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Changes in plasma levels of gonadal steroids during spawning behaviour in territorial male demoiselles Chromis dispilus (Pisces: Pomacentridae) sampled underwater.

Males of the demoiselle Chromis dispilus (a species endemic to New Zealand) were hand-netted from nest territories by scuba divers and blood sampled in situ. Reproductive condition and behaviour prior to capture were compared with plasma levels of testosterone (T) and 17 alpha,20 beta dihydroxy-4-pregnen-3-one (17,20 beta P). Spawning cycles of about 7 days, consisting of 1-2 days of spawning activity followed by 4-5 days of brooding behaviour, were observed. Plasma levels of T and 17,20 beta P were elevated (4-6 and 1-1.5 ng.ml-1, respectively) in fish that were engaged in courtship display, spawning, or guarding freshly spawned eggs. Levels of both hormones fell to less than 1 (T) and less than 0.4 ng.ml-1 (17,20 beta P) at the end of spawning episodes and remained low through the subsequent brooding period. T levels increased slightly toward the end of the brooding phase. Samples taken at 2- to 3-hr intervals during daylight hours showed similar correlation of steroid levels and activity; however, there was a marked fall in 17,20 beta P in spawning fish with the onset of shelter seeking at dusk. Values returned to high levels with the resumption of spawning the next day. Plasma steroid levels were not consistently correlated with reproductive condition (degree of spermiation). It is concluded that there is a strong correlation between plasma T and 17,20 beta P levels and the male spawning cycle, but not brooding or territorial behaviour per se.

Animals↗

Endocrine and behavioral responses to aggression and social dominance in the green anole lizard, Anolis carolinensis.

Adult males of the small arboreal iguanid lizard, Anolis carolinensis, will fight and form social dominance hierarchies when placed in habitats with limited resources. The relationships between time since initial aggressive interaction, relative social dominance, reproductive activity, and corticosterone and androgen levels were determined for 34 pairs of lizards. A discriminant analysis established a "dominance index" which indicated that over 90% of the difference between individuals who had won or lost aggressive interactions (putative social dominants and subordinates) was attributable to a single discriminant function reflecting altered body color, perch site selection, and circulating androgen. Animals that had darker body color also selected lower perch sites and had depressed rates of courtship relative to winners of fights and were thus designated as social subordinates. These animals also had levels of circulating androgen significantly lower than that of dominants, but circulating corticosterone was not significantly affected. Winners of fights showed a dramatic surge in circulating androgen at 1 hr but returned to near control values by 1 week; losers, however, showed depressed circulating androgen levels at 1 week.

Aggression↗

Thyroid and adrenal function in the ring dove (Streptopelia risoria) during food deprivation and a breeding cycle.

The concentration of plasma triiodothyronine (T3), thyroxine (T4), corticosterone, and glucose as well as body weight and food intake were measured in pairs of ring doves throughout a breeding cycle and during food deprivation. During courtship and incubation the levels of plasma T4 were significantly higher in males than in females. A transitory decrease in food intake in both sexes was measured at the onset of incubation but was not associated with a change in body weight or in levels of plasma thyroid hormones. After the eggs had hatched, food intake increased but was associated with a reduction in body weight and concentrations of plasma T3 and T4, although plasma corticosterone and glucose both increased. Food deprivation for 48 hr resulted in a significant fall in the concentration of plasma T3 and an increase in plasma corticosterone. These changes suggest that all the food was not being digested by the adult birds during brooding but was almost exclusively regurgitated to feed the squabs.

Adrenal Glands↗

Prostaglandin induction of spawning behavior in Cichlasoma bimaculatum (Pisces cichlidae).

Prostaglandin (PG) stimulates female spawning behavior in goldfish and in some other teleosts in which female reproductive behaviors consist of postovulatory oviposition acts. This study examined the effects of PG on female sexual behavior in a teleost fish, Cichlasoma bimaculatum, in which female reproductive behaviors involve both preovulatory courtship and substrate cleaning behaviors, and post-ovulatory oviposition behavior. In females of established pairs, PGF2 alpha injection (5 micrograms, im) at any stage of the spawning cycle, or in the parental phase, rapidly induced substrate cleaning which soon merged into oviposition behavior (without egg release). These results support a role for PG in oviposition behavior of Cichlasoma. However, indomethacin (1 mg, ip), a PG synthesis inhibitor, did not block oviposition in ovulated females which had begun to spawn. Indomethacin may not have lowered PG levels sufficiently. Alternatively, as shown by J.J. Polder (1971, Neth. J. Zool. 21, 265), oviposition behavior may be induced or maintained by other factors associated with the spawning situation.

