Inferring performance in the songs of dark-eyed juncos (Junco hyemalis)

Within bird species, songs differ in their attractiveness tofemales or effectiveness in male–male interactions. Somesongs are more difficult to sing than others, and receiversmay use a singer's performance of difficult songs as a meansfor evaluating the quality of the singer. The concept of songperformance aims at quantifying how physiologically demandingare different songs. Using variation between song types of dark-eyedjuncos, Junco hyemalis, we show that some song traits tradeoff with costly aspects of song output—short intervalsbetween syllables or loud sound amplitude—suggesting thatthose traits are difficult to sing. First, after controllingfor other traits, long syllables require longer intervals forrecovery. This supports the idea that a measure of "respiratoryperformance" could be based on the relative lengths of syllablesand intervals. Second, some syllable traits trade off stronglywith sound amplitude, suggesting that these traits may be difficultto sing at high amplitudes. The ratio of frequency bandwidthand trill rate has been used to infer performance in other birdspecies, but we found no evidence that frequency bandwidth tradesoff with any aspect of song output in the junco. The negativeassociation of bandwidth with trill rate may instead be a passiveconsequence of syllable length, with longer syllables randomlyaccumulating frequency modulation. We conclude that bird receiversmay best evaluate how well a song is performed if they integratemultiple cues and discuss how researchers may similarly devisemeasures of song performance.     

The distribution and turnover of 35S methionine as influenced by diet in the dark-eyed Junco (Junco hyemalis)

1. Dark-eyed Juncos (Junco hyemalis) were maintained on one of three semi-synthetic diets, differing only in their sulphur amino acid content. 2. 35S-methionine was administered intraperitoneally after 3 days on their respective diets and the animals were killed 24 or 96 hr post-injection. 3. Faecal loss of 35S-methionine was significantly lower in the low methionine/high cysteine (HC) group after 24 hr when compared with other two dietary groups (4.69 vs 5.53 and 5.47%) indicating a whole-body conservation of the limiting amino acid. 4. Incorporation of 35S-methionine in hepatic and intestinal tissues was also greater in the HC group, even as the animals were losing body weight.     

Comparative morphology of dark-eyed juncos Junco hyemalis breeding at two elevations: a common aviary experiment

Morphologies of bird species often vary along elevation gradients, yet causes of the variation have not been examined experimentally. We investigated variation in morphological traits of the dark‐eyed junco Junco hyemalis, breeding at 1,000 m a.s.l. (low‐elevation; i.e. low) and 2,000 m asl (high‐elevation; i.e. high) in the Rocky Mountains, Canada. Eight morphological traits were measured in free‐living birds. We found two consistent differences in populations between elevations: at high‐elevation sites, females had longer wings and males had longer tails than birds from low‐ elevation sites. Other age‐ and gender‐ specific results were observed in free‐living birds between elevations: tarsi were shorter in high‐elevation second year (SY) females and after second year (ASY) males, beak lengths were slightly longer in low‐elevation SY females, and high‐elevation ASY females tended to have lower fat than low‐elevation ASY females. Morphological differences may result from genetic differences between elevations, or phenotypic flexibility resulting from exposure to the different environmental conditions. To identify which mechanism caused the difference in morphometrics, hand‐reared birds from low‐ and high‐elevation habitats were raised in identical conditions with unlimited access to high quality food until they had replaced all feathers. The traits measured in the lab (wing and rectrix length, weight and fat score) tended to increase in magnitude compared to field values. Juncos from high‐ and low‐elevations had similar responses to the aviary environment, with one exception: males from high‐elevation sites had greater weight gain relative to free‐living juncos than males from low‐elevation sites. Thus, morphological traits in dark‐eyed juncos were phenotypically flexible, capable of growing larger in the laboratory environment. However, there were also persistent genetic or perinatal/maternal differences underlying population responses that prevented traits from converging under aviary conditions. As a result, trait size differences between high‐ and low‐elevation populations were maintained or exacerbated in the common aviary environment.     

