Corticosterone-binding proteins and behavioral effects of high plasma levels of corticosterone during the breeding period in the pied flycatcher

In the pied flycatcher there exists an anomaly in the relationship between cortical histology and plasma levels of corticosterone during the breeding period. In an attempt to study this anomaly, binding capacity and binding affinity of plasma corticosterone-binding proteins (CBP) were studied in free-living pied flycatchers during the early and late parts of the breeding period. Binding capacity of CBP showed a significant decrease with the progress of the breeding season in both males and females. During the early parts of the breeding season binding capacity was significantly higher in males than in females. No difference between sexes was observed during the nestling period. In males there also was a seasonal decrease in the binding affinity of CBP. The results show that there is a good relationship between periods with high plasma levels of corticosterone and its binding capacity in the blood. A second study showed that an experimentally increased plasma level of corticosterone during the nestling period drastically reduced reproductive success. Parents given silastic implants containing corticosterone fed their nestlings less frequently and produced significantly fewer fledglings than did controls. Unlike the control birds, the body weight of the corticosterone-implanted birds did not decrease during the nestling period. Birds given corticosterone implants in which one small hole had been punched, in order to facilitate diffusion of corticosterone, all abandoned their territories and, consequently, these parents produced no fledglings. Thus, the results show that an elevation of plasma levels of corticosterone in adult pied flycatchers during the nestling period affects parental as well as territorial behavior.     

Why do experienced birds reproduce better? Possible endocrine mechanisms in a long-lived seabird, the common tern

The influence of age or breeding experience on reproductive success in vertebrates is well known but physiological mechanisms came into focus just recently. Assessing hormone levels could allow insights into these mechanisms and reproductive strategies in long-lived species. Prolactin and corticosterone are two hormones involved in breeding decisions: high prolactin values are necessary for expressing breeding behavior whereas corticosterone is related to activity or stress. We analyzed baseline prolactin and corticosterone under field conditions in common terns (Sterna hirundo) between 2006 and 2010. We took 760 blood samples of 346 known birds 9-14days after their clutch completion, obtained via blood-sucking bugs (Dipetalogaster maximus), a non-invasive method with negligible stress for the birds. Many individuals were sampled repeatedly during the study period allowing investigation of hormone change on individual level. Prolactin levels increased during the early breeding career, which was confirmed on individual level, whereas corticosterone levels increased mostly in experienced birds, more pronounced in males. Low hormone levels during the first years of breeding could indicate a reduced ability of the endocrine system to secret hormones or it might express a down-regulation to limit parental expenditure. Higher corticosterone values of males could be related to increased foraging activity. Amongst the oldest birds, prolactin seemed to increase in males but not in females. This possible consequence of female senescence might be compensated by high values of male mates. Body mass showed only a weak positive correlation with prolactin level and no correlation with corticosterone concentration.     

The effects of an "El Niño" southern oscillation event on reproduction in male and female blue-footed boobies, Sula nebouxii

