Heterosexual housing increases the retention of courtship behavior following castration and elevates metabolic capacity in limbic brain nuclei in male whiptail lizards,Cnemidophorus inornatus

In male vertebrates the display of courtship behavior depends on the presence of testicular androgens. However, social experiences in adulthood can alter the hormonal dependence of courtship behavior in a variety of species, and we have previously proposed that these behavioral changes are linked to changes in neural metabolic capacity (cytochrome oxidase activity). Here we investigated the effects of prior social experience (housing with females vs housing in isolation) on the retention of courtship behavior following gonadectomy and on cytochrome oxidase (CO) activity in male little striped whiptail lizards, Cnemidophorus inornatus. In Experiment 1, we found that males that were previously housed with females (HWF males) continued to display courtship behavior longer after castration than males previously housed in isolation (ISOLATE males). This is similar to the behavioral plasticity found in rodents and cats. On the other hand, courtship behavior while gonadally intact was indistinguishable between HWF and ISOLATE males. Because all males were housed individually following castration, the difference is due to different social experiences prior to castration. In Experiment 2, we found that gonadally intact HWF males had significantly elevated CO activity in the preoptic area, amygdala, and anterior and ventromedial hypothalamic areas relative to intact ISOLATE males. No significant differences in metabolism were found in the lateral septum, lateral hypothalamus, and habenula or in hindlimb muscle, suggesting that the increase in metabolism is specific to brain nuclei involved in courtship behavior. Altogether, this demonstrates that elevations in metabolic capacity correlate with experience-dependent increases in robustness to castration.     

Release of chemicals by prostaglandin-treated female fathead minnows, Pimephales promelas, that stimulate male courtship

Intramuscular injection of PGF2 alpha tromethamine salt (Sigma) did not induce sexual behavior in female fathead minnows, Pimephales promelas, when in the presence of male conspecific. However, coresident males exhibited an increased frequency of courtship behavior, consisting of approach and leading, to females following PGF2 alpha treatment. When presented with both saline-injected and PGF2 alpha-injected females, males showed increased courtship only to PGF2 alpha-treated females. Isolated males also showed increased leading behavior following the introduction of water exposed to PGF2 alpha-treated females, but not with water exposed to PGF2 alpha-treated males, untreated females or males, or PGF2 alpha alone. Hence, PGF2 alpha appears to induce the release of a female-specific chemical in P. promelas that triggers courting behavior in conspecific males.     

Hormonal control of male courtship behavior and female attractivity in the garter snake (Thamnophis sirtalis sirtalis)

Mating behavior in both intact and gonadectomized garter snakes (Thamnophis sirtalis sirtalis) was measured following hormone administration. Male courtship was androgen-dependent; subcutaneous implants of crystalline testosterone propionate (TP) pellets induced mating behavior within 2 days in both intact, reproductively inactive males and castrated males. Female attractivity, as measured by male courtship of the female, was stimulated by exogenous estrogen; 20 μg/day of estradiol benzoate (EB) was the minimum effective dose for stimulating female attractivity in both intact, reproductively inactive females and ovariectomized females. TP-implanted males selectively courted EB-primed females in both sequential and simultaneous (choice) mating tests. It is probable that males use estrogen-dependent olfactory cues produced by the females to discriminate between hormone- and vehicle-injected females.     

The postnatal demasculinization of sexual behavior in the Japanese quail (Coturnix coturnix japonica)

Three experiments were performed to analyze the time course of demasculinization in the Japanese quail and to test the activating and organizing effects of estradiol (E2) in adult sexually active birds. In Experiment 1, males and females were castrated at the age of 1 day or 1, 2, 4, and 6 weeks and treated as adults with testosterone (T). The age of castration had no effect on behavior and morphology in males. Plasma gonadotrophins (LH and FSH) were, however, higher in males castrated at or before than in those castrated after 2 weeks of age. This suggests that postnatal testicular secretions have organizing effects on the pituitary activity. Females which were castrated before 1 week of age were less sensitive to the activating effects of T than males, but were not fully demasculinized. The demasculinization of different reproductive characteristics such as male sexual behavior, cloacal gland size, and weight of the syringeal muscles is achieved in females at different times posthatching. In Experiment 2, castration of male and female quail at the ages of 4 days or 4 weeks confirmed that postnatal ovarian secretions contribute to the full behavioral and morphological demasculinization of females. It is easier to elicit mounting in T-treated females when they are tested in their home cage instead of a test arena. This difference was not observed in males. During Experiment 3, it was impossible to demasculinize sexually active adult males or females by treatment with Silastic implants of E2. E2 did not maintain sexual behavior in ovariectomized females showing male sexual behavior when treated with T but maintained the behavior in males.     

