Rapid elevations in both steroid hormones and vocal signaling during playback challenge: a field experiment in Gulf toadfish

It is well known that plasma androgens are rapidly released in response to aggressive or sexual stimuli in a broad array of vertebrates. However, experimental work on behavioral functions of rapid androgen elevation is rare. A combination of field-based behavioral experiments and lab-based neuroendocrinological approaches is beginning to show how steroid hormones rapidly regulate the expression of vocal communication signals in Gulf toadfish (Opsanus beta). Male toadfish emit multiharmonic "boatwhistles" and shorter-duration, broadband "grunts" during intraspecific communication. Neurophysiology experiments demonstrate that androgens and glucocorticoids rapidly modify vocal motor patterning in male toadfish. In this study, we simulated territorial intrusions (vocal "challenges") with acoustic playbacks to toadfish in the field, and observed simultaneous, rapid (within 5-20 min) changes in vocalizations and steroid hormones. Both plasma androgens and vocal activity increased following the presentation of pure tones that mimic the duration of natural boatwhistles (275 ms), while they remained unchanged following playbacks of tone stimuli that mimic the duration of grunts (75 ms) or the upper-range of boatwhistles (475 ms). Circulating glucocorticoids were elevated in calling vs. non-calling males but were unaffected by playback stimuli, suggesting a role in the energetics of vocalization. These results strongly suggest that one function of rapid androgen elevation in response to social challenge is to mediate similarly rapid changes in territorial vocal signaling. Given the conserved organization of neuroendocrine and vocal motor systems, rapid steroid action on vocalization mechanisms may be true of other vocal vertebrates as well, including birds and mammals.     

Plasma levels of androgens and cortisol in relation to breeding behavior in parental male bluegill sunfish, Lepomis macrochirus

Like many teleosts, male bluegill sunfish (Lepomis macrochirus) provide sole parental care. To understand some of the proximate costs of parental care, we measured body condition and plasma levels of testosterone (T), 11-ketotestosterone (11KT) and cortisol in nesting bluegill males during pre-spawning, spawning and parental care stages. T and 11KT were at their highest mean levels during the pre-spawning period and decreased to lower levels early during the parental care period before rising again when the eggs hatched. Cortisol levels fluctuated across the breeding stages, but there was a noticeable increase from low levels on the day of spawning during the first 2 days of parental care when egg fanning is most intense. Levels of all hormones varied considerably among males, with androgen levels often correlating positively with a male's body condition. We also demonstrate, using a brood reduction experiment and repeated sampling of known individuals, that the presence of eggs affects hormone levels shortly after eggs hatch. Parental males in better body condition had higher levels of androgens during parental care. Males that were known to renest later in the season also had higher androgen levels and were in better body condition during the first nesting bout than males only known to have nested once. However, circulating levels of cortisol did not differ significantly between these groups. We discuss our findings in the context of proximate and ultimate costs of parental care and propose several reasons why elevated androgen levels may not be as incompatible with the expression of paternal care in male teleosts, as compared with avian and mammalian fathers.     

Preference for the presence of substrate in male cichlid fish: Effects of social dominance and context

Many cichlid species dig spawning pits or nests in soft bottoms and exhibit many substrate oriented activities. Despite this fact being of general knowledge, captive cichlids in laboratory and aquaculture set-ups are often kept in the absence of a soft substrate that they can manipulate. This raises a potential welfare issue, depending on how the substrate is valued by cichlid fish. The aim of this study is to assess the importance of substrate for male Mozambique tilapia (Oreochromis mossambicus) in social and non-social contexts. Preferences were established as a measure of time spent in two choice compartments, before and after the presence of a female. Locomotory activity, social interactions and substrate-related behaviours were recorded. Results show that dominant males prefer the area with substrate regardless of social context, and that female's presence strengthens this preference. The same preference is not apparent in the subordinate males, except for foraging. To draw conclusions on the importance of substrate to subordinates, preferences should also be assessed in agonistic contexts, during which substrate may serve to displace aggression. These results, together with related previous studies, show that the lack of substrate is particularly deleterious in a reproductive context, and thus it is likely to decrease the welfare state of breeding males of Mozambique tilapia.     