Steroid hormones and immune function: experimental studies in wild and captive dark-eyed juncos (Junco hyemalis)

Monogamous and polygynous male songbirds generally differ in their breeding season profiles of circulating testosterone. Testosterone level spikes early in the breeding season of monogamists and then declines, but it remains high in polygynists. Male dark-eyed juncos (Junco hyemalis) are socially monogamous and exhibit the usual pattern, but experimental maintenance of high testosterone throughout the breeding season alters normal behavior and physiology and affects various components of annual reproductive success but not overall annual success. Because stabilizing selection predicts that alteration of naturally existing phenotypes should reduce lifetime reproductive success, we asked whether prolonged testosterone exposure might impair immune function and perhaps thereby reduce life span. We assessed immune function in captive and wild male juncos that we treated with either testosterone-filled or empty Silastic implants. Results indicate that prolonged elevation of testosterone suppresses antibody production in captive males and cell-mediated immunity in wild males. Together these results suggest that testosterone-treated males may be more susceptible to disease or parasitic infection. As earlier studies have shown, levels of corticosterone as well as testosterone are higher in testosterone-treated males, so it is unclear whether the immune suppression we observed is due to testosterone's direct effects on immunity or testosterone's influence on glucocorticoid production. We discuss results in the context of recent hypotheses regarding life-history theory and potential endocrine-immune interactions.     

Effects of captivity and testosterone on the volumes of four brain regions in the dark-eyed junco (Junco hyemalis)

This study investigates the effects of captivity and testosterone treatment on the volumes of brain regions involved in processing visual and spatial information in adult dark-eyed juncos (Junco hyemalis). We treated captive and free-living male juncos with either testosterone-filled or empty implants. Captive juncos had a smaller hippocampal formation (HF) (both in absolute volume and relative to telencephalon) than free-living birds, regardless of hormone treatment. Testosterone-treated males (both captive and free-living) had a smaller telencephalon and nucleus rotundus, but not a smaller HF or ectostriatum, than controls. We found that free-living testosterone-treated males had larger home ranges than free-living controls in agreement with earlier experiments, but we found no corresponding difference in HF volume. We discuss the implications of the effect of captivity on HF volume for past and future laboratory experiments.     

Relative abundance of males to females affects behaviour, condition and immune function in a captive population of dark-eyed juncos Junco hyemalis

Relative numbers of males and females in breeding groups may vary from expected values owing to a variety of factors. To determine how sex ratio might influence individual phenotypes in a captive population of dark-eyed juncos Junco hyemalis during the breeding season, we established three treatment groups: a male-biased (2:1), equal (1:1), and female-biased group (1:2). Within-group density (birds/m²) was constant across groups. We assessed the frequency of flight chases (a proxy for social instability), measured changes in body mass and pectoral muscle condition, assayed plasma levels of testosterone (T) and compared cell-mediated immunity of individuals. We found significantly more chases in the male-biased group than in the female-biased group. Birds in the male-biased group lost more mass and displayed poorer pectoral-muscle condition than birds in the equal group. Cell-mediated immune responses were reduced in individuals in the male-biased group in comparison to the female-biased group. Plasma T levels in both sexes did not vary with sex ratio. Collectively, these results suggest that during the breeding season, social instability is greater in male-biased populations, and instability may lead to decreased general health and vigour.     

