Hormonal correlates of parental behavior in yellow-eyed penguins (Megadyptes antipodes)

Penguins show varying degrees of brood reduction behavior, from obligate brood reducers to brood maximizers, and we hypothesize that this is associated with differences in prolactin secretion. To address this hypothesis, we determined the breeding season prolactin profile of the yellow-eyed penguin (Megadyptes antipodes) for comparison with those of other penguin species found in the literature. We also measured sex steroid plasma concentrations to better characterize the reproductive cycle of the species. Plasma concentrations of prolactin increased from early in the season, reaching a peak during late incubation, and remained elevated up to the guard period. This general pattern was similar to that of other penguins for which we have corresponding data. However, we found that throughout the laying period, prolactin titers in yellow-eyed penguins remained elevated while they fell to basal levels after the laying of the first egg in macaroni penguins, which corresponds to differences in incubation behavior during this time. We conclude, therefore, that differences in the brood reduction behavior in penguins, may be reflected in the pattern of PRL concentrations around the time of egg laying.     

Exogenous corticosterone and nest abandonment: A study in a long-lived bird, the Adélie penguin

Breeding individuals enter an emergency life-history stage when their body reserves reach a minimum threshold. Consequently, they redirect current activity toward survival, leading to egg abandonment in birds. Corticosterone (CORT) is known to promote this stage. How and to what extent CORT triggers egg abandonment when breeding is associated with prolonged fasting, however, requires further investigation.We manipulated free-living male Adélie penguins with CORT-pellets before their laying period. We then examined their behavioral response with respect to nest abandonment in parallel with their prolactin levels (regulating parental care), and the subsequent effects of treatment on breeding success in relieved birds.Exogenous CORT triggered nest abandonment in 60% of the treated penguins ~ 14 days after treatment and induced a concomitant decline in prolactin levels. Interestingly, prolactin levels in treated penguins that did not abandon their nest were higher at the point of implantation and also after being relieved by females, when compared with abandoning penguins. Among successful birds, the treatment did not affect the number of chicks, nor the brood mass.Our results show the involvement of CORT in the decision-making process regarding egg abandonment in Adélie penguins when incubation is associated with a natural long fast. However, we suggest that CORT alone is not sufficient to trigger nest abandonment but that 1) prolactin levels need to reach a low threshold value, and 2) a rise in proteolysis (i.e. utilization of protein as main energy substrate) seems also to be required.     

Environmental regulation of the reproductive system in a flexibly breeding Sonoran Desert bird, the Rufous-winged Sparrow, Aimophila carpalis

We investigated reproductive regulation in male Rufous-winged Sparrows, Aimophila carpalis, a Sonoran Desert passerine that breeds after irregular summer rains. Field and captive data demonstrate that increased photoperiod stimulates testicular development in March and maintains it until early September. Free-living birds caught in July and placed on captive long days (16L: 8D) maintained developed testes for up to 7 months, and free-living birds caught in September, during testicular regression, redeveloped testes when placed on captive long days, indicating that these birds were still photosensitive. Captive birds on long days maintained testicular development when exposed to temperatures mimicking those occurring during regression in free-living birds. In free-living birds, testicular development was observed during spring and summer, but unless this was associated with rainfall, breeding (indicated by juveniles) did not occur. Large increases in plasma luteinizing hormone (LH) in free-living males were correlated with heavy rainfall in July/August, when the birds bred, and in November, when they did not breed. In captive birds, plasma LH concentrations were unresponsive to photoperiodic changes, but may have responded to social cues. Plasma prolactin concentrations were directly correlated with photoperiod in free-living birds, but an effect of photoperiod on prolactin secretion was not seen in captive birds. It is concluded that male Rufous-winged Sparrows use long photoperiods to stimulate and maintain testicular development, but exposure to long photoperiods does not terminate breeding by inducing absolute photorefractoriness. The specific timing of reproductive behaviors is apparently determined by elevated plasma LH coinciding with long day stimulated gonad development.     