Effects of intracranial implantation of dihydrotestosterone on sexual behavior in male Cnemidophorus inornatus, a direct sexual ancestor of a parthenogenetic lizard

Dihydrotestosterone was implanted directly into the brain of castrated male Cnemidophorus inornatus, a direct sexual ancestor of the parthenogenetic species C. uniparens. Only implants located in the anterior hypothalamus--preoptic area (AH-POA) induced male-typical sexual behavior. Implants in other brain regions, including the ventromedial hypothalamus, failed to elicit courtship or copulatory behavior. Radioimmunoassay revealed no significant difference in the concentrations of circulating androgens between the responding and nonresponding animals. Previous data from this laboratory demonstrated that the AH-POA controls male-like pseudosexual behavior in C. uniparens. The current results support the hypotheses that (i) the AH-POA is the major area of hormone action in the brain controlling male-typical sexual behavior in C. inornatus as in other vertebrates and (ii) the neural circuits controlling male-typical behavior have been conserved in the evolution of the parthenogen C. uniparens.     

SEX DIFFERENCES IN MATE RECOGNITION AND CONSPECIFIC PREFERENCE IN SPECIES WITH MUTUAL MATE CHOICE

Sexual isolation is often assumed to arise because choosy females recognize and reject heterospecific males as mates. Yet in taxa in which both males and females are choosy, males might also recognize and reject heterospecific females. Here, we asked about the relative contribution of the sexes to the strong sexual isolation found in limnetic–benthic species pairs of threespine sticklebacks, which show mutual mate choice. We asked whether males and females of the two species recognize conspecifics and also prefer to mate with them. We found evidence for mate recognition by both sexes but only females prefer conspecifics. The nature of male courtship depended on which species of female they were courting, indicating that males recognized conspecific females and differentiated them from heterospecifics. However, males courted both species of females with equal vigor and changed courtship in a manner that would increase the chance of mating with heterospecifics. Females both recognized conspecifics and strongly preferred them. They responded very little to heterospecific male courtship and almost never mated with them. Therefore, males are likely to undermine sexual isolation, but females uphold it. Despite mutual mate choice and mate recognition in both sexes, females are primarily responsible for sexual isolation in these taxa.     

Preference for conspecifics evolves earlier in males than females in a sexually dimorphic radiation of fishes

Speciation by sexual selection is generally modeled as the coevolution of female preferences and elaborate male ornaments leading to behavioral (sexual) reproductive isolation. One prediction of these models is that female preference for conspecific males should evolve earlier than male preference for conspecific females in sexually dimorphic species with male ornaments. We tested that prediction in darters, a diverse group of freshwater fishes with sexually dimorphic ornamentation. Focusing on the earliest stages of divergence, we tested preference for conspecific mates in males and females of seven closely related species pairs. Contrary to expectation, male preference for conspecific females was significantly greater than female preference for conspecific males. Males in four of the 14 species significantly preferred conspecific females; whereas, females in no species significantly preferred conspecific males. Relationships between the strength of preference for conspecifics and genetic distance revealed no difference in slope between males and females, but a significant difference in intercept, also suggesting that male preference evolves earlier than females’. Our results are consistent with other recent studies in darters and suggest that the coevolution of female preferences and male ornaments may not best explain the earliest stages of behavioral isolation in this lineage.     

Males,but not females,contribute to sexual isolation between two sympatric species of <Emphasis Type="Italic">Gambusia</Emphasis>

Reproductive isolation restricts genetic exchange between species. Various pre- and post-mating barriers, such as behavior, physiology and gametic incompatibility, have been shown to evolve in sympatry. In certain scenarios, isolation can be asymmetrical, where species differentially prefer conspecifics. We examined sexual isolation via conspecific mate preference between Gambusia affinis and G. geiseri in both sexes. To investigate male contribution to sexual isolation, we compared the number of mating attempts (gonopodial thrusts) directed at either a conspecific or a heterospecific female, in both species. We also examined sperm priming and expenditure in males in the presence of conspecific or heterospecific females. We then measured female preference for either a conspecific or heterospecific male, in both species. We found that males of both species preferred to mate with conspecific females, but showed no difference in sperm production or expenditure between conspecific and heterospecific females. Females of both species did not prefer conspecific over heterospecific males. Our results suggest that sexual isolation might be mediated by male mate choice in this system and not female choice, suggesting that there is asymmetrical reproductive isolation between the sexes in G. affinis and G. geiseri, but symmetrical species isolation.     

