Social cues and hormone levels in male Octodon degus (Rodentia): a field test of the Challenge Hypothesis

Social interactions are important factors determining and regulating individual behaviors. Testosterone has been related to agonistic interactions, while glucocorticoids have been related to social stress, especially during interactions of dominance. We compared testosterone and cortisol concentrations in male degus (Octodon degus, Rodentia) under laboratory conditions without male social interactions, with data from wild males in nature. Under natural conditions, males should present higher levels of testosterone during the breeding season due to social interactions (Challenge Hypothesis). Alternatively, intense social instability could act as a stressing environment, raising glucocorticoids, which inhibit testosterone concentrations. Our results show a significant increase in agonistic interactions between males during the breeding season, and disappearance of non-agonistic male interactions during this period. Hormone levels in breeding season show nonsignificant differences between laboratory groups, but testosterone concentrations in field males were significantly higher than in laboratory males. Testosterone levels were similar among pre-breeding and breeding periods, but in field animals the concentration was approximately 30% higher than in laboratory degus. In field animals, we found two different mating strategies: resident males, with territorial behavior, and transient males, displayed an opportunistic approach to females. Finally, cortisol presents a similar pattern in both laboratory and field animals; pre-breeding values of cortisol are higher than during the breeding season. This suggests that social interactions in O. degus activate a rise in testosterone, supporting the Challenge Hypothesis, and could be considered as partial support of the Social Stress Hypothesis.     

Changes in social behavior of the male golden hamster accompanying photoperiodic changes in reproduction

Testicular regression and decreased serum testosterone levels result when male hamsters are placed in a short-day photoperiod, and these changes are reversed after reexposure to long-days. The present study was undertaken to determine whether these physiological changes lead to modifications in social behavior in the male hamster. Animals in a nonstimulatory short-day (LD 6:18) or a stimulatory long-day (LD 14:10) photoperiod were observed every 3 weeks for 21 weeks. After 9 weeks, half of the LD 6:18 males were returned to LD 14:10 and showed rapid testicular recrudescence. The other half remained in LD 6:18 and showed a slower rate of testicular recrudescence. Assessment of aggression was accomplished by placing an intruder in the home cage of a test animal and recording latency to attack, decisive encounters, number of attacks, and number of aggressive postures. Dominance was evaluated in a neutral arena with opponents of different weights, using the same parameters. Scent-marking was recorded in response to both bedding from an intact male and bedding from an ovariectomized female in a clean cage. Short-day animals undergoing testicular regression showed heightened levels of aggression and an upward shift in their dominance status. Alteration of the photoperiod did not affect the pattern or number of scent marks. As animals underwent testicular recrudescence, there was a return to lower levels of aggression and to a lower dominance status. Males undergoing rapid testicular recrudescence also showed the most rapid decline in aggressive behaviors. These results suggest that levels of agonistic behavior in the male hamster are inversely related to serum testosterone levels. Scent-marking appears to be unreleated to androgen levels in the male hamster and its role in hamster social behavior remains unclear.     

Seasonal changes in and effects of familiarity on agonistic behaviors of rat-like hamsters (Cricetulus triton)

The seasonal changes in agonistic behaviors and effects of familiarity on agonistic behaviors in wild-caught adult rat-like hamsters (Cricetulus triton) were observed in dyadic encounters in a neutral arena. The aggression of opposite- and same-sex encounters became higher or remained the same during the non-breeding season. This indicates that the hamsters were solitary during both seasons. Familiarity increased the aggression in male–male encounters and decreased the aggression in female–female encounters during both seasons. Familiarity also increased the aggression in female–male encounters during the non-breeding season and had no effect on the aggression in female–male encounters during the breeding season. These results may be related to the hamsters social structure. The more agonistic acts both male and female hamsters had, the more frequently they marked using flank glands during both seasons. This implies that flank gland marking can be used to advertise status and can be assessed by opponents to reduce the agonistic costs.     