Ghrelin gene in cichlid fish is modulated by sex and development

We have identified the complementary and genomic DNA sequences of the novel growth hormone-releasing peptide, ghrelin, in the cichlid fish (tilapia Oreochromis niloticus). The tilapia ghrelin precursor cDNA was 855 bp long, consisting of 119 bp of 5(')-untranslated region, 324 bp open reading frame, and 412 bp of 3(')-untranslated region. The serine residues at positions 2 and 3 of the "active core" (GSSF) of the ghrelin mature peptide are conserved between tilapia and the mammalian species. The tilapia ghrelin gene has four exons and three introns and resembles the structure of other known ghrelin genes. RT-PCR analysis revealed ghrelin mRNA predominantly expressed in the stomach but absent in the brain, pituitary, heart, kidney, ovary, and testis. Real-time PCR analysis showed an age/body size dependent increase in gastric ghrelin, which stagnated at 7 cm body size (onset of maturation). Ghrelin mRNA levels were unchanged in food-deprived sexually mature animals but were significantly higher in females compared to males. The present study shows that the structure of ghrelin peptide is highly conserved, and the reported differences in somatic and gonadal growth in tilapia could be a consequence of age- and sex-related synthesis of gastric ghrelin.     

The effects of social context on the hormonal and behavioral responsiveness of human fathers

We tested first-time fathers with their 22-month old toddlers to determine whether social context variables such as pre-test absence from the child and presence of the mother affected physiological measures associated with paternal responsiveness. Heart rate and blood pressure readings as well as blood samples to determine prolactin, testosterone and cortisol levels were taken before and after the 30-min father–toddler interactions. Fathers were tested on a day when they were away from their child for several hours before testing (‘without-child’ day) and on another day where they remained with their child throughout the day (‘with-child’ day). Most measures decreased over the 30-min test period but relative decreases were context-dependent. Men maintained higher prolactin levels when they were away from their children longer before testing on the ‘without-child’ day. Cortisol levels decreased during both tests and they decreased more on the ‘with-child’ day for men who had spent more time alone with their toddler before the test. Heart-rate and diastolic (but not systolic) blood pressure decreased more on the ‘with-child’ day than on the ‘without-child’ day. Fathers' testosterone levels decreased when their partners were less involved in the interactions. Compared to men with high responsiveness ratings on both days, men whose responsiveness increased after being away from their child on the ‘without-child’ day maintained higher systolic blood pressure and had a greater decrease in testosterone levels. We conclude that context may be more important in determining fathers' physiological responses to child contact than has previously been appreciated, particularly for some individuals.     

Reproduction in context: Field testing a laboratory model of socially controlled sex change in Lythrypnus dalli (Gilbert)

Social interactions can have profound effects on reproduction and the proximate mechanisms involved are just beginning to be understood. Lythrypnus dalli, the bluebanded goby, is an ideal organism for analyzing the dynamics of socially controlled sex change both in the laboratory and field. As with most research species, the majority of its behavioural and physiologic study has been performed in the laboratory. The goal of our study was to induce sex change of L. dalli in a more natural environment and compare field dynamics with our laboratory-based model. Groups of L. dalli, composed of one large male and three females of varying sizes, were introduced into artificial habitats in the field. After male removal, the dominant, largest female underwent protogynous sex change in the majority of the groups. Within 15 days, 9 of 15 of the dominant females (focal fish) successfully fertilized eggs as males, compared to 13 of 17 in the laboratory. Focal fish displayed the distinctive temporal sequence of behaviour changes consisting of a dominance, quiescent, and courtship phase. In addition, focal fish had gonads, genital papillae, and accessory gonadal structures with morphology in between that of females and males. Those fish that fertilized eggs had this transitional morphology, but were functionally male. Steroids of focal fish were assayed by water sample, and morning samples of free 11-ketotestosterone (11-KT) positively correlated with the percent of male tissue in the gonad, with the size of the accessory gonadal structure but not the genitalia (genital papilla), and with aggressive displacement behaviour on the last day before the fish were sacrificed. These morphological, physiological, and behavioural patterns parallel those seen in the laboratory. Lower rates of behaviour and the dramatic effects of ambient temperature in the field provide insights as to how the environmental context modifies the behaviour and, subsequently, the reproductive function of individuals within a social group.     

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.     

