Failure of a variety of antiestrogens to mimic estrogen action in the induction of sexual receptivity in a female lizard

The purpose of the present study was to determine whether nonsteroidal antiestrogens could act as estrogens by inducing or facilitating estrogen-induced sexual receptivity in a female lizard (Anolis carolinensis). Experiments were conducted to (1) test the ability of enclomiphene (ENC) and zuclomiphene (ZUC) to induce sexual receptivity in estrogen-untreated ovariectomized females; (2) to determine the effect of ENC and ZUC pretreatment on E₂B induction of sexual receptivity; and (3) to examine the ability of a variety of antiestrogens to act as estrogens by inducing sexual receptivity in females pretreated with a behaviorally ineffective estrogen treatment regimen.     

Effects of ovariectomy and estrogen replacement on attractivity and receptivity in the red-sided garter snake (Thamnophis sirtalis parietalis)

Activation of courtship behavior in male red-sided garter snakes, Thamnophis sirtalis parietalis, is independent of the presence of sex steroids. The only consistent treatment that stimulates courtship behavior in males is prolonged exposure to low temperature followed by subsequent warming, mimicking the emergence from hibernation. We investigated whether attractivity and receptivity in female red-sided garter snakes is similarly steroid independent.Female red-sided garter snakes are attractive when they emerge from hibernation and are courted by males; most mate within an hour of emergence. In a series of experiments, groups of females were either ovariectomized (OVEX) in the late spring, fall or while in hibernation. They were tested for attractivity and receptivity upon emergence from hibernation. Females OVEX in the spring were unattractive whereas those OVEX in fall or while in hibernation were attractive. Thus, attractivity appears determined the year before emergence and is dependent on the presence of the ovaries. All OVEX females were unreceptive upon emergence. OVEX females were also given replacement estradiol (E) treatment (either in Silastic capsules or single injections) at various points of their annual cycle. The only treatment that resulted in reinstating receptivity in OVEX females was the injection of E (20 g) one hour prior to emergence. The effectiveness of E in reinstating receptivity was time dependent: the longer the period between emergence and injection, the less effective the same dosage was in stimulating receptive behavior.These experiments suggest that sexual behavior in female red-sided garter snakes is, unlike males, dependent on the presence sex steroid hormones. Although E is naturally at its lowest seasonal level upon emergence, the concentration is sufficient to stimulate receptivity. However, it appears that temperature regulates a time-limited window of sensitivity to E.     

Sexual receptivity of Anastrepha obliqua (Diptera: Tephritidae) wild females affected by nutrition and egg load

The influence of feeding, egg load and stage of ovarian development on the sexual receptivity of some insect species has been reported. This work investigated the effects of different brewer's yeast concentrations in the diet offered to females of West Indian fruit fly, Anastrepha obliqua, on their sexual receptivity. Wild females were fed on artificial diets containing 2.0 g brewer's yeast per 100 mL of water (low‐yeast diet, LY diet), 6.5 g yeast per 100 mL of water (high‐yeast diet, HY diet) or only sucrose (Su diet) and then placed with the males. Female sexual receptivity tests were carried out in acrylic boxes in the laboratory under controlled conditions (temperature, humidity and light). Females fed on the HY diet were more sexually receptive to male courtship than the other females. Additionally, when fed on the same diet (HY diet), those females with higher egg load were more sexually receptive. The data suggest that the ingestion of a protein source is essential for increasing sexual receptivity and that egg load is the most important factor affecting sexual receptivity of A. obliqua females.     

Signalling and Sexual Conflict: Female Spiders Use Stridulation to Inform Males of Sexual Receptivity

The use of acoustic signals by males during courtship and mating is well known. Nevertheless, their association with female unwillingness to mate is much less studied. In spiders, stridulation during sexual interactions is relatively common in some groups, but mainly restricted to males. In the pholcid spider Holocnemus pluchei, both sexes have stridulatory organs. The aims of the present work were (1) to establish whether female stridulation occurs during intra‐ and inter‐sexual interactions, (2) to determine whether female reproductive status affects the likelihood that she will stridulate and (3) to determine whether female stridulation is influenced by male sexual behaviour. We found that female stridulation usually occurs both during intrasexual interactions and, most frequently, during intersexual interactions. Females with more previous matings stridulated more frequently. Stridulation intensity was higher in females that did not accept new copulations compared with those that copulated. Female stridulation did not vary in elaborated and non‐elaborated courtship. Thus, females use stridulation to communicate levels of sexual receptivity. It is also possible that females use stridulation to indirectly assess male ability to persist and persuade.     

