Seasonal changes of serum sex steroids concentration and aromatase activity of gonad and brain in red-spotted grouper (Epinephelus akaara)

Levels of serum sex steroids (estradiol-17beta, E2; testosterone, T; 11-ketotestosterone, 11-KT) in male, female and natural sex-reversing red-spotted grouper (Epinephelus akaara), and aromatase activity of gonad and brain in both male and female were investigated throughout an annually reproductive cycle. In females, serum E2 and T peaked during vitellogenesis, but in males and natural sex-reversing fish, 11-KT, T and E2 reached peak during spermatogenesis. In addition, in females, serum 11-KT levels (monthly means: 0.32 +/- 0.03 ng/ml) which were very low did not significantly fluctuate during the annual reproductive cycle. In breeding season, females displayed higher E2 levels than males and sex-reversing fish, while males and sex-reversing fish showed higher 11-KT levels and, to a lesser extent, higher T levels than females. Furthermore, the changing pattern of sex steroids in males was similar to that in natural sex-reversing fish, and a second peak of serum androgens 11-KT and T appeared in December both in male and natural sex-reversing fish; significantly higher serum 11-KT levels were observed in natural sex-reversing fish than that in females from December to April. In females, but not in males, aromatase activity of brain and gonad demonstrated significantly seasonal changes (exhibiting a peak in breeding season); moreover, aromatase activity in females was higher than that in males. Furthermore, significantly lower aromatase activity in testis was observed in breeding season, in contrast to that in ovary. Taken together, the present findings indicated that changes of serum sex steroids levels and aromatase activity in red-spotted grouper were closely associated with sex inversion. In addition, the present results also suggested that sex inversion in red-spotted grouper peaked mainly from December to March.     

Effects of constant short and long photoperiod regimes on the spawning performance and sex steroid levels of female and male sea bass

Under constant short photoperiod, the spawning time of 2-year-old sea bass Dicentrarchus labrax was advanced as compared to controls, whereas spawnings were delayed under constant long photoperiod. High plasma levels of 17β-oestradiol (E_2/) and testosterone (T) in females were coincident with the appearance of vitellogenic oocytes in the ovary, while high levels of 11-ketotestosterone (11-KT) and T in males were coincident with the presence of spermiating males. Although plasma levels of E₂ in females and 11-KT in males were low during the remainder of the cycle, levels of T were always >1 ng ml⁻¹ in both sexes, suggesting that T could play an important role during the initial stages of gonadal development. The profiles of E₂ and T in females and 11-KT and T in males exposed to constant short days were similar to those in the control group, but fish which were maintained under constant long photoperiods showed a bimodal pattern of these steroids. The results obtained from fish exposed to constant photoperiod regimes provide further evidence that an endogenous process could be operating to control the reproduction of sea bass.     

Equivalent whole-body concentrations of 11-ketotestosterone in female and male coral goby Gobiodon erythrospilus, a bidirectional sex-changing fish

The relationship between whole-body concentrations of 11-ketotestosterone (11-KT) and sexual function was examined in the coral goby Gobiodon erythrospilus , a bi-directional sex-changing fish. 11-KT occurred in both female and male G. erythrospilus , but levels were not always higher in males than in females within heterosexual pairs, and were not related to the stage of gonadal development of individual fish. These results suggest that comparable 11-KT levels in both sexes may allow serial adult sex change to take place in bi-directional sex-changing species, such as Gobiodon spp.     

The involvement of sex steroid hormones in downstream and upstream migratory behavior of masu salmon

From May through July when masu salmon, Oncorhynchus masou, commence downstream migration under natural conditions, yearling precocious male masu salmon (resident form) showed higher GSI and plasma levels of testosterone (T) and 11-ketotestosterone (11-KT) in contrast to immature smolts (migratory form). From March through September coinciding with the upstream migration period, 2-year-old male and female adults also showed higher GSI and plasma levels of T, estradiol-17beta (E(2)) 11-KT, 17alpha-hydroxyprogesterone and 17alpha,20beta-dihydroxy-4-pregnene-3-one (DHP). In order to test the effects of steroid hormones on migratory behaviors, silascone tube capsules containing 500 microg of T, E(2), 11-KT, DHP, or a vehicle was implanted into smolts, castrated precocious males, or immature parr, and downstream and upstream behavior were observed in artificial raceways in spring and autumn. Downstream behavior of smolts was inhibited significantly by T, E(2) and 11-KT. Upstream behavior was stimulated by T and 11-KT in castrated precocious males and stimulated by T, E(2) and 11-KT in immature parr. These results indicate that T, E(2) and 11-KT are the factors regulating downstream and upstream migratory behavior. In particular, because of its changing patterns in plasma and significant effects, T, the common precursor hormone of E(2) (female) and 11-KT (male), is considered to play central roles in both types of behavior.     