Song Variation in a Recently Founded Population of the Dark-Eyed Junco (Junco hyemalis)

During natural colonization events, songs are expected to change as a result of both selection and drift-like processes. We studied song variation within a small isolated population of dark-eyed juncos, Junco hyemalis , which became established in an unusual environment on the University of California at San Diego (UCSD) campus in the early 1980's, and compared this variation with the native range. At UCSD, the average repertoire size is 4.2 song types/male, which is similar to that in the native range. There is a low level of song-type sharing across the population. Typically a male shares at least one song type with on average 40% of his neighbors, and song-type sharing decreases rapidly with distance. Thus song-type diversity is high, and in this respect appears similar to that in ancestral mountain populations. These results suggest a moderate to large number of males founded and/or subsequently immigrated into the UCSD population, but several selection mechanisms, if coupled with a high cultural mutation rate, may also favor high song-type diversity. When combined with results from a previous study, songs are significantly shorter at UCSD than in the mountain population we studied. It appears that founder effects have not influenced the evolution of songs in this population. More generally, if song-type diversity and other features of song result from various social and natural selection pressures, many of these pressures may have remained similar between the native and the recently established population.     

Experimentally-elevated testosterone, female parental care, and reproductive success in a songbird, the Dark-eyed Junco (Junco hyemalis)

In male dark-eyed juncos (Junco hyemalis), experimentally elevated testosterone (T) decreases male parental care and offspring survival, but results in higher overall fitness through greater mating success. To help address the ensuing question of what prevents selection from favoring higher levels of T in male juncos, we manipulated T in female juncos. A previous study demonstrated no effect of experimentally elevated T on female incubation behavior, suggesting that female parental behavior might be insensitive to T. In this study we asked whether experimentally elevated T mediates other female parental behaviors and whether variation in T-mediated parental behavior might influence reproductive success. We videotaped free-living control- and T-females during nesting to quantify brooding behavior when young were 3 days old and provisioning behavior when young were 6 days old. Nest defense was measured by quantifying responses to a mounted predator placed near the nest. Reproductive success was assessed via fecundity, nestling quality, and nest survival. T-females spent less time than control females brooding but did not differ in provisioning rate. T-females performed fewer dives at the predator mount and, unlike controls, failed to increase defense as nesting progressed. T-females also had lower daily nest survival and lower nest success (odds of producing at least one fledgling). We conclude that some aspects of female parental behavior are sensitive to experimentally elevated T while others are not and consider the implications for the evolution of T-mediated characters in both sexes.     

Effects of testosterone and photoperiodic condition on song production and vocal control region volumes in adult male dark-eyed juncos (Junco hyemalis)

In seasonally breeding male oscines, song learning and expression are controlled by brain regions (vocal control regions, VCRs) that exhibit seasonal neural plasticity in adulthood. Several VCRs contain androgen receptors, and gonadal androgens play important roles in the control of seasonal structural and functional changes of VCRs. Recent studies also found that adult VCRs are influenced by factors other than gonadal hormones, including photoperiod, but the relative importance of these factors and their mechanisms of action are poorly understood. To address this issue, we investigated the contributions of photoperiod and testicular androgens to the regulation of VCR volumes and to the control of song expression in adult dark-eyed juncos, Junco hyemalis. Exposing castrated (CX) photosensitive males to long days (LD) enhanced their high vocal center (HVc) volumes compared to those of males held on short days (SD). These volumes were not further increased by concurrent testosterone (T) treatment, revealing a marked and gonadal androgen-independent stimulatory influence of photoperiod on the size of this brain region. HVc sizes were smaller in LD-exposed photorefractory than photosensitive males irrespective of whether birds were intact or had been castrated before photoperiodic manipulations, but HVc sizes increased in response to T treatment in intact photorefractory males. Thus, LD exposure can increase HVc volumes in the absence of gonadal T, but large volume induction in photorefractory males requires elevated plasma T levels. Testosterone treatment of SD-exposed photosensitive males increased HVc, but not Area X, MAN, or RA volumes. Only T-treated males sang and this treatment given to castrates was equally effective behaviorally when administered to photosensitive, photostimulated, or photorefractory juncos. This result indicates that the stimulating influence of LD exposure on HVc volumes is insufficient to induce song in the absence of elevated plasma T levels.     