Short days and exogenous melatonin increase aggression of male Syrian hamsters (Mesocricetus auratus)

Many nontropical rodent species rely on photoperiod as a primary cue to coordinate seasonally appropriate changes in physiology and behavior. Among these changes, some species of rodents demonstrate increased aggression in short, "winter-like" compared with long "summer-like" day lengths. The precise neuroendocrine mechanisms mediating changes in aggression, however, remain largely unknown. The goal of the present study was to examine the effects of photoperiod and exogenous melatonin on resident-intruder aggression in male Syrian hamsters (Mesocricetus auratus). In Experiment 1, male Syrian hamsters were housed in long (LD 14:10) or short (LD 10:14) days for 10 weeks. In Experiment 2, hamsters were housed in long days and half of the animals were given daily subcutaneous melatonin injections (15 microg/day in 0.1 ml saline) 2 h before lights out for 10 consecutive days to simulate a short-day pattern of melatonin secretion, while the remaining animals received injections of the vehicle alone. Animals in both experiments were then tested using a resident-intruder model of aggression and the number of attacks, duration of attacks, and latency to initial attack were recorded. In Experiment 1, short-day hamsters underwent gonadal regression and displayed increased aggression compared with long-day animals. In Experiment 2, melatonin treatment also increased aggression compared with control hamsters without affecting circulating testosterone. Collectively, the results of the present study demonstrate that exposure to short days or short day-like patterns of melatonin increase aggression in male Syrian hamsters. In addition, these results suggest that photoperiodic changes in aggression provide an important, ecologically relevant model with which to study the neuroendocrine mechanisms underlying aggression in rodents.     

Seasonal changes in testosterone and corticosterone levels in four social classes of a desert dwelling sociable rodent

Animals have to adjust their physiology to seasonal changes, in response to variation in food availability, social tactics and reproduction. I compared basal corticosterone and testosterone levels in free ranging striped mouse from a desert habitat, comparing between the sexes, breeding and philopatric non-breeding individuals, and between the breeding and the non-breeding season. I expected differences between breeders and non-breeders and between seasons with high and low food availability. Basal serum corticosterone was measured from 132 different individuals and serum testosterone from 176 different individuals of free living striped mice. Corticosterone and testosterone levels were independent of age, body weight and not influenced by carrying a transmitter. The levels of corticosterone and testosterone declined by approximately 50% from the breeding to the non-breeding season in breeding females as well as non-breeding males and females. In contrast, breeding males showed much lower corticosterone levels during the breeding season than all other classes, and were the only class that showed an increase of corticosterone from the breeding to the non-breeding season. As a result, breeding males had similar corticosterone levels as other social classes during the non-breeding season. During the breeding season, breeding males had much higher testosterone levels than other classes, which decreased significantly from the breeding to the non-breeding season. My results support the prediction that corticosterone decreases during periods of low food abundance. Variation in the pattern of hormonal secretion in striped mice might assist them to cope with seasonal changes in energy demand in a desert habitat.     

Spring and autumn territoriality in song sparrows: same behavior, different mechanisms?

Vertebrates show a diverse array of social behaviors associatedwith territoriality. Field and laboratory experiments indicatethat underlying themes—including mechanisms—mayexist. For example in birds, extensive evidence over many decadeshas implicated a role for testosterone in the activation ofterritorial aggression in reproductive contexts. Territorialityat other times of the year appeared to be independent of gonadalhormone control. One obvious question is—why this diversityof control mechanisms for an apparently similar behavior? Controlof testosterone secretion during the breeding season must balancethe need to compete with other males (that tends to increasetestosterone secretion), and the need to provide parental care(that requires lower testosterone concentrations). Regulationof aggressive behaviors by testosterone in the non-breedingseason may incur substantial costs. A series of experimentson the male song sparrow, Melospiza melodia morphna, of westernWashington State have revealed possible mechanisms to avoidthese costs. Song sparrows are sedentary and defend territoriesin both breeding and non-breeding seasons. Dominance interactions,territorial aggression and song during the non-breeding seasonare essentially identical to those during the breeding season.Although in the non-breeding season plasma testosterone andestradiol levels are very low, treatment with an aromatase inhibitordecreases aggression and simultaneous implantation of estradiolrestores territorial behavior. These data suggest that the mechanismby which testosterone regulates territorial behavior at theneural level remains intact throughout the year. How the hormonalmessage to activate such behavior gets to the brain in differentseason does, however, appear to be different.     

