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Season- and age-related variation in neural cGnRH-I and cGnRH-II immunoreactivity in house sparrows (Passer domesticus).

Vertebrates exhibit several forms of gonadotropin-releasing hormone (GnRH). In birds, chicken GnRH-I (cGnRH-I) is the primary hypophysiotropic form that regulates the hypothalamo-pituitary-gonad (HPG) axis. In seasonally breeding songbirds, cGnRH-I immunoreactivity is primarily in the preoptic septo-infidibular regions and varies greatly between the breeding and non-breeding season. Chicken GnRH-II (cGnRH-II) is more widely distributed throughout the avian brain, but cGnRH-II immunoreactivity has been found in the median eminence of quail. Thus, cGnRH-II may function in addition to cGnRH-I in the regulation of the HPG axis. If so, we predicted that cGnRH-II immunoreactivity should exhibit seasonal and age related variation similar to that observed for cGnRH-I. In this study, we compared hypothalamic immunoreactivity of cGnRH-I and -II (ir-cGnRH-I and ir-cGnRH-II) between breeding and non-breeding adult and juvenile house sparrows (Passer domesticus). We used immunocytochemistry with highly specific antibodies developed for quail to examine brains of free-living house sparrows collected in the breeding and non-breeding season. Results showed a greater number of ir-cGnRH-I cell bodies and beaded fibres in breeding adult birds compared to non-breeding adult and juvenile birds. Moreover, there were significantly more ir-cGnRH-II fibres in the preoptic area of breeding adult birds compared to non-breeding adult and juvenile birds. These data suggest that cGnRH-II undergoes seasonal- and age-related changes similar to cGnRH-I and implicate a role in regulating reproduction.

Aging↗

Development of stress reactivity in white-crowned sparrow nestlings: total corticosterone response increases with age, while free corticosterone response remains low.

Activation of the adrenocortical response to stress during development can have fundamental consequences over the lifetime of the organism; as such, many organisms are less responsive to stress during critical developmental periods. In this study, we evaluated stress reactivity in nestling white-crowned sparrows, examining corticosterone and binding globulin levels in response to both restraint stress and ACTH challenge. Restraint stress induced a significant corticosterone response in both 4 to 6- and 7 to 9-day-old nestlings, but not in the youngest group (1-3 days); this non-significant increase in corticosterone in the youngest birds resembles the mammalian hyporesponsive period, wherein young animals are resistant to most stressors. Binding globulin levels appear to extend this period of low reactivity: when free corticosterone levels were calculated, only the oldest age group (7-9 day) showed a significant response to restraint. ACTH challenge data revealed that all ages of white-crowned sparrow nestlings responded to exogenous ACTH treatment with significant elevation of corticosterone, although early-stage nestlings did not reach adult levels of response.

Adrenal Cortex↗

Hormonal, behavioral, and thermoregulatory responses to bacterial lipopolysaccharide in captive and free-living white-crowned sparrows (Zonotrichia leucophrys gambelii).

Exposing vertebrates to pathogenic organisms or inflammatory stimuli, such as bacterial lipopolysaccharide (LPS), activates the immune system and triggers the acute phase response. This response involves fever, alterations in neuroendocrine circuits, such as hypothalamo-pituitary-adrenal (HPA) and -gonadal (HPG) axes, and stereotypical sickness behaviors that include lethargy, anorexia, adipsia, and a disinterest in social activities. We investigated the hormonal, behavioral, and thermoregulatory effects of acute LPS treatment in a seasonally breeding songbird, the white-crowned sparrow (Zonotrichia leucophrys gambelii) using laboratory and field experiments. Captive male and female sparrows were housed on short (8L:16D) or long (20L:4D) day lengths and injected subcutaneously with LPS or saline (control). LPS treatment activated the HPA axis, causing a rapid increase in plasma corticosterone titers over 24 h compared to controls. Suppression of the HPG axis occurred in long-day LPS birds as measured by a decline in luteinizing hormone levels. Instead of a rise in body temperature, LPS-injected birds experienced short-term hypothermia compared to controls. Birds treated with LPS decreased activity and reduced food and water intake, resulting in weight loss. LPS males on long days experienced more weight loss than LPS males on short days, but this seasonal effect was not observed in females. These results paralleled seasonal differences in body condition, suggesting that modulation of the acute phase response is linked to energy reserves. In free-living males, LPS treatment decreased song and several measures of territorial aggression. These studies highlight immune-endocrine-behavior interrelationships that may proximately mediate life-history tradeoffs between reproduction and defense against pathogens.