Androgen control of vocal control region volumes in a wild migratory songbird (Junco hyemalis) is region and possibly age dependent

Previous laboratory studies have shown that photoperiodic adult songbirds experience seasonal variations in singing frequency that correlate with plasma androgen levels, as well as changes in the brain regions that control singing (vocal control regions). The present study investigates naturally occurring seasonal changes in the sizes of these regions in a wild migratory species (dark-eyed junco, Junco hyemalis), with samples from adolescence to post-breeding fall migration. In adult males, the volumes of the vocal control regions area X and the higher vocal center (HVC) were large during the breeding season when birds were singing and androgen levels were high, and decreased in size after the breeding season when singing had stopped and androgen levels were low. HVC volume in adolescent males caught in the fall (no singing), when plasma androgen levels were low, was smaller than in breeding adults, thereby following the seasonal pattern of change in plasma androgen levels. In adolescent males, however, area X volume was the same as in breeding adults. Thus, area X size in adolescent male juncos may be testosterone independent. The seasonal pattern of robust nucleus of the archistriatum volume was similar to that of the HVC. The volumes of neither the magnocellular nucleus of the anterior neostriatum nor the nucleus rotundus, a control region, differed seasonally. Castration of breeding adult males caused both area X and HVC volumes to decrease compared to castrated controls with testosterone replacement, indicating that maintenance of these two region volumes is testosterone dependent in adults. © 1997 John Wiley & Sons, Inc. J Neurobiol 32: 391–402, 1997.     

Short-term fasting affects locomotor activity,corticosterone, and corticosterone binding globulin in a migratory songbird

Unpredictable events such as severe storms lead to an increase in circulating corticosterone (CORT) in breeding birds. This increase is often accompanied by elevations in foraging and irruptive behavior. We were interested in determining if acute food restriction (such as might occur during inclement weather) is a sufficient cue to elicit an increase in locomotor activity, increase CORT secretion, and/or decrease circulating levels of corticosterone binding globulin (CBG) in white-crowned sparrows (Zonotrichia leucophrys gambelii). Male Z.l. gambelii were housed individually in environmental chambers on long days (LD 20:4) to simulate breeding season daylength. Birds were fed ad libitum, and on select days, food was removed 2 h after lights on (fasted treatment), or was removed and replaced (control). We analyzed CORT and CBG levels after 1, 2, 6, 22 (lights on), and 23 h under fasted and control conditions. We also measured activity during the 23-h experiment. Activity levels were increased under fasted conditions during the daytime relative to control conditions, but activity levels did not differ between treatments during the night. Fasting as little as 1, 2, and 6 h significantly increased total CORT levels above baseline (control), although after 22 h, total CORT levels under fasted conditions matched those under control conditions. Plasma CBG decreased after the 22-h fast, and remained low after the 23-h fast. This change was sufficient to significantly elevate free CORT levels in fasted birds relative to ad libitum food conditions, despite the lack of difference in total CORT levels.     

The role of central and peripheral temperature changes in the regulation of thermal polypnea in the chicken

The temperature of free-flowing blood in the aorta, the skin temperature of the comb, and the temperature of the skin under the feathers, of nine male chickens were measured, before, during and after exposure to a hot environment (45°C). Respiratory frequencies were also recorded, and no evidence was obtained that thermal polypnea occured before the deep-body temperature increased. In some birds, the highest respiratory frequency was noted after the air temperature was lowered. It is suggested that, although an increase in skin temperature is not involved in the initiation of thermal polypnea, changes in skin temperature may modify the respiratory pattern once thermal polypnea is established.     

