Sexual behaviour of female rats subjected to early protein energy malnutrition

Young rats (left with the mother until weaned) were exposed from birth to the age of 49 days to a low protein (malnutritional) diet. At 80 days the young females were ovariectomized and were subjected to standard long-term treatment with oestradiol and progesterone. The females' sexual behaviour was studied in interaction tests with an intact male. It was evaluated according to a scale of sexual responsiveness comprising copulatory (a lordosis posture) and precopulatory (a presenting posture, hopping, ritualized darting) patterns. All the experimental females displayed a high copulatory readiness and precopulatory behaviour; a tendency to lower sexual responsiveness, interpreted as diminished sensitivity to the given doses of the hormones, was manifested only after their repeated administration. Compared with the control females, the number of experimental animals in which the given doses of the hormones induced complete precopulatory behaviour--ritualized darting--was smaller. The results contrast with findings on the sexual behaviour of males subjected to malnutrition from an early age.     

Initiation of copulatory behavior in castrated male rats injected with critically adjusted doses of testosterone

In castrated male rats it was possible by means of administration of adequate testosterone propionate doses to induce a state in which the males did not initiate copulatory behavior with a stimulus female displaying no (Lordotic female) or weak (Presenting female) intensity of precopulatory behavior. On the other hand, such males started to copulate and finished copulatory series when a stimulus female exhibiting the complete pattern of precopulatory behavior (Darting female) was offered. The males “prepared” in this manner displayed regularly precopulatory behavior directed toward all the stimulus females. The requisite testosterone doses differed individually. However, in the majority of cases they were in the range of 500–700 μg of testosterone injected once a week.     

Estradiol plus progesterone treatment and precopulatory behavior in prepubertally ovariectomized female rats: dose-response relationships

The copulatory and precopulatory behavioral repertoire expressed in terms of the intensity scale of sexual responsiveness was investigated in female rats ovariectomized on the 30th day after birth and treated for 9 weeks, once weekly, with estradiol dipropionate (5, 10, 15, 20, and 30 micrograms per animal) plus progesterone (in the range from 0.0 to 2.4 mg per animal). It was found that the higher the dosage of estradiol and progesterone used, the more complete the precopulatory behavioral pattern that was induced. Further, the same level of sexual responsiveness was achieved by various combinations of both hormones used. The dose-response relationships are less complicated in prepubertally ovariectomized females in comparison with those spayed as adults. Finally, the most complete pattern, characterized by ritualized darting, was not achieved at all.     

The regulation of precopulatory behavior by ovarian hormones in the female Mongolian gerbil

The hormonal regulation of precopulatory behavior in the female Mongolian gerbil was studied using two groups (N = 6) of sexually experienced females. A novel testing procedure was used which involved females living continuously with test males for several days. The test males showed either full sexual behavior (copulating males, C) or only precopulatory behavior (noncopulating males, NC). Experiment 1 investigated changes during the estrous cycle and following ovariectomy in females. Experiment 2 studied the effects of hormonal treatment of these ovariectomized females with 6 micrograms estradiol benzoate (EB) followed by 0.4 mg progesterone (P) or by 0.04 ml arachis oil. When tested with NC males, females displayed a greater range of precopulatory behavior. The patterns could be classified into three groups according to the manner of response to ovariectomy and hormone treatment. Group I patterns (approach, leave, and olfactory investigation of the male's head) were affected by neither ovariectomy nor EB treatment relative to Day 3 levels (Day 3, day preceding estrus; Day 4, estrus), but they were increased to estrous levels by EB and P. Group II patterns (darting, foot-stomping, and the present and piloerection postures) appeared only during estrus, did not appear after ovariectomy, and reappeared only after sequential EB and P treatment. Group III patterns (investigation of the male's anogenital area, allogrooming, ventral gland marking, and sand-rolling) were reduced relative to both estrus and Day 3 levels by ovariectomy and increased above Day 3 levels by EB alone; EB and P treatment further increased Group III patterns to the level of estrus. It is suggested that female precopulatory behavior patterns differ in their responsiveness to ovarian hormones. Estrogen appears to affect those patterns associated with the earliest stages of estrus (Group III).     

