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.     

Seasonal and diel transitions in physiology and behavior in the migratory dark-eyed junco

Body mass, fat stores, activities of lipogenic and lipolytic enzymes, and plasma corticosterone were measured throughout seasonal and diel transitions from fall through spring encompassing the non-migratory stages of early and mid winter, the prealternate molt, and the spring migratory stage in captive dark-eyed juncos to determine the physiological mechanisms underlying adaptations for migration. On a seasonal basis, lipid enzymes and corticosterone varied little throughout the stages even though the birds underwent dramatic alterations in mass, fat deposition, behavior, and activation of the reproductive axis. By contrast, diel changes were found in lipogenesis, lipolysis, muscle lipoprotein lipase, and plasma corticosterone when comparing birds in the two phases of spring migration--active flight and resting, as during times of stopover. In these two phases of migration, coordination of the lipogenic and lipolytic systems appear to maximize storage of fatty acids during rest and delivery/utilization during flight. Diel patterns of corticosterone revealed fairly consistent peaks during the night time (23:00) throughout the nonmigratory period. The profile of this pattern altered during the migratory period with variation between the flight and resting phases. In sum, the results from these captive studies offer a new approach for studying the regulation of migratory physiology in free-living birds.     

Age structure and age‐related differences in molt status and fuel deposition of Dunlins during the nonbreeding season at Chongming Dongtan in east China

ABSTRACT Although most shorebirds exhibit deferred migration and deferred breeding during their first summer, Dunlins (Calidris alpina) migrate to breeding areas and breed during their first summer. First‐year and adult Dunlins should, therefore, have similar fueling and molt patterns if energetic and physiological constraints are responsible for deferred migration. From 2006 to 2008, we examined the age structure of Dunlins during the nonbreeding season at Chongming Dongtan, an estuarine wetland in the Yangtze River estuary in east China, and examined the effects of date, age, and molt status on fuel deposition during migration and during the winter. The Dunlin population at Chongming Dongtan was composed primarily of first‐year birds. Most adults and first‐year birds arrived together in late August. Regression analyses indicated that age, date, and molt status affected fuel deposition (as indicated by body mass) of Dunlins. Adults had significantly greater fuel deposits than first‐year Dunlins near the end of northward migration (May: adults 70.8 ± 6.4 g, first‐year 63.8 ± 8.0 g) and at the start of southward migration (September: adults 50.2 ± 6.1 g, first‐year 47.2 ± 4.9 g). Adults also had significantly higher fuel deposition rates than first‐year Dunlins during northward migration. Nonetheless, first‐year Dunlins migrate and breed in their first summer. Thus, other factors, such as migration distance and body size, may be more important in determining if first‐year shorebirds defer migration during their first spring and summer. During boreal spring and autumn, first‐year Dunlins in active body molt had greater body mass than those that had not initiated body molt or those in suspended molt, and premigratory fuel deposits for northward migration were greatest after prealternate molt was completed. These results suggest that body molt requires additional fuel deposits and imposes a constraint on fuel deposition for migratory flights.     

Demographic variation in timing and intensity of feather molt in migratory Fork‐tailed Flycatchers (Tyrannus s. savana)

Understanding the annual cycle of migratory birds is imperative for evaluating the evolution of life‐history strategies and developing effective conservation strategies. Yet, we still know little about the annual cycle of migratory birds that breed at south‐temperate latitudes of South America. We aged, sexed, and determined the progression and intensity of body, remige, and rectrix molt of migratory Fork‐tailed Flycatchers (Tyrannus s. savana) at breeding sites in southern South America and at wintering sites in northern South America. Molt of both body and flight feathers occurred primarily during the winter. In early winter, a similar proportion of young and adult flycatchers molted remiges and rectrices, but remige molt intensity (number of remiges molting) was greater and primary molt progression (mean primary feather molting) more advanced in adults. In late winter, remige molt intensity and primary molt progression did not differ between age groups. We found no difference between males and females either in the proportion of individuals molting in winter or in the intensity or progress of remige molt. Our results suggest that the nominate subspecies of Fork‐tailed Flycatcher undergoes one complete, annual molt on the wintering grounds, and represents the first comprehensive evaluation of molt timing of a migratory New World flycatcher that overwinters in the tropics. Given that breeding, molt, and migration represent three key events in the annual cycle of migratory birds, knowledge of the timing of these events is the first step toward understanding the possible tradeoffs migratory birds face throughout the year.     