Effectiveness of exposure to longday followed by melatonin treatment on semen characteristics of Damascus male goats during breeding and non-breeding seasons

Eight mature Damascus male goats were randomly divided into two equal groups. The first group served as control and the second group was used to study the effect of exposure to longday (16L/8D) followed by melatonin treatment (2mg/(animal/day)) on semen characteristics and blood hormonal levels during breeding and non-breeding seasons. During breeding season, longday exposure followed by melatonin treatments resulted in significant increases (P<0.01) in means of sperm motility, ejaculate volume, sperm concentration, total sperm output, total functional sperm fraction and blood testosterone concentration, while means of reaction time, dead sperm, abnormal sperm and blood triiodothyronine (T(3)) concentration were significantly decreased (P<0.01). During non-breeding season, longday exposure followed by melatonin treatment exhibited improvements in some reproductive parameters by reducing (P<0.05) reaction time and percentage of dead sperm (P<0.01) and increasing (P<0.05) total functional sperm fraction. Furthermore, longday exposure followed by melatonin treatment resulted in significant increases (P<0.01) in both of blood testosterone and T(3) concentrations. It is concluded that longday exposure followed by melatonin treatment successfully improve semen characteristics of Damascus male goats during breeding and non-breeding seasons. However, the beneficial effects were more pronounced during the breeding season.     

Estrous female goats use testosterone-dependent cues to assess mates

In a promiscuous species like the domestic goat (Capra hircus), in which maternal investment is greater than paternal investment, a female may mate selectively with a more-fit male to improve her reproductive fitness. Testosterone (T) controls a large suite of male-typical behaviors and morphological characteristics. High T concentrations may be energetically costly or even detrimental to survival; thus, preventing lower quality males from falsely advertising their fitness. Three preference studies were conducted to examine if females use T-dependent cues to assess potential mates. For Experiment 1, females were given a choice between a pair of morphologically similar males, bucks (intact males) and stags (post-pubertally castrated males), during the breeding and non-breeding seasons. In both seasons, females preferred the bucks compared to stags. In Experiment 2, females were given a choice between bucks, stags and wethers (pre-pubertally castrated males) during the non-breeding season. For some comparisons, castrated males received 25 mg testosterone propionate (TP) or were untreated. Females preferred TP-treated males compared to untreated males and showed no preference when given a choice between either two TP-treated or two untreated males. In Experiment 3, females were given a choice between a pair of bucks and a pair of stags treated with 25 mg TP during monthly tests in the breeding season. At each monthly test, females preferred the males with higher T concentrations near the time of the behavior test. These studies suggest that females use T-dependent cues to assess potential mates, and T concentrations may indicate a male's overall fitness.     

Effects of short days on aromatization and accumulation of nuclear estrogen receptors in the hamster brain

Exposure of hamsters to short days increases sensitivity to the negative feedback effects of testosterone (T) but decreases responsiveness to the behavioral effects of the hormone. Since T is metabolized in the brain to 5 alpha-dihydrotestosterone (DHT) and estradiol, which differentially affect gonadotropin secretion and sex behavior, it is reasonable to postulate that daylength can modulate neural responses by quantitative or qualitative alterations in T metabolism and subsequent receptor binding of active hormone. Experiments reported here focused on aromatization and the nuclear accumulation of estrogen receptors. Adult male hamsters were maintained for 6-12 wk in long (14:10 LD) or short (8:16 LD) daily photoperiods. Both intact and castrated animals were used to assess direct effects of short days versus changes due to short-day-induced testicular regression. Discretely dissected regions of the brain (preoptic area, POA; hypothalamus, HTH; and corticomedial amygdala, CMA) or limbic blocks (LIM) comprised of all three regions were assayed for estrogen-synthesizing activity (aromatase) and estrogen-binding activity (receptors). Aromatase was estimated in vitro by conversion of [7-(3)H] androstenedione to [3H] estrogen and in vivo by measuring increases in nuclear estrogen receptor levels after injection of aromatizable androgen. Receptor-binding activity was assayed in crude cytosolic and nuclear extracts by incubating samples with [3H] estradiol +/- 100-fold excess inert estradiol, and separating free and bound steroids by Sephadex LH-20 gel filtration. When aromatase was assayed in homogenates prepared from discrete brain regions of individual hamsters, significantly lower activity was found in the HTH of short-day animals than in long-day controls. This effect was seen in both intact and castrated animals, which indicates that it was not mediated by the testis. Decreased enzyme activity in the POA and CMA of short-day hamsters was not significant, nor was there an effect of castration independent of short days. Low levels of nuclear estrogen receptors were present in LIM of intact males, but these were reduced after castration or concomitant with testicular regression after short-day exposure. This suggests that the hamster testis normally secretes estrogen or aromatizable androgen. A single injection of estradiol or aromatizable androgen (T or androstenedione) increased nuclear receptors in LIM of castrated animals. Cytosolic receptors were not different in short-day vs. long-day hamsters, nor were there differences in nuclear receptor levels after a single estradiol injection.(ABSTRACT TRUNCATED AT 400 WORDS)     