SEX POLYMORPHISM IN SILENE ACAULIS (CARYOPHYLLACEAE) AND THE POSSIBLE ROLE OF SEXUAL SELECTION IN MAINTAINING FEMALES

Pollen accumulation rates and their relationship to stigma morphology and pollinator visitation behavior were compared between female and hermaphrodite sex morphs of the alpine plant Silene acaulis var. subacaulescens (Caryophyllaceae). Although stigmas of female and perfect flowers collected comparable pollen loads, stigmas of female flowers became receptive earlier in anthesis and therefore recruited a larger number of pollen tubes. Together with early receptivity young female flowers had a larger stigmatic area with longer papillae than did perfect flowers. Pollinator behavior also differed between morphs, with bumble bees spending more time probing female than perfect flowers. Differences in stigma receptivity schedules of female and perfect flowers have consequences for different opportunities for sexual selection in the two sex morphs. Female flowers, by providing a more effective gametophytic screen, have the potential to produce higher quality offspring. This is proposed as a further compensatory advantage maintaining females, with a single fitness function, in populations containing hermaphrodites which have both male and female fitness functions.     

Behaviors associated with mating success in the “purring” wolf spider Gladicosa gulosa

Studies of sexual selection in animal systems frequently use courtship and receptivity behaviors as metrics for mating success, and this is generally built upon data associating such behaviors with mating. Spider systems are becoming increasingly popular in sexual selection research, and the wolf spider Gladicosa gulosa has the potential to help answer unique questions in animal communication. However, this species is understudied, and no quantitative data exist that demonstrate which male and female behaviors may be indicators of mating success. We paired male and female G. gulosa and recorded the behaviors involved in their courtship and mating. We found a number of significant behaviors from both males and females, and we suggest which behaviors/variables are perhaps most predictive of mating success. Beyond building a data‐based foundation for future work, we also suggest that the relationship of certain male and female variables may shed light on some of the more unique features of this species.     

Effects of male mating interval on spermatophore formation,transfer, and subsequent female receptivity and fecundity in the sorghum plant bug,Stenotus rubrovittatus

Males of the sorghum plant bug, Stenotus rubrovittatus (Matsumura) (Heteroptera: Miridae), transfer a spermatophore to females during copulation. After a 1‐day interval between the first and second copulation, males transferred both sperm and a spermatophore to females during the second copulation. However, when male mating interval was <1 h, they transferred sperm but no spermatophores to females during the second copulation. Therefore, the male mating interval probably produces two types of mated females, those with and those without a spermatophore. Mated females of S. rubrovittatus do not remate for at least 3 days after mating, even when courted, and lay more eggs than virgin females at the beginning of the oviposition period. The effects of spermatophores on female sexual receptivity and fecundity were examined using mated females with or without a spermatophore. Only one of the 40 (2.5%) mated females with a spermatophore remated, whereas 10 of the 26 (38.5%) without a spermatophore remated. Furthermore, mated females with a spermatophore laid more eggs than those without a spermatophore. These results suggest that spermatophores participate in reducing female sexual receptivity and enhancing female fecundity in S. rubrovittatus.     

Effects of soil resources on expression of a sexual conflict over timing of stigma receptivity in a mixed‐mating plant

While environmental factors strongly influence plant growth and reproduction, less is known about environmental effects on sexual selection and sexual conflict. In this study on mixed‐mating Collinsia heterophylla we investigated whether soil resource environment affected traits associated with sexual conflict. In C. heterophylla a sexual conflict over timing of stigma receptivity occurs. Early stigma receptivity benefits pollen parents by securing paternity while late stigma receptivity benefits female fitness in terms of increased seed production. We performed hand‐pollinations combining recipients and donors grown either in high or low resource environments and asked whether these treatments influenced sexual conflict traits – recipient‐ and donor‐based influence on timing of stigma receptivity – and conflict costs related to reduced early seed production. We also asked whether resource environment affected eight traits related to general fitness and mating system. Sexual conflict‐associated traits – timing of stigma receptivity and seed production – were generally unaffected by resource environment. While no universal effect of resources was detected, we did observe donor‐specific responses to environment, suggesting that environment can nonetheless contribute to variation in timing of stigma receptivity. Recipients grown under low resources showed pronounced differences among donors for number of seeds per capsule, indicating that recipients favour some donors over others under resource‐low conditions. Moreover, high resources increased number of flowers but reduced pollen germination rate, while other traits were unaffected, indicating variation in the response to resource environment for fitness‐ and mating system‐traits. Our results suggest that even though soil resource environment had a low impact on the sexual conflict traits and related costs in C. heterophylla, it generated variability in pollen donor‐influence on this trait and in recipient sorting among donors. Thus, it is possible that both sexual conflict and sexual selection is affected by environmental factors not only in animals but also in plants.     