Radio transmitters do not affect the body condition of Savannah Sparrows during the fall premigratory period

ABSTRACT Radio telemetry can be a valuable tool for studying the behavior, physiology, and demography of birds. We tested the assumption that radio transmitters have no adverse effects on body condition in an island population of Savannah Sparrows (Passerculus sandwichensis). To assess possible changes in condition, 20 radiotagged and 25 nontagged Savannah Sparrows were captured and recaptured throughout the postfledging period. We used four measures of condition: mass, an index of fat free dry mass (measured via heavy water dilution), pectoral muscle depth (measured via ultrasound imaging), and an index of fat mass (measured via heavy water dilution). Using both a generalized linear modeling framework and paired design, we found no significant differences in the body condition of radiotagged and nontagged adults and juveniles. Thus, our results provide evidence that radiotransmitters have no effect on the condition of Savannah Sparrows during the premigratory period.     

Consequences of elevating plasma testosterone in females of a socially monogamous songbird: evidence of constraints on male evolution?

To explore whether selection for testosterone-mediated traits in males might be constrained by costs of higher testosterone to females, we examined the effects of experimental elevation of plasma testosterone on physiological, reproductive, and behavioral parameters in a female songbird, the dark-eyed junco (Junco hyemalis). We used subcutaneous implants to elevate testosterone (T) in captive and free-living female juncos. In captive birds, we measured the effects of high T on body mass, feather molt, and brood patch formation. In the field, we monitored its effects on the timing of egg laying, clutch size, egg size, egg steroid levels, incubation, and nest-defense behavior. Females implanted with testosterone (T-females) had significantly higher circulating levels of testosterone than did control females (C-females). Captive T-females had lower body mass, were less likely to develop brood patches, and delayed feather molt relative to C-females. Among free-living females, the interval between nest completion and appearance of the first egg was longer for T-females than for C-females and egg yolk concentrations of testosterone were higher, but there were no significant differences in estradiol levels, clutch size, or egg size. Incubation and nest defense behavior were also similar between T- and C-females. Our results suggest that selection on males for higher testosterone might initially lead to a correlated response in females producing changes in body mass and feather molt, both of which could be detrimental. Other possible female responses would be delayed onset of reproduction, which might reduce reproductive success, and higher yolk testosterone, which might have either positive or negative effects on offspring development. We found no reason to expect reduced parental behavior by females as a negative fitness consequence of selection for higher testosterone in males.     

Differential Regulation of Adipokines May Influence Migratory Behavior in the White-Throated Sparrow (Zonotrichia albicollis)

White-throated sparrows increase fat deposits during pre-migratory periods and rely on these fat stores to fuel migration. Adipose tissue produces hormones and signaling factors in a rhythmic fashion and may be controlled by a clock in adipose tissue or driven by a master clock in the brain. The master clock may convey photoperiodic information from the environment to adipose tissue to facilitate pre-migratory fattening, and adipose tissue may, in turn, release adipokines to indicate the extent of fat energy stores. Here, we present evidence that a change in signal from the adipokines adiponectin and visfatin may act to indicate body condition, thereby influencing an individual''s decision to commence migratory flight, or to delay until adequate fat stores are acquired. We quantified plasma adiponectin and visfatin levels across the day in captive birds held under constant photoperiod. The circadian profiles of plasma adiponectin in non-migrating birds were approximately inverse the profiles from migrating birds. Adiponectin levels were positively correlated to body fat, and body fat was inversely related to the appearance of nocturnal migratory restlessness. Visfatin levels were constant across the day and did not correlate with fat deposits; however, a reduction in plasma visfatin concentration occurred during the migratory period. The data suggest that a significant change in the biological control of adipokine expression exists between the two migratory conditions and we propose a role for adiponectin, visfatin and adipose clocks in the regulation of migratory behaviors.     

Extraocular light perception in photoperiodic responses of the White-crowned sparrow (Zonotrichia leucophrys) and of the Golden-crowned sparrow (Z. atricapilla)

Summary Testicular maturation, migratory restlessness (Zugunruhe) and fattening are induced in many species of birds by long day-lengths in Spring. We have tested the hypothesis that extraocular photoreceptors located in the brain are involved in mediating these photoperiodic responses in White-crowned sparrows (Zonotrichia leucophrys) and Golden-crowned sparrows (Z. atricapilla). Our approach consisted in reducing the amount of light penetrating to the brain with either black India ink injected under the skin of the head (Golden-crowned sparrows) or by covering the entire head (except eyes and beak) with a black collodion hood (White-crowned sparrows). Birds treated in these ways showed significantly less testicular growth, Zugunruhe and premigratory fattening when placed under a 16L-8D photoperiod than control birds which did not have their brains shielded from light. However, even when the bird brains were shielded from light and although light intensity was close to threshold level, some testicular growth, Zugunruhe and fattening did occur. We conclude that extraocular photoreceptors are involved in the control of the three photoperiodic responses studied, but that the eyes are possibly of significance as well.Supported by NASA Headquarters grant NGR-05-020-391 to Dr. C. S. Pittendrigh     