Decapitated females as a tool in the analysis of Drosophila behaviour

Decapitated females provide a method of arranging variable stimulus environments to evaluate components of behaviour involved in sexual and frequency dependent selection. Conspecific matings are bypassed via male preference for normal females per se. Presence of conspecific decapitatated females induces male to increase insemination of alien females. Discrimination mechanisms, including sign stimuli, are the least labile, and excitability is the most labile component of courtship. D. subpalustris males with abnormal stimulation use a qualitatively different courtship element not present in their normal repertoire. An efficiency index (courtship/insemination) shows D. palustris males, with more heterogenous stimulation in their courtship, more effective in overcoming female reluctance. Sexual isolation may differ operationally from sexual selection in the necessity for particular stimuli rather than more stimulation.     

Increases in the circulating testosterone of maturing male guinea pigs appear neither necessary nor sufficient for heightened maternally directed sexual and social/courtship behavior

Periadolescent male guinea pigs housed continuously with their mother since birth exhibit little maternally directed sexual behavior. However, if rehoused apart from the mother for 24 h, they show elevations in circulating testosterone concentrations and display frequent sexual responses and increased social/courtship behavior upon reunion with her. We investigated the role of testosterone in the disinhibition of maternally directed sexual and social/courtship behavior. Subcutaneous implants of testosterone (Experiment 1) did not trigger maternally directed sexual behavior or an increase in social/courtship behavior among males housed continuously with their mothers. Further, neither blocking androgen receptors (Experiment 2) nor preventing the surge in testosterone (Experiment 3) prevented males housed without the mother from exhibiting increased maternally directed sexual and social/courtship behavior upon reunion. These findings indicate that the increase in testosterone that males exhibit when rehoused apart from the mother is neither sufficient nor necessary for the disinhibition of maternally directed sexual and social/courtship behavior observed when mother and son are reunited.     

Reproductive suppression in eusocial Cryptomys darnavensis colonies: socially-induced infertility in females

Inhibition of reproduction occurs in colonies of the Damaraland mole-rat ( Cryptomys damarensis ), where one female and usually one male are reproductively active. They remain the sole reproductive animals thoughout their stay in the colony, which in the field may exceed five years.
An 18-month study on a captive colony of C. damarensis shows that non-reproductive females remain anovulatory and always have concentrations of urinary progesterone lower than that of the reproductive female 10.7 8.8 nmols/mmol creatinine (n = 85), although their progesterone concentrations are slightly elevated when the reproductive female is early in pregnancy. In contrast the reproductively active female has elevated concentrations of progesterone 63.3 70.1 nmols/mmol creatinine (n= 14).
Standard histological together with immunohistochemical examination of the ovarian structure shows that follicular development in the non-reproductive females is halted at varying stages prior to ovulation and that the unruptured follicles luteinize. These unruptured luteinized follicles stain positive for 3β hydroxysteroid dehydrogenase—the enzyme catalysing the synthesis of progesterone from pregnenolone. It is possible that the low levels of circulating progesterone produced by the luteinized follicles in the non-reproductive females in the colony are sufficient to feedback on the hypothalamo-pituitary axis and prevent the surge of LH necessary for ovulation.
Behavioural studies provide corroborative evidence for sexual suppression. Thus the non-reproductive females in the colony are never involved in courtship or copulatory behaviour. nor do they actively solicit males in the colony. The reproductive female, however, plays an active role in mate selection and is the initiator in courtship behaviour.
The suppression of ovulation in non-reproductivc females is maintained for as long as there is a reproductive female in the colony.     

Hormonal regulation of male reproductive behavior in the lizard Anolis sagrei: a test of the aromatization hypothesis

This study examined the hypothesis that aromatization of testosterone (T) to estradiol (E) is required to activate reproductive behavior in castrated male lizards (Anolis sagrei). Adult, reproductively active males were assigned to an intact control group or to one of four treatment groups. Treatment males were castrated and 1 week later three of the four castrated groups were implanted with subcutaneous pellets containing either 0.05 mg of E, 0.5 mg of T, or 0.5 mg of dihydrotestosterone (DHT). Two weeks after pellet implantation, males were tested with stimulus males, and 2 days later were tested with stimulus females. Behavioral tests were of 15-min duration and were videotaped. Significantly fewer E-treated castrates erected a crest in tests with stimulus males than did intact males. In tests with stimulus females, significantly fewer E-treated castrates displayed, neck-gripped, and intromitted than did intact males. Estradiol-treated castrates also showed significantly less display behavior than did intact males. However, aggressive and sexual behavior of DHT-treated castrates was not significantly different from that of intact males. The same was true for T-treated castrates with the exception that display behavior in tests with stimulus females was reduced compared to that of intact males. The results suggest that aromatization of T to E is not required for induction of androgen-dependent reproductive behavior in this lizard.     