Male Mating Interest Varies with Female Fecundity in <Emphasis Type="Italic">Pan troglodytes schweinfurthii</Emphasis> of Kanyawara,Kibale National Park

Female chimpanzees mate promiscuously during a period of extended receptivity marked by prominent sexual swelling. Recent studies of wild chimpanzees indicate that subtle variations in swelling size could act as a reliable cue of female fertilization potential both within and between cycles (Emery and Whitten Behavioral Ecology and Sociobiology, 54, 340–351, 2003; Deschner et al. Hormones and Behavior, 46, 204–215, 2004). Copulation rates increase during the periovulatory period and during conception cycles (Deschner et al. Hormones and Behavior, 46, 204–215, 2004; Emery Thompson American Journal of Primatology, 67, 137–158, 2005a), suggesting that males may be able to assess female fertilization potential. We asked whether facultative timing of copulation in Kanyawara chimpanzees was due to increased male mating interest or to increased female proceptivity during the most fecund days. We assessed multiple measures of male mating effort in cycles aligned relative to the day of detumescence and compared periovulatory days to other days of maximal swelling, and conception cycles to nonconception cycles. The rate and proportion of male initiative in soliciting sexual behavior increased during periods of highest fertilization potential. Males were also more likely to interrupt copulations, associate with estrous females, and compete with other males when females were most likely to conceive. Females initiated copulations more frequently during conception cycles but did not visibly shift mating behavior within cycles. Our results support the hypothesis that male chimpanzees have the ability to assess the profitability of mating attempts, a trait that may act as a counter-adaptation to female strategies to obscure paternity. We discuss potential cues and the implications for female reproductive strategies.     

Intracerebral prostaglandin E2: Effects upon sexual behavior,open field activity and body temperature in ovariectomized female rats

A single 27 gauge implant of PGE₂ into the periventricular region of the hypothalamus resulted in a significant increase in sexual receptivity in estrogen primed, ovariectomized female rats. Open field activity levels were only slightly decreased while rectal body temperature increased significantly over control values. It is postulated that the effects upon sexual receptivity might be mediated by PGE₂ stimulated LRF release.     

Cycles of mating behaviour, oestrogen and progesterone in the thick-tailed bushbaby (Galago crassicaudatus crassicaudatus) under laboratory conditions

Vaginal smears and blood samples were taken throughout the reproductive cycle of female Galago c. crassicaudatus. Blood plasma was assayed for oestradiol and progesterone, and vaginal smears were initially classified dioestrus or vaginal oestrus. During vaginal oestrus the females were tested daily for sexual receptivity by being placed with a male. Those days on which the male achieved intromission were reclassified as behavioural oestrus. During dioestrus the females were tested weekly with males. Female receptivity increased and then declined across a 6-day period of behavioural oestrus during the 44-day cycle. Fully cornified smears were characteristic of the period of maximal receptivity and oestradiol secretion. The luteal phase lasted 24 days with a plasma progesterone peak midway through dioestrus.     