Sexual interaction in adult laboratory rats: importance of female's stimuli and of male's experience

Precopulatory behaviour of adult male rats towards the passively receptive and towards the ovariectomized females was observed in dyadic interaction, first, at the time when the males were sexually inexperienced and later on after they had gained small and uniform copulatory experience (with a darting female). It was found that precopulatory activity of the sexually inexperienced males towards the ovariectomized female was "higher and richer" than that of males exposed to the passively receptive female. When the males were sexually experienced the frequency of precopulatory behavior increased and the patterns changed only in the animals exposed to the passively receptive female. Precopulatory activity of males weakened during the interaction under all experimental conditions. It is remarkable, however, that the weakening of precopulatory activity of sexually inexperienced males exposed to the ovariectomized female was reinforced again after the female temporarily exhibited behaviour typical of the male. We concluded that both the quality of the female's precopulatory behaviour and the quantity of previous male's experience are the decisive factors in successful interaction.     

Transition from precopulatory to copulatory behaviour in male rats with lesions in medial preoptic area: dependence on precopulatory pattern of female

In the present study, both the precopulatory behaviour and the copulatory readiness of male rats following the bilateral medial preoptic area lesions was compared with their intact states. In behavioural testing, the intensity of female precopulatory behaviour was used as an experimental variable. The results showed that the natural threshold of copulatory readiness of males was increased in the lesioned state, the animals were more dependent on the soliciting patterns of the female. However, all the males exhibited conspicuous precopulatory behaviour towards the stimulus females used. Apparently, further brain structures participate in the regulation of sexual behaviour of males, above all, in activation or maintenance of precopulatory activity.     

Role of the medial preoptic area in sexual behaviour of the male rat: a study using repeated cycloheximide infusions

The effect of cycloheximide (CHX), an inhibitor of proteosynthesis, on sexual behaviour was studied in adult male rats in which it was infused into the medial preoptic area of the hypothalamus (MPOA). Sexual interaction took place under control and modification of the precopulatory behaviour of the female. Among the various bilaterally infused amounts of CHX--20 micrograms in 1 microliter, 40 micrograms in 1 microliter, 80 micrograms in 2 microliters--the effect of the largest dose corresponded to a hypothetical state of copulatory readiness of the males; 2 h after administration the males were not capable of initiating copulatory behaviour with a passively receptive female, but did so successfully with a highly soliciting female. The dependence of copulatory readiness of the males on the precopulatory behaviour of the females was confirmed repeatedly after this dose of CHX. At the same time, the precopulatory activity of the males towards a passively receptive female was unimpaired. The effect of CHX was reversible; 48 h after infusion the males displayed high copulatory readiness. Only half the males (n = 7) given bilateral infusions of 80 micrograms CHX fulfilled criterion of copulatory performance. Histological control demonstrated that the MPOA was affected bilaterally by CHX infusion in 10 males out of 14. The results are discussed from the aspect of participation of the MPOA in the regulation of male sexual behaviour. The method allows changes in sexual behaviour to be studied in the same individual in a chronic experiment.     

Castration, sex steroids, and heterosexual behavior in adult male laboratory-housed stumptailed macaques (Macaca arctoides)

The sociosexual behaviors of six stable male-female pairs of stumptailed monkeys were studied in half-hour pair tests. Their performance before and after castration of the males was compared. The effects of replacement therapy with sex steroids on male-female interaction were studied. Also the effects of new females as sexual partners were investigated. Castration caused a significant decrease in sexual behavior. Individual males could display ejaculatory behavior up to about 1 year postcastration. Dihydrotestosterone propionate (75 mg/week/male) alone or in combination with estradiol benzoate (0.9 or 3 mg/week/male) was not effective in restoring sexual behavior to precastration levels in the three castrated males tested. Replacement therapy with testosterone propionate (75 or 10 mg/week/male) was effective in restoring copulatory behavior in half of the castrated males. In some males the introduction of a new female caused an increase in sexual activity, usually when sexual activity with their familiar partner was low. This occurred both in the castration condition and in the steroid treatment period, suggesting, that low activity was caused by low "motivation" and not by the inability to perform.     

Sniffing behaviour, copulatory readiness, copulatory performance and reproductive morphology of adult male rats weaned prematurely

Sexual behaviour of adult male rats weaned prematurely--on day 15 after birth--and of males weaned at the optimal age--on day 28--was compared at two life periods: at the age of 90 days and when the males were one year old. Sexual behaviour was evaluated by means of three characteristics: a) sniffing behaviour, b) copulatory readiness, c) copulatory performance. The differences in sniffing behaviour between prematurely and normally weaned males were assessed in terms of their responsiveness to sccents of unrelated adult males or oestrous females and from the point of view of both housing conditions after weaning and the animals' sexual experience. No differences were found in copulatory readiness as well as in copulatory performance between the two groups of males tested in their interaction with a female exhibiting complete precopulatory pattern. The morphological examination revealed no differences in the weight of reproductive organs and in spermatozoa development. We conclude that the sexual behaviour as well as reproductive organs of male rats are very resistant to early separation of pups from the mother which contrasts with some earlier findings.     