Gonadal hormones modulate the display of conditioned defeat in male Syrian hamsters

It has been widely reported that gonadal hormones influence the display of aggression in Syrian hamsters; conversely, much less is known about whether gonadal hormones modulate submissive/defensive behaviors in these animals. Following social defeat, male hamsters no longer display normal territorial aggression but instead display submissive/defensive behavior in the presence of a smaller opponent, a phenomenon we have termed conditioned defeat (CD). The purpose of the present study was to examine the effect of gonadal hormones on the display of CD in male hamsters. In Experiment 1, males were castrated or sham-operated. The castrated males were significantly more submissive following social defeat relative to their intact counterparts. The increased submissive behavior in the castrated males during CD testing was particularly surprising, given the fact that they were attacked significantly less during CD training. In Experiment 2a, males were castrated and given hormone replacement. Castrated males treated with testosterone or dihydrotestosterone displayed significantly less submissive behavior following social defeat than did those treated with cholesterol or estradiol. Finally, in Experiment 2b, there was no effect of hormone replacement on aggressive behavior in non-defeated hamsters suggesting that the decrease in submissive behavior in males treated with dihydrotestosterone or testosterone is specific to being previously defeated. Taken together the data indicate that the presence of androgens reduces the display of submission in defeated male hamsters. More importantly, these findings suggest that androgens may have a protective effect against the development of depression-like or anxiety-like behaviors following exposure to an ethologically relevant stressor.     

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.     

Effect of season and photoperiod on the follicle-stimulating hormone receptors in a subtropical bird

Annual changes in and photoperiodic influence oh the weight of gonads, a parameter of gonadal activity, are much smaller in female birds than in males. Effect of season and photoperiod on the follicle-stimulating hormone receptors in the testis or ovary was studied using a subtropical weaver finch. The number of follicle-stimulating hormone binding sites per unit testicular weight showed a peak in the non-breeding phase; while the total number of binding sites per two testes was maximal in the breeding phase and minimal in the regressive phase. In contrast, seasonal changes in follicle-stimulating hormone binding sites in the ovary were less marked. Exposure to short-day during the breeding phase induced marked decreases in the numbers of binding sites per unit testicular weight and per two testes. These numbers markedly increased after transfer to long-day during the non-breeding phase. However, there was no significant effect of short-day or long-day exposure on follicle-stimulating hormone binding sites in the ovary. These results suggest that photoperiod is an effective environmental factor in the regulation of follicle-stimulating hormone receptors in the testis and the effect is manifested by pronounced changes in the testicular weight during annual breeding cycle.     

Regulation of seasonal reproductive activity in the stallion, ram and hamster

This review considers seasonal reproduction in male animals with emphasis on the stallion, ram and hamster. The pineal hormone melatonin is the common link between photoperiod and reproduction. An increase in the daily diurnal period of melatonin secretion is associated with a decrease in GnRH release in long-day breeders, but an increase in GnRH release in short-day breeders. Melatonin influences GnRH release within or close to the mediobasal hypothalamus in rams; whereas melatonin receptors have not been found in the hypothalamus of horses. Prolactin release is positively correlated with daylength. Prolactin concentrations are consequently low during the breeding season of sheep and high during the breeding season of horses and hamsters. Prolactin stimulates testicular function in rams. Seasonal changes in GnRH release in the horse are regulated by changes in a GnRH-inhibitory opioidergic tone. Opioids are at least, in part, responsible for the decrease in testicular function during winter. An opioidergic inhibition of LH release is present during the breeding season in rams; but dopaminergic pathways inhibit LH release during long daylight hours. A dopaminergic inhibition of LH release does not exist in stallions.     