Prolactin's mediative role in male parenting in parentally experienced marmosets (Callithrix jacchus)

Prolactin has been implicated in promoting paternal care behaviors but little evidence of causality has been found to date except for birds and fish. This study was designed to examine the possible causal relationships between prolactin and male parenting behaviors, reproductive hormones, and physical changes in cooperatively breeding common marmosets, Callithrix jacchus. Fifteen parentally experienced fathers were studied over three consecutive infant care periods during two weeks prior and three weeks following their mates' parturition under three-treatment conditions: normal control pregnancy, decreased prolactin and elevated prolactin. The treatments significantly altered the serum prolactin levels in the fathers. Using three methods of determining a father's level of parental care: infant carrying, family effort and responsiveness to infant stimulus tests, we found that only the male response to infant stimuli was altered by the hormone treatments. Lowering prolactin significantly reduced male responsiveness to infant stimuli but elevating prolactin showed the same effect. Hormonal sampling indicated that testosterone levels showed an inverse relationship to prolactin levels during a normal peripartum period and prolactin treatment reduced this relationship. Prepartum estradiol levels were significantly elevated during the lowered prolactin treatment and estradiol was significantly lowered postpartum with the elevated prolactin treatment. Father's weight decreased significantly by the third week of infant care during the normal treatment. Males in the elevated prolactin treatment lost little or no weight from prepartum while in the lowered prolactin treatment showed the most weight loss. The present findings did not distinguish a direct causal relationship of prolactin on behavior in experienced fathers but did find an interaction with other hormones and weight gain.     

Endocrine correlates of parental care in an Antarctic winter breeding seabird, the emperor penguin, Aptenodytes forsteri

Plasma levels of luteinizing hormone (LH) and prolactin associated with parental behavior were measured in the Antarctic winter breeding emperor penguin, Aptenodytes forsteri. Males exclusively incubate the egg while females exclusively brood the nonhomeothermic young; both sexes alternate in rearing the homeothermic young. Birds were sampled on arrival from the sea through egg laying, incubation, and brooding. All parent birds lost their chicks at the end of the brooding period due to harsh weather but sampling continued. In females, LH titers dropped after egg laying but levels were restored when the birds returned from the sea to brood the chicks and were not depressed by high prolactin levels. Plasma prolactin remained low in males captured on arrival and kept until the free-living males finished incubation. In breeders, prolactin secretion increased during the prelaying period when day length decreased. Prolactin levels stayed elevated in males during incubation and in brooding females returning after a 2-month absence at sea. Prolactin values were higher in brooding females than in males ending incubation or returning in late brooding. These levels did not drop after chick loss, and the sexual difference in prolactin values was maintained after breeding failure. In emperor penguins, increased prolactin secretion appears to be triggered around the time of egg laying and continues, driven by an endogenous mechanism, through incubation and brooding until rearing is completed. Prolactin secretion independent of external stimuli may have evolved in pelagic seabirds to maintain parental care despite long absences at sea from the breeding colony.     

Avian malaria parasites (

There is concern that avian malaria may be partly responsible for fluctuations in yellow-eyed penguin (Megadyptes antipodes) populations in New Zealand. Recent findings, however, have provided no evidence of avian malaria parasites infecting yellow-eyed penguins on the Otago Peninsula, raising questions as to whether this area is currently free of such parasites. To test this possibility we collected blood samples from 109 individuals of five non-native bird species known to carry malarial parasites elsewhere in New Zealand. Molecular screening by polymerase chain reaction revealed 6% of the sampled birds were positive for malarial parasites, indicating that a local reservoir of infection is present. Sequence data revealed a generalist strain of Plasmodium is present, one that infects a number of native and non-native bird species elsewhere in the country. The absence of this generalist strain in yellow-eyed penguins, some of which were sampled during the same period as the current study, may be due to low levels of mosquito vectors of disease during the study period, low densities of non-native birds around yellow-eyed penguin colonies, or infected penguins dying before they could be sampled. Continued monitoring of mosquito populations and the factors that affect their densities should be included in the future management of native birds in this area.     