Behavioral and physiological plasticity: Rapid changes during social ascent in an African cichlid fish

In many vertebrates, reproduction is regulated by social interactions in which dominant males control access to females and food. Subordinate males that displace dominant individuals must rapidly adopt behavioral and physiological traits of the higher rank to gain reproductive success. To understand the process of phenotypic plasticity during social ascent, we analyzed the temporal expression pattern of dominance behaviors and circulating androgen levels when socially-suppressed males of an African cichlid fish Astatotilapia burtoni ascended in status. These experiments tested a prediction of the ‘challenge hypothesis’ that, during periods of social instability, male androgen levels are higher than during socially stable times. We found that socially and reproductively suppressed males perform territorial and reproductive behaviors within minutes of an opportunity to ascend in status, and that animals switch from initial expression of territorial behaviors to more reproductive behaviors during territory establishment. Following this rapid response, social stability may be achieved within 1-3 days of social ascent. Consistent with predictions of the ‘challenge hypothesis’, circulating 11-ketotestosterone (11-KT) levels were elevated within 30 min following social opportunity, coincident with increased aggressive behavior. However, territorial behaviors and serum 11-KT levels were then dissociated by 72 h after social ascent, suggesting either rapid social stability and/or increased physiological potential for androgen production. This behavioral and physiological plasticity in male A. burtoni suggests that perception of social opportunity triggers a suite of quick changes to facilitate rapid transition towards reproductive success, and reveals important features of social ascent not previously recognized.     

Sex differences and similarities in the neuroendocrine regulation of social behavior in an African cichlid fish

An individual's position in a social hierarchy profoundly affects behavior and physiology through interactions with community members, yet little is known about how the brain contributes to status differences between and within the social states or sexes. We aimed to determine sex-specific attributes of social status by comparing circulating sex steroid hormones and neural gene expression of sex steroid receptors in dominant and subordinate male and female Astatotilapia burtoni, a highly social African cichlid fish. We found that testosterone and 17β-estradiol levels are higher in males regardless of status and dominant individuals regardless of sex. Progesterone was found to be higher in dominant individuals regardless of sex. Based on pharmacological manipulations in males and females, progesterone appears to be a common mechanism for promoting courtship in dominant individuals. We also examined expression of androgen receptors, estrogen receptor α, and the progesterone receptor in five brain regions that are important for social behavior. Most of the differences in brain sex steroid receptor expression were due to sex rather than status. Our results suggest that the parvocellular preoptic area is a core region for mediating sex differences through androgen and estrogen receptor expression, whereas the progesterone receptor may mediate sex and status behaviors in the putative homologs of the nucleus accumbens and ventromedial hypothalamus. Overall our results suggest sex differences and similarities in the regulation of social dominance by gonadal hormones and their receptors in the brain.     

Social competition affects electric signal plasticity and steroid levels in the gymnotiform fish Brachyhypopomus gauderio

Sexually-selected communication signals can be used by competing males to settle contests without incurring the costs of fighting. Steroid regulation of these signals can render them as reliable indicators of a male's physiological state. We investigated how plasticity in electrocommunication signals is driven by social competition for mates, mediated by steroid hormones, and subject to the effects of past social experience. We measured the electric waveform's amplitude and duration and steroid hormone levels of male gymnotiform electric fish (Brachyhypopomus gauderio) following week-long periods of social isolation, and low or high social competition. To quantify the effect of social history on the modulation of the electric signal, six groups of six males experienced all three social conditions but in different order. We found that males differentially modulate their electric signals depending on the order they experienced these conditions. Thus, past social interactions affect both present and future social electric signals. Cortisol levels and the amplitude of the electric signal appeared to track the intensity of competition, while androgen levels and the duration of the electric signal only responded to the presence (low and high competition) or absence (isolation) of a social environment (low and high androgens respectively). In addition, cortisol levels were related to the body size of the males at high social competition. Taken together, these findings suggest that the capacity of males to modulate their signals in response to social competition is regulated by steroids.     