Body condition and feather molt of a migratory shorebird during the non‐breeding season

Migratory shorebirds have some of the highest fat loads among birds, especially species which migrate long distances. The upland sandpiper Bartramia longicauda makes long‐distance migrations twice a year, but variation in body condition or timing of feather molt during the non‐breeding season has not been studied. Molt is an important part of the annual cycle of migratory birds because feather condition determines flight performance during migration, and long‐distance movements are energetically costly. However, variation in body condition during molt has been poorly studied. The objective of our field study was to examine the timing and patterns of feather molt of a long distance migratory shorebird during the non‐breeding season and test for relationships with body size, fat depots, mass, and sex. Field work was conducted at four ranches in the Northern Campos of Uruguay (Paysandú and Salto Departments). We captured and marked 62 sandpipers in a 2‐month period (Nov–Jan) during four non‐breeding seasons (2008–2012). Sex was determined by genetic analyses of blood samples taken at capture. Molt was measured in captured birds using rank scores based on published standards. Body mass and tarsus length measurements showed female‐biased sexual size dimorphism with males smaller than females. Size‐corrected body mass (body condition) showed a U‐shaped relationship with the day of the season, indicating that birds arrived at non‐breeding grounds in relatively good condition. Arriving in good body condition at non‐breeding grounds is probably important because of the energetic demands due to physiological adjustments after migration and the costs of feather molt.     

Changes in the blood and plasma volume of Harris' sparrows during postnuptial molt

In captive Harris' sparrows (Zonotrichia querula), plasma and blood volume increased significantly during stages 2 and 3 of postnuptial molt, when extensive feather replacement was taking place in the wings, tail and body tracts. Erythrocyte number did not increase significantly until molt was essentially complete. Consequently the 6% mean maximum reduction in hematocrit that was observed during molt appeared to be largely the result of plasma volume expansion. This plasma volume expansion was not due to changes in the osmolality or concentration of proteins within the blood plasma since both decreased significantly during intense molt.     

Dietary thyroxine induces molt in chickens (Gallus gallus domesticus)

Thyroxine increases during a molt in wild and captive birds, and thyroidectomy prevents induction of molt. This trial examined the effect of dietary thyroxine on molt induction molt in chickens (laying hens, 59 weeks of age). In a completely randomized design (n=15 hens/replication; 6 replications/treatment), hens were randomly assigned to either a traditional molting program consisting of feed withdrawal (FWD), or to diets containing 40 mg thyroxine/kg diet (HT), 20 mg thyroxine/kg diet (LT), or 40 mg thyroxine from thyroactive iodinated casein/kg diet (TIC). The molting treatment lasted 7-13 d, until egg production reached 0%. After molt induction, birds had ad libitum access to the same diet, until egg production was re-initiated and maximized ( approximately 56 d). All treatments induced molt, based upon cessation of egg laying and regression of ovary and oviduct. Birds on FWD treatment lost more body weight during the molting period, but gained more after molt compared to thyroxine treatments (P<0.01 for each), although all body weights were similar when egg production was maximized. Data demonstrate that oral thyroxine, in purified or non-purified form, induces a molt and may enhance animal well-being by reducing the need for FWD.     

Catabolic enzyme activities in the pectoralis muscle of premigratory and migratory juvenile Reed Warblers Acrocephalus scirpaceus (Herm.)

Summary Differences in catabolic capacity in the pectoralis major muscle of premigratory and migratory Reed Warblers were examined. The oxidative capacity was greater in the migratory birds, probably reflecting a training effect caused by the increased locomotion activity just prior to migration.Fatty acid oxidation was already high in the premigratory birds, possibly reflecting a saving of carbohydrate for anabolic purposes (feather growth) during the moult. Glycolytic capacity was slightly increased in the migratory birds, suggesting an increased carbohydrate oxidation during migration in spite of the great contribution of fatty acid oxidation.     