Endocrine and morphological correlates of reproduction in the draughtboard shark Cephaloscyllium laticeps (Elasmobranchii: Scyliorhinidae)

This study examined the endocrine and reproductive correlates of reproduction in 636 female and 468 male draughtboard sharks (Cephaloscyllium laticeps) captured from southeastern Australia. Females were oviparous and displayed a single external-type ovary with a maximum follicle diameter of 35 mm. Vitellogenesis commenced at a follicle diameter of 10 mm. Females showed a constant overlap between follicular recruitment, ovarian growth, and egg laying. The male reproductive tract consisted of paired testes with spermatocysts undergoing diametric development. Plasma levels of the presumptive gonadal steroids, testosterone (T), 17beta-estradiol (E2), progesterone (P4), and 11-ketotestosterone (11-KT; males only) were correlated with morphological developmental stages of the gonads. In females, E2 increased as the follicle developed before declining as the follicle reached maturity. T remained low during the first stages of ovarian growth and increased as the follicle reached maturity. P4 showed a peak just before ovulation. In males, T was the only hormone that varied with maturity, increasing in adults; E2 and P4 were present at low plasma concentrations in males and did not change with stage of gonadal development. 11-KT was undetectable at all times. Endocrine changes in draughtboard sharks were consistent with hormonal correlates reported for other species and suggest roles for E2( in females) and T (in both sexes) in gametogenesis and P4 in maturational events in females.     

Inter-male aggression with regard to polygynous mating system in Pampean grassland mouse, Akodon azarae (Cricetidae: Sigmodontinae)

Based on the hypothesis that, in Akodon azarae, polygyny operates through female defence, we studied inter-male aggression in order to test the following predictions: during the breeding period (1) resident males are more aggressive than intruder males in the presence of females (FP), and (2) aggressive behaviour is independent of male condition (resident or intruder) in the absence of females (FA). To test our predictions, we used the resident male behavioural response towards an intruder male in relation to FP or FA. We conducted 30 encounters in FP and 27 in FA in 0.79-m² round enclosures placed in the Espinal Reservation. Our results support the prediction that, in FP, the intensity of aggressive behaviour exhibited by males varied in relation to resident or intruder condition. Resident males showed high levels of aggression towards intruders, and intruders exhibited the greatest values of submissive behaviours with residents. In FA, the intensity of aggressive behaviour did not vary in relation to resident or intruder condition. Both resident and intruder males exhibited low aggressive behaviour and inter-male encounters resulted mainly in non interactive behaviours. Our results support the hypothesis that, in A. azarae, the polygynous mating system operates through female defence.     

Male-male competition and female choice in brown trout

In some salmonid species, the females have been assumed to choose their mates on the size of the male's adipose fin. This hypothesis was tested in a stream water aquarium, in which 19 brown trout, Salmo trutta, females were allowed to choose between two males of the same body size but with different adipose fin sizes. The two males were separated from each other in cages. After the female had started to prepare her nest close to one of them, the males were released and allowed to fight each other for the opportunity to spawn. Out of 19 females, 14 prepared a nest closest to the male with the larger adipose fin. However, only six of the 14 females spawned with this male. Males that spawned were more dominant (i.e. were more likely to win fights). When the female spawned with the male she chose, he was less aggressive towards her than when she spawned with the other male. There were no significant differences in the plasma levels of testosterone (T) and 11-ketotestosterone (11-KT) between the chosen males and those not chosen. However, the dominant males had significantly higher plasma levels of T and 11-KT both before and after the experiment. The results support the view that female brown trout exhibit mate choice, but their choice is overruled by male-male competition. Copyright 1999 The Association for the Study of Animal Behaviour.     