Testosterone and 11-ketotestosterone have different regulatory effects on electric communication signals of male Brachyhypopomus gauderio

The communication signals of electric fish can be dynamic, varying between the sexes on a circadian rhythm and in response to social and environmental cues. In the gymnotiform fish Brachyhypopomus gauderio waveform shape of the electric organ discharge (EOD) is regulated by steroid and peptide hormones. Furthermore, EOD amplitude and duration change on different timescales and in response to different social stimuli, suggesting that they are regulated by different mechanisms. Little is known about how androgen and peptide hormone systems interact to regulate signal waveform. We investigated the relationship between the androgens testosterone (T) and 11-ketotestosterone (11-KT), the melanocortin peptide hormone α-MSH, and their roles in regulating EOD waveform of male B. gauderio. Males were implanted with androgen (T, 11-KT, or blank), and injected with α-MSH before and at the peak of androgen effect. We compared the effects of androgen implants and social interactions by giving males a size-matched male stimulus with which they could interact electrically. Social stimuli and both androgens increased EOD duration, but only social stimuli and 11-KT elevated amplitude. However, no androgen enhanced EOD amplitude to the extent of a social stimulus, suggesting that a yet unidentified hormonal pathway regulates this signal parameter. Additionally, both androgens increased response of EOD duration to α-MSH, but only 11-KT increased response of EOD amplitude to α-MSH. Social stimuli had no effect on EOD response to α-MSH. The finding that EOD amplitude is preferentially regulated by 11-KT in B. gauderio may provide the basis for independent control of amplitude and duration.     

Long-term effects of adolescent stress on neophobic behaviors in zebra finches are modulated by social context when in adulthood

Experiencing stress during adolescence can increase neophobic behaviors in adulthood, but most tests have been conducted in the absence of conspecifics. Conspecifics can modulate responses to stressors, for example by acting as ‘social buffers’ to attenuate the aversive appraisal of stressors. Here, we investigate the long-term effects of adolescent stress on the behavioral responses to novel stimuli (a mild stressor) across social contexts in an affiliative passerine bird, the zebra finch. During early (days 40–60) or late (days 65–85) adolescence the birds (n = 66) were dosed with either saline or the hormone corticosterone (CORT). CORT was given in order to mimic a physiological stress response and saline was given as a control. In adulthood, the birds' behavioral responses to a novel environment were recorded in both the presence and absence of conspecifics. An acute CORT response was also quantified in adolescence and adulthood. Our findings show clear evidence of social context mediating any long-term effects of adolescent stress. In the presence of familiar conspecifics no treatment effects were detected. Individually, birds dosed with CORT in early adolescence were slower to enter a novel environment, spent more time perching in the same novel environment, and, if female, engaged in more risk assessment. Birds dosed in late adolescence were unaffected. No treatment effects were detected on CORT, but adolescents had a higher CORT concentration than adults. Our results are the first to suggest that familiar conspecifics in adulthood can buffer the long-term effects of stress that occurred during early adolescence.     

Differential effects of 11-ketotestosterone on dimorphic traits in a teleost with alternative male reproductive morphs

Species with dimorphic males typically have one morph (type I) associated with territoriality and courtship, and another morph (type II) associated with cuckoldry. In teleost fishes, type I males generally have higher levels of the androgen 11-ketotestosterone (KT) than type II males. Is KT causal to phenotypic differences between morphs? We investigated this question in the midshipman fish (Porichthys notatus) in which type I males have detectable levels of KT during the breeding season, whereas levels in type II males are usually undetectable. Type I midshipman will either cuckold or hold territories and court females, whereas type II males are only known to cuckold. Acoustic courtship by type I's is supported by the sonic motor nucleus, which innervates a sound-producing sonic muscle. Type I males have larger sonic motor nuclei and larger sonic muscles than type II males, consistent with the more dynamic vocal repertoire of type I's. Here, we tested whether intraperitoneal KT implants in adult type II males would induce type I male-like traits in brain, sonic muscle, and behavior. Type II's treated with KT did not differ from blank-implanted type II's in sonic motor nucleus volume. Sonic muscle mass increased in KT-implanted type II's, but did not reach the relative mass naturally observed in type I's. While neither territoriality nor courtship were induced, cuckoldry behavior intensified in KT-implanted type II's. Thus, for some but not all characters, KT exaggerated the expression of already existing type II male traits rather than inducing the type I male-like traits of territoriality, courtship, and an expansive vocal motor system.     

