital stimulation between individuals of the same sex, both males and females, adults and infants. These homosexual interactions occurred in positive emotional contexts, were accompanied by positive emotional facial expressions, and often were evidently elictied by the observation of heterosexual behavior. Infants' first copulations were performed with adults of the same or opposite sex who actively aided them. This suggests that homosexual and heterosexual encounters function as training for adult sexual roles, and that homosexual behavior is a basic primate pattern not exclusive to man.

Age Factors↗

Multiple ejaculations during prolonged sexual tests and lack of resultant serum testosterone increases in Male Stumptail Macaques (M. arctoides).

The sexual behavior of four male and four female adult stumptail macaques was observed in standardized 3-hr heterosexual pair tests. The males achieved from 11 to 19 ejaculations during a single test, thus apparently exceeding any other primate studied under laboratory conditions for the ability to display multiple ejaculatory patterns in relatively short periods of time. The stage of the menstrual cycle was not found to be related to the performance of the male or female, although this variable was not analyzed in depth. Progressive increases in the interejaculatory interval (IEI) occurred for all four males for the initial two to four ejaculations of the series, and then either a plateau or a transient decrease in this measure occurred. Since the data varied considerably for IEI as well as for other quantitative measures of sexual behavior for a given pair, individual data rather than group averages are presented. Blood samples drawn immediately before and after behavioral testing as well as at comparable intervals 1 week prior to behavioral testing were analyzed for testosterone concentration by radioimmunoassay for each male. Serum testosterone values were not found to be influenced by multiple ejaculations. Instead, decreased levels of serum testosterone were encountered on the second blood sample of the day for most males, regardless of whether sexual behavior occurred. The possibility that the decreases were related to stress effects of handling is discussed. Because of this complication, it could be only tentatively concluded that repeated copulation to ejaculation over a 3-hr period did not result in appreciable changes in testosterone levels.

Animals↗

Bilateral injections of a selective mu-receptor agonist (morphiceptin) into the medial preoptic nucleus produces a marked delay in the initiation of sexual behavior in the male rat.

We examined the putative functional role of the medial preoptic nucleus mu-receptor population in the expression of male copulatory behavior in sexually vigorous Long-Evans rats. In the first experiment, three doses of morphiceptin (10, 500, and 1000 ng) a selective mu-receptor agonist injected bilaterally into the medial preoptic nucleus, produced a marked delay in the initiation of male copulatory behavior compared to saline injected controls. These injections significantly lengthened intromission and mount latencies while having no appreciable effect on any other parameter of male copulatory behavior. In a separate experiment, the transient inhibition of the expression of male copulatory behavior was completely abolished following pretreatment of naloxone 20 minutes prior to bilateral injections of morphiceptin (1000 ng) into the medial preoptic nucleus. Collectively, these results suggest that the delay in the initiation of copulation that is commonly observed following peripheral or central injections of opioids is mediated at least in part by mu receptors located within the medial preoptic nucleus.

Analgesics↗

Sexual behavior mutants revisited: molecular and cellular basis of Drosophila mating.

The study of Drosophila melanogaster by a combination of forward genetics with specific mutants, and reverse genetics, in which a given gene is expressed in an appropriate brain area to test its effect on behavior, provides a unique opportunity to explore the causal relationship between a particular gene, its function in the cell and the behavioral outcome at the organismic level. Enhanced male-to-male courtship has been shown to occur as a result of mutations in several different genes. For example, the Voila mutant exhibits intense GAL4 reporter expression in the tarsal gustatory sensilla, suggesting the importance of tapping by a male on the female abdomen with his forelegs. Feminization of parts of the antennal lobe and mushroom body by targeted expression of a female-determining gene transformer+ (tra+) drives the male to court other males. Mutations in the tra target gene fruitless (fru), which is expressed in the antennal lobe as well as the suboesophageal ganglion (the gustatory inputs are processed here), also induce homosexual courtship in males. These results suggest that sensory inputs mediated and/or processed by the tarsal receptors, suboesophageal ganglion, antennal lobe and mushroom body contribute to the regulation of male-female courtship. Mosaic analysis localized the neural center for male courtship behavior to the posterior dorsal brain, in which the sensory information processed by the aforementioned neural structures may be integrated. Another mosaic study mapped the neural center for female sexual behavior, as measured by her receptiveness to copulation, to the anterior dorsal brain. The issue as to how the mutations that reduce female sexual receptiveness, e.g. dissatisfaction (dsf), spinster (spin) and chaste (cht), affect the structure and/or function of this neural center deserves to be addressed urgently.

Animals↗

The effect of vegetation density on male mate guarding and extra-territorial forays in the yellow-breasted chat (Icteria virens).