Animals↗

The effects of early and late androgen treatments on the behavior of Sarotherodon mossambicus (Pisces: Cichlidae).

Groups of Sarotherodon mossambicus were treated with androgen by immersion or oral treatment at various stages of development. Fish were allowed to mature and the effects of treatment on gonadal and behavioral differentiation were examined. The effects of treatment on gonadal differentiation were assessed by determining the sex ratio for each group. Three treatments resulted in sex ratios significantly different from the 1:1 sex ratio obtained in untreated fish. Behavioral differences were detected between groups of males in three measures of territorial defense and aggression, and differences were detected between groups of females in two measures of male-female courtship interaction. A second experiment determined that early-treated females were more sensitive to a second androgen treatment later in life than females not exposed to androgen during development. A number of sex-reversed genetic females functioning as males were detected in two treatment groups with predominantly male sex ratios. There were no differences in the behavior of sex-reversed and non-sex-reversed male fish from the same treatment group. This study establishes that hormone treatments administered during development influence behavioral differentiation in a teleost.

Age Factors↗

Seasonality of adult male Japanese macaques (Macaca fuscata): androgens and behavior in a confined troop.

Seasonal variations in levels of serum testosterone, dihydrotestosterone (DHT), reproductive behavior, and social behavior were investigated in 12 adult males (5 to 20+ years of age) of the Oregon troop of Japanese macaques (Macaca fuscata). Blood samples were collected at 2- to 4-month intervals, and behaviors were monitored twice weekly over a 15-month period. Significant seasonal variations in levels of testosterone and DHT, and in frequencies of mount series, ejaculations, number of female partners, displays, courtship, and aggression were observed. Seasonal variations in reproductive and social behaviors did not correlate with seasonal variations in androgen levels because seasonal increases in these behaviors followed seasonal increases in the androgens with a 1- to 2-month delay. However, significant correlations between increased androgen levels and the onset of mating activity occur when mean monthly frequencies of mount series are shifted 1 to 2 months earlier to coincide with the rise in serum androgen levels. The frequency of adult male play and male-male mounting increased significantly when androgen levels were low. We suggest that photoperiod changes may function as a proximate cue in male Japanese macaques which induces a state of biological readiness for mating, and the behavioral consequences (i.e., mating) are then dependent upon the presence of receptive females.

Agonistic Behavior↗

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.

Animals↗

Altered endocrine and behavioral responses with reproductive aging in the male Japanese quail.

Experiments were conducted to further characterize the age-related decline in reproductive capability previously described in the male Japanese quail. Behavioral testing of a large number of males in several age groups confirmed earlier results that showed a significant reduction in courtship and mating behavior by 80 weeks of age. Intact males, 78 weeks of age, that showed no reproductive behavior were given Silastic implants containing testosterone. This treatment restored behavior in 80% of the males. Gonadal regression induced by exposure to shortened photoperiod followed by transfer to a stimulatory photoperiod did not successfully restore the endocrine or behavioral components of reproduction. Males of two age groups, 42 and 208 weeks of age, were categorized as behaviorally active or inactive and then castrated. Eight weeks later they were given implants containing testosterone. Older males required longer exposure time to the exogenous steroid and in some cases, an additional implant to restore mating behavior as compared to younger males. The presence or absence of behavioral activity prior to castration did not affect the length of time required for restoration of behavior. Challenge with exogenous luteinizing hormone-releasing hormone revealed that there was reduced release of luteinizing hormone in aged, behaviorally inactive males as compared to young males. Results of these experiments in addition to previous data give evidence that age-related changes occur at the level of the hypothalamus and pituitary as well as at the level of the gonads.

Aging↗

Hormonal control of sex differences in ultrasound production by hamsters.

Male and female hamsters differ in the stimulus control of the ultrasounds they produce during courtship and mating. In particular, untreated males show greater increases in ultrasound rate after exposure to stimulus females than after contact with other males. Conversely, estrous females are more responsive to stimulus males than females. This sex difference reflects both organizational and activational effects of gonadal hormones. Thus, responses to early castration or treatment with testosterone propionate (TP), estradiol benzoate (EB), or dihydrotestosterone propionate suggest that the development of male-like patterns of ultrasound production is facilitated by perinatal exposure to aromatizable androgen. However, even neonatally feminized subjects will show male-like calling if tested during adult treatment with TP. In contrast, the same subjects respond like naturally estrous females during adult treatment with EB plus progesterone (P). The contrasting responses of neonatally feminized subjects to later TP and EB + P treatments suggest that female hamsters retain a greater capacity for heterotypical patterns of ultrasound production than do males. This obviously differs from the common observation of greater "bipotentiality" for mating behavior in males. In turn, this suggests that the mechanisms controlling sexual bipotentiality are specific to their target behaviors, yielding distinct patterns of hormonal control for at least ultrasound production and lordosis.