Induction of sexual receptivity in the female lizard, Anolis carolinensis: Effects of estrogen and the antiestrogen CI-628

Daily behavioral testing revealed that there is a latency period of at least 48 hr from the administration of a single injection of estradiol benzoate (EB) to the first significant increase in female sexual receptivity in the ovariectomized female lizard, Anolis carolinensis. This latency period did not vary with dosage of EB used in these experiments (i.e., 0.8, 1.4, and 4.0 μg) nor with method of injection (subcutaneous vs intraperitoneal for dose of 1.4 μg EB). Following a single EB injection, female sexual receptivity increased after the 48-hr latency period, reached an observed peak from Day 3 to Day 6, and thereafter declined to pretreatment levels by Day 19. Although both 1.4 and 4.0 μg of EB produced higher levels of female sexual receptivity than did treatment with 0.8 μg of EB, results obtained with 4.0 μg EB did not differ from those obtained with 1.4 μg EB. Administration of the nonsteroidal antiestrogen CI-628 (80 μg) at either 4 or 24 hr following a single subcutaneous injection of 1.4 μg EB significantly reduced subsequent female sexual receptivity. These results suggest that there is a critical length of time during which estrogen must act on the brain and support the concept of an estrogen “maintenance” effect during this priming period.     

Male-Induced Sociosexual Behavior by Vaginal Secretions in Macaca arctoides

Odor communication in Old World monkeys and apes is controversial, because most females have evolved visual and behavioral cues to signal fertility, e.g., sexual swellings. Female stump-tailed macaques (Macaca arctoides) do not have swellings, and mediation of chemical communication likely occurs because males engage in sexual behavior mostly throughout the periovulatory phase of the menstrual cycle. We tested whether vaginal secretions from different cycle phases, with saline solution as a control, promote changes in the frequency of male genital exploration, copulation, and coercive behavior toward females different from the donors, while female donors were apart from the group. Males explored more female genitals when exposed to follicular, periovulatory, and early luteal secretions in comparison to saline or menstrual or late luteal secretions. The increase in coercive behavior after exposure to follicular and periovulatory secretions most likely was a male response to the lack of cooperation of target females in engaging in copulation, as the latter were not receptive during the tests. The strength of male response to vaginal secretions varied significantly as a result of individual variability between donor females, yet the variability does not correlate either to dominance rank or to female age. Exploratory behavior of males correlates significantly with their social rank. Our results suggest that vaginal secretions are among the cues that male Macaca arctoides use to acknowledge the reproductive status of females in the absence of visual signals.     

Effects of male paragonial gland secretion on sexual receptivity and courtship behaviour of female Drosophila melanogaster

Female Drosophila melanogaster show a switch-off in sexual receptivity and an increase in oviposition rate following impregnation. These effects are also observed in unfertilized virgin females which receive transplants of intact male paragonia into the abdominal body cavity. Virgin Amherst wildtype females show a repertoire of responses to male courtship characterized by high rates of kicking and fending and a low rate of genital extrusion. Fertilized females show low rates of kicking and fending but a high rate of genital extrusion. Virgin female hosts receiving paragonial gland implants show an altered pattern of courtship rejection with low rates of kicking and fending. Studies on these host females, and on females homozygous for the mutant allele female-sterile, suggest that the expression of genital extrusion depends, in addition to the paragonial gland substance, upon additional information input from abdominal stretch receptors or from the ovary.     

Induction of sexual receptivity in the female broad-headed skink, Eumeces laticeps, by estradiol-17 beta

Sexual receptivity in the female scincid lizard Eumeces laticeps occurs naturally only during the spring breeding season, which is also when maximal follicular development occurs. The presumption that high estrogen levels are coincidentally present and the need for a reliable method of inducing sexual receptivity for behavioral studies prompted tests of the hypothesis that estrogen induces sexual behavior. A series of experiments established that estradiol-17 beta induces sexual behavior. A series of experiments established that estradiol-17 beta induces sexual receptivity within 4 days when injected every other day at 2.0 micrograms in 20 microliters peanut oil in intact or ovariectomized females. In behavioral tests conducted during August, all control females (intact or ovariectomized injected with vehicle only) rejected courtship whereas all females receiving estrogen copulated. Estrogen injections also induced a statistically significant change from rejection to receptivity within individuals. Initial attempts to implant estradiol-17 beta in Silastic tubes killed all females so treated.     

Sexual signalling by females: do unmated females increase their signalling effort?

Theory predicts that females should invest least in mate searching when young, but increase their effort with age if they remain unmated. Few studies have examined variation in female sexual signalling. Female Dawson''s burrowing bees (Amegilla dawsoni) search for males by signalling their receptivity on emergence, but many leave the emergence site unmated and must attract males at feeding sites. Female bees prevented from mating on emergence had more extreme versions of cuticular hydrocarbon profiles that make them attractive to males, lending empirical evidence of adaptive shifts in female mating effort.     