This study attempted to determine endocrine correlates of reproductive success in relation to major deleterious environmental conditions. In 1992, an El Ni?o southern oscillation event resulted in complete reproductive failure in a colony of blue-footed boobies, Sula nebouxi, on Isla Isabel in the Pacific Ocean off San Blas, Nayarit, Mexico (21.5 degrees N, 105.5 degrees W). In 1993, the El Ni?o event had waned and reproductive success was high. The mean sea surface temperature in 1992 was 26.69 degrees, the warmest year for 11 years of data (mean, 25.63 degrees ). In 1993, mean sea surface temperature was 25.75 degrees. Plasma levels of testosterone were highest during the egg-laying period in 1993 and declined markedly during incubation. There were no differences between males and females. Comparisons of testosterone levels between 1992 and 1993 (egg-laying time point removed for 1993) showed no significant differences. Thus reproductive failure during an El Ni?o year was not related to testosterone levels. Baseline plasma levels of corticosterone did not change over the nesting cycle in either sex. There was a trend for plasma levels of corticosterone to be higher in males and females during the earlier stages of breeding in 1992 compared with 1993, and if all levels were combined within years then females showed significantly higher plasma levels of corticosterone in the El Ni?o year. Plasma levels of corticosterone showed marked increases following capture and handling in both sexes and at every stage of the breeding cycle in each year. There was no variation in the adrenocortical responses to stress with year or stage of nesting in males. However, in females, maximum corticosterone levels were greatest during the parental phase of 1992, the El Ni?o year, when all nests ultimately failed. Comparisons of the dynamics of corticosterone changes during the capture stress protocol revealed no correlations with body mass in 1992 or 1993. These data suggest that although massive reproductive failure in the El Ni?o year was not related to testosterone levels, baseline circulating concentrations of corticosterone may have a role in inhibiting onset of breeding. In contrast, after the nesting cycle has been initiated, increased adrenocortical sensitivity to acute stress may be involved in nest abandonment.     

Steroid levels and reproductive cycle of the Galápagos tortoise, Geochelone nigra, living under seminatural conditions on Santa Cruz Island (Galápagos)

The Galápagos Islands are home to 11 subspecies of large terrestrial tortoises (Geochelone nigra). All Galápagos tortoises are considered endangered and approximately 12,000 animals still exist. Until now, the reproductive cycle of the Galápagos tortoise has been studied only in captive animals, and no data from free-ranging tortoises have been available. During a one-year period, blood samples were collected from male and female G. nigra living under seminatural conditions on Santa Cruz Island, Galápagos. Plasma steroid hormones were measured by radioimmunoassays (RIAs). In males, plasma testosterone and corticosterone increased a few months before the onset of the mating season. Peak levels were observed while most copulations occurred and environmental temperatures were highest. Both testosterone and corticosterone showed low levels during the cold and dry nesting season and high levels during the hot and rainy mating season. In females, testosterone and corticosterone also rose during the hot and rainy mating season. Both hormones peaked during the second half of the mating season and decreased during the cooler dry season. Female estradiol levels increased at the onset of the mating season, reaching the highest level at the peak of the mating season, which coincided with the highest annual temperatures measured. Estradiol slowly decreased within the next months and rapidly dropped at the onset of the nesting season when temperatures decreased. Progesterone levels were high close to the time of ovulation and showed clearly elevated levels at the beginning of the nesting season after some females had laid their first clutch. Progesterone decreased during the nesting season, when ambient temperatures began to decrease, and reached minimal levels in the postbreeding period shortly before the onset of the next mating season. There were significant annual variations in plasma testosterone in both males and females. Plasma corticosterone was generally higher in males than in females and varied throughout the year in both sexes.     

Effect of age, breeding experience and senescence on corticosterone and prolactin levels in a long-lived seabird: the wandering albatross

Although several studies have described age-specific pattern of breeding performance, physiological and endocrine mechanisms underlying the variation in reproductive success in relation to age are poorly understood. We described baseline levels of: (1) corticosterone, which can trigger nest desertion when secreted at high levels, and (2) prolactin, a hormone known to trigger parental behaviour in incubating known-aged (7-39 years old) wandering albatrosses (Diomedea exulans). In this long-lived species, reproductive performance increases with age and breeding experience before it stabilizes in middle-aged individuals and declines in senescent individuals. We found that breeding experience was a much better statistical predictor of hormone levels than age. Baseline corticosterone levels slightly increased with advancing experience and reached maximal levels about the sixth breeding attempt. Similarly, prolactin levels were positively correlated with breeding experience in males. No such relationship was found in females, but first-time breeding females had lower prolactin levels than experienced females. This parallel increases in hormonal levels with advancing experience could result from improvements of skills, development of an endocrine system adapted to energetic constraints of reproduction or may mirror a higher investment in reproduction. Corticosterone levels decreased in senescent birds, but such a decline was not observed for prolactin. Low corticosterone levels in senescent birds could be associated with a lower ability to secrete/sustain elevated corticosterone levels or with a decreased parental expenditure.     