Social cues attenuate photoresponsiveness of the male reproductive system in Siberian hamsters (Phodopus sungorus)

Transfer of adult Siberian hamsters (Phodopus sungorus) from long (16 h light and 8 h dark, 16L:8D) to short (8L:16D) daily photoperiods induces an involution of the gonads and a cessation of reproductive behavior 8 to 10 weeks later. However, when male and female long-day hamsters were paired on transfer to short photoperiods, the males' gonads did not undergo the typical short-day response. Similarly, when male long-day hamsters were paired with refractory females (i.e., females housed in short photoperiods for at least 28 weeks so that they became unresponsive to short photoperiods), the response of the males' reproductive system to short photoperiods also was attenuated. Thus, social cues can override or delay the effects of photoperiod on the testes of this species. These results suggest that the inhibitory effects of long durations of melatonin secretion (in response to short photoperiods) on the male hypothalamic-pituitary-gonadal axis may be attenuated by social cues such as contact with the opposite sex.     

Continuous light after a long-day treatment is equivalent to melatonin implants to stimulate testosterone secretion in Alpine male goats

In rams, artificial long days followed by continuous light stimulate testosterone secretion during the non-breeding season. The objective of this study was to determine whether artificial long days followed by continuous light could stimulate testosterone secretion in Alpine bucks as well as in those exposed to long days followed by a melatonin treatment. All bucks were kept in shaded open pens. Control males were exposed to natural photoperiod conditions (n=5). Males of the two experimental groups were exposed to 2.5 months of long days from 1 December (n=5 each). On 16 February, one group of males was exposed to 24 h of light per day until 30 June; the other group was exposed to natural variations of photoperiod and received two s.c. melatonin implants. Testicular weight was determined every 2 weeks, and the plasma testosterone concentrations once a week. In the control and the two photoperiodic-treated groups, a treatment×time interaction was detected for testicular weight and plasma testosterone concentrations (P<0.001). In control bucks, testicular weight increased from January and peaked in June, whereas in both photoperiodic-treated groups, this variable increased from January, but peaked in April, when the values were higher than in controls (P<0.05). In the control group, plasma testosterone concentrations remained low from January to June, whereas in both photoperiodic-treated groups, this variable remained low from January to March; thereafter, these levels increased in both photoperiodic-treated groups, and were higher than controls in April and May (P<0.05). We conclude that continuous light after a long-day treatment stimulate testosterone secretion in Alpine male goats during the non-breeding season as well as the long days followed by a melatonin treatment. Therefore, continuous light could replace the implants of melatonin.     

Individual differences in aggressiveness of female hamsters: response to intact and castrated males and to females

The aggressive behaviour of female hamsters was studied while they were housed in large enclosures with males and in brief tests with males or females. Some females are not aggressive with any male, whereas others are very aggressive toward all males in both testing conditions. Females that are not aggressive toward intact males may be very aggressive toward castrated males or females. When the animals are housed together for long periods of time, males dominate only if they are much heavier. Male dominance takes a relatively long time to establish and often there is an equivocal period characterized by reversals of dominance. Female dominance is rapidly established. Unless the male is much heavier, the female determines the presence or absence of agonistic behaviour.     

黑线仓鼠殴斗行为模式及其与生理状态的关系

本文以分布于鲁西南山区的黑线仓鼠雄体为对象,通过测定重复遭遇战(repeated encounters)对黑线仓
鼠体重、生理指标等参数的作用,以期阐明社群冲突(social conflict)对黑线仓鼠生理状态的影响。研究结果表
明:在4 周的重复遭遇战过程中,参与冲突个体的体重增长率略有降低,但未达到显著水平;粪便肾上腺皮质
激素(GCs)含量呈现波动性变化,在整个遭遇战过程中,优势个体与从属个体的GCs 含量交替显著升高,与
冲突时间及个体社群地位均有关;优势个体保持较高的睾酮水平,利于其增强攻击行为和获得优势地位;从属
组雄体的肾上腺显著增大,但生殖腺间差异不显著;HPA 轴对HPG 轴具有显著的抑制效应,肾上腺分泌的皮质
醇可显著抑制个体的睾酮分泌,二者呈显著的负相关关系。这些数据表明,黑线仓鼠雄体可通过斗殴行为建立
明确的优势- 从属关系,睾酮可促进个体优势地位的形成并受到肾上腺皮质醇抑制;HPA 轴对HPG 轴的抑制未
能在器官指数方面得到证实。从属个体受到胁迫,对优势雄性黑线仓鼠野外生存和繁殖有重要意义。     