Estradiol effects on behavior and serum oxytocin are modified by social status and polymorphisms in the serotonin transporter gene in female rhesus monkeys

Despite the well-documented relation between estradiol (E2) and behavior, exposure to stressors may modify sensitivity to E2. The effects of E2 on behavior are, in part, likely related to their modulation of the serotonin (5HT) and oxytocin systems. The short allele (s-variant) polymorphism found in the promoter region of the SLC6A4 gene that encodes the 5HT transporter (5HTT) modulates responsivity to stressors. The current study used ovariectomized adult female rhesus monkeys to evaluate how exposure to the psychosocial stressor of social subordination and polymorphisms in the gene encoding 5HTT influence the behavioral effects of E2 and immunoreactive serum oxytocin. Dominant females had higher levels of oxytocin than subordinate animals even though E2 increased immunoreactive serum oxytocin in all females. E2 increased affiliative behaviors in all animals, with even more of these prosocial behaviors directed at dominant females. S-variant females, regardless of social status, were more aggressive toward more subordinate cage mates and these behaviors too were increased by E2. Subordinate s-variant females are most often involved in agonistic behavior, less affiliative behavior, and were less responsive to the anxiolytic action of E2. The results show that the short allele of the 5HTT gene synergizes with psychosocial stress exposure to affect the behavioral efficacy of E2 while confirming the actions of E2 for producing generalized behavioral arousal in females. Whether differences in the central action of 5HT and/or oxytocin are responsible for this effect requires further study.     

The physiological effects of social status in the cooperatively breeding cichlid Neolamprologus pulcher

The physiological effects of social rank were examined in three different experiments with Neolamprologus pulcher a cooperatively breeding cichlid, endemic to Lake Tanganyika, East Africa. The effects of rank on physiology between pairs of dominant and subordinate size‐matched fish (experiment 1) and among groups of four size‐matched fish (experiment 2) were examined. A third experiment mimicked the natural social structure in the wild; pairs were observed with other group members including breeders. The effect of social position was investigated on growth rates, liver concentrations of adenosine triphospate (ATP), lipids, proteins, creatine phosphate (CrP), glucose and glycogen as well as plasma cortisol. In naturalistic group settings, dominants displayed higher levels of liver protein and plasma cortisol. In the absence of breeders, dominant individuals (of helper pairs) had higher liver glycogen levels and dominant fish (held in groups of four) grew most. These results support previous cooperatively breeding mammal studies and suggest that dominant individuals experience higher cortisol levels as well as higher growth rates.     

The effects of short-term resistance training on endocrine function in men and women

This investigation examined hormonal adaptations to acute resistance exercise and determined whether training adaptations are observed within an 8-week period in untrained men and women. The protocol consisted of a 1-week pre-conditioning orientation phase followed by 8 weeks of heavy resistance training. Three lower-limb exercises for the quadriceps femoris muscle group (squat, leg press, knee extension) were performed twice a week (Monday and Friday) with every other Wednesday used for maximal dynamic 1 RM strength testing. Blood samples were obtained pre-exercise (Pre-Ex), immediately post-exercise (IP), and 5 min post-exercise (5-P) during the first week of training (T-1), after 6 weeks (T-2) and 8 weeks (T-3) of training to determine blood concentrations of whole-blood lactate (LAC), serum total testosterone (TT), sex-hormone binding globulin (SHBG), cortisol (CORT) and growth hormone (GH). Serum TT concentrations were significantly (P ≤ 0.05) higher for men at all time points measured. Men did not demonstrate an increase due to exercise until T-2. An increase in pre-exercise concentrations of TT were observed both for men and women at T-2 and T-3. No differences were observed for CORT between men and women; increases in CORT above pre-exercise values were observed for men at all training phases and at T-2 and T-3 for women. A reduction in CORT concentrations at rest was observed both in men and women at T-3. Women demonstrated higher pre-exercise GH values than men at all training phases; no changes with training were observed for GH concentrations. Exercise-induced increases in GH above pre-exercise values were observed at all phases of training. Women demonstrated higher serum concentrations of SHBG at all time points. No exercise-induced increases were observed in men over the training period but women increased SHBG with exercise at T-3. SHBG concentrations in women were also significantly higher at T-2 and T-3 when compared to T-1 values. Increases in LAC concentrations due to exercise were observed both for men and women for all training phases but no significant differences were observed with training. These data illustrate that untrained individuals may exhibit early-phase endocrine adaptations during a resistance training program. These hormonal adaptations may influence and help to mediate other adaptations in the nervous system and muscle fibers, which have been shown to be very responsive in the early phase of strength adaptations with resistance training. Accepted: 11 December 1997     

The effect of social isolation on androgen and corticosteroid levels in a cichlid fish (Haplochromis burtoni) and in swordtails (Xiphophorus helleri)

The blood androgen and glucocorticoid levels of male Haplochromis burtoni and Xiphophorus helleri were measured in socially isolated individuals (8 and 4 weeks, respectively) and nonisolated controls. In both species social isolation leads to significant reductions of mean concentrations and variances, both for androgens and corticosteroids. It is known for H. burtoni—and new evidence for X. helleri is presented—that social isolation reduces aggressivity in nonescalating situations. It is proposed that social stimuli in both species stimulate the testes to produce more androgenic hormones, and that this can lead to higher levels of aggression. At least in tropical fish species the androgenic optimum for defending a territory or a social status may be much more controlled by social stimuli than previously realized.     