Effect of elevated summer temperatures on gonadal steroid production, vitellogenesis and egg quality in female Atlantic salmon

Sex change in the coral-dwelling goby Gobiodon histrio was induced by placing two adult fish of the same sex on a coral colony. The sex change of individual fish was confirmed using histology, and whole-body concentrations of the gonadal steroids testosterone (T), 11-ketotestosterone (11-KT), and 17β-oestradiol (E₂) were examined. The results show that T, 11-KT and E₂ occurred in both female and male G. histrio . E₂ concentration in females was twice that in males, while concentrations of T did not differ between the sexes. Contrary to predictions, concentrations of T and E₂ did not differ between fish that changed sex and those that did not. Most samples had 11-KT concentrations below minimum levels of detection (  i.e. <0·15 ng ml⁻¹) and were therefore not analysed statistically. The results suggest that: (i) specific activation or de-activation of the T–E₂ (aromatase) pathway is a probable candidate for mediating serial adult sex change in G. histrio , and (ii) low levels of 11-KT may be important in allowing serial adult sex change in G. histrio .     

A multidisciplinary study on social status and the relationship between inter-individual variation in hormone levels and agonistic behavior in a Neotropical cichlid fish

Social animals with hierarchal dominance systems are particularly susceptible to their social environment. There, interactions with conspecifics and hierarchal position can greatly affect an individual's behavior, physiology and reproductive success. Our experimental model, Cichlasoma dimerus, is a serially-monogamous Neotropical cichlid fish with a hierarchical social system, established and sustained through agonistic interactions. In this work, we aimed to describe C. dimerus social structure and its association with hormonal profiles and testicular cellular composition. We recorded and quantified agonistic interactions from the territorial pair, i.e. the top ranked male and female, and the lowest ranked male of stable social groups. Plasma levels of 11-ketotestosterone (11-KT), testosterone, 17β-estradiol (E₂) and cortisol were measured by ELISA. Results show that territorial pairs cooperatively guarded the territory, but rarely attacked in synchrony. Territorial males had higher testosterone and 11-KT plasma levels than non-territorial males, while E₂ and an index of its metabolization from testosterone were higher in non-territorial males. No difference was observed in cortisol levels. Plasma 11-KT and an index of the conversion of testosterone to 11-KT, positively correlated with the frequency of aggressiveness, while E₂ showed the opposite pattern. Territorial males had a higher gonadosomatic index than non-territorial males. The quantification of testicular cellular types revealed that the percentage of spermatocytes and spermatids was higher in non-territorial males, while territorial males showed a greater percentage of spermatozoa. Thus, C. dimerus male social position within a stable hierarchy is associated with distinct behaviors, steroid levels and testicular degree of development.     

Contrasting seasonal modulation of the stress response in male and female rainbow trout

Between June and September the magnitude of the plasma cortisol response of maturing male and female rainbow trout to confinement was indistinguishable. A progressive increase in confinement-induced cortisol levels in both sexes occurred during May to September, associated with the seasonal rise in water temperature. Between September and January a reduction of >50% in the magnitude of the cortisol response to confinement in male fish (but no decline in females) coincided with declining water temperature and significant increases in plasma 11-ketotestosterone and elevated plasma testosterone levels. Plasma oestradiol-17β levels were significantly greater in females than males throughout the study period and this difference was maximal between September and January. However, plasma testosterone was also elevated in females during this period and levels overall were higher than those in male fish. Previous studies have shown oestradiol-17β and testosterone to have diametrically opposed effects on stress responsiveness in trout, with the former enhancing, and the latter suppressing, the cortisol response to a stressor. The relative roles of androgens, estrogen and water temperature in modulating the stress responsiveness of rainbow trout are discussed.     