The post-copulation withdrawal response in female baboons: A functional analysis

Rapid withdrawal of females from males at the end of the copulatory sequence (prior to male dismounting) characterizes several primate species. The purpose of this paper is to make a preliminary investigation into possible functional aspects of these “copulatory darts.” Two hypotheses are proposed; (1) females use darts to aid competing males to locate the source of copulation calls; and (2) females dart in order to promote re-mating with the same male. Both hypotheses postulate that darts may thus enhance male-male competition, although acquisition of other benefits may be the primary drive to darting according to the re-mating hypothesis. Using data from 157 copulations collected from six females spread across four groups in a wild population of chacma baboonsPapio cynocephalus ursinus, darting behaviour is described. Within females, darting distance is highly variable. Darting distance is greatest at the time of ovulation (and maximal sexual swelling), and there is some evidence that darting may also be more frequent, and cover greater distances, when the mating male is adult rather than juvenile. While darting behaviour appears unrelated to the presence or absence of ejaculation, it tends to correlate positively with the duration of the female copulation call. These preliminary results are consistent with a mediating role in male-male competition and therefore provide support for both hypotheses.     

Precopulatory behaviour of male rats: developmental aspects and dependence on female's solicitation

Sexual communication between sex partners can involve just one sensory modality or a combination, depending upon such factors as species, habitat, and context. Ethological aspects of sexual communication has been widely documented in rodents. In rats, sexual communication between male and female varies according to the production of signals by a female that signal receptivity, proceptivity, and attractivity. However, in the laboratory experiment, such approach is often neglected. In the present study, two types of stimulus female--Lordotic and Darting--were used with the aim to examine developmental changes in precopulatory behaviour of males. Besides the dependence of the male's precopulatory repertoire on the strength of proceptive stimuli emitted by the female was studied. Male rats ranging from 30 to 175 days of age were observed under the dyadic interaction. It was found: (a) Precopulatory behaviour of the 30-day-old males was not clear-cut, the males devoted more time to social investigation of adult (Darting) female. (b) All the 40-day-old males exhibited precopulatory behaviour in the range of the repertoire displayed by adult animals. (c) Precopulatory activity of juvenile (45-day-old) as well as of adult (90-day-old) males exposed to Lordotic female was significantly lower as compared with that exhibited by males toward Darting female. (d) Copulatory readiness of males increased with the age, in fact, all animals aged from 75-135 day were able to pass from the precopulatory into the copulatory phase of sexual interaction. (e) Although the 175-day-old males exhibited pronounced precopulatory activity, they did not initiate copulations. The implications of each of these findings are discussed from the point of view of both the developmental aspects and the stimulus-response relationships. To sum up, the development of normal flow of sexual behaviour of male rats proceeds simultaneously with the development of physiological and morphological parameters. The appearance of precopulatory behaviour is less dependent on the internal (hormonal) readiness than copulatory behaviour. The connection of both phases, i.e. precopulatory and copulatory, is terminated at about Day 75, and is the key moment of sexual interaction from the point of view of reproductive success. In fact, a successful course of sexual interaction is codetermined by the intensity of behavioural stimuli and/or by the completeness of proceptive patterns provided by the female partner. It is clear that the reproductive process cannot be completed without some degree of communication. The deterioration of sexual interaction found in males aged 175 day seems to be the consequence of their absolute heterosexual abstinence.(ABSTRACT TRUNCATED AT 400 WORDS)     

Effect of visual cues from female on the postejaculatory operant behavior of male rat

Bar-press response reinforced by the contact with estrous female were investigated in male rats in two experimental situations, when the visual communication with female was prevented or when it was permitted. After the instrumental responses directly preceded by ejaculations, depending on the increasing duration of response latency, three categories of male behavior were observed: avoiding of the contact with female, precopulatory behavior, or copulatory behavior. The enabling the visual communication with a female caused the significant increase of occurrence of copulatory behavior and significant decrease of contact-avoiding behavior, whereas the precopulatory behavior showed unsignificant increase. It is suggested that when prevented to see the female male rats can perform the instrumental response even when the sexual arousal is to low for precopulatory behavior. Such unrewarded responses lead to the appearance of frustration. On the contrary, when the female is visible the instrumental response is delayed up to the moment when the visual cues coming from female can evoke at least precopulatory behavior.     