Environmental and endocrine control of reproduction in the song sparrow, Melospiza melodia. I. Temporal organization of the breeding cycle

Endocrinologic investigations of free-living populations of song sparrows, Melospiza melodia, have revealed temporal patterns of secretion of reproductive hormones that differ from those of other monogamous avian species. Males arrive in the breeding area in March whereas females arrive 1-2 weeks later. In males the periods of territory establishment and attraction of a mate are characterized by high circulating levels of luteinizing hormone (LH) and testosterone (T), whereas testis mass is low, and growth of the cloacal protuberance (CPL) is just beginning. In April, testes and CPL develop rapidly reaching a peak in early May when females are laying eggs, and when most copulations occur. Plasma LH and T decline in early April, but increase for a second time in late April and early May coincident with the egg-laying period. Thereafter, circulating LH and T decline during the parental phase, but not to basal levels. Although there is an increase in LH during the egg-laying period for the second clutch, there is no change in T levels. Testis mass, CPL, plasma LH, and T all decline to basal levels simultaneously in late July and August. The high levels of LH and T in March, followed by a temporary decline and resurgence in April and May, indicate that environmental factors in addition to the well known effects of increasing day length can regulate secretion of these hormones. In females, plasma levels of T and dihydrotestosterone (DHT) are also elevated in March and early April, and decline to low levels as the nesting phase progresses. A further decline, to basal levels, occurs in August during the molt. LH and estradiol (E2) titers in females show two peaks coincident with the egg-laying periods for each clutch. Plasma levels of corticosterone (B) increase during the breeding season in males, but not females. In both sexes B levels are basal during the moult and increase in October after moult is completed. Body mass and fat depot decline in males as the nesting phase progresses, and then increase dramatically after breeding is terminated. As expected, females show two peaks of body mass and fat depot coincident with the two periods of egg laying. The postbreeding increases in body mass and fat depot are much less pronounced in females than in males.     

The prolactin response to an acute stressor in relation to parental care and corticosterone in a short-lived bird, the Eurasian hoopoe

Prolactin plays an important role in mediating parental care in birds, but little is known about changes in prolactin levels when animals disrupt their reproductive behaviour during emergency life-history stages. We investigated the variation of prolactin levels with breeding stage, sex, body condition and as a response to a standardized acute stressor in a small short-lived bird, the Eurasian hoopoe Upupa epops under natural field conditions. We found higher baseline levels of prolactin in females during the brooding phase than in their mates which feed them and their chicks at this stage. Moreover, this is the first report of a differential prolactin stress-response between sexes with contrasting parental care within a breeding phase. Capture, handling and restraint induced a clear decrease of prolactin levels which was less pronounced in females at the very early stage of brooding compared to females in later stages. In contrast, the prolactin stress response in males remained nearly constant over the breeding stages and was stronger than in females. Baseline levels of prolactin, but not handling-induced levels, were positively correlated with body condition. We found a weak relationship between the decrease in prolactin due to acute handling stress and handling-induced levels of corticosterone. Taken together, both baseline and stress response levels of prolactin were related to the amount of parental care, although we found no relationship with reproductive success. It appears that the response to an acute stressor in prolactin levels is finely tuned to parental duties and investment. Hence, prolactin appears to be involved in mediating the trade-off between current reproduction versus self-maintenance and future reproduction.     