Acute-Phase Reaction↗

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.

Animals↗

Reproductive state, but not testosterone, reduces immune function in male house sparrows (Passer domesticus).

The immune system requires energetic and nutritional resources to optimally defend organisms against pathogens and parasites. Because resources are typically limited, immune function may require a trade-off with other physiologically demanding activities. Here, we examined whether photoperiodically induced seasonal states (breeding, molting, or nonbreeding) affected the cutaneous immune response of captive male house sparrows (Passer domesticus). To assess immune function in these birds, we injected the mitogen phytohemagglutinin (PHA) into the patagium and measured the resulting wing web swelling. Molting and nonbreeding birds had similar immune responses to PHA injection. However, males in a breeding state showed lower immune responses than both molting and nonbreeding birds even though they did not actually breed. We tested whether this decrease in the PHA swelling response in birds in a breeding state was due to elevated plasma concentrations of testosterone (T) by administering T to birds in a nonbreeding state. Contrary to some evidence in the literature, T did not suppress the response to PHA in house sparrows. Our data show that passerine birds show seasonal modulation in immune function, even in benign environmental conditions. However, even though T is often cited as a strong immunosuppressant, it is not fully responsible for this seasonal modulation.

Analysis of Variance↗

Fitness correlates of song repertoire size in free-living song sparrows (Melospiza melodia).

Models of sexual selection propose that exaggerated secondary sexual ornaments indicate a male's own fitness and the fitness of his offspring. These hypotheses have rarely been thoroughly tested in free-living individuals because overall fitness, as opposed to fitness components, is difficult to measure. We used 20 years of data from song sparrows (Melospiza melodia) inhabiting Mandarte Island, British Columbia, Canada, to test whether a male's song repertoire size, a secondary sexual trait, predicted overall measures of male or offspring fitness. Males with larger song repertoires contributed more independent and recruited offspring, and independent and recruited grandoffspring, to Mandarte's population. This was because these males lived longer and reared a greater proportion of hatched chicks to independence from parental care, not because females mated to males with larger repertoires laid or hatched more eggs. Furthermore, independent offspring of males with larger repertoires were more likely to recruit and then to leave more grandoffspring than were offspring of males with small repertoires. Although we cannot distinguish whether observed fitness variation reflected genetic or environmental effects on males or their offspring, these data suggest that female song sparrows would gain immediate and intergenerational fitness benefits by pairing with males with large song repertoires.

Animals↗

Hamilton and Zuk meet heterozygosity? Song repertoire size indicates inbreeding and immunity in song sparrows (Melospiza melodia).

Hamilton and Zuk's influential hypothesis of parasite-mediated sexual selection proposes that exaggerated secondary sexual ornaments indicate a male's addictive genetic immunity to parasites. However, genetic correlated of ornaments and immunity have rarely been explicitly identified. Evidence supporting Hamilton and Zuk's hypothesis has instead been gathered by looking for positive phenotypic correlations between ornamentation and immunity; such correlations are assumed to reflect causal, addictive relationships between these traits. We show that in a song sparrows, Melospiza melodia, male's song repertoire size, a secondary sexual trait, increased with his cell-mediated immune response (CMI) to an experimental challenge. However, this phenotypic correlation could be explained because both repertoire size and CMI declined with a male's inbreeding level. Repertoire size therefore primarily indicated a male's relative heterozygosity, a non-addictive genetic predictor of immunity. Caution may therefore be required when interpreting phenotypic correlations as support for Hamilton and Zuk's addictive model of sexual selection. However, our results suggest that female song sparrows choosing with large repertoires would on average acquire more outbred and therefore more heterozygous mates. Such genetic dominance effects on ornamentation are likely to influence evolutionary trajectories of female choice, and should be explicitly incorporated into genetic models of sexual selection.

