Water and energy economy of an omnivorous bird: population differences in the Rufous-collared Sparrow (Zonotrichia capensis)

We investigated the intraspecific variation in basal metabolic rate (BMR) and total evaporative water loss (TEWL) in the omnivorous passerine Zonotrichia capensis from two populations inhabiting regions with different precipitation regimes and aridity indices. Values of TEWL in birds from the semi-arid region were significantly lower than those found in sparrows from the mesic region. TEWL in birds from the semi-arid site was 74% of the expectation based on body mass for passerines from mesic areas and similar to the allometric expectation for passerines from arid environments. In sparrows from the mesic area, TEWL was higher than predicted by their body mass for passerines from arid environments (133%), but very close (97%) to the expectation for passerines from mesic areas. BMR values were 25% lower in sparrows from the semi-arid region. The lower TEWL and BMR of birds from the semi-arid region may be a physiological adjustment that allows them to cope with fewer resources and/or water. We propose that the lower endogenous heat production in birds from the semi-arid environment may decrease their water requirements.     

High rates of extra‐pair paternity in two equatorial populations of rufous‐collared sparrow,Zonotrichia capensis

The latitudinal increase in extra‐pair paternity (EPP) rates in birds suggests broad selective benefits to low EPP rates in the tropics. However, we have few EPP data from tropical birds, particularly from species with close relatives at high latitudes. Here, we report EPP rates in two resident equatorial populations of rufous‐collared sparrow Zonotrichia capensis, a genus well‐represented at high latitudes. We found 64% and 60% of broods contained extra‐pair offspring, and 42% and 52% of all young were extra‐pair. EPP rates were similar in these populations, despite clear differences in elevation, temperature, rainfall, and breeding season length. These findings provide evidence that EPP rates in tropical birds can be as high as those observed in temperate birds, and suggest that the selective pressures acting on EPP rates vary markedly across tropical birds.     

Migratory sleeplessness in the white-crowned sparrow (Zonotrichia leucophrys gambelii)

Twice a year, normally diurnal songbirds engage in long-distance nocturnal migrations between their wintering and breeding grounds. If and how songbirds sleep during these periods of increased activity has remained a mystery. We used a combination of electrophysiological recording and neurobehavioral testing to characterize seasonal changes in sleep and cognition in captive white-crowned sparrows (Zonotrichia leucophrys gambelii) across nonmigratory and migratory seasons. Compared to sparrows in a nonmigratory state, migratory sparrows spent approximately two-thirds less time sleeping. Despite reducing sleep during migration, accuracy and responding on a repeated-acquisition task remained at a high level in sparrows in a migratory state. This resistance to sleep loss during the prolonged migratory season is in direct contrast to the decline in accuracy and responding observed following as little as one night of experimenter-induced sleep restriction in the same birds during the nonmigratory season. Our results suggest that despite being adversely affected by sleep loss during the nonmigratory season, songbirds exhibit an unprecedented capacity to reduce sleep during migration for long periods of time without associated deficits in cognitive function. Understanding the mechanisms that mediate migratory sleeplessness may provide insights into the etiology of changes in sleep and behavior in seasonal mood disorders, as well as into the functions of sleep itself.     

Endocrine correlates of alternative phenotypes in the white-throated sparrow (Zonotrichia albicollis)

Many vertebrate species exhibit alternative phenotypes (or morphs), in which one sex displays phenotypic variation equal to or greater than the variation between the sexes. Males in such species typically display differences in reproductive strategies and morphology. Steroid hormones such as testosterone are known modulators of reproductive behavior and morphology and therefore are obvious candidates for the mediation of phenotypic differences between morphs. We conducted a year-round study in the white-throated sparrow (Zonotrichia albicollis) that exhibits alternative phenotypes in plumage coloration and behavior in both sexes: during the breeding season, white-striped males and females are more aggressive and have higher song rates than tan-striped individuals. At the beginning of the breeding season, free-living white-striped males had higher plasma testosterone concentrations than tan-striped males. However, this finding might have been due to different social experiences because captive male morphs sampled at similar times of year did not differ in testosterone concentrations. Captive white-striped males had larger testis and cloacal protuberance sizes than tan-striped males, which might be related to the divergent mating strategies of the morphs. Male morphs showed similar increases in luteinizing hormone following injections of gonadotropin-releasing hormone, but white-striped males showed larger increases in testosterone, indicating differences between morphs in gonadal testosterone production. Females had low concentrations of testosterone, and morphs did not differ. Plasma dehydroepiandrosterone (DHEA) concentrations were elevated in both sexes and morphs during the breeding and non-breeding seasons. These data do not support the hypothesis that testosterone activates behavioral differences between alternative phenotypes in the white-throated sparrow. Alternative testable hypotheses include hormonal effects during early development and direct genetic effects.     