Courtship song recognition in the Drosophila melanogaster complex: heterospecific songs make females receptive in D. melanogaster, but not in D. sechellia

Abstract. The courtship song emitted by male wing vibration has been regarded as one of the most important signals in sexual isolation in the species of the Drosophila melanogaster complex. Inter- and intraspecific crosses were observed using males whose wings were removed (mute) or females whose aristae were removed (deaf). Females of D. melanogaster, D. simulans , and D. mauritiana mated with heterospecific males in the song-present condition (cross between normal females and winged males) more often than in the no-song condition (cross between normal females and wingless males or between aristaless females and winged males) or they showed no preference between the two conditions. It is possible that in these females heterospecific courtship songs play a role as if they were conspecific. In contrast, the females of D. sechellia mated with D. melanogaster or D. simulans males in the no-song condition more often than in the song-present condition, suggesting that they reject males with heterospecific song. Female mate recognition depending on the courtship song in D. melanogaster, D. simulans , and D. mauritiana is considered to be relatively broader and that in D. sechellia narrower.     

Evolutionary Changes in Dopaminergic Modulation of Courtship Behavior in Cnemidophorus Whiptail Lizards

Preoptic dopamine release is integral to the display of copulatory behaviors in male mammals and birds. However, while the anatomical distributions of the dopamine synthesizing enzyme tyrosine hydroxylase are similar among vertebrates, evolutionary changes in the functional role of dopamine are poorly understood. In this study, we tested whether a dopamine D1 receptor agonist would facilitate the display of courtship and copulatory behaviors in two related Cnemidophorine lizards (Cnemidophorus inornatus and Cnemidophorus uniparens). Cnemidophorus lizards offer a unique system to study evolutionary changes in functionality because ancestral (e.g., C. inornatus) and descendant (e.g., C. uniparens) species can be studied in parallel. Cnemidophorus uniparens is an all-female, parthenogenetic species and is the triploid descendant of the sexual and diploid species C. inornatus. Here we report that in castrated male C. inornatus and ovariectomized C. uniparens a dopamine D1 agonist increased the proportion of individuals mounting and decreased the latency to mount. Moreover, there was a species difference in sensitivity to the agonist: Mounting was elicited at a lower dose in C. uniparens than in C. inornatus. One possible explanation for this heightened sensitivity in the triploid parthenogen is that, by virtue of the increased ploidy, the parthenogen has elevated levels of D1 receptor in limbic brain areas modulating courtship behavior.     

Gonadal hormones and sexual behavior in groups of adult talapoin monkeys (Miopithecus talapoin).

Three heterosexual groups of six to eight monkeys were studied; all females were ovariectomized, whereas males were either intact or castrated. Aggressive hierarchies were evident in all groups, with females generally outranking males. When females were treated with estradiol, all males looked more frequently at the latters' sexual skin swellings, but only one male who was both dominant and intact copulated with them. Thus, either castration or low rank resulted in decreased levels of sexual behavior in male talapoins. The sexual behavior of dominant castrated males was restored by testosterone therapy, whereas subordinate castrates never copulated, even after large doses of testosterone, though penile erections and ejaculatory reflex (during masturbation) were restored. Following removal of a dominant male, the sexual behavior of the next male in rank was restored, provided he was not castrated and untreated. In contrast to males, female talapoins showed no consistent correlation between their rank and sexual activity. Estradiol therapy was without overall effect upon the frequency of female mounting behavior, though some females mounted and presented to one another more often. Estradiol treatment also caused females to present to males more frequently, but only to those that were sexually active (i.e., who mounted females).     