The glucocorticoid stress response is attenuated but unrelated to reproductive investment during parental care in a teleost fish

We investigated whether circulating glucocorticoids and androgens are correlated with reproductive investment in smallmouth bass (Micropterus dolomieu), a teleost fish with sole paternal care. Circulating cortisol and androgens prior to and 25 min following a standardized 3 min emersion stressor were quantified for non-reproductive and parental fish across the parental care period. To experimentally investigate the influence of reproductive investment on endocrine parameters, we manipulated brood size (reduced, enlarged, sham-treated, or unmanipulated) 24 h prior to sampling parental fish. We predicted that fish guarding offspring would exhibit increased androgens and baseline cortisol levels, and an attenuated cortisol response to the stressor when compared with non-reproductive individuals. We further predicted that these effects would scale with reproductive investment. As predicted, parental care-providing fish exhibited lower post-stress plasma cortisol concentrations than non-reproductive fish. This difference was strongest early during parental care. However, no differences in baseline or post-stress cortisol concentrations were detected among parents guarding offspring with varying brood sizes. There was, however, a trend for parental fish to exhibit an increased cortisol response following brood manipulation, regardless of the direction of change in brood size, a response that likely reflected disturbance. No differences were found in baseline cortisol concentrations. Circulating androgens were found to be highest during early parental care, and no differences were found among parents guarding manipulated broods. Collectively, these findings demonstrate that the endocrine stress response is affected by reproductive status, but the response in this model species does not appear to be scaled according to reproductive investment as predicted by life-history theory.     

Seasonal steroid hormone levels and their relation to reproduction in the Western Diamond-backed Rattlesnake, Crotalus atrox (Serpentes: Viperidae)

We report seasonal variation in steroid hormone levels in blood samples from free-ranging Western Diamond-backed Rattlesnakes (Crotalus atrox), and the relationship between these hormones and events in the reproductive cycle. At a field site in the Sonoran Desert of south-central Arizona, we collected monthly blood samples over the course of two active seasons from 17 radiotelemetered females, and over three active seasons from 103 randomly encountered males. We used radioimmunoassay to measure plasma levels of 17beta-estradiol, progesterone, corticosterone, and testosterone in samples from females, and corticosterone and testosterone in samples from males. Non-reproductive females have consistently low levels of circulating 17beta-estradiol, progesterone, and testosterone throughout the year. In reproductive females, 17beta-estradiol levels increase dramatically and testosterone levels increase modestly during vitellogenesis in April and May, while progesterone levels increase dramatically at ovulation in June and then steadily decline until parturition in August. Corticosterone levels appear relatively constant in non-reproductive females, whereas reproductive females show increased levels at the end of gestation. Plasma testosterone levels in males are low in early summer and are elevated during spring and late summer, corresponding to the two mating periods of C. atrox. Plasma corticosterone levels in males did not vary seasonally and were not related to testosterone levels.     

Corticosterone stress response and plasma metabolite levels during breeding and molt in a free-living migratory songbird, the wood thrush (Hylocichla mustelina)

Many birds face energetic trade-offs between different life history stages, such as reproductive effort, feather molt and the non-breeding period. Little is known about how physiological measures of condition (corticosterone, plasma metabolites) in free-living birds change from nesting stages to the post-breeding molt period or whether this is influenced by prior reproductive effort. We evaluated whether corticosterone (CORT) and plasma metabolite levels vary with date, nest stage and sex in a free-living migratory songbird, the wood thrush (Hylocichla mustelina). We also tested whether (1) baseline CORT levels early in the season were predictive of subsequent reproductive success and (2) whether prior reproductive effort influenced CORT levels and blood metabolites during molt. Baseline CORT levels decreased with date during both the incubation stage and nestling stage, but did not vary significantly across stage of breeding season. Stress-induced CORT declined with date during incubation and varied significantly across breeding stage, with lower levels during feather molt. Profiles of the metabolites of β-hydroxybutyrate, glycerol, and triglyceride did not vary significantly with date or breeding stage. Only triglycerides varied significantly with sex, with females having higher levels than males. Reproductive output was highly variable (0-10 fledglings per season) but baseline CORT levels in females during the first incubation period of the season was not related to subsequent reproductive output. Prior reproductive effort, measured as the cumulative number of young hatched during the breeding season, was positively related to stress-induced CORT during molt. High reproductive effort in wood thrush appears to have physiological carry-over effects into the molt period which could potentially affect rate of molt and preparation for fall migration.     