Differences in prolactin levels between three alternative male reproductive tactics in striped mice (Rhabdomys pumilio)

In male fishes, birds and mammals, increased prolactin secretion is thought to play a role in species showing paternal behaviours. This hypothesis was investigated in the striped mouse (Rhabdomys pumilio). This paper compares serum prolactin levels in 71 free-living male striped mice following three different reproductive tactics: (i) paternal group-living breeders, (ii) alloparental philopatric group-living males, and (iii) roaming non-paternal solitary males. Prolactin levels of breeding males were significantly higher than that of roamers. Alloparental philopatric males had low prolactin levels, which concur with studies of cooperatively breeding mammals, but contrasts with studies of cooperatively breeding birds. Both breeding males and females showed a decrease in prolactin levels after the breeding season, but not alloparental philopatric males. Prolactin levels were correlated with neither corticosterone levels nor age. These results are in agreement with the hypothesis that prolactin is one proximate mechanism of male reproductive tactics, possibly regulating differences in male parental care.     

Spontaneous egg or chick abandonment in energy-depleted king penguins: a role for corticosterone and prolactin?

Various exogenous or endogenous factors may induce an emergency response in birds, redirecting current activity towards survival. In fasting, breeding penguins, the achievement of a critical energy depletion was suggested to induce egg abandonment and departure to sea for re-feeding. How such a behavioral shift is hormonally controlled remains unknown. The possible involvement of corticosterone and prolactin was examined by characterizing the nutritional and hormonal states of king penguins at egg abandonment. Further, we tested if these states differ according to whether an egg or a chick is abandoned, and according to the timing of breeding. In every case of abandonment, birds were in phase III fasting characterized by accelerated protein catabolism. However, body condition at egg abandonment was lower in early than in late breeders, suggesting that king penguins are willing to tolerate a larger energy depletion when their potential breeding success is high. At egg and chick abandonment, plasma corticosterone levels were, respectively, increased by 2- and 4-fold, whereas plasma prolactin levels were, respectively, depressed by 3- and 1.4-fold. The increase in plasma corticosterone and the decrease in plasma prolactin could be involved in the control of abandonment by, respectively, stimulating the drive to re-feed and diminishing the drive to incubate or brood. The smaller decrease in prolactin levels and the greater increase in corticosterone levels observed at chick vs egg abandonment suggest that, in addition to nutritionally-related stimuli, tactile or audible stimuli from the egg or chick could intervene in the endocrine control of abandonment.     

Hormones associated with non-maternal infant care: a review of mammalian and avian studies

Hormonal changes during non-maternal infant care have been demonstrated in many cooperatively breeding bird species, some monogamous rodents and two species of New World primates. Coevolution of hormones and social traits may have provided for the different breeding systems that occur today. Several hormones have been shown to covary with the breeding systems of vertebrates. Elevated levels of the hormone prolactin with male parenting behaviours are common to many birds, rodents and the callitrichid monkeys Callithrix jacchus and Saguinus oedipus. In birds, prolactin may be elevated in both male and female breeders during various stages of nest building, egg laying, incubating and feeding of young. Testosterone levels appear to have an inverse relationship to prolactin levels during infant care in birds and rodents, but this relationship has not been examined for primates. In cooperatively breeding birds, helpers who remain at the nest also have elevated levels of prolactin when displaying parental care behaviours. Prolactin levels are elevated in helper callitrichid monkeys during the postpartum period. Monogamous male rodents demonstrate elevated prolactin levels with parental care behaviour but, in contrast to the birds, the mechanisms mediating prolactin increase appear to differ for male and female rodents. Two factors may influence male parental behaviours and hormonal changes: stimuli from the pregnant female and stimuli from the newborn pups; whereas maternal behaviours are influenced by the maternal hormones of the female and the pup stimuli. An experiential factor may also influence male parental behaviours. Neuropeptides such as oxytocin and vasopressin appear to be involved in male rodent parental care and there may be an interaction between a series of hormones and neurosecretions and stimuli from mates and pups. Studies of Saguinus oedipus, the cotton-top tamarin, suggest that prolactin levels are responsive to stimuli from contact with infants and the level of infant care experience influences the levels of prolactin with male infant care. Father tamarins also have elevated levels of prolactin before the birth of infants suggesting that cues from the pregnant female are important. Prolactin's role in parental care may have evolved from prolactin's role in other reproductive functions. Hormonal regulation of non-maternal care may occur due to a complex interaction of many hormones and neurotransmitters. Studies described here should provide the impetus for further work on parental care hormones in a wide variety of primates.     