Subspecies differences in hormonal and behavioral responses after group formation in squirrel monkeys

The behavioral and hormonal responses of squirrel monkeys of the Bolivian and Guyanese subspecies were compared after a group formation procedure. Two groups of each subspecies, consisting of five females and three males (later reduced to two) were observed daily during the week of group formation and for nine weeks following removal of a single male from each group. Measures of plasma cortisol were examined in the females after the initial group formation and after the groups were reduced by one male. The levels of plasma testosterone were assessed in all the males during the initial week of group formation. Linear dominance hierarchies were determined both within and across the sexes in both subspecies. The frequency and directionality of low-level aggressive interactions indicated that females of the Bolivian subspecies were dominant to the males, while the males of the Guyanese subspecies ranked over all the females. Additionally, the Bolivian squirrel monkey females showed an elevation of plasma cortisol on the day of group formation, which declined 48 hr later, then reelevated after the groups were reduced by one male and declined gradually over a nine-week period. Guyanese females showed little change in cortisol levels during both periods. This suggests fundamental hormonal, as well as behavioral, differences between the two subspecies. The change in plasma testosterone levels in the males during the initial week of group formation was positively correlated with social status. Furthermore, differences in the dynamics within individual groups for each subspecies were reflected by the levels of plasma cortisol of the female members.     

Androgen correlates of socially induced changes in the electric organ discharge waveform of a mormyrid fish

Weakly electric fish from the family Mormyridae produce pulsatile electric organ discharges (EODs) for use in communication. For many species, male EODs are seasonally longer in duration than those of females, and among males, there are also individual differences in EOD duration. While EOD elongation can be induced by the administration of exogenous androgens, androgen levels have never before been assessed under natural or seminatural conditions. By simulating the conditions occurring during the breeding season in the laboratory, we provide evidence of a sex difference in EOD duration as well as document levels of circulating androgens in males. In this study, we analyzed the nature of social influences on male EOD duration and plasma androgen levels in Brienomyrus brachyistius. Individual males, first housed with a single female and then placed into social groups consisting of three males and three females, showed status-dependent changes in EOD duration. Top-ranking males experienced a relatively large increase in EOD duration. Second-ranking males experienced a more modest increase, and low-ranking males experienced a decrease in EOD duration. These changes were paralleled by differences in circulating levels of plasma 11-ketotestosterone (11-KT), but not testosterone, suggesting that the changes in EOD duration may have been mediated by changes in plasma 11-KT levels. Thus, it appears that EOD duration is an accurate indicator of male status, which is under social and hormonal control.     

Endocrine and metabolic responses to stress in a laboratory population of the tropical damselfish Acanthochromis polyacanthus

The effects of confinement and exercise on the stress response of the spiny damselfish Acanthochromis polyacanthus were investigated in a laboratory stock of fish. Cultured spiny damselfish had basal plasma cortisol values (<16 ng ml⁻¹) similar to those found in wild fish, and basal plasma glucose and lactate levels that were similar to those found in other teleosts. Plasma cortisol concentrations increased in response to stress with a latency period of 5–10 min. Removal of the stressor resulted in partial recovery of cortisol levels by 24 h. Plasma glucose levels increased in response to stress in all experiments with significant increases occurring within 15 min of the imposition of stress. Elevations in plasma glucose concentrations were not initially reflected in changes in liver or muscle glycogen content, with significant reductions in liver glycogen concentrations only occurring in response to extended periods of stress. In contrast to many temperate species, plasma lactate concentrations did not consistently increase in response to stress, suggesting that the stress response in spiny damselfish is not strongly characterized by anaerobiosis.     

Serum cortisol concentrations of single-housed and isosexually pair-housed adult rhesus macaques

Possible social distress was evaluated in 20 adult rhesus macaques housed in compatible isosexual pairs (5 female pairs, 5 male pairs) for the purpose of social environmental enrichment. Serum cortisol concentrations of paired animals were compared with serum cortisol concentrations of individually housed adult rhesus macaques of both sexes (5 females, 5 males). In both sexes, cortisol concentrations of paired animals (means 10 females = 19.5 +/- 2.9 micrograms/dl; means 10 males = 17.5 +/- 4.6 micrograms/dl) showed no significant difference (p always greater than 0.1) with those of single animals (means 5 females = 20.5 +/- 2.1 micrograms/dl; means 5 males = 15.9 +/- 2.6 micrograms/dl). Both in male and in female pairs, dominant partners had cortisol concentrations that were equivalent to those of their subordinate counterparts. It was concluded that neither female nor male adult rhesus macaques experience more distress when sharing a cage with a compatible partner of the same sex than when living alone.     