Chromosome polymorphism due to pericentric inversion in Zonotrichia capensis (Emberizidae-Passeriformes-Aves)

Chromosomal polymorphism involving chromosome pairs 3 and 5 problably due to a pericentric inversion is reported for 60 specimens of Zonotrichia capensis (Passeriformes-Aves). The frequency of seven different karyotypes, due to the fact that chromosome 3 is present in the 3^st and 3^sm conditions and chromosome 5 in the 5^st and 5^m conditions, and the significance of this polymorphism are discussed for the genus Zonotrichia and for other genera from the same family (Emberizidae).     

Plasma corticosterone increases during migratory restlessness in the captive white-crowned sparrow Zonotrichia leucophrys gambelli

Plasma corticosterone increases during the period of spring migration in a variety of bird species. Long-distance migrants show elevations in corticosterone specifically in association with the stage of flight, suggesting that corticosterone may support flight-related processes, for example, locomotor activity and/or energy mobilization. The pattern of corticosterone secretion as it relates to migratory flight has hitherto not been clearly described in migrants that frequently interrupt flight to refuel, for example, the Gambel's white-crowned sparrow (Zonotrichia leucophrys gambelii). The Gambel's white-crowned sparrow fuels by day and expresses peak migratory activity during the first few hours of night. To determine if plasma corticosterone increases in association with the stage of migratory flight also in this short-bout migrant, we induced captive white-crowned sparrows to enter into the migratory condition by placing photosensitive birds on long days (16L:8D) and then evaluated birds for plasma corticosterone and locomotor activity during four time points of the day. Patterns found in long-day birds were compared to those observed in short-day controls (8L:16D). Differences in energy metabolism as determined from plasma metabolites were also evaluated. We found that locomotor activity and corticosterone were significantly elevated at the onset of the dark period, but only in long-day birds. Plasma beta-hydroxybutyrate (a ketone body) was also elevated. Thus, findings suggest that plasma corticosterone and ketogenesis increase in association with migratory restlessness in a short-bout migrant. In fact, corticosterone may play a regulatory role, because it shows a trend to increase already before night-time activity.     

Electron microscopic and experimental studies of the pineal organ in the white-crowned sparrow,Zonotrichia leucophrys gambelii

Summary The structure of the pineal organ of Zonotrichia leucophrys gambelii, as revealed by light- and electron-microscopy, resembles that of Passer domesticus (Oksche and Kirschstein, 1969; Ueck, 1970). The typical cellular element is the pinealocyte with certain basic structural features of the pineal photoreceptors of lower vertebrates (see Oksche, 1971). However, instead of the characteristic, cone-like outer segments, there are, as in other species of birds, only bulbous cilia with ectopic whorls of lamellae. This structure of the outer segment is, in a sense, contrary to the demonstration of synaptoid contacts, numerous unmyelinated, and occasional myelinated nerve fibers by electron microscopy. In Nissl preparations it was possible to demonstrate typical nerve cells. The pinealocytes of Z. l. gambelii are secretory; their Golgi complex forms granulated vesicles (800–1,400 Å in diameter) that belong to the group of granular inclusions characteristic of monoamines. Autonomie nerve fibers course within the connective tissue capsule of the pineal organ. In many pinealocytes of Z. l. gambelii, the granular endoplasmic reticulum contains extensively expanded cisternae that are filled with a flocculent material and closely associated with bundles of filaments. In a number of cases such loop-like structures are selectively stainable with aldehyde fuchsin. It was not possible to demonstrate specific secretory activity in the supporting cells. Extirpation of the pineal organ in Z. l. gambelii had no definitely detectable influence on the photoperiodic control of testicular growth.Aves, Passeriformes, Fringillidae.Supported by grants from the Deutsche Forschungsgemeinschaft to Professor Oksche and by the National Science Foundation (GB 11905) to Professor Farner. A part of this investigation was effected while Professor Kobayashi held a Visiting Professorship at the University of Giessen.     