Photoperiodic influences on sexual behavior in male Syrian hamsters

The effect of photoperiodic conditions on sexual behavior was investigated in male Syrian hamsters that were either gonadally intact, or castrated and treated with low doses of testosterone throughout the experiment. Hamsters were exposed to long (LD 16:8) or short (LD 8:16) days for 7 weeks; for the next 8 weeks, either they were exposed to an intermediate daylength (LD 12:12), or daylength conditions remained unchanged. Sexual behavior was affected by photoperiod conditions in both gonadally intact animals and testosterone-treated castrates, but to different degrees. Intact males exposed to short days for 15 weeks exhibited gonadal regression, and their copulatory performance was impaired. The percentage of animals that intromitted or ejaculated was significantly reduced. Additional measures of sexual performance among the copulating males were also affected. In contrast, among the castrates with testosterone clamped at low but stable levels, the proportion of males that mounted, intromitted, or ejaculated was not affected by photoperiod. However, among the males that continued to copulate, sexual performance changes were present in the short-day castrates that resembled those displayed by the intact males. We infer that these behavioral effects in both hormonal conditions reflect primarily a difficulty in the attainment of intromission. Gonadal regression alone cannot easily account for the behavioral deficits of the intact males, because circulating testosterone levels at the end of the experiment were not significantly different between the gonadally intact hamsters and the castrated, testosterone-treated hamsters exposed continuously to short days. Males transferred from either long or short days to the intermediate-daylength condition responded behaviorally to this photoperiod as if it were a short day, that is, their ejaculatory frequency declined. We conclude that male hamsters exposed to photoinhibitory daylengths exhibit deficits in their sexual behavior, not only because endogenous levels of testosterone decrease, but also because the substrates on which this hormone acts become less responsive. We hypothesize that under physiological conditions, the episodic secretion of testosterone imposes constraints on the maintenance or restoration of copulation, and that the potent behavioral effects achieved by constant-release implants of testosterone may mask the presence of photoperiodically induced alterations in the hamster's sensitivity to this gonadal hormone.     

Architecture of the dorsal and ventral lobes of the prostate of the Syrian hamster, Mesocricetus auratus, after regrowth from short day-induced regression

The structures of the dorsal and ventral lobes of the prostate of 4 groups of Syrian hamsters were studied by a stereological approach. Groups studied were young hamsters (kept in long-day photoperiods for 4 weeks) in their first breeding season, older hamsters (kept in short-day photoperiods for 24 weeks so that their prostates had regressed and regrown) in their second breeding season, and two groups of older hamsters in an extended first breeding season (either because they were kept in long days for 24 weeks or because they were pinealectomized before being put into short-day photoperiods for 24 weeks). There were very few differences between groups, but generally dorsal prostates of older hamsters in their second breeding season closely resembled those of young hamsters in their first breeding season. More differences were noted between either of these two groups and older hamsters in an extended first breeding season than between these two groups. The differences noted generally involved increases in the amount of smooth muscle in the walls of secretory tubules in the dorsal prostates of hamsters in an extended first breeding season. This may be associated with the fact that these glands had not regressed and regrown. Ventral prostates were very similar in all 4 groups, which may reflect the fact that they normally regress very little in short days.     

Impact of season and social challenge on testosterone and corticosterone levels in a year-round territorial bird

Plasma testosterone increases during breeding in many male vertebrates and has long been implicated in the promotion of aggressive behaviors relating to territory and mate defense. Males of some species also defend territories outside of the breeding period. For example, the European nuthatch (Sitta europaea) defends an all-purpose territory throughout the year. To contribute to the growing literature regarding the hormonal correlates of non-breeding territoriality, we investigated the seasonal testosterone and corticosterone profile of male (and female) nuthatches and determined how observed hormone patterns relate to expression of territorial aggression. Given that non-breeding territoriality in the nuthatch relates to the reproductive context (i.e., defense of a future breeding site), we predicted that males would exhibit surges in plasma testosterone throughout the year. However, we found that males showed elevated testosterone levels only during breeding. Thus, testosterone of gonadal origin does not appear to be involved in the expression of non-breeding territoriality. Interestingly, territorial behaviors of male nuthatches were stronger in spring than in autumn, suggesting that in year-round territorial species, breeding-related testosterone elevations may upregulate male-male aggression above non-breeding levels. In females, plasma testosterone was largely undetectable. We also examined effects of simulated territorial intrusions (STIs) on testosterone and corticosterone levels of breeding males. We found that STIs did not elicit a testosterone response, but caused a dramatic increase in plasma corticosterone. These data support the hypothesis that corticosterone rather than testosterone may play a role in the support of behavior and/or physiology during acute territorial encounters in single-brooded species.