A STOCHASTIC SIMULATION STUDY ON SPECIATION BY SEXUAL SELECTION

A two-locus multi-allele sexual isolation model incorporating mutation and genetic drift which was first proposed by Nei et al. (1983) is studied here. One locus controls the male mating character, and the other controls female receptivity. All females are assumed to have equal mating success. Therefore, the frequencies of female receptivity alleles are changed by mutation, drift, and hitchhiking with male character alleles. Without hitchhiking, development of sexual isolation between allopatric populations proceeds faster in smaller populations, as expected. The hitchhiking effect, by triggering the mutual reinforcement of mating behavior of both sexes (or the runaway process, Fisher [1958]), speeds up the evolution of sexual isolation significantly. For populations with 2Nv ≤ 0.2 (N = population size, ν = mutation rate), the rates of divergence all approach the maximum possible rate. Sympatric sexual isolation develops quite frequently if two favorable conditions are met: 1) There is no selection on female phenotype (except in some limited cases), and 2) The population size is large enough to carry several female receptivity alleles. Because of stochastic factors, these alleles may lead to the formation of two discrete groups of females, each group receptive to males of different mating characters. The formation of sympatric sexually-isolated groups is also aided significantly, at the incipient stage, by the runaway process.     

Chimpanzee genital swelling and its role in the pattern of sociosexual behavior

Behavioral observations were made on thirteen female and seven male adult group-living chimpanzees (Pan troglodytes). The behavioral data were analyzed as a function of the day of the females' menstrual cycles to explore the possible interrelationship between genital swelling and sociosexual behavior of female chimpanzees. Copulatory behavior was confined almost entirely to the period of genital swelling and the occurrence of male-to-female genital inspection (both female- and male-initiated) was negatively correlated with the days from swelling onset, i.e., as the presumed day of ovulation approached, genital inspection decreased, while copulatory behavior increased. In addition, more females groomed their male cagemates during the luteal phase than in the follicular phase of their cycles, whereas male-to-female grooming was positively correlated with the progress of the cycle, with peaks during the time of swelling onset and menstruation. The profile of sociosexual behavior observed throughout the menstrual cycle suggested that, although chimpanzees exhibit an extended period of sexual receptivity and genital swelling, the presumed fertile period is not concealed. The role of genital swelling in chimpanzees is discussed in relation to the possible hormonal effect on female sexuality and the evolution of chimpanzee mating strategies. © 1992 Wiley-Liss, Inc.     

Sensory cues mediating mating-induced potentiation of sexual receptivity in female rats

Repeated mating of estradiol-primed female rats increases sexual receptivity. Two studies were conducted to determine the contribution of vaginal--cervical stimulation (VCS) to this increase. In the first study, female rats were repeatedly mated for 165 min. The vaginas of half of the females were covered with tape (masked) to prevent intromissions by the males. The remaining females were unmasked. Only females receiving intromissions (unmasked) showed a significant increase in sexual receptivity during repeated mating, suggesting that VCS from intromissions is necessary for repeated mating to increase sexual receptivity. In the second experiment, female rats received either experimentally administered VCS or control scapular stimulation administered with a plastic probe 1 h prior to testing for sexual receptivity. VCS applied in this manner significantly increased sexual receptivity. Together, these findings suggest that VCS from intromissions is one of the primary factors responsible for increases in sexual receptivity following repeated mating.     

Estrogen treatment during development alters adult partner preference and reproductive behavior in female laboratory rats

There is broad acceptance for the idea that during development estradiol ‘organizes’ many aspects of reproductive behavior including partner preferences in the laboratory rat. With respect to partner preference, this idea is drawn from studies where estrogen action was in someway blocked, either through aromatase or estrogen receptor inhibition, during development in male rats. The lack of estrogens neonatally results in a decrease in the male rat's preference for females. In this study, the effect of early postnatal estradiol treatment on the partner preferences of female rats was examined as a further test of the hypothesis that male-typical partner preference is dependent upon early exposure to estrogens. Our principal finding was that increased postnatal estradiol exposure during development affected partner preference in the expected direction, and this effect was seen under several adult hormonal and behavioral testing conditions. Female rats that received exogenous estradiol during development spent more time with an estrous female and less time with a sexually active male than did cholesterol treated females. The estradiol treatment also disrupted normal female sexual behavior, receptivity, and proceptivity.