Seasonal Changes in Follicle-Stimulating Hormone in Free-Living Great Tits

We describe the annual cycle in plasma levels of FSH in free-living male and female great tits from southwest Sweden. Both juvenile (here defined as first-time breeders or birds <1 year old) and adult great tits, of both sexes, showed clear annual cycles with three periods of elevated FSH levels: period from territorial establishment till end of breeding (mid-March to June), October (a time when the birds break photorefractoriness), and winter (January–February). Significant differences between ages and between sexes occurred only during March and April (period of territorial establishment and gonadal recrudescens). Male FSH levels increased significantly as early as between early February and early March. Levels continued to increase until mid-April when maximal values had been reached in paired males. Territorial, but unpaired, males had significantly lower plasma levels of FSH in mid-April than did territorial and paired males. After this, FSH levels did not change until levels had decreased to basal in early July. On average, females did not show a vernal increase in FSH levels until early/mid April. However, just as in males, female FSH levels were dependent upon whether she was paired or unpaired. Females having a mate in mid-April had significantly higher FSH levels than did single females. Contrary to the males, females showed a transitory FSH peak during egg-laying. Females showed no differences between other breeding stages, until basal levels were reached during incubation of the second clutch in July. Adult birds (of both sexes) had significantly higher plasma levels of FSH than juvenile birds of the same sex during the period March–April. Furthermore, during this period adult males continually had higher FSH levels than did adult females, and juvenile males continually had higher levels than did juvenile females. In all groups a second period of elevated FSH levels occurred during early October, a time when the great tits break photorefractoriness. All groups showed basal levels during November and December. By January all four groups had increased their circulating levels of FSH to moderately high. This elevated level was maintained during February and was at the same level as that observed in October.     

Circulating corticosterone levels in breeding blue tits Parus caeruleus differ between island and mainland populations and between habitats

Little is known about whether adaptations to an insular life also involve adaptations in basal corticosterone levels or in the adrenocortical stress response, thus being part of a genetically based island syndrome. However, differences in corticosterone between island and mainland may also be a direct phenotypic response to differences in environmental conditions or may depend on individual characteristics of the animal such as body condition or parental investment. In this paper, we investigated whether insular (Island of Corsica) and mainland (nearby Southern France) blue tits Parus caeruleus populations differed in baseline and handling-stress induced corticosterone levels during the breeding season as a response to biological changes of insular biota. We also examined whether corticosterone levels of both mainland and insular blue tits differed between birds living in two different habitats (summergreen and evergreen oak woods) that differ in food availability and whether individual characteristics affected corticosterone levels. We found (a) differences in baseline corticosterone plasma levels between Corsica and the mainland, independent of regional differences in fat scores, (b) a regional difference in the relationship between corticosterone levels and brood size, (c) a difference in the rapidity of onset of the stress response to handling between habitats, independent of region, and (d) a negative relationship between body fat stores and baseline corticosterone levels independent of region. Reduced baseline corticosterone levels on Corsica may be a component of the insular syndrome, allowing birds to be less aggressive and to enhance parental investment despite higher breeding densities. We suggest that baseline corticosterone levels are only elevated if food availability affects directly the parents. However, when conditions deteriorate unexpectedly (as mimicked by handling stress), food allocation between parents and offspring needs to be re-adjusted in favor of the parents, possibly by increased circulating corticosterone levels. The switch to self-maintenance seems to be modified by the amount of body energy stores.     