Physiological and biochemical responses to dietary protein in the omnivore passerine Zonotrichia capensis (Emberizidae)

We studied the physiological, biochemical and morphological responses of the omnivore sparrow Zonotrichia capensis, a small opportunistic passerine from Central Chile acclimated to high- and low-protein diets. After 4 weeks of acclimation to 30% (high-protein group) or 7% (low-protein group) dietary casein, we collected urine and plasma for nitrogen waste production and osmometry analysis, and measured gross renal morphology. Plasma osmolality and hematocrit were not significantly affected by dietary treatment, but urine osmolality was higher in the high-protein group than in the low-protein group. Kidney and heart masses were higher in animals acclimated to the high-protein diet. Mean total nitrogen waste was significantly higher in the high-protein group, but the proportions of nitrogen excreted as uric acid, urea and ammonia were unaffected by diet. Our data suggest that the response of Z. capensis to an increase in dietary protein content is through greater amounts of total nitrogen excretion and hypertrophy of kidney structures, without any modification of the proportion of excretory compounds.     

Haplotype-based genomic sequencing of a chromosomal polymorphism in the white-throated sparrow (Zonotrichia albicollis)

Inversion polymorphisms have been linked to a variety of fundamental biological and evolutionary processes. Yet few studies have used large-scale genomic sequencing to directly compare the haplotypes associated with the standard and inverted chromosome arrangements. Here we describe the targeted genomic sequencing and comparison of haplotypes representing alternative arrangements of a common inversion polymorphism linked to a suite of phenotypes in the white-throated sparrow (Zonotrichia albicollis). More than 7.4 Mb of genomic sequence was generated and assembled from both the standard (ZAL2) and inverted (ZAL2(m)) arrangements. Sequencing of a pair of inversion breakpoints led to the identification of a ZAL2-specific segmental duplication, as well as evidence of breakpoint reusage. Comparison of the haplotype-based sequence assemblies revealed low genetic differentiation outside versus inside the inversion indicative of historical patterns of gene flow and suppressed recombination between ZAL2 and ZAL2(m). Finally, despite ZAL2(m) being maintained in a near constant state of heterozygosity, no signatures of genetic degeneration were detected on this chromosome. Overall, these results provide important insights into the genomic attributes of an inversion polymorphism linked to mate choice and variation in social behavior.     

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     

Investigations of the butylcholinesterase-containing cells of the adenohypophysis of the white-crowned sparrow,Zonotrichia leucophrys gambelii

Summary In the adenohypophysis of Zonotrichia leucophrys gambelii two types of cells with butylcholinesterase(BuChE) activity can be demonstrated histochemically. Type I occurs in the cephalic lobe of the pars distalis and in the pars tuberalis; it consists of oval and round cells. It is a distinctive cell type that is identical with the amphophilic cells described by Matsuo, Vitums, King and Farner (1969). Whereas castration or inhibition of thyroid gland activity causes only minor changes in these cells, blocking of adrenal cortex activity, or adrenalectomy, causes conspicuous hyperplasia and hypertrophy suggesting that these cells are involved in the production or release of ACTH. The second type, which occurs in both cephalic and caudal lobes, consists mostly of irregularly formed cells. Various observations indicate that it is a composite group, consisting, at least in part, of degenerating cells.These investigations were supported by grants from the National Institutes of Health (S ROI NB 06812) and the National Science Foundation (GB 5969) to Professor Farner.     

Interaction of testosterone, corticosterone and corticosterone binding globulin in the white-throated sparrow (Zonotrichia albicollis)

Plasma binding globulins bind steroid hormones and are thought to regulate hormone access to tissues. Mammals have both sex steroid binding globulin (SSBG) and corticosteroid binding globulin (CBG). Birds, however, have no detectable SSBG, leading to the early conclusion that birds have no plasma regulation of sex steroids. CBG, however, can bind androgens with relatively high affinity. In birds, therefore, the control of androgenic effects may be tightly regulated by glucocorticoid physiology because glucocorticoids compete with androgens for CBG binding sites. We report levels of total testosterone (T), total corticosterone, CBG, and estimated free T in the males, the more aggressive morph had higher levels of total T; female morphs did not differ. Approximately 96% of T was bound to CBG, but a lack of morph or sex-specific differences in corticosterone titers or CBG capacity caused patterns of free T to mirror those of total T. While CBG has the potential to greatly influence T availability to tissues, in this species interactions between T, CBG and corticosterone do not appear to alter general patterns of T availability to tissues.     