Complex postbreeding molt strategies in a songbird migrating along the East Asian Flyway,the Pallas’s Grasshopper Warbler Locustella certhiola

Molt strategies have received relatively little attention in current ornithology, and knowledge concerning the evolution, variability and extent of molt is sparse in many bird species. This is especially true for East Asian Locustella species where assumptions on molt patterns are based on incomplete information. We provide evidence indicating a complex postbreeding molt strategy and variable molt extent among the Pallas's Grasshopper Warbler Locustella certhiola, based on data from six ringing sites situated along its flyway from the breeding grounds to the wintering areas. Detailed study revealed for the first time that in most individuals wing feather molt proceeds from the center both toward the body and the wing‐tip, a molt pattern known as divergent molt (which is rare among Palearctic passerines). In the Russian Far East, where both breeding birds and passage migrants occur, a third of the adult birds were molting in late summer. In Central Siberia, at the northwestern limit of its distribution, adult individuals commenced their primary molt partly divergently and partly with unknown sequence. During migration in Mongolia, only descendantly (i.e., from the body toward the wing‐tip) molting birds were observed, while further south in Korea, Hong Kong, and Thailand the proportion of potential eccentric and divergent feather renewal was not identifiable since the renewed feathers were already fully grown as expected. We found an increase in the mean number of molted primaries during the progress of the autumn migration. Moderate body mass levels and low‐fat and muscle scores were observed in molting adult birds, without any remarkable increase in the later season. According to optimality models, we suggest that an extremely short season of high food abundance in tall grass habitats and a largely overland route allow autumn migration with low fuel loads combined with molt migration in at least a part of the population. This study highlights the importance of further studying molt strategy as well as stopover behavior decisions and the trade‐offs among migratory birds that are now facing a panoply of anthropogenic threats along their flyways.     

Regulation of the hypothalamic-pituitary-adrenal axis in free-living pigeons

We studied feral free-living pigeons (Columba livia) to determine whether either unstressed or stress-induced corticosterone release was altered during a prebasic molt. The pigeons were at various stages of molt throughout the study, but corticosterone responses in molting and nonmolting birds did not differ. This was further reflected in equivalent adrenal responses to exogenous adrenocorticotropic hormone (ACTH), suggesting equivalent steroidogenic capacity of adrenal tissues during both physiological states. There was a slight change, however, in pituitary regulation during molt. Whereas exogenous arginine vasotocin (AVT) elevated corticosterone levels in nonmolting birds, during molt an equivalent dose of AVT was ineffective, suggesting that the pituitary is less sensitive to an AVT signal during molt. AVT also appears to be more effective than corticotropin-releasing factor at eliciting ACTH release in pigeons. Overall, these data indicate that pigeons regulate their corticosterone release differently during molt than other avian species studied to date.     

Hypothalamic and hormonal control of photoperiodically induced vernal functions in the white-crowned sparrow,Zonotrichia leucophrys gambelii

Summary The effects of implantation of testosterone propionate (TP) in various sites in the hypothalamus on the photoperiodically induced vernal premigratory functions in the White-crowned Sparrows were investigated in order to assess the role of the hypothalamo-hypophysial-testicular axis in the induction of these responses.Implantation of glass capillary tubes containing TP in the basal infundibular nucleus (IN), in the median eminence, or in the pars distalis inhibited the photoperiodically induced increase in plasma levels of luteinizing hormone (LH), as measured by radioimmunoassay, and testicular growth. The effective implants significantly lowered the levels of LH in birds held on nonstimulatory short days. These TP implants apparently inhibited release from the pars distalis of both LH and follicle-stimulating hormone (FSH). It is concluded that the site of sensitivity in the negative feedback by testosterone is either the basal IN or the pars distalis, or both. The implants of TP that inhibited the increase in plasma LH and testicular growth completely did not prevent the birds from fattening.These investigations were supported, in part, by research grants from the National Institutes of Health (HD-6527) and National Science Foundation (BMS 79-13933) to Professor Donald S. Farner. This paper is based on a dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy from the University of WashingtonThe author is grateful to Professor Donald S. Farner for his guidance throughout the course of these investigations. The assistance of Mr. Philip W. Mattocks, Jr. in performing radioimmunoassays is sincerely appreciated     

Response of the haematocrit to body condition changes in Northern Bald Ibis Geronticus eremita

We study the usefulness of the haematocrit as a predictor of body condition in birds, using a captive population of the endangered species Northern Bald Ibis (Geronticus eremita). This population is 14% of the worldwide captive population, which is far greater than the known free-living population. The haematocrit, body mass and body condition index responded in the same sense to two different nutritional periods, and there was a statistically significant relationship between changes in condition index and haematocrit of individual birds between the two periods. We discuss the relationship of these parameters with subcutaneous fat and muscle mass, and analyse the individuals' nutritional status in each of the periods studied. The conclusion was that the haematocrit is sensitive to variations in body condition since it responds to mass-loss processes corresponding to phases of mobilization of fat reserves, a situation prior to the mobilization of muscle proteins when there is a manifest deterioration of the individual's aspect.     