Serum 11-ketotestosterone and testosterone concentrations associated with reproduction in male bluegill (Lepomis macrochirus: Centrarchidae)

Male bluegill (Lepomis macrochirus) display a complex reproductive behavior involving two alternative life history pathways: delay of sexual maturation to become "parentals" or precocious maturation as "cuckolders." The purpose of our study was to investigate the association of two androgens, 11-ketotestosterone (11KT) and testosterone (T), with reproduction in these two types of males. Radioimmunoassay techniques were used to measure daily levels of the two androgens in the blood serum of parental male bluegill captured during the prespawning, spawning, and nesting periods throughout the reproductive season. Dramatic changes in the levels of 11KT and T were observed among parental males during these periods. Peaks occurred at the onset of spawning activity during each breeding bout. Compared to spawning parental males, spawning cuckolder males had significantly lower serum levels of 11KT. In contrast, the serum levels of T among parental and cuckolder males were not significantly different. These findings suggest that the elevated levels of 11KT are associated with the behaviors displayed by spawning parental males. The levels of T, however, seem to be associated with the occurrence of a phenomenon common to both parental and cuckolder males, such as development of gonads and/or spermiation.     

Plasma glucocorticoid concentrations and body mass in ground squirrels: seasonal variation and circannual organization

We examined variation in plasma glucocorticoid concentrations of free-living Belding's ground squirrels (Spermophilus beldingi) and captive golden-mantled ground squirrels (Spermophilus lateralis) housed in constant environmental conditions. Plasma corticosterone concentrations were not significantly correlated with time held captive in traps prior to collection of blood samples in males or non-breeding females, but these variables were significantly correlated in breeding females during the pre-mating, lactation, and post-lactation periods. Among male S. beldingi, plasma corticosterone concentrations increased over the course of the active season, and were significantly higher in non-breeding than breeding individuals. Corticosterone concentrations also increased in non-breeding females throughout the active period. In breeding females, baseline (non-stress) corticosterone concentrations, determined from blood samples collected within 3 min of capture, increased during gestation and declined during lactation, whereas stress values of corticosterone, determined from blood samples collected within 4-11 min of capture, increased during gestation and then again during the period after young emerged from the natal burrow. Changes in plasma corticosterone concentrations of S. beldingi paralleled changes in body mass. Among S. lateralis, plasma concentrations of cortisol were elevated when males and females were in reproductive condition and lower in reproductively quiescent squirrels. The annual peak in plasma cortisol concentrations occurred just prior to increases in body mass associated with pre-hibernation fattening. Collectively, these results suggest that concentrations of circulating glucocorticoids fluctuate with circannual rhythmicity in conjunction with annual cycles of change in body mass and activity; within these circannual cycles glucocorticoid concentrations are influenced by stress and breeding status.     

Changes in baseline and stress-induced glucocorticoid levels during the active period in free-ranging male and female little brown myotis, Myotis lucifugus (Chiroptera: Vespertilionidae)

Baseline and stress-responsive glucocorticoid (GC) levels were characterized during the active period in free-ranging male and reproductive female little brown myotis (Myotis lucifugus). Bats were trapped and blood was sampled within 3 min of capture at two maternity sites during the summer and at one swarming site prior to hibernation in New England. Both GC hormones, cortisol and corticosterone, were detected, with cortisol accounting for an average of approximately 95% of total circulating GCs. Samples collected at the dusk emergence and after the first return from feeding showed significant seasonal differences across the active period (early pregnancy, mid-to-late pregnancy, lactation [and comparable mid-summer times for males], and pre-hibernation) within and between each sex. Elevated baseline values were found in mid-to-late pregnancy females at emergence, and in both males and females at the swarming site compared to other groups. Female GC values during mid-to-late pregnancy and during the pre-hibernation period were greater than those for males. Significantly higher GC levels following 15 min of restraint were exhibited by all animals in the summer and prior to hibernation. There was little variation between groups or sexes in the total GC levels reached following restraint. Taken together, these results suggest that: (1) GCs may be involved in the increased feeding and/or fat deposition characteristic of pregnancy and the pre-hibernation period, (2) GCs may be related to mating and to the generally increased levels of activity that occur during the pre-hibernation period, and (3) regardless of sex or reproductive condition, all animals maximally respond to restraint stress.     