Light-microscope studies of the cytology of the adenohypophysis of the white-crowned sparrow,Zonotrichia leucophrys gambelii

Summary Adenohypophyses from more than two hundred white-crowned sparrows of both sexes and different ages, and from different periods of their annual reproductive cycle, have been used for this investigation. In addition to examination of these normal birds, we have also studied the adenohypophyses of 23 castrates and 24 controls held in different photoperiodic conditions.Cytologically the pars distalis of the adenohypophysis of the white-crowned sparrow is typically avian with distinct cephalic and caudal lobes, each with characteristic cell-types.Four basic cell-types, the acidophils, basophils, amphophils, and chromophobes, have been identified in the pars distalis by means of Matsuo tetrachrome and Matsuo modified PAS-methyl blue staining methods.Three types of acidophils, orange, red, and small, are confined to the caudal lobe of the pars distalis. Their possible functions are discussed.Light basophils (PAS-light red cells) and deep basophils (PAS-deep red cells) are equally distributed in both lobes. It is suggested that basophils may be involved in gonadotropic function since their appearance correlates well with the annual gonadal cycle and photoperiodic stimulation of gonadal growth and with the results of castration.The amphophils or PAS-purple cells (aldehyde-fuchsin positive) are found only in the cephalic lobe. Their probable function is discussed.Two types of chromophobes, specific and ordinary chromophobes, have been observed. The specific chromophobes are found only in the cephalic lobe and are similar to the Kernhaufen described by Romeis (1940). The ordinary chromophobes are similar to those of the pars distalis of other avian species and of mammals.The castration cells are found in both lobes of the photosensitive castrates under natural photoperiodic conditions as well as in those subjected artificially to photostimulation (20-hour daily photoperiods). Similar cells have also been observed in the pars tuberalis of the castrated photostimulated birds.The relations of the rostral and caudal groups of the portal vessels to the cell-types found in the cephalic and caudal lobes are discussed.Dedicated to Professor Dr. Y. Kato, Department of Anatomy of the Domestic Animals, Kyushu University, Fukuoka, Japan, and to President Dr. H. Mimura, Shinshu University, Matsumoto, Japan, in honor of their retirement.The investigation reported herein was supported by a research grant (5RO 1-HEO7240 NEUA) from the National Institutes of Health to Professor Vitums, by funds for biological and medical research made available by State of Washington Initiative Measure No. 171 to Professor Vitums; by a Research Career Development Award (5K3 AM-18, 370) from the National Institutes of Arthritis and Metabolic Diseases to Professor King; and by a research grant (5RO 1 NB 06 187) from the National Institutes of Health to Professor Farner. The senior author is greatful to Professor Dr. Hideo Murai and Doctor Yasukuni Watanabe, Department of Animal Science, Shinshu University,Ina,Japan, for their cooperation and support in this investigation. We wish to thank Mrs. Sumiko Sumida for technical assistance, and Miss Kathleen Reinhardt for the preparation of the drawings.     

Circadian and masking control of migratory restlessness in Gambel's white-crowned sparrow (Zonotrichia leucophrys gambelii)

Avian migration is a seasonal activity that requires intricate timing on both an annual and daily basis. With increasing evidence for endogenous regulation of daily activities in migrant species, we tested whether a circadian oscillator may be involved with the expressions of daily locomotor activities and specific behaviors of the long-distance migrant, Gambel's white-crowned sparrow (Zonotrichia leucophrys gambelii). Our previous studies have identified both daytime and nighttime behavioral patterns under a photoperiod of 18L:6D. In 2 separate trials, birds in the vernal migratory life-history stage were exposed to constant dim light, (DD)dim, and constant bright light, LL, while locomotor activity and behavioral observations were collected. Under (DD)dim, the daytime behaviors that included active and quiescent components observed under 18L:6D were lost as migratory restlessness, the intense nighttime activity, persisted nonstop for 36.4 h. Furthermore, the specific behaviors of migratory restlessness that are normally confined to the dark phase of 18L:6D, beak-up and beak-up flight, were expressed also during the subjective day of (DD)dim. Birds exposed to LL retained similar patterns of activity to the 18L:6D condition for 3 days, after which they became arrhythmic. Behavioral observations of intense locomotor activity observed during the subjective night of LL revealed no beak-up and beak-up flight. Thus, the complete expression of migratory restlessness that includes beak-up and beak-up flight may not be regulated by a circadian oscillator but instigated by very low light intensity. Locomotor activity and associated daytime behaviors appear to be influenced by a circadian oscillator, given their persistent patterns under LL. Therefore, we suggest that the separate components of migratory behavior are regulated differentially by a circadian oscillator and ambient lighting conditions.