Endocrine control of sexual behavior in sneaker males of the peacock blenny Salaria pavo: effects of castration, aromatase inhibition, testosterone and estradiol

The effects of castration and sex steroid manipulations on the expression of sexual behavior were investigated in a small fish, the peacock blenny, Salaria pavo. In this species, large males defend nests and attract females while small "sneaker" males reproduce by imitating the female morphology and courtship behavior in order to approach nests during spawning events and parasitically fertilize eggs. Sneakers switch into nest holders in their second breeding season, thus displaying both male and female-like sexual behavior during their lifetime. We tested the effects of castration and of an aromatase inhibitor (Fadrozole, F), testosterone (T) or 17beta-estradiol (E(2)) implants on the expression of male and female-like behavior in sneakers. Sneakers were either sham-operated, castrated or castrated and implanted with vehicle, F, T+F or E(2)+F. Seven days after the treatment, sneakers were placed in a tank with a nesting male, two ripe females and an available nest. Castrated fish had lower levels of circulating T and increased the time spent displaying female typical nuptial coloration. T implants had the opposite effect, inhibiting the expression of female-like behavior and coloration. E(2) implants had no significant effect on the display of sexual behavior but the frequency of aggressive displays decreased. The results agree with previous findings in sneakers of S. pavo that demonstrated an inhibition of female-like behavior by 11-ketotestosterone (11-KT). The reported increase in T and 11-KT production when sneakers change into nest holders may thus contribute to behaviorally defeminize sneakers. Contrarily, both T and E(2) failed to promote male-like behavior, suggesting that behavioral masculization during tactic switching depends on other neuroendocrine mechanisms or that the time length of the experiment was insufficient to induce male-like behavioral changes in sneakers.     

Lack of species-specific primer effects of odours from female Atlantic salmon, Salmo salar, and brown trout, Salmo trutta

We exposed, in two successive spawning seasons, individually placed precocious male Atlantic salmon ( Salmo salar ) and brown trout ( Salmo trutta ) parr to odour stimuli (ovarian fluid and urine mix) from ovulated conspecific or heterospecific anadromous females. Atlantic salmon parr had significantly higher plasma concentrations of the hormones 17α,20β-dihydroxy-4-pregnen-3-one (17,20β-P), 11-ketotestosterone (11-KT) and testosterone (T) after exposure to odours from conspecific females or from brown trout females compared to parr exposed to a control solution (0.9% NaCl). We did not observe any significant differences between the hormone levels in salmon parr exposed to the two female odours. The salmon parr exposed to conspecific odours had significantly higher volumes of strippable milt compared to the controls, but we did not find any significant differences when comparing the effect of the two female odours. Brown trout parr had significantly higher plasma 17,20β-P levels following exposure to heterospecific female odours compared to control males, but there was no significant difference between males exposed to the different female odours. We did not observe any significant differences in plasma levels of T and 11-KT and in milt volumes between exposed and control trout. Taken together, the results from both tested species indicate that the potency of heterospecific stimuli in stimulating increased plasma sex steroid hormone levels in male parr was as strong as stimuli from conspecific females. The results are discussed in connection to observed hybridisation between the two sympatric species.     

Hormonal and behavioral correlates of morphological variation in an Amazonian electric fish (Sternarchogiton nattereri: Apteronotidae)

The weakly electric fish from the main channel of the Amazon river, Sternarchogiton nattereri, offers a striking case of morphological variation. Females and most males are toothless, or present only few minute teeth on the mandible, whereas some males exhibit exaggerated, spike-like teeth that project externally from the snout and chin. Androgens are known to influence the expression of sexually dimorphic traits, and might be involved in tooth emergence. In this study we assess the relationship in S. nattereri between morphological variation, 11 ketotestosterone (11-KT) and testosterone (T). We also examine relationships of morphology and androgen levels with electric organ discharge (EOD) frequency, reproductive condition, and seasonality. Our main finding is that male morph categories differed significantly in plasma concentrations of 11-KT, with toothed males showing higher levels of 11-KT than toothless males. By contrast, we did not detect statistical differences in T levels among male morph categories. Reproductive condition, as measured by gonadosomatic indexes (GSI), differed across two sample years, increased as the season progressed, and was higher in toothed males than in non-toothed males. EOD frequency was higher in toothed males than in either toothless males or females. Taken together, our findings suggest that S. nattereri male sexual characters are regulated by 11-KT levels, and that both morphology and androgens interact with reproductive condition and EOD frequency in ways that vary within and across reproductive seasons.     