Glucocorticoid response to food availability in breeding barn swallows (Hirundo rustica)

Parents feeding altricial nestlings have to trade-off the competing demands of self-maintenance and reproductive investment over their lifetime. Corticosterone, a glucocorticoid hormone released by birds in response to stressors, might play a key role in regulating parental investment when conditions unexpectedly deteriorate. However, birds breeding in unpredictable environmental conditions have been shown not to increase circulating levels of corticosterone as a response to bad weather to avoid nest abandonment when investment in offspring is high or when the probability of re-nesting is low. We investigated whether parent barn swallows Hirundo rustica, a passerine bird whose aerial insect food varies greatly in abundance depending on weather, also belongs to those species or whether it responds with an activation of the hypothalamo-pituitary-adrenal axis to natural variation in insect availability. We correlated plasma corticosterone levels of parents with weather conditions, the availability of aerial insects and parental body condition. Plasma corticosterone concentrations increased when mean daytime temperature declined, and consequently insect availability decreased and body condition of the parents deteriorated. Low temperatures also had a negative effect on body mass of the nestlings and there was a negative relationship between circulating corticosterone of parents and body mass of nestlings. We conclude that corticosterone is probably involved in the regulation of parental investment.     

Effects of trapping and subsequent short-term confinement stress on plasma corticosterone in the brown treesnake (Boiga irregularis) on Guam.

The effects of capture in a live trap and subsequent handling stress on plasma concentrations of corticosterone and other sex steroids were examined in wild male and female brown treesnakes (Boiga irregularis), an introduced species on Guam that has been implicated in the extirpation or decline of many of that island's vertebrate species. Males and females that spent 1 night in a trap had plasma levels of corticosterone about four and two times higher, respectively, than those of the respective free-ranging controls. Mean plasma levels of corticosterone of snakes that had spent 3 nights in a trap were intermediate between, but not significantly different from, those of snakes that had spent 1 night in a trap and free-ranging snakes, suggesting that some acclimation to capture occurred during this period. Snakes that were taken from traps and held in collecting bags for 10 min and 2 h prior to blood sampling had levels of corticosterone about two and three times higher, respectively, than those of control snakes that were taken from traps and bled immediately. Concentrations of plasma corticosterone in free-ranging females were about two times higher than those of males but were well within the range of basal levels observed in other reptiles. Few snakes of potential reproductive size were reproductive (males: 1 of 35; females: 2 of 33), and plasma concentrations of testosterone and progesterone in nonreproductive males and females, respectively, were accordingly low. The possible relationship between corticosterone and these sex steroids, therefore, could not be adequately assessed, although there was a positive relationship between plasma progesterone and corticosterone in the nonreproductive females. Nonetheless, as a prerequisite for studies on the seasonal hormonal cycles of this species on Guam, our observations raise the possibility that the stress caused by trapping could affect the levels of other sex steroids and that, therefore, such studies should use free-ranging individuals.     

Endocrine changes during natural spawning in the white sucker, Catostomus commersoni. I. Gonadotropin, growth hormone, and thyroid hormones

White suckers (Catostomus commersoni; Cypriniformes, Teleosteii) spawning in a small stream in central Alberta were captured during different stages of their spawning migrations in 1981 and 1982, blood was sampled, and the fish were examined to determine their reproductive condition. Blood samples were analyzed for gonadotropin (GtH), growth hormone (GH), triiodothyronine (T3), and thyroxine (T4) by radioimmunoassay. GtH levels in both sexes were lowest prior to the onset of spawning, increased significantly in spawning males, females in which germinal vesicle migration had begun, and ovulated females and then dropped significantly in spent fish of both sexes. GH was lowest in prespawning females, increased significantly at ovulation, and remained high in spent females. In contrast, GH levels in males were relatively constant throughout spawning. In both sexes, highest T4 levels were found in prespawning fish, and T4 decreased significantly in spent fish. Although a similar decline was seen in T3 in 1981, in 1982 there were no T3 changes associated with changes in reproductive condition. No significant diurnal variations were detected in the levels of GtH or T3; T4 levels appeared to vary on a diurnal basis in prespawning males only. Spawning activity in both sexes therefore appears to be associated with increases in GtH occurring at ovulation in females and at the initiation of spawning in males. GH levels may also be related to reproductive condition in females, but not in males. The relationship of thyroid hormone levels to reproductive condition is less clear, however, and these levels may reflect both endocrine and environmental influences on thyroid function.     