Expression of annual cycles in preen wax composition in red knots: constraints on the changing phenotype

Birds living in seasonal environments change physiology and behavior in correspondence to temporally changing environmental supplies, demands and opportunities. We recently reported that the chemical composition of uropygial gland secretions of sandpipers (Scolopacidae, order Charadriformes) changes during the breeding season from mixtures of monoesters to diesters, which fulfill specific functions related to incubation. A proper temporal match between the expression of diester preen waxes and incubation requires a flexible organization of the trait. Here we analyze the possible degrees of flexibility with reference to the functionality of better-understood molt and body mass cycles of free-living and captive red knots (Calidris canutus). The relative flexibility of seasonal cycles in preen wax composition was examined by two experimental perturbations: (1) giving birds restricted access to food and (2) monitoring them long-term under a constant photoperiodic regime. We found that wax type cannot change instantaneously, but that changing the type of wax is under similar organizational time constraints as the replacement of feathers. Just as molt and mass cycles, the seasonal rhythm of diester secretion appeared to be under endogenous control: most birds placed in a constant photoperiod still maintained seasonally changing preen waxes. Diester preen wax secretion was synchronized with the peak in body mass in spring, but became less well expressed under constant photoperiodic conditions and when food availability was limited.     

Flexibility and constraints in the molt schedule of long‐distance migratory shorebirds: causes and consequences

Molt is a major component of the annual cycle of birds, the timing and extent of which can affect body condition, survival, and future reproductive success through carry‐over effects. The way in which molt is fitted into the annual cycle seems to be a somewhat neglected area which is both of interest and of importance. Study of the causes of annual variation in the timing of molt and its potential consequence in long‐distance migratory birds was examined using the Curlew Sandpiper, Calidris ferruginea, as a model species. Using the maximum likelihood molt models of Underhill and Zucchini (1988, Ibis 130:358–372), the relationship between annual variability in the start dates of molt at the population level with conditions on the breeding area was explored. Adult males typically started early in years when temperature in June on the Arctic breeding grounds were high compared to cold years while adult females molted later in years of high breeding success and/or warm July temperature and vice versa. When molt started later, the duration was often shorter, indicating that late completion of molt might have fitness consequences, probably jeopardizing survival. Evidence of this was seen in the low body condition of birds in years when molt was completed late. The results indicate that these migratory shorebirds follow a fine‐tuned annual life cycle, and disturbances at a certain stage can alter next biological events through carry‐over effects.     

Daily and photoperiod variations of basal and stress-induced corticosterone concentrations in house sparrows (Passer domesticus)

Corticosterone concentrations were measured in captive house sparrows (Passer domesticus) and found to vary both daily and with different photoperiods. Basal corticosterone was highest during the dark hours of the daily cycle and lowest during the light hours. This trend remained constant when the birds were held on short-day and long-day light cycles, and while the birds were undergoing a prebasic molt. At all times, corticosterone concentrations significantly increased in response to the stress of handling and restraint. Stress-induced corticosterone concentrations, however, only reflected a daily rhythm when the birds were held on short-days. Furthermore, even though mean basal corticosterone concentrations were equivalent over the short-day, long-day, and molt, total corticosterone output in response to stress was lower in molting birds, especially at night. Therefore, these data indicate that captive house sparrows modulate corticosterone in daily cycles that change in response to photoperiod.     

Hypothalamic and hormonal control of the photoperiodically induced vernal functions in the White-crowned Sparrow,Zonotrichia leucophrys gambelii

To assess the role of the hypothalamo-hypophysial complex in the control of photoperiodically induced vernal premigratory responses in the White-crowned Sparrow, the effects of hypothalamic lesions and systemic administration of several hormones on these responses were investigated. Lesions that destroyed the posterior median eminence (PME) or the entire median eminence (ME) inhibited photoperiodically induced testicular growth, premigratory fattening and Zugunruhe. Lesions in the basal infundibular nucleus (IN) that resulted in complete inhibition of testicular growth abolished Zugunruhe, but allowed varying degrees of fattening. The systemic administration of prolactin, testosterone propionate (TP) or the combination thereof in the PME-lesioned birds induced fattening similar to that observed in photostimulated controls but did not induce Zugunruhe. It is concluded that testosterone and prolactin are the most important hormones involved in the control of vernal premigratory fattening. The role of these hormones in the induction of vernal Zugunruhe is not positively proven.