Extra-pair paternity is widely documented in birds, but the behaviors that lead to extra-pair copulations (EPCs) have been less well studied, particularly in territorial songbirds. We studied the behaviors associated with extra-territorial forays (ETFs) and male mate guarding in a socially monogamous, but genetically promiscuous, neotropical migrant passerine, the yellow-breasted chat (Icteria virens). Focal observations and radiotelemetry revealed that both males and females engaged in ETFs. 65% of the females in our study engaged in at least one foray onto a neighboring territory. 50% of males also were observed engaging in ETFs, but males were much more conspicuous during these intrusions compared to females. Females preferred to remain close to the ground in areas where vegetation was the densest. Female behavior was difficult to observe during ETFs but females sometimes interacted with neighboring extra-pair males. Males attempted to guard their mates by remaining close to them and following them during extra-territorial intrusions. We observed instances in which a male attacked his mate and appeared to herd her back to his territory. However a male's ability to maintain close proximity to his mate was significantly and negatively correlated with vegetation density. Our results suggest that the behaviors which lead to extra-pair encounters are influenced by the behavior of all participants and are modified by the characteristics of the habitat.

Animals↗

Masculine copulatory behavior is facilitated by intrathecally administered muscarine.

Behavioral experiments were conducted to examine the role of the cholinergic receptor-agonist muscarine or its antagonist homatropine on the mating behavior of sexually experienced male rats. Male copulatory behavior was recorded after intrathecally administered saline, muscarine (7.5 microg), or homatropine (25 microg). Changes in copulatory behavior were assessed by the following parameters: intromission latency, intromission frequency, intercopulatory interval, ejaculation latency, and postejaculatory interval. Intromission frequency, intercopulatory interval, and ejaculation latency were decreased significantly by muscarine. Intrathecal homatropine decreased the number of copulating animals (five out of 13). In the five animals that were able to ejaculate after homatropine, intromission latency, intercopulatory interval, and ejaculation latency increased significantly. The effects of both drugs on locomotion were also tested. Muscarine induced no significant changes in locomotion compared with saline. A significant increase in locomotion was found after homatropine treatment. These results suggest that acetylcholine, acting at spinal-cord muscarinic receptors, may be involved in ejaculation.

Animals↗

Sex pheromone mimicry in the early spider orchid (ophrys sphegodes): patterns of hydrocarbons as the key mechanism for pollination by sexual deception.

We investigated the female-produced sex pheromone of the solitary bee Andrena nigroaenea and compared it with floral scent of the sexually deceptive orchid Ophrys sphegodes which is pollinated by Andrena nigroaenea males. We identified physiologically and behaviorally active compounds by gas chromatography with electroantennographic detection, gas chromatography-mass spectrometry, and behavioral tests in the field. Dummies scented with cuticle extracts of virgin females or of O. sphegodes labellum extracts elicited significantly more male reactions than odorless dummies. Therefore, copulation behavior eliciting semiochemicals are located on the surface of the females' cuticle and the surface of the flowers. Within bee and orchid samples, n-alkanes and n-alkenes, aldehydes, esters, all-trans-farnesol and all-trans-farnesyl hexanoate triggered electroantennographic responses in male antennae. Most of the alkanes and alkenes occurred in similar patterns both in the bees and orchids. O. sphegodes leaf extracts contained mostly the same compounds but in different proportions. In behavioral tests with synthetic compounds, blends of alkenes triggered significantly more approaches and pounces of the males whereas alkanes were not more attractive than odorless dummies. Since alkanes and alkenes together were most attractive, we conclude they constitute the bees' sex pheromone as well as the pseudocopulation-behavior releasing orchid-odor bouquet.

Adaptation, Biological↗

Location of the reproductive timer in the male cricket Gryllus bimaculatus DeGeer as revealed by local cooling of the central nervous system.

The location of the reproductive timer for the post-copulatory, time-fixed, sexually refractory stage was investigated in the male cricket Gryllus bimaculatus. This stage was defined as the interval between spermatophore protrusion and recommencement of copulation or a calling song. To inactivate the central nervous system locally and reversibly, different body regions were cooled to 10 degrees C for 20-30 min after spermatophore protrusion. A behavioural test then measured the duration of the refractory stage after males recovered from cooling. Males with the head, thorax and anterior abdomen cooled did not show a lengthening of that stage. In contrast, males with the entire abdomen or even the posterior abdominal segments containing only the 6th and terminal (7th-11th) abdominal ganglia showed a lengthening of the refractory stage up to, but not exceeding, the cooling duration. When 20-min cooling was interposed twice after spermatophore protrusion, the refractory stage was lengthened by about 40 min, indicating that interposed cooling did not reset the timer. These results are in agreement with our previous hypothesis that the reproductive timer for the refractory stage in the male cricket is located in the posterior abdominal ganglia, possibly within the terminal abdominal ganglion.

Abdomen↗

No apparent indirect genetic benefits to female red flour beetles preferring males with intense copulatory courtship.

Whether females gain indirect genetic benefits through mate choice is a controversial issue since this requires additive genetic variance in the preferred male traits. Condition dependence could maintain the necessary genetic variance by linking the expression of male traits to the supposedly large number of genes affecting condition. Copulating males of the red flour beetle Tribolium castaneum rub their legs along the female elytra. Females favor males with vigorous rubbing through cryptic female choice. We measured the repeatability and heritability of this behavior and assessed its potential use as indicator of viability and condition. We found genetic variance in larval to adult survival and in the rate of leg rubbing in males. However, the rate of leg rubbing was not related to offspring survival or condition dependent. The genetic variance in leg rubbing was mostly non-additive with very low narrow sense heritability. Therefore, we failed to document any indirect genetic benefits to choosy females through viability of their offspring or attractiveness of their sons.

Animals↗