Fitness effects of female mate choice: preferred males are detrimental for Drosophila melanogaster females

The evolution of female mate choice, broadly defined to include any female behaviour or morphology which biases matings towards certain male phenotypes, is traditionally thought to result from direct or indirect benefits which females acquire when mating with preferred males. In contrast, new models have shown that female mate choice can be generated by sexual conflict, where preferred males may cause a fitness depression in females. Several studies have shown that female Drosophila melanogaster bias matings towards large males. Here, we use male size as a proxy for male attractiveness and test how female fitness is affected by reproducing with large or small males, under two different male densities. Females housed with large males had reduced lifespan and aged at an accelerated rate compared with females housed with small males, and increased male density depressed female fitness further. These fitness differences were due to effects on several different fitness components. Female fitness covaried negatively with male courtship rate, which suggests a cost of courtship. Mating rate increased with male size, whereas female fitness peaked at an intermediate mating rate. Our results suggest that female mate choice in D. melanogaster is, at least in part, a by-product of sexual conflict over the mating rate.     

Individual variation in intensity of sexual behaviors in captive male Cnemidophorus inornatus.

The present studies investigated the source of individual variation in intensity of sexual behaviors in captive male whiptail lizards, Cnemidophorus inornatus. No correlation was found between an individual's circulating concentration of dihydrotestosterone, testosterone, or corticosterone at the time of capture or in the laboratory and their level of sexual behaviors observed in the laboratory. A large percentage of males that initially exhibited low intensity courtship remained low intensity courters, although some became more reliable courters following 6 months of acclimation to the laboratory. Similarly, following castration and androgen replacement, most low intensity courters continued to exhibit weak and infrequent sexual behaviors. The data suggest that individual variation in sexual behaviors exhibited by captive male C. inornatus is not due to (i) low circulating concentrations of androgens, (ii) elevated circulating concentrations of corticosterone, or (iii) different profiles of testicular steroidogenesis. Rather, the source of differences may lie in (i) an inability to respond to androgens, (ii) an inability to exhibit sexual behavior, or (iii) non-hormonal stress related to captivity.     

Effects of group stability, male-male aggression, and male courtship behaviour on environmentally-induced ovarian recrudescence in the lizard Anolis carolinensis

Reproductively inactive, winter dormant female Anolis carolinensis were exposed to a stimulatory environmental regime while in socially stable or unstable male-female groups. In a second experiment, winter dormant females were exposed to varying amounts of male courtship and male-male aggression. Results indicate that (1) the presence of a stable dominance hierarchy among group males is necessary for male facilitation of environmentally-induced ovarian recrudescence (OR) (an unstable dominance hierarchy inhibits OR), (2) the behaviour patterns responsible for the facilitation and inhibition of environmentally-induced OR are courtship and male-male aggression, respectively, and (3) ovarian development is graded in accordance to the amount of male courtship to which the female is exposed. It is proposed that male courtship insures normal gonadotropin (GT) secretion while the absence of male courtship results in subnormal GT secretion and the presence of male-male aggression suppresses environmentally-induced GT secretions. The functional significance of the inhibition of environmentally-induced OR by male-male aggression and its possible relation to differential vernal emergence in temperate-zone reptiles is discussed.     

Androgenic control of male-typical behavior,morphology and sex recognition is independent of the mode of sex determination: A case study on Lichtenfelder's gecko (Eublepharidae: Goniurosaurus lichtenfelderi)

Previous work on lizards has shown that many sexually dimorphic traits depend on testosterone (T), but the details of this control can vary among species. Here, we tested the role of T on the expression of morphological, physiological, and behavioral traits in Lichtenfelder's gecko (Goniurosaurus lichtenfelderi), from the lizard family Eublepharidae notable for interspecific variation in sexually dimorphic traits and the mode of sex determination. Experiments included three groups of males (intact control, surgically castrated, castrated with T replacement) and two groups of females (intact control, T supplemented). In males, castration caused reductions in 1) the size of hemipenes, 2) offensive aggression, 3) male sexual behavior in a neutral arena, 4) activity of precloacal glands, and 5) loss of male chemical cues for sex recognition. These reductions were not observed in castrated males with T replacement. Interestingly, castrated males performed sexual behavior in their home cages, which shows that the effect of T depends on the environmental context. Notably, tail vibration, previously reported as a courtship behavior in other eublepharids, is displayed by males of G. lichtenfelderi during interactions with conspecifics of both sexes, suggesting an evolutionary shift in the meaning of this signal. In females, T induced growth of hemipenes and male-typical courtship but did not induce precloacal pore activity, aggression, or mounting. In comparison to previous reports on Eublepharis macularius, our results indicate that effects of T do not depend on the mode of sex determination. Further, our results extend our understanding of the complexity of control of male traits and illustrate how lability in the effects of T can be a general mechanism causing evolutionary changes in the components of suites of functionally correlated traits.