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Seasonal regulation of membrane and intracellular corticosteroid receptors in the house sparrow brain.

A number of studies have demonstrated seasonal regulation of the adrenocortical response to stress, or of corticosteroid binding globulins, but very few studies have examined seasonal regulation of corticosteroid receptor levels. As a result, there have been few attempts to produce an integrated picture of seasonal plasticity of the stress response. We measured baseline and stress-induced corticosterone (CORT), corticosteroid binding globulin and neuronal cytosolic and membrane corticosteroid receptor levels in male and female, wild-caught house sparrows (Passer domesticus) during three different seasons over the annual cycle (nesting, molting and winter). We identified three neuronal corticosteroid receptors in the house sparrow brain: two intracellular receptors and one membrane-associated receptor. Little is known about corticosteroid receptors in neuronal membranes of avian and mammalian species, but we found that the levels of membrane corticosteroid receptors varied seasonally, being lowest during the nesting season. Cytosolic corticosteroid receptor numbers (both low and high affinity receptors) also varied seasonally. In contrast to the membrane bound receptors, however, the numbers of low and high affinity cytosolic receptors were lowest during winter. In addition, mean levels of total basal and stress-induced CORT in the plasma varied seasonally. Both basal and stress-induced levels of total CORT were significantly higher during nesting than during winter or molt. Finally, corticosteroid binding globulin levels in plasma were also seasonally regulated, in a pattern similar to total CORT, so that estimated free CORT levels did not vary between seasons. These data indicate that multiple components of the stress response are seasonally regulated in birds obtained from wild populations. Interactions between these regulated components provide a basis for seasonal differences in behavioural and physiological responses to stress.

Animals↗

Effects of acute treatment with 8-OH-DPAT and fluoxetine on aggressive behaviour in male song sparrows (Melospiza melodia morphna).

The role of serotonin in modulating male aggressive behaviour was investigated in male song sparrows, Melospiza melodia morphna, using two different serotonergic drugs, fluoxetine and 8-OH-DPAT. Fluoxetine is a selective serotonin reuptake inhibitor of the neuronal reuptake pump increasing synaptic concentrations of serotonin, and 8-OH-DPAT is a specific serotonin (5-HT1A) receptor agonist. The serotonergic control of aggression in passerines has not been previously investigated. We examined these behaviours within a controlled setting using a laboratory simulated territorial intrusion, with a hierarchical scale to quantify male-male aggressive behaviour. Utilizing this scale, we quantified the extent of male aggressive behaviour in two experiments. In experiment 1, song sparrows were given 100 micro l, s.c. injections of either fluoxetine (10 mg/kg) or 8-OH-DPAT (1 mg/kg). Experiment 2 was a dose-response study using three doses of 8-OH-DPAT (0.1, 1 and 10 mg/kg). In both studies, aggressive behaviour was measured 1 h after injection for 10 min in response to the presence of a novel male decoy combined with playback of conspecific song. Both drugs significantly reduced male aggressive behaviour, and 8-OH-DPAT did so in a dose-dependent manner. The effect of the two drugs upon general activity was also measured using infra-red perch hop detectors. Activity levels were not effected by either fluoxetine or 8-OH-DPAT at all of the respective doses, indicating that the reduction in aggressive behaviour was specific. These results demonstrate that, in a passerine species, the serotonergic system negatively regulates male-male aggressive behaviour. These results further demonstrate that aggression can be effectively studied in a laboratory setting and natural aggressive responses can be elicited using this method.

8-Hydroxy-2-(di-n-propylamino)tetralin↗

Sexual variation in heritability and genetic correlations of morphological traits in house sparrow (Passer domesticus).

Estimates of genetic components are important for our understanding of how individual characteristics are transferred between generations. We show that the level of heritability varies between 0.12 and 0.68 in six morphological traits in house sparrows (Passer domesticus L.) in northern Norway. Positive and negative genetic correlations were present among traits, suggesting evolutionary constraints on the evolution of some of these characters. A sexual difference in the amount of heritable genetic variation was found in tarsus length, wing length, bill depth and body condition index, with generally higher heritability in females. In addition, the structure of the genetic variance-covariance matrix for the traits differed between the sexes. Genetic correlations between males and females for the morphological traits were however large and not significantly different from one, indicating that sex-specific responses to selection will be influenced by intersexual differences in selection differentials. Despite this, some traits had heritability above 0.1 in females, even after conditioning on the additive genetic covariance between sexes and the additive genetic variances in males. Moreover, a meta-analysis indicated that higher heritability in females than in males may be common in birds. Thus, this indicates sexual differences in the genetic architecture of birds. Consequently, as in house sparrows, the evolutionary responses to selection will often be larger in females than males. Hence, our results suggest that sex-specific additive genetic variances and covariances, although ignored in most studies, should be included when making predictions of evolutionary changes from standard quantitative genetic models.

Animals↗

Spatial and temporal variation of passer Per2 gene expression in two distinct cell groups of the suprachiasmatic hypothalamus in the house sparrow (Passer domesticus).