Male and female cooperatively breeding fish provide support for the "Challenge Hypothesis"

The idea that territorial aggression is regulated by androgensand that aggression itself can modulate androgen levels is wellestablished in males. In many species, females also displayaggressive behavior, yet little work has been conducted on theeffects of female aggression on hormone levels. In this study,we compared the effects of a simulated territory intrusion (amethod for testing the Challenge Hypothesis) on males and femalesof the fish, Neolamprologus pulcher. This cichlid fish fromLake Tanganyika is a particularly useful species to examinesex differences in the behavioral mediation of hormones as breedingpairs remain in a territory year round and both sexes defendthis territory against conspecific and heterospecific intruders.In our study, both sexes indeed aggressively defended theirterritory against a simulated territory intruder. In responseto intruders, both males and females displayed elevated levelsof circulating 11-ketotestosterone, but only females exhibitedincreases in testosterone. Neither aggressing male nor femalefish showed changes in estradiol levels compared to control(nonaggressing) fish. Residents were more aggressive than theintruders and won most of the interactions. However, residents(or winners) did not show higher hormone levels than intruders(or losers). We suggest that aggression commonly modulates androgenlevels in both male and female teleost fish.     

Territorial aggression and hormones during the non-breeding season in a tropical bird

The hormonal control of territorial aggression in male and female vertebrates outside the breeding season is still unresolved. Most vertebrates have regressed gonads when not breeding and do not secrete high levels of sex steroids. However, recent studies implicate estrogens in the regulation of non-breeding territoriality in some bird species. One possible source of steroids during the non-breeding season could be the adrenal glands that are known to produce sex steroid precursors such as dehydroepiandrosterone (DHEA). We studied tropical, year-round territorial spotted antbirds (Hylophylax n. naevioides) and asked (1). whether both males and females are aggressive in the non-breeding season and (2). whether DHEA is detectable in the plasma at that time. We conducted simulated territorial intrusions (STIs) with live decoys to male and female free-living spotted antbirds in central Panama. Non-breeding males and females displayed robust aggressive responses to STIs, and responded more intensely to decoys of their own sex. In both sexes, plasma DHEA concentrations were detectable and higher than levels of testosterone (T) and 17beta-estradiol (E(2)). In males, plasma DHEA concentrations were positively correlated with STI duration. Next, we conducted STIs in captive non-breeding birds. Captive males and females displayed robust aggressive behavior. Plasma DHEA concentrations were detectable in both sexes, whereas T was non-detectable (E(2) was not measured). Plasma DHEA concentrations of males were positively correlated with aggressive vocalizations and appeared to increase with longer STI durations. We conclude that male and female spotted antbirds can produce DHEA during the non-breeding season and DHEA may serve as a precursor of sex steroids for the regulation of year-round territorial behavior in both sexes.     

Endocrine correlates of intra-specific variation in the mating system of the St. Peter's fish (Sarotherodon galilaeus)

The Challenge Hypothesis postulates that androgen levels are a function of the social environment in which the individual is living. Thus, it is predicted that in polygynous males that engage in social interactions, androgen levels should be higher than in monogamous animals that engage in parental care. In this study, we tested this hypothesis at the intra-specific level using a teleost species, Sarotherodon galilaeus, which exhibits a wide variation in its mating system. Experimental groups of individually marked fish were formed in large ponds with different operational sex-ratios (OSR) to study the effects of partner availability on blood plasma levels of sex steroids [11-ketotestosterone (11-KT), testosterone (T), and 17,20beta-dihydroxy-4-pregnen-3-one (17,20beta-P)] and gonadosomatic index (GSI). Polygyny mostly occurred in the female biased OSR groups. 17,20beta-P and gonadosomatic index did not differ among OSR groups. However, 11-KT was high in male biased OSR and positively correlated with aggressive challenges, thereby supporting the central postulate of the Challenge Hypothesis. The results of T were the inverse of those of 11-KT, probably because 11-KT is metabolized from T. 11-KT levels of polygynous males did not differ neither from those of monogamous males, nor from those of males that participated in parental care. These results do not support the expected relationships between polygyny, parental care, and androgen levels. The differences from expectations for 11-KT may be related to the fact that in S. galilaeus, the mating and the parenting phase are not clearly separated and thus, males may still fight and court while they are brooding.