Auditory stimulation of reproductive function in male Rufous-winged Sparrows, Aimophila carpalis

Prolonged exposure to conspecific song stimulates gonadal function and reproductive hormone secretion in female birds but few studies have investigated the physiological effects of conspecific song exposure on males outside of short-term, aggressive interactions. We exposed male Rufous-winged Sparrows, Aimophila carpalis, either to conspecific song (CS Song), to heterospecific song (Black-throated Sparrow, Amphispiza bilineata; HS Song), or to no recorded song (No Song) for 59 consecutive days (two h per day). Birds were exposed to short days (8L:16D) for the first 21 days of treatment and were then transferred to long days (13L:11D) for the remaining 38 days. During long day exposure, CS Song birds experienced faster growth of testes than HS Song and No Song birds. HS Song birds also grew their testes faster than No Song birds. Plasma luteinizing hormone (LH) and testosterone did not differ between CS Song and No Song birds. However, plasma LH was higher in HS Song birds compared to other groups. There were no differences in hypothalamic immunocytochemical labeling for gonadotropin-releasing hormone, its precursor proGnRH, or gonadotropin-inhibitory hormone, nor were there differences in two song control nuclei volumes (HVC and RA) between CS Song and No Song treatment groups. Furthermore, we found no effect of heterospecific song on free-living Rufous-winged Sparrow aggressive behaviors. These data indicate that long-term exposure to auditory stimuli, such as song, can influence the reproductive system of male songbirds and different types of auditory stimuli can have differential effects on reproductive function.     

Comparison of population numbers of yellow-eyed penguins,

During a comprehensive survey in 1999, 2000 and 2001, we investigated the number of breeding yellow-eyed penguin pairs on Stewart Island, where cats are present, and on adjacent cat-free islands. We found 79 pairs of yellow-eyed penguin breeding in 19 discrete locations on Stewart Island (4.2 pairs per location), and 99 pairs breeding in 10 discrete locations on all cat-free islands (9.9 pairs per location). Large-scale humaninduced habitat modifications have not occurred on Stewart Island, nor on any of its adjacent offshore islands. While the extensive coastline of Stewart Island (673 km) offers potentially large areas of breeding habitat for penguins, the highest number of breeding pairs were found on the smaller, predator-free Codfish Island (25 km coastline), where a total of 61 breeding pairs were recorded. On Stewart Island, where mustelids do not occur, only feral cats can pose a serious threat to penguin offspring. Results from this study suggest that feral cats may prey on yellow-eyed penguins on Stewart Island. Further work is necessary to investigate whether the observed low numbers of yellow-eyed penguins on Stewart Island are caused by feral cat predation. If so, it may be possible to develop appropriate measures to protect this penguin species from a population decline.     

Straight Line Foraging in Yellow-Eyed Penguins: New Insights into Cascading Fisheries Effects and Orientation Capabilities of Marine Predators

Free-ranging marine predators rarely search for prey along straight lines because dynamic ocean processes usually require complex search strategies. If linear movement patterns occur they are usually associated with travelling events or migratory behaviour. However, recent fine scale tracking of flying seabirds has revealed straight-line movements while birds followed fishing vessels. Unlike flying seabirds, penguins are not known to target and follow fishing vessels. Yet yellow-eyed penguins from New Zealand often exhibit directed movement patterns while searching for prey at the seafloor, a behaviour that seems to contradict common movement ecology theories. While deploying GPS dive loggers on yellow-eyed penguins from the Otago Peninsula we found that the birds frequently followed straight lines for several kilometres with little horizontal deviation. In several cases individuals swam up and down the same line, while some of the lines were followed by more than one individual. Using a remote operated vehicle (ROV) we found a highly visible furrow on the seafloor most likely caused by an otter board of a demersal fish trawl, which ran in a straight line exactly matching the trajectory of a recent line identified from penguin tracks. We noted high abundances of benthic scavengers associated with fisheries-related bottom disturbance. While our data demonstrate the acute way-finding capabilities of benthic foraging yellow-eyed penguins, they also highlight how hidden cascading effects of coastal fisheries may alter behaviour and potentially even population dynamics of marine predators, an often overlooked fact in the examination of fisheries’ impacts.     