Animals↗

Aromatase inhibition depresses ultrasound production and copulation in male hamsters.

Rates of ultrasound production and copulatory behavior were observed in castrated male hamsters maintained on 100 micrograms/day of injected testosterone propionate (TP). Groups matched on their initial levels of behavior received either continued treatment with TP alone, or TP together with 6 mg/day injections of the aromatase inhibitor 1,4,6-androstatriene-3,17-dione (ATD). Testing at 11-15 days after the start of these treatments revealed deficits in the sexual behaviors of the subjects in the latter group. Specifically, these males showed lower rates of ultrasound production and intromission during, as opposed to before, treatment with ATD. These results support previous work suggesting that aromatization plays significant roles in the mediation of androgenic effects on both the courtship and copulatory behaviors of male hamsters.

Androstatrienes↗

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.

Animals↗

Activation of sexual behavior by implantation of testosterone propionate and estradiol benzoate into the preoptic area of the male Japanese quail (Coturnix japonica).

Intracranial implantation of minute pellets of gonadal steroids was performed to determine neuroanatomical loci at which steroids activate sexual behavior in the Japanese quail (Coturnix japonica). In this species, systemic treatment of castrated males with either testosterone propionate (TP) or estradiol benzoate (EB) restores male-typical copulatory behavior (head grabbing, mounting, and cloacal contact movements). In addition, EB activates female-typical receptive behavior (crouching). Adult male castrated quail were implanted intracranially with 300-micrograms pellets containing TP, EB, or cholesterol (CHOL) and behavior was tested with intact males and females. Either TP or EB pellets in the preoptic area (POA) activated male-typical copulatory behavior. Mounting was specifically activated without concomitant activation of other steroid-sensitive sexual and courtship behaviors. TP and EB implants in adjacent nuclei containing receptors for these steroids and CHOL implants in POA had no effect on male-typical copulatory behavior. Eighteen percent of all males tested for female-typical receptivity crouched, but no specific effect of EB was seen at any site. The similarity of the POA sites for activation of mounting by TP and EB is consistent with the hypothesis that cells within the POA aromatize testosterone to estrogens, which directly stimulate the cellular processes leading to behavioral activation.

Animals↗

Vasoactive intestinal polypeptide-like immunoreactivity during reproduction in doves: influence of experience and number of offspring.

To assess the possibility that vasoactive intestinal polypeptide (VIP) plays a role in naturally occurring changes in prolactin secretion in ring doves, we used immunohistochemical techniques to measure VIP-like immunoreactivity in the brain as a function of stage of the reproductive cycle. Differences between parental and nonparental birds in VIP profiles were detected in the ventral portion of the infundibular region. More specifically, there is an increase in cell size and staining intensity in the ventral infundibulum of breeding birds compared to simultaneously processed tissue taken from control animals. In both sexes, an increase in size of VIP-like immunoreactive cells is detectable during courtship and early incubation, anticipating the increase in plasma prolactin levels. VIP cell size remains elevated from about Incubation Day 14 to Brooding Day 14, and a steady decrease is observed during the remaining posthatching period, as squab begin to feed independently. Compared to parents rearing one squab, those with two young have a prolonged interval of increased infundibular VIP immunoreactivity. Furthermore, doves with no previous experience of a breeding cycle exhibit prolonged VIP-like immunoreactivity compared to experienced parents, paralleling previously described differences between these groups in parental behavior.

Animals↗

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.

Animals↗

The vomeronasal organ: primary role in mouse chemosensory gender recognition.