Animals↗

Terminal investment induced by immune challenge and fitness traits associated with major histocompatibility complex in the house sparrow.

The terminal investment hypothesis predicts that individuals should invest more in their present reproduction if they are less likely to survive to future reproductive events. Infections, which reduce viability, may be used by individuals as a cue of a diminishing residual reproductive value and could therefore theoretically trigger an intensification of breeding effort. We tested this hypothesis in a natural population of house sparrows (Passer domesticus). We manipulated the immune system of breeding females by injecting them with a vaccine against the Paramyxo virus, the agent of Newcastle disease. Females were captured and treated immediately after completion of their first clutch either with the vaccine (NDV) or with phosphate buffered saline (PBS). The entire clutch was subsequently removed. We also screened Mhc class I genes of females to assess possible genotype-by-immune treatment interactions on reproductive investment. Our results indicate that vaccinated females were more likely to lay replacement clutches and that the difference in number of eggs between first and replacement clutches was greater for NDV females than for controls. In addition, chick size, both in terms of tarsus length and body mass, was affected by immune activation but in interaction with nestling age and female body mass, respectively. Mhc genotype-by-immune treatment interactions were never significant; however, allelic diversity was positively correlated with nestling survival. These results show that immune system activation is potentially used as a cue of reduced survival prospect and appears to induce a costly terminal investment behavior, and Mhc diversity might be under selection in a natural population of house sparrows.

Animals↗

Seasonal modulation of sickness behavior in free-living northwestern song sparrows (Melospiza melodia morphna).

A variety of vertebrate species modulate immune function on a seasonal basis to cope with seasonal energy deficits and competing life-history demands, such as reproduction. Most studies to date have focused upon seasonal variation of cellular and humoral immunity, while neglecting behavioral responses to infection. These behavioral strategies are collectively termed sickness behaviors and are hypothesized to divert energy away from normal activities to combat and overcome infection. Sickness behavior can be triggered experimentally by injecting bacterial lipopolysaccharide (LPS). In this study, we provide the first evidence for seasonal modulation of sickness behavior in a free-living animal. Male song sparrows of western Washington state (Melospiza melodia morphna) are sedentary and territorial year round, except for a brief time during molt. Treatment with LPS decreased territorial aggressive behavior of males in the winter (nonbreeding), but not in the spring (breeding). Subjects were recaptured approx. 25 h after treatment. Recaptured LPS males in the winter lost more body mass than saline-injected controls while LPS males in the spring did not. These data indicate that birds in breeding condition were relatively insensitive to the effects of LPS. On a proximate level, suppression of sickness behavior during breeding is likely mediated by seasonal differences in energy allocation, as wintering sparrows were significantly heavier and had larger subcutaneous fat reserves and lower baseline corticosterone levels than breeding birds. Ultimately, suppression of sickness behavior may represent an allocation strategy to balance current reproductive opportunities with the life-history costs of self-defense.

Aggression↗

Effects of thyroxine (T4) or triiodothyronine (T3) replacement therapy on the programming of seasonal reproduction and postnuptial molt in thyroidectomized male American tree sparrows (Spizella arborea) exposed to long days.

This study tested the hypothesis that T3 (triiodothyronine) is the tissue-active "seasonality" hormone by determining whether T3 could mimic T4 (thyroxine) and program photostimulated thyroidectomized (THX) male American tree sparrows (Spizella arborea) for three components of seasonality (i.e., full-blown testicular growth, photorefractoriness, and postnuptial molt). Photosensitive males were radiothyroidectomized, transferred to long days 4 weeks later, and administered 14 daily injections (s.c.) of alkaline saline (V) containing 0.1, 1, or 10 micrograms T4 or T3. THX and thyroid-intact (THI) controls received only V. After 5 additional weeks on long days, all birds were tested for photosensitivity/photorefractoriness. Periodically during the experiment, primary flight feathers were scored for molt, and testis length was monitored by laparotomy. As an independent measure of reproductive (i.e., photosensitive vs. photorefractory) state, hypothalami collected at the end of the experiment were assayed for cGnRH-I (chicken gonadotropin-releasing hormone I) content. Like THI controls, THX males administered 1 or 10 micrograms T4 exhibited full-blown testicular growth and then regression, initiated molt, and had low hypothalamic cGnRH-I, indicating that photostimulated birds that received mid- or high-dose T4 replacement therapy had been programmed for all three components of seasonality. On the other hand, both THX controls and THX males administered low-dose (0.1 microgram) T3 replacement therapy exhibited only modest testicular growth, signifying that neither group had been programmed for any component of seasonality. By contrast, photostimulated THX males that received 0.1 microgram T4, or 1 or 10 micrograms T3, were programmed for testicular growth, but not for photorefractoriness or molt. Collectively, these results show that subcutaneously administered T3 mimicked T4 imperfectly and suggest either that T3 does not program photostimulated male tree sparrows for photorefractoriness and postnuptial molt, or that T3 does not cross the blood-brain barrier as efficiently as does T4.