High levels of LH and testosterone in a tropical seabird with an elaborate courtship display

For birds breeding in temperate areas, territoriality and courtship co-occur during spring and are mirrored by elevated plasma levels of luteinizing hormone (LH) and testosterone. By contrast, several tropical species show no major changes in LH and testosterone throughout the year and plasma levels are often low. This study describes changes in body condition, LH and testosterone levels throughout breeding in a tropical seabird, the magnificent frigatebird (Fregata magnificens). Although bi-parental care is the rule in frigatebirds, these seabirds are unique among seabirds in having a lek-like mating system where males perform an elaborate courtship display during which an extravagant sexual ornament (gular pouch) is inflated. Body condition peaked during the displaying stage in males and declined gradually in both sexes, being the lowest during chick-rearing. LH and testosterone titers were the highest in displaying males with a fully inflated gular pouch (means: 23.6 ng/ml for LH and 6.2 ng/ml for testosterone) and remained low (means: 3.8 ng/ml for LH and 0.3 ng/ml for testosterone) during incubation and chick-rearing in both sexes. Body condition was positively related to testosterone levels in displaying males. In male frigatebirds, high levels of testosterone are not associated with aggression but are probably required to display the inflated gular pouch and to perform an elaborate courtship display. Because males invest in incubation and chick-rearing, testosterone has to return to low levels, to permit parental care. This hormonal pattern differs greatly with that of the few tropical seabirds studied so far and is probably related to the unique reproductive strategy of frigatebirds.     

Relation of glucocorticosteroids and testosterone to the annual cycle of free-living degus in semiarid central Chile.

We investigated seasonal patterns of plasma glucocorticosteroids (GCs) in both sexes and testosterone (T) in males in relation to the annual cycle in central Chile of a natural population of the degu (Octodon degus), a caviomorph rodent. We wanted to find out which GCs are present in degus, whether their seasonal variation suggests suppressive or synergistic interrelationships with T, and whether seasonal variation in GC levels indicates a relationship with energy mobilization and demands of reproduction. Degus mated in late autumn, and female body mass increased in pregnancy and remained high during lactation and throughout spring. Over the subsequent period of summer drought both sexes declined to a minimal body mass before the next mating season. Cortisol appears to be the principal GC in degus. In fact cortisol levels were so high that the extremely low levels of corticosterone measured were probably largely due to the cross-reactivity of our corticosterone antiserum with cortisol. Titers of cortisol in females exceeded 1000 ng/ml at lactation in the spring of 2 years; cortisol declined greatly following lactation and during the summer and reached its lowest mean level of about 500 ng/ml at mating. Males were more difficult to capture than females and thus our sampling was limited, but male cortisol levels were similar to those of females during the times of year when we measured them. Male T levels remained within a low range all year, but at mating, when mean T was highest (0.16 ng/ml) and when most males had detectable T, degus showed their lowest cortisol levels. The minimal cortisol level of males during mating represents a possible suppressive effect of T, as described in other mammals. At the time of their spring emergence, 60% of juvenile males had detectable T levels comparable to those of adults, suggesting important organizational effects of T at that time in their maturation. Peak cortisol titers in both sexes were associated with lactation in females, when energy mobilization, production, and body mass were at their greatest.