Four experiments were conducted to determine the chemosensory modality that supports ultrasonic courtship vocalizations by male mice to females and to chemosignals from females. Both removal of the olfactory bulbs (Experiment 1) and removal of the vomeronasal organ (Experiments 2-4) produced similar deficits in the pattern of ultrasonic vocalizations elicited by conspecifics or their odors. Removal of the vomeronasal organ did not impair the ability to locate food buried under cage shavings. These results are consistent with the notion that the analysis of food related odors is subserved by olfaction and that vocalizations to sex chemosignals are elicited primarily by stimulation of the vomeronasal organ/accessory olfactory bulb. Removal of the vomeronasal organ did not induce seminal vesicle regression or lower plasma immunoreactive testosterone levels (Experiment 2) nor was an attempt to restore vocalizations with exogenous testosterone successful (Experiment 4). Thus the altered vocalization pattern following removal of the vomeronasal organ does not appear to arise as a motivational deficit mediated by androgens. Experiments 2 and 3 demonstrated that, in the absence of the vomeronasal organ, stimulation of other sensory systems can, to some extent, maintain the male's tendency to vocalize more to females or their odors than to males or their odors. However, this responsiveness to females may rely upon additional behavioral cues. Previous experience also plays a considerable role in the response to chemosensory gender cues by males who lack their vomeronasal organs. Removal of the vomeronasal organ prior to adult heterosexual encounters (Experiment 3) virtually eliminated the male's responsiveness to either anesthetized females or their chemosignals. Hence males require adult heterosexual experience with a functioning vomeronasal organ before other chemosensory systems acquire the ability to mediate gender recognition as measured by ultrasonic vocalizations.

Animals↗

Effects of forebrain lesions on spawning behaviour in the male goldfish.

Lesions were stereotaxically placed in medial nuclei of the ventral telencephalon and preoptic area of male goldfish. Only lesions in the area ventralis telencephali pars supracommissuralis (Vs) and posterior pars ventralis (pVv) were effective in reducing the proportion of males spawning, as compared to sham groups, both 5 days (Experiment 1) and up to 4 weeks (Experiment 2) postoperatively. Spawning consistency over 7 weekly tests was negatively correlated with the volume of Vs-pVv destruction. Two-thirds of Vs-pVv lesioned males spawned on at least one of their weekly tests, with latencies for the onset of each courtship behaviour similar to those of control fish, partial performance of the spawning sequence was rare. These results suggest that lesion of the supracommissural telencephalon (Vs-pVv region) blocks the initiation of spawning behaviour in the male goldfish, perhaps by lowering reproductive motivation specifically or by interfering with the perception of sexual, particularly olfactory, cues.

Animals↗

Ultrasonic vocalizations in immature gerbils: emission rate and structural changes after neonatal exposure to androgen.

Emission rates of stereotyped ultrasonic vocalizations in sexually immature Mongolian gerbils are sexually dimorphic. Calling rates in interacting pairs of females are characteristically higher than in male dyads. Juvenile vocalizations produced during paired encounters show a similar spectrographic structure to adult courtship calls. Three experiments examined the developmental effects of a single injection of 100 micrograms testosterone propionate given to 0-12 h old female gerbils on total emission rates, spectrographic structure of ultrasonic vocalizations and display of other social-sexual play behaviors during encounters between immature androgenized females and stimulus partners. Reduced total rates of juvenile ultrasound emission were recorded in TP-treated females paired with stimulus males or females when compared with control male-male or female-female pairs in Experiment 1 and confirmed in Experiment 2. Male-typical play mounting was increased in pairs containing androgenized females in Experiment 2. In Experiment 3, emission rates of female-type Warble sounds were significantly decreased and the rate of male-type Steep curvilinear and Long rectilinear calls significantly increased in androgenized female and stimulus animal dyads compared with control animal pairs. New spectrographic forms of ultrasound, with readily discernible single or multiple upper harmonics, were recorded in androgenized female-stimulus partner interactions. We conclude that androgenization of female neonates (a) masculinized and also defeminized sexually dimorphic ultrasound emission rates, and virilized nonvocal behavior, suggesting a direct effect on organization of the central nervous system; (b) influenced the sonagraphic structure of calls. It is proposed that organization of juvenile vocal rates in gerbils primarily involves a number of androgen sensitive mechanisms in the central and peripheral nervous systems. However, androgenization also appeared to have an indirect role on calling rates by affecting treated female-stimulus partner relationships.

Age Factors↗

Mating increases plasma levels of prostaglandin F2 alpha in female garter snakes.

Plasma levels of prostaglandin F2 alpha (PGF2 alpha) in female red-sided garter snakes (Thamnophis sirtalis parietalis) were measured at intervals after mating or exposure to males. PGF2 alpha levels increased significantly within 15 minutes of mating and peaked 6-24 hr after mating. Females that did not mate, but received similar amounts of male courtship, had levels of PGF2 alpha significantly lower than those of females that mated. These results extend previous findings that unmated female garter snakes injected with PGF2 alpha exhibit sexual behavior characteristic of females that have mated. Together these data indicate that female garter snakes elaborate PGF2 alpha in response to stimuli associated with mating and that PGF2 alpha has a functional role in inducing post-mating declines in sexual behavior of this species.

Animals↗