Low sex steroids, high steroid receptors: Increasing the sensitivity of the nonreproductive brain

Male aggressive behavior is generally regulated by testosterone (T). In most temperate breeding males, aggressive behavior is only expressed during the reproductive period. At this time circulating T concentrations, brain steroid receptors, and steroid metabolic enzymes are elevated in many species relative to the nonreproductive period. Many tropical birds, however, display aggressive behavior both during the breeding and the nonbreeding season, but plasma levels of T can remain low throughout the year and show little seasonal fluctuation. Studies on the year-round territorial spotted antbird (Hylophylax n. naevioides) suggest that T nevertheless regulates aggressive behavior in both the breeding and nonbreeding season. We hypothesize that to regulate aggressive behaviors during the nonbreeding season, when T is at its minimum, male spotted antbirds increase brain sensitivity to steroids. This can be achieved by locally up-regulating androgen receptors (ARs), estrogen receptors (ERs), or the enzyme aromatase (AROM) that converts T into estradiol. We therefore compared mRNA expression of AR, ERalpha, and AROM in free- living male spotted antbirds across reproductive and nonreproductive seasons in two brain regions known to regulate both reproductive and aggressive behaviors. mRNA expression of ERalpha in the preoptic area and AR in the nucleus taeniae were elevated in male spotted antbirds during the nonbreeding season when circulating T concentrations were low. This unusual seasonal receptor regulation may represent a means for the year-round regulation of vertebrate aggressive behavior via steroids by increasing the brain's sensitivity to sex steroids during the nonbreeding season.     

Diet of feral house cats Felis catus, ferrets Mustela furo and stoats M. erminea in grassland surrounding yellow-eyed penguin Megadyptes antipodes breeding areas, South Island, New Zealand

Spring and summer diets of feral house cats ( Felis catus ), ferrets ( Mustela furo ) and stoats ( M. erminea ) were studied in grassland surrounding breeding areas of yellow-eyed penguins ( Megadyptes antipodes ), a regionally threatened native species. All three predator species ate large numbers of young rabbits ( Oryctolagus cuniculus ) and birds. Stoats also relied heavily on mice ( Mus musculus ). Use of rabbits increased in rank order of increasing predator size, and male stoats ate more lagomorphs than female stoats. Diet differences may reflect character displacement as a result of exploitation competition, but interference competition or predation may force the smaller species to exploit micro-habitats with increased ground cover and consequent increased availability of smaller prey. Reduction of predation of native species like yellow-eyed penguins by decreasing or increasing staple mammal prey numbers of the introduced predators may provide lasting conservation benefits, but could also trigger diet changes that increase risk to endangered wildlife.     

Importance of Montane Riparian Forest and Influence of Wildfire on Nest-Site Selection of Ground-Nesting Birds