Animals↗

Brain aromatase, 5 alpha-reductase, and 5 beta-reductase change seasonally in wild male song sparrows: relationship to aggressive and sexual behavior.

In many species, territoriality is expressed only during the breeding season, when plasma testosterone (T) is elevated. In contrast, in song sparrows (Melospiza melodia morphna), males are highly territorial during the breeding (spring) and nonbreeding (autumn) seasons, but not during molt (late summer). In autumn, plasma sex steroids are basal, and castration has no effect on aggression. However, inhibition of aromatase reduces nonbreeding aggression, suggesting that neural steroid metabolism may regulate aggressive behavior. In wild male song sparrows, we examined the neural distribution of aromatase mRNA and seasonal changes in the activities of aromatase, 5 alpha-, and 5 beta-reductase, enzymes that convert T to 17 beta-estradiol, 5 alpha-dihydrotestosterone (5 alpha-DHT, a potent androgen), or 5 beta-DHT (an inactive metabolite), respectively. Enzyme activities were measured in the diencephalon, ventromedial telencephalon (vmTEL, which includes avian amygdala), caudomedial neostriatum (NCM), and the hippocampus of birds captured during spring, molt, or autumn. Aromatase and 5 beta-reductase changed seasonally in a region-specific manner. Aromatase in the diencephalon was higher in spring than in molt and autumn, similar to seasonal changes in male sexual behavior. Aromatase activity in the vmTEL was high in both spring and autumn but significantly reduced at molt, similar to seasonal changes in aggression. 5 beta-Reductase was not elevated during molt, suggesting that low aggression during molt is not a result of increased inactivation of androgens. These data highlight the relevance of neural steroid metabolism to the expression of natural behaviors by free-living animals.

Aggression↗

Social inertia in white-throated sparrows results from recognition of opponents.

Social inertia is a term for the stability of dominance relationships despite changes in the intrinsic dominating abilities of opponents. In a standard test for social inertia, low-ranking birds in an established hierarchy receive implants with testosterone (treated) and high-ranking birds receive empty implants (untreated). Social inertia occurs when the treated birds remain subordinate to untreated opponents in these groups, despite evidence that similarly treated birds dominate untreated strangers. In previous demonstrations of social inertia, however, treated and untreated birds were returned to their original aviaries and tested with familiar opponents, and thus the effects of familiarity with the location and those of familiarity with opponents were not separated. To address this issue, we investigated social inertia in 16 groups of white-throated sparrows Zonotrichia albicollis. When low-ranking treated birds were placed in new aviaries with familiar high-ranking, untreated opponents (treatment S, same opponents), dominance relationships showed social inertia. When such birds were placed in new aviaries with unfamiliar opponents (treatment N, new opponents), testosterone influenced dominance. When groups of high-ranking, untreated birds acquainted with each other were placed with unfamiliar treated opponents (treatment G, grouped dominants), 'coat-tail' effects (dominance by association with high-ranking individuals) sometimes outweighed the effects of testosterone. Social inertia in this species is thus a result of familiarity with opponents, rather than familiarity with locations of encounters. Measurements of aggressive tendencies confirmed a previous report that social inertia suppresses activation of aggression by testosterone. White-throated sparrows can thus recognize their opponents, and this ability affects the expression of both dominance and aggression. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Territory tenure in song sparrows is related to song sharing with neighbours, but not to repertoire size.