Sexual orientation, proceptivity, and receptivity in the male rat as a function of neonatal hormonal manipulation

The influence of neonatal androgen on the potential to exhibit feminine sexual behavior was investigated. Male rats castrated on Day 0 but not those castrated on Day 4 or later showed hop/darting, ear wiggling, and lordotic behavior in response to treatment with estrogen and progesterone in adulthood at a frequency equal to that of females. Neonatal treatment with testosterone propionate (1 mg/rat for 4 days) abolished the capacity to show these behaviors. In subsequent experiments, involving castration of male rats at 0 or 4 hr after cesarean delivery, the effect of the postnatal surge of testicular secretions on the expression of female sexual behavior was investigated. No differences were seen in the frequency of hop/darting, ear wiggling, and receptivity between males castrated immediately or 4 hr after delivery. In a preference test where the experimental male could choose between an estrous female and a sexually active male, the neonatally castrated males preferred the company of a male when treated with estrogen and progesterone. The implantation of testosterone resulted in a preference for an estrous female. It was concluded that testicular secretions in the newborn male influence adult sexual orientation and suppress the ability to show proceptive and receptive behaviors.     

Paternity analyses in wild‐caught and laboratory‐reared Caribbean cricket females reveal the influence of mating environment on post‐copulatory sexual selection

Polyandry is ubiquitous in insects and provides the conditions necessary for male‐ and female‐driven forms of post‐copulatory sexual selection to arise. Populations of Amphiacusta sanctaecrucis exhibit significant divergence in portions of the male genitalia that are inserted directly into the female reproductive tract, suggesting that males may exercise some post‐copulatory control over fertilization success. We examine the potential for male–male and male–female post‐copulatory interactions to influence paternity in wild‐caught females of A. sanctaecrucis and contrast our findings with those obtained from females reared in a high‐density laboratory environment. We find that female A. sanctaecrucis exercise control by mating multiple times (females mount males), but that male–male post‐copulatory interactions may influence paternity success. Moreover, post‐copulatory interactions that affect reproductive success of males are not independent of mating environment: clutches of wild‐caught females exhibit higher sire diversity and lower paternity skew than clutches of laboratory‐reared females. There was no strong evidence for last male precedence in either case. Most attempts at disentangling the contributions of male–male and male–female interactions towards post‐copulatory sexual selection have been undertaken in a laboratory setting and may not capture the full context in which they take place – such as the relationship between premating and post‐mating interactions. Our results reinforce the importance of designing studies that can capture the multifaceted nature of sexual selection for elucidating the role of post‐copulatory sexual selection in driving the evolution of male and female reproductive traits, especially when different components (e.g. precopulatory and post‐copulatory interactions) do not exert independent effects on reproductive outcomes.     

Experimental evolution exposes female and male responses to sexual selection and conflict in Tribolium castaneum

Between-individual variance in potential reproductive rate theoretically creates a load in reproducing populations by driving sexual selection of male traits for winning competitions, and female traits for resisting the costs of multiple mating. Here, using replicated experimental evolution under divergent operational sex ratios (OSR, 9:1 or 1:6 ♀:♂) we empirically identified the parallel reproductive fitness consequences for females and males in the promiscuous flour beetle Tribolium castaneum. Our results revealed clear evidence that sexual conflict resides within the T. castaneum mating system. After 20 generations of selection, females from female-biased OSRs became vulnerable to multiple mating, and showed a steep decrease in reproductive fitness with an increasing number of control males. In contrast, females from male-biased OSRs showed no change in reproductive fitness, irrespective of male numbers. The divergence in reproductive output was not explained by variation in female mortality. Parallel assays revealed that males also responded to experimental evolution: individuals from male-biased OSRs obtained 27% greater reproductive success across 7-day competition for females with a control male rival, compared to males from the female-biased lines. Subsequent assays suggest that these differences were not due to postcopulatory sperm competitiveness, but to precopulatory/copulatory competitive male mating behavior.     

Drosophila female precopulatory behavior is modulated by ecdysteroids

The effect of ecdysteroid signaling on Drosophila female precopulatory behavior was investigated using two types of mutants with either globally reduced ecdysteroid availability or reduced expression of ecdysone receptors in fruitless neurons, known to control sexual behavior. While being courted by males, mutant females performed significantly less full ovipositor extrusion behavior to reject male copulation attempts. Ecdysteroid depleted females (ecdysoneless(1)) performed male-like courtship behaviors, including unilateral wing extension and song production with patterns very similar to male courtship song. These results support the hypothesis that ecdysteroids modulate female sexual behavior, perhaps acting as a regulator of sexual motivation, and as a component affecting the performance of sex specific behavior patterns.     