Abstract: We studied breeding populations of 2 coexisting ground-nesting birds, the red-faced warbler (Cardellina rubrifrons) and yellow-eyed junco (Junco phaeonotus), in high-elevation (>2,000 m) forested drainages of the Santa Catalina Mountains, Arizona, USA. From 2004 to 2005, we 1) estimated density and nesting success of breeding populations of red-faced warblers and yellow-eyed juncos, 2) identified nest-site characteristics for each species (i.e., used sites vs. random plots), 3) compared nest-site characteristics between the 2 species, and 4) examined effects of a recent (2003) wildfire on distribution of nests of both species. In addition, we estimated the areal extent of montane riparian forest (the preferred breeding habitat of both species) within high-elevation forests of the Santa Catalina Mountains. We found that red-faced warblers and yellow-eyed juncos were the 2 most common ground-nesting birds within our study area with an average density of 2.4 and 1.4 singing males/ha, respectively, along drainage bottoms. Compared to random plots, most red-faced warbler and yellow-eyed junco nests were located close (≤30 m) to drainage bottoms within a strip of montane riparian forest characterized by abundant brush, small woody debris, ferns, and forbs (both species), high number and diversity of saplings and small trees (red-faced warblers), and abundant shrubs and downed logs and less canopy cover (yellow-eyed juncos). Although both species nested in close proximity within montane riparian forest, nest-site characteristics differed between the 2 species, especially at finer spatial scales. For example, most yellow-eyed juncos nested adjacent to grass (principally Muhlenbergia spp.), whereas red-faced warblers nested adjacent to a variety of plant species, including grass, bigtooth maple (Acer grandidentatum), white fir (Abies concolor), and Douglas-fir (Pseudotsuga menziesii). Both red-faced warblers and yellow-eyed juncos avoided nesting in areas burned during a recent wildfire. In addition, nesting success was low for red-faced warblers (13%) and yellow-eyed juncos (19%) following the wildfire, suggesting an indirect negative effect of fire on breeding populations in the short-term. Montane riparian forest appears to provide important breeding habitat for red-faced warblers and yellow-eyed juncos. However, little research or conservation planning has been directed toward montane riparian forest in the region, even though this forest type is limited in its areal extent (<11% of high-elevation forest in the Santa Catalina Mountains) and increasingly threatened by disturbance. Results from our study can be used to facilitate the management and conservation of breeding populations of red-faced warblers and yellow-eyed juncos in high-elevation forests of the southwestern United States.     

Testosterone alters genomic responses to song and monoaminergic innervation of auditory areas in a seasonally breeding songbird

Behavioral responses to social stimuli often vary according to endocrine state. Our previous work has suggested that such changes in behavior may be due in part to hormone‐dependent sensory processing. In the auditory forebrain of female white‐throated sparrows, expression of the immediate early gene ZENK (egr‐1) is higher in response to conspecific song than to a control sound only when plasma estradiol reaches breeding‐typical levels. Estradiol also increases the number of detectable noradrenergic neurons in the locus coeruleus and the density of noradrenergic and serotonergic fibers innervating auditory areas. We hypothesize, therefore, that reproductive hormones alter auditory responses by acting on monoaminergic systems. This possibility has not been examined in males. Here, we treated non‐breeding male white‐throated sparrows with testosterone to mimic breeding‐typical levels and then exposed them to conspecific male song or frequency‐matched tones. We observed selective ZENK responses in the caudomedial nidopallium only in the testosterone‐treated males. Responses in another auditory area, the caudomedial mesopallium, were selective regardless of hormone treatment. Testosterone treatment reduced serotonergic fiber density in the auditory forebrain, thalamus, and midbrain, and although it increased the number of noradrenergic neurons detected in the locus coeruleus, it reduced noradrenergic fiber density in the auditory midbrain. Thus, whereas we previously reported that estradiol enhances monoaminergic innervation of the auditory pathway in females, we show here that testosterone decreases it in males. Mechanisms underlying testosterone‐dependent selectivity of the ZENK response may differ from estradiol‐dependent ones.© 2013 Wiley Periodicals, Inc. Develop Neurobiol 73: 455–468, 2013     

Should I stay or should I go? Hormonal control of nest abandonment in a long-lived bird,the Adélie penguin

According to life-history theory, long-lived birds should favor their survival over the current reproductive attempt, when breeding becomes too costly. In seabirds, incubation is often associated with spontaneous long-term fasting. Below a threshold in body reserves, hormonal and metabolic shift characteristics of a switch from lipid to protein utilization (phase III, PIII) occur. These metabolic changes are paralleled by nest abandonment and stimulation of refeeding behavior. Parental behavior is then under control of two hormones with opposite effects: corticosterone (CORT) and prolactin which stimulate foraging and incubation behavior, respectively.The aim of this study was to determine the respective role of these two hormones in nest abandonment by Adélie penguins. To this end, plasma hormone levels were measured before egg-laying and at departure from the colony (i.e. when birds were relieved by their partner or abandoned their nest), and related to nutritional state and incubation success.We found that males abandoning their nest in PIII presented high CORT levels and low prolactin levels. Interestingly, males which presented high plasma levels of prolactin in PIII did not abandon. We show that although CORT is the first hormone to be affected by prolonged energy constraints, the combined effects of high CORT and low prolactin levels are necessary for parents to favor self-maintenance and abandon the nest. We provide insights into time-course changes of the endocrine profile as PIII proceeds and report that reaching proteolytic late fasting is not sufficient to induce nest abandonment in a long-lived bird.     