Song repertoires may be a product of sexual selection and several studies have reported correlations of repertoire size and reproductive success in male songbirds. This hypothesis and the reported correlations, however, are not sufficient to explain the observation that most species have small song repertoire sizes (usually fewer than 10, often fewer than five song types). We examined a second important aspect of a male's song repertoire, the extent to which he shares songs with his neighbours. Song sharing has not been measured in previous studies and it may be partially confounded with repertoire size. We hypothesized that in song sparrows, Melospiza melodia, song sharing rather than repertoire size per se is crucial for male territorial success. Our longitudinal study of 45 song sparrows followed from their first year on territory showed that the number of songs a bird shares with his neighbourhood group is a better predictor of lifetime territory tenure than is his repertoire size. We also found that song sharing increases with repertoire size up to but not beyond eight to nine song types, which are the most common repertoire sizes in the population (range in our sample 5-13). This partial confound of song sharing and repertoire size may account for some earlier findings of territory tenure-repertoire size correlations in this species and other species having small- or medium-sized repertoires. Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗

Effect of day length on luteinizing hormone beta-subunit mRNA and subsequent gonadal growth in the white-crowned sparrow, Zonotrichia leucophrys gambelii.

We have applied a cDNA probe for chicken luteinizing hormone beta-subunit to adenohypophysial preparations collected from Gambell's white-crowned sparrows. In situ hybridization showed luteinizing hormone (LH) beta-subunit mRNA distributed throughout the adenohypophysis in a manner identical to that of LH visualized by immunocytochemistry. Northern blot analysis showed a single band (0.8 kbp) identical to that from Japanese quail. Dot blot analysis was used to quantify changes in LH beta-subunit mRNA during a photoperiodically induced gonadal cycle in white-crowned sparrows. Transfer of adult males from short days (8L 16D) to long days (20L 4D) resulted in rapid testicular growth accompanied by increases in plasma levels of LH and body and liver mass. By Day 30 of photostimulation, mRNA levels in the adenohypophysis were significantly higher than those in birds on short days. In photorefractory birds, i.e., when gonads were regressed (60 days of long days), mRNA levels were significantly lower than on Day 30 of photostimulation. The increase in mRNA during photostimulation was accompanied by a highly significant increase in plasma concentrations of LH. The binding of iodinated LH to testes peaked at Day 15 of photostimulation and declined by Day 30. Curiously, peak production of testosterone and cAMP (induced by incubation with gonadotropins in vitro) were highest at Day 30. By 60 days of photostimulation, when birds were photorefractory, plasma levels of LH, binding of LH to the testes, and testosterone and cAMP production were greatly reduced. These data suggest that photoperiodically induced gonadal development and regression is mediated in part by regulation of mRNA levels for LH beta-subunit in the adenohypophysis.

Animals↗

Time of thyroidectomy variably affects seasonality in female American tree sparrows (Spizella arborea).

Female American tree sparrows (Spizella arborea) were injected with Na131I before, at, or after the onset of photostimulation in order to study the effect of time of thyroidectomy on three components of seasonality: thyroid-dependent photoperiodic ovarian growth, photorefractoriness, and postnuptial (prebasic) molt. Thyroidectomy before or at the onset of photostimulation abolished all components of seasonality; birds exhibited only minor thyroid-independent photoperiodic ovarian growth. Thyroidectomy on day 7 of photostimulation blocked the transition from photosensitivity to photorefractoriness; although birds showed thyroid-dependent photoperiodic ovarian growth, they neither exhibited ovarian regression nor initiated postnuptial molt. Thyroidectomy on day 14, 21, or 28 of photostimulation had no remarkable effect on any component of seasonality. We conclude that separate mechanisms control photoperiodic ovarian growth and photorefractoriness/molt and that, early during photostimulation, the thyroid has a codependent role in programming female tree sparrows for vernal as well as autumnal seasonal events.

Animals↗

Regulation of territorial behavior in the sedentary song sparrow, Melospiza melodia morphna.