The effects of polychlorinated biphenyls on plasma steroid levels and hepatic microsomal enzymes in fish

Aroclor 1254 was administered intraperitoneally (25 mg kg body wt⁻¹ in 1 ml of arachis oil) at weekly intervals for 4 weeks to trout and carp; arachis oil was used as the control. Activities of the following hepatic microsomal enzymes, aminopyrine demethylase, p-nitroreductase, UDP-glucuronyl-transferase and 1-leucyl-β-naphthylamide splitting enzyme were measured in both species; cytochrome P₄₅₀ and microsomal protein contents were also determined.
The changes in the levels of androgens, oestrogens and corticoid hormones were measured in the circulating blood of control and treated groups at weekly intervals. The blood was obtained by cardiac puncture.
Results indicated (a) a significant increase in the activities of all the enzymes measured except 1-leucyl-β-naphthylamide splitting enzyme, (b) cytochrome P₄₅₀ and microsomal protein contents were increased in trout, but not in carp, (c) a significant reduction in the plasma levels of androgens, oestrogens and corticoids in the treated groups, particularly at the end of the fourth week and (d) there was a correlation between increased enzyme activities and a decrease in plasma hormone levels.     

Hormonal correlates of social status and courtship display in the cooperatively lekking lance-tailed manakin

Though cooperative behavior has long been a focus of evolutionary biology, the proximate hormonal mechanisms underlying cooperative interactions remain poorly understood. Lance-tailed manakins (Chiroxiphia lanceolata) are tropical passerines that form long-term male-male partnerships and cooperate in paired male courtship displays. To elucidate patterns of natural hormonal variation in relation to cooperation and reproductive behavior, we examined circulating androgen levels of male lance-tailed manakins in relation to social status, display behavior, and time of year. We found significantly higher circulating androgen levels in alpha-ranked (breeding) males compared to non-alpha adult males in the population. Beta males, which participated in courtship displays but did not copulate, had androgen levels indistinguishable from those of unpaired adult males that never displayed for females, suggesting that an elevated concentration of plasma testosterone in tropical lekking birds may be associated primarily with copulatory behavior or other status-specific traits, and not the performance of courtship display. Androgens decreased throughout the breeding season for males of all status categories. Interestingly, alphas that displayed for females in the observation session prior to sampling had lower androgen levels than alphas that did not display for females. This pattern may result from female discrimination against alpha males at display areas with high levels of social conflict among males, as social disruption is linked to elevated testosterone in many species. However, recent change of a display partner was not related to alpha androgen levels. We discuss alternative explanations and the possible implications of these results, and generate several testable predictions for future investigations.     

A comparison of the annual changes in testicular activity and serum androgen levels between the early and delayed maturing groups of male Cottus hangiongensis

Synopsis Annual changes in testicular activity and concentration of two serum androgens were monitored in two groups of the river-sculpin Cottus hangiongensis collected from the upper and lower reaches of a river at southern Hokkaido, Japan. One of them (early maturing group) underwent testicular maturation with aberrant spermatids and spermatid masses produced during the reproductive cycle. Moreover, regular seasonal changes in serum testosterone and 11-ketotestosterone concentrations were observed. On the other hand, in the other group (delayed maturing group), although body size of the fish was large enough to undergo reproduction, annual changes in gonadosomatic index and testicular activity did not vary much. During the months of active testicular development in the early maturing group, spermatogenesis was observed to begin in some regions of the testes of delayed maturing fish, but always resulted in the formation of aberrant spermatids and spermatid masses. Moreover, concentration of serum androgens did not significantly vary throughout the year. Results suggest that low androgen production is a proximal factor for delayed sexual maturity in the delayed maturing group, and that the occurrence of aberrant spermatids and spermatid masses during spermatogenesis is not linked to the delayed maturity.