Vocal control region sizes of an adult female songbird change seasonally in the absence of detectable circulating testosterone concentrations

Previous research established that in several species of seasonally breeding oscine birds, brain areas [vocal control regions (VCRs)] that control vocal behavior learning and expression exhibit seasonal plasticity, being larger during than outside the reproductive period. In adult males, this seasonal decrease correlates with circulating testosterone (T) concentrations. VCRs contain androgen receptors and T plays an important role in neural plasticity and in the control of singing behavior. In behaviorally dimorphic species, VCRs are larger in males than females and change seasonally also in females, but the dependency of these changes on circulating T levels in females has not been established. In free-living adult dark-eyed juncos (Junco hyemalis), a species in which females do not normally sing, the sizes of three VCRs (high vocal center, robust nucleus of the archistriatum, and Area X) were larger in males than females and decreased between summer and fall in both sexes. In males, this decrease was associated with changes in circulating T concentrations. Females, however, had on average undetectable T levels throughout the breeding season. Seasonal changes in VCR volumes in adult females may depend on very low (below detection limit) circulating T concentrations, on nonandrogenic plasma steroids, on androgen (or androgen metabolites) produced in brain tissues, and/or on nonsteroidal factors such as photoperiod or social interactions with conspecific birds.     

Should I stay or should I go? Hormonal control of nest abandonment in a long-lived bird, the Adélie penguin

According to life-history theory, long-lived birds should favor their survival over the current reproductive attempt, when breeding becomes too costly. In seabirds, incubation is often associated with spontaneous long-term fasting. Below a threshold in body reserves, hormonal and metabolic shift characteristics of a switch from lipid to protein utilization (phase III, PIII) occur. These metabolic changes are paralleled by nest abandonment and stimulation of refeeding behavior. Parental behavior is then under control of two hormones with opposite effects: corticosterone (CORT) and prolactin which stimulate foraging and incubation behavior, respectively.The aim of this study was to determine the respective role of these two hormones in nest abandonment by Adélie penguins. To this end, plasma hormone levels were measured before egg-laying and at departure from the colony (i.e. when birds were relieved by their partner or abandoned their nest), and related to nutritional state and incubation success.We found that males abandoning their nest in PIII presented high CORT levels and low prolactin levels. Interestingly, males which presented high plasma levels of prolactin in PIII did not abandon. We show that although CORT is the first hormone to be affected by prolonged energy constraints, the combined effects of high CORT and low prolactin levels are necessary for parents to favor self-maintenance and abandon the nest. We provide insights into time-course changes of the endocrine profile as PIII proceeds and report that reaching proteolytic late fasting is not sufficient to induce nest abandonment in a long-lived bird.     

Social regulation of plasma estradiol concentration in a female anuran

The behavior of an individual within a social aggregation profoundly influences behavior and physiology of other animals within the aggregation in such a way that these social interactions can enhance reproductive success, survival and fitness. This phenomenon is particularly important during the breeding season when males and female must synchronize their reproductive efforts. We examined whether exposure to conspecific social cues can elevate sex steroid levels, specifically estradiol and androgens, in female túngara frogs (Physalaemus pustulosus). We compared plasma estradiol and androgen concentrations in wild-caught females before and after exposure to either natural mate choruses or random tones. After exposure to mate choruses for 10 consecutive nights, estradiol concentrations were significantly elevated whereas there was no significant elevation in estradiol concentrations in the group of females exposed to random tones for 10 nights. Plasma androgen concentrations were not significantly changed after exposure to either natural mate choruses or random tones for 10 consecutive nights. Social modulation of estradiol concentrations may be important in maintaining a female's reproductive state while males are chorusing. To our knowledge, this is the first study to demonstrate social regulation of estradiol concentration in female anurans.