Appetite regulation: The central role of melatonin in Danio rerio

Melatonin is the hormonal mediator of photoperiodic information to the central nervous system in vertebrates and allows the regulation of energy homeostasis through the establishment of a proper balance between energy intake and energy expenditure. The aim of this study was to evaluate the role of melatonin in appetite central control analyzing the involvement of this hormone in the regulation of feeding behavior in the zebrafish Danio rerio. For this purpose, the effect of two different melatonin doses (100 nM and 1 μM) administered for 10 days, via water, to zebrafish adults was evaluated at both physiological and molecular level and the effect of melatonin was considered in relation to the most prominent systems involved in appetite regulation. For the first time, in fact, melatonin control of food intake by the modulation of leptin, MC4R, ghrelin, NPY and CB1 gene expression was evaluated.The results obtained indicate that melatonin significantly reduces food intake and the reduction is in agreement with the changes observed at molecular level. A significant increase in genes codifying for molecules involved in feeding inhibition, such as leptin and MC4R, and a significant reduction in the major orexigenic signals including ghrelin, NPY and CB1 are showed here.Taken together these results support the idea that melatonin falls fully into the complex network of signals that regulate food intake thus playing a key role in central appetite regulation.     

Sex and migratory strategy influence corticosterone levels in winter-grown feathers,with positive breeding effects in a migratory pelagic seabird

To overcome unpredictable stressful transitory events, animals trigger an allostatic response involving the hypothalamic–pituitary–adrenal cortex. This hormonal response, which involves the release of glucocorticoids which in turn mediate between the main physiological mechanisms that regulate the energetic demands and resource allocation trade-off with behavioural responses to environmental perturbations and may ultimately lead to variation in fitness. We have used the Cory’s shearwater Calonectris borealis, a sexually dimorphic pelagic seabird with a partial migratory strategy, as a model bird species to analyse a number of traits related to the stress response. We investigated whether the activation of a stressful response, mediated by corticosterone, during the wintering period (1) correlated with the previous breeding success, (2) was affected by the migratory behaviour of male birds and (3) had consequences in the fitness of the birds. Corticosterone levels in feathers grown overwinter were analysed in 61 adult birds during three consecutive migratory periods (2009–2012) and in 14 immature birds in the wintering period 2010–2011. Moreover, the levels of corticosterone were analysed in experimental birds which were freed from their reproductive duties and compared with control birds which raised fledglings to the end of the breeding period. The results show that the levels of corticosterone were sex dependent, differed between years and were affected by the migratory strategy performed by the birds. The activation of the stressful response over the wintering period generated residual carry-over effects that positively affected the reproductive output in the subsequent breeding stage, a phenomenon previously undescribed in a long-lived pelagic seabird. Our study provides evidence that the analysis of corticosterone from feathers is a useful tool to evaluate carry-over effects in birds far away from breeding sites, opening new possibilities for future studies in this field.     

The Effects of Experimentally Elevated Testosterone and Food Deprivation on Food Consumption and Prey Size Preferences in Male Dark-Eyed Juncos (Junco hyemalis, Emberizidae: Passeriformes)

Numerous studies have shown that the experimental elevation of circulating levels of testosterone reduces parental behaviour in male birds, particularly the provisioning of young. The mechanisms responsible for this change in behaviour are not fully understood. In this study, we examine the effects of elevated testosterone on food consumption and prey selection, both of which have potential consequences for nestling provisioning behaviour. We manipulated testosterone and performed two experiments on a captive, non-breeding population of male dark-eyed juncos ( Junco hyemalis ) on long day-lengths. In the first experiment, we subjected juncos to 3 h of food deprivation and compared food consumption and prey size selection by males with elevated testosterone (testosterone males) to that of control males. Testosterone males consumed more food than control males and showed a preference for larger prey. In a second experiment in which small prey were more abundant than large prey, food consumption and prey size preferences did not differ between testosterone and control males. We also manipulated the duration of food deprivation in the second experiment. Males of both treatments consumed more small prey under conditions of mild (1 h) or moderate (5 h) food deprivation and consumed more large prey under conditions of intermediate (3 h) food deprivation. We discuss our results and the effects that testosterone has on self-maintenance behaviour and male parental effort.