Male song sparrows (Melospiza melodia morphna) of western Washington State show year-round territoriality. Although territorial aggression during the breeding season was accompanied by high circulating levels of luteinizing hormone (LH) and testosterone (T), similar aggression in the non-breeding season occurred when gonadal hormone levels were basal. Experimental removal of territorial males in autumn resulted in new males taking over the territory within a few days. These males had basal levels of LH and T despite establishing a new territory and the accompanying high level of aggression. There was also no difference in plasma levels of corticosterone in replacement versus established territorial males suggesting that social stress was not a factor. To test the possibility that low levels of gonadal steroids, or potentially some gonadal hormone not measured here, might regulate territorial aggression in the non-breeding season, free-living male song sparrows were castrated in early autumn and their aggressive territorial behavior was quantified by a simulated territorial intrusion (STI). Castrates showed the same aggressive responses to STI as controls indicating that establishment and maintenance of a territory was not dependent upon hormones of gonadal origin. Castrated males retained territories throughout the autumn and winter and even into the following breeding season. Further experiments were conducted to establish a role, if any, for T in territorial behavior in this species. During the breeding season when males had high levels of T, the responses to STI continued after the stimuli (caged conspecific male and tape-recorded songs) were withdrawn. Some males continued to patrol the territory and sang for long periods after STI. In contrast, males exposed to STI in autumn showed a strong aggressive response during the challenge, but then territorial aggression abated rapidly when the stimuli were withdrawn. Implants of T into males in autumn reinstated the high level of aggression after withdrawal of STI in a manner similar to that seen during the breeding season. These data suggest that in M. m. morphna T does not activate territorial aggression per se, but may increase the intensity of an aggressive response to STI, and persistence of the behavior after STI is withdrawn.

Aggression↗

Corticosterone treatment has no effect on reproductive hormones or aggressive behavior in free-living male tree sparrows, Spizella arborea.

We examined the effect of corticosterone on plasma levels of reproductive hormones (testosterone, dihydrotestosterone, and luteinizing hormone) and territorial defense behavior in male tree sparrows, Spizella arborea. Birds receiving Silastic implants filled with corticosterone (B) had significantly higher plasma levels of B than control birds, which received empty implants, and exhibited pectoral muscle wastage and a decrease in body mass. We evaluated the hormonal and agonistic responses of the two implanted groups of birds using a simulated territorial intrusion (STI) 2 to 4 days after they were implanted. Corticosterone-treated and control birds did not differ in their circulating levels of reproductive hormones or in their behavioral responses to STI (latency to respond to intrusion, number of songs, and closest approach to a decoy and tape recording). Unlike previous studies of north temperate passerines, high physiological levels of exogenous B had no effect either on circulating levels of reproductive hormones or on territorial behaviors associated with breeding. Nonetheless, untreated tree sparrows do mount a robust adrenocortical response, having a two- to fourfold increase in plasma B levels during a 1-h period of capture. Thus, adrenocortical responsiveness is maintained in these birds, but elevated levels of glucocorticoids do not suppress reproductive hormones or associated behaviors. We believe that this hormonal and behavioral refractoriness to glucocorticoids-or uncoupling of the stress response from the reproductive axis-may be advantageous for species having extreme temporal constraints on their breeding schedules.

Adrenal Cortex↗

Circadian entrainment by feeding cycles in house sparrows, Passer domesticus.

We studied the potential zeitgeber qualities of periodic food availability on the circadian rhythms of locomotor and feeding activity of house sparrows. The birds were initially held in a LD-cycle of 12:12 h, with food restricted to the light phase. After transfer to constant dim light, the birds remained entrained by the restricted feeding schedule. Following an exposure to food ad libitum conditions, the rhythms could be resynchronized by the feeding cycle. Shortening of the zeitgeber period to 23.5 h resulted in the loss of entrainment in most birds, whereas a longer zeitgeber period of 25 h re-entrained the rhythms of most birds. Although these results prove that periodic food availability can act as a zeitgeber for the circadian rhythms of house sparrows, several features of our data indicate that restricted feeding is only a weak zeitgeber. The pattern of feeding activity prior to the daily time of food access shown under some experimental conditions suggests that anticipation is due to a positive phase-angle difference of the birds' normal circadian system rather than being caused by a separate pacemaker.

Animals↗