Gonadal influences on sexual behavior in the male musk shrew (Suncus murinus)

In this series of experiments the hormonal bases for male copulatory behavior in the musk shrew (Suncus murinus) were examined. Male musk shrews failed to show copulatory behavior after castration. Testosterone replacement fully reinstated sexual behavior. Males castrated at birth, and tested as adults after receiving testosterone implants, did not show male-typical sexual behavior. It appears that the gonads are essential for the regulation of male sexual behavior in this primitive mammal.     

Psychobiology of sexual behavior in a whiptail lizard, Cnemidophorus inornatus

This study investigated the effects of social environment on gonadal recrudescence and sexual behavior in male and female Little Striped Whiptail lizards (Cnemidophorus inornatus). The presence of sexually active males facilitates ovarian recrudescence in conspecific females. Similarly, the presence of reproductively active females facilitates testicular recrudescence in conspecific males. Males housed with females, however, had lower average circulating concentrations of testosterone and dihydrotestosterone, and higher average concentrations of corticosterone compared to intact males housed in isolation. In other studies, the presence of reproductively active females partially restored courtship behavior in castrated males compared to castrated males housed in isolation. Despite the stimulatory effects of females on castrates, exogenous androgens are required for complete restoration of all components of sexual behavior in male C. inornatus. Females are receptive to male courtship and copulatory behavior only during the vitellogenic stages; females in previtellogenic or postovulatory ovarian stages aggressively reject male courtship advances. These findings demonstrate reciprocal effects of sexual behaviors of males and females upon each other's reproductive behavior and physiology.     

A time-sequence functional analysis of mating behaviour and genital coupling in Drosophila: role of cryptic female choice and male sex-drive in the evolution of male genitalia

Male genitalia in Drosophila exemplify strikingly rapid and divergent evolution, whereas female genitalia are relatively invariable. Whereas precopulatory and post-copulatory sexual selection has been invoked to explain this trend, the functional significance of genital structures during copulation remains obscure. We used time-sequence analysis to study the functional significance of external genitalic structures during the course of copulation, between D. melanogaster and D. simulans. This functional analysis has provided new information that reveals the importance of male-driven copulatory mechanics and strategies in the rapid diversification of genitalia. The posterior process, which is a recently evolved sexual character and present only in males of the melanogaster clade, plays a crucial role in mounting as well as in genital coupling. Whereas there is ample evidence for precopulatory and/or post-copulatory female choice, we show here that during copulation there is little or no physical female choice, consequently, males determine copulation duration. We also found subtle differences in copulatory mechanics between very closely related species. We propose that variation in male usage of novel genitalic structures and shifts in copulatory behaviour have played an important role in the diversification of genitalia in species of the Drosophila subgroup.     

Precopulatory Sexual Cannibalism Causes Increase Egg Case Production,Hatching Success,and Female Attractiveness to Males

Precopulatory sexual cannibalism is an extreme form of sexual conflict that can entail significant costs to the cannibalized individual and a variety of costs and benefits to the cannibal itself. Characterizing these costs and benefits is fundamental to our understanding of how this behavior evolves. Using the spider Agelenopsis pennsylvanica, we tested the reproductive consequences of precopulatory sexual cannibalism by staging cannibalization events and comparing the performance of experimental cannibals against natural cannibals (i.e., those that cannibalized on their own) and non‐cannibals. We found two performance benefits associated with precopulatory sexual cannibalism: first, experimental cannibals were more likely to produce egg cases than non‐cannibals, and second, egg cases from experimental cannibals and natural cannibals were significantly more likely to hatch than those produced by non‐cannibals. We then tested whether males were more likely to approach the webs of experimental cannibals vs. non‐cannibalistic control females. Our data demonstrate that sexual cannibalism increases female attractiveness to males. Although this result seems counterintuitive, in fact, rates of precopulatory sexual cannibalism were much lower in females that had already cannibalized their first male: 38% of sexually naïve females engaged in precopulatory sexual cannibalism, whereas only 5% of females engaged in cannibalism a second time. Thus, males that approach cannibals receive two benefits: they are less likely to be cannibalized precopula, and they have the possibility of mating with females that have a higher probability of producing viable egg cases. Taken together, our data suggest that precopulatory sexual cannibalism affords females numerous benefits and may have a hand in shaping male mate choice decisions.