In mammals, the major pacemaker controlling circadian rhythmicity is located in the hypothalamic suprachiasmatic nuclei. Although there is evidence for the presence of a hypothalamic circadian oscillator in birds from lesioning studies, neuroanatomical, neurochemical and functional investigations have failed to identify its exact location. Two cell groups in the avian hypothalamus have been shown to bear characteristics of the mammalian suprachiasmatic nucleus: the suprachiasmatic nucleus and the lateral hypothalamic retinorecipient nucleus. We cloned an avian period homologue (pPer2) and investigated the temporal and spatial expression pattern of this gene in the house sparrow hypothalamus using in situ hybridization. Applying quantitative morphometry, we found rhythmic expression of pPer2 during light-dark as well as in constant conditions in the suprachiasmatic nucleus and in the lateral hypothalamus. The temporal and spatial distribution of pPer2 expression in the suprachiasmatic nucleus suggest a longitudinal compartmentalization of the nucleus with period gene expression being initiated in the most rostral portion of the suprachiasmatic nucleus before lights on. In the lateral hypothalamus, phasing of pPer2-rhythmicity appeared different from the suprachiasmatic nucleus. The major difference between light-dark and constant conditions was a decrease in the amplitude of pPer2 rhythmicity in the suprachiasmatic nucleus. Our data demonstrate that, unlike in mammals, Per gene expression in the suprachiasmatic hypothalamus of the house sparrow is not confined to a single cell group, indicating a more complex organization of the circadian oscillator in the hypothalamus of birds.

Animals↗

Auditory representation of autogenous song in the song system of white-crowned sparrows.

The HVc (hyperstriatum ventrale, pars caudale) is a forebrain nucleus in the motor pathway for the control of song. Neurons in the HVc also exhibit auditory responses. A subset of these auditory neurons in the white-crowned sparrow (Zonotrichia leucophrys) have been shown to be highly selective for the individual bird's own (autogenous) song. By using multiunit recording techniques to sample from a large population, we demonstrate that the entire population of auditory neurons in the HVc is selective for autogenous song. The selectivity of these neurons must reflect the song-learning process, for the acoustic parameters of a sparrow's song are acquired by learning. By testing with laboratory-reared birds, we show that HVc auditory neurons prefer autogenous song over the tutor model to which the birds were exposed early in life. Thus, these neurons must be specified at or after the time song crystallizes. Since song is learned by reference to auditory feedback, HVc auditory neurons may guide the development of the motor program for song. The maintenance of a precise auditory representation of autogenous song into adulthood can contribute to the ability to distinguish the fine differences among conspecific songs.

Journal Article↗

Correlation of song learning and territory establishment strategies in the song sparrow.

In a field study, we show that a young song sparrow (i) selects his songs from three or four older birds who have neighboring territories, (ii) preferentially learns song types that these tutor neighbors share, and (iii) ultimately sets up his territory next to, or replaces, one of these tutor neighbors. The consequence of this song learning strategy is that the young bird's song repertoire represents the "logical intersection" of the song repertoires of his tutor neighbors. We argue that this repertoire is optimally designed for mimicry (sounding like your neighbors) and for communication between neighbors (song sparrows address or reply to a neighbor with a song they share with that neighbor).

Journal Article↗

Digestive responses during food restriction and realimentation in nestling house sparrows (Passer domesticus).

We used nestling house sparrows (Passer domesticus) under laboratory conditions to test for modulation of digestive efficiencies during periods of low and high food intake and tested the hypothesis that nestlings would exhibit compensatory changes in digestive efficiency following a period of food restriction. During the low intake period, nestlings were held at constant body mass for 48 h beginning on either day 3 or day 6 of life by feeding them at 50% of control rations. After 48 h of food restriction, nestlings were fed as much as they could consume, allowing the nestlings restricted at day 6 (early restriction not assessed) to consume 14% more food than control nestlings. For nestlings restricted at day 6 apparent dry mass assimilation of the entire diet was found to be 5% and 8% lower during food restriction and realimentation, respectively, compared with control nestlings that were not under- or overfed. There were no significant differences in radiolabeled starch assimilation efficiencies between control and restricted nestlings. Starch assimilation efficiencies remained constant from 3 d of age onward in control nestlings. Total starch extracted was lower during food restriction but reached a rate similar to that of control nestlings during the realimentation period. Passage times (time of first defecation, mean retention time, and mode passage time) measured with an indigestible marker were longer during food restriction and shorter during realimentation, relative to control nestlings. During realimentation there was no difference in intestinal rates of hydrolysis or mediated uptake of L-leucine compared with control nestlings. The main effect of changing food intake was apparently to alter flow rate, and hence retention time, causing slight changes in digestive efficiency. Thus, nestlings did not exhibit compensatory changes in digestion rates as implied by the hypothesis. Our finding of a lower dry mass assimilation efficiency and similar total starch assimilation during realimentation (relative to controls) helps explain why nestling house sparrows do not display compensatory growth, despite higher food intake. Our results indicate that the gut has little spare capacity to deal with increased food intake during growth following food restriction.

Adaptation, Physiological↗

Effects of house sparrow age and arbovirus infection on attraction of mosquitoes.

To test the hypothesis that arbovirus infection or advancing age increases the attractiveness of avian hosts to vector mosquitoes, we used an olfactometer that measures the response of mosquitoes to the passive, vertical diffusion of host-related cues. St. Louis encephalitis or western equine encephalomyelitis virus infection in house sparrows (Passer domesticus (L.], regardless of age, had no detectable effect on Culex quinquefasciatus Say or Cx. tarsalis Coquillett attraction, respectively, at close range. Sparrow age, however, was associated positively with increased Cx. quinquefasciatus attraction; i.e., nestlings were significantly less attractive than adults, and there was a trend for increased mosquito attraction as nestling age increased. When birds were examined in groups, attraction was not equal to the sum of the individual birds in the group. Nestlings, either alone or in a group, were less attractive than a single adult. One adult plus three older nestlings were more attractive than one adult with three young nestlings, but no more attractive than one adult alone. We speculate on the epidemiological significance of these laboratory results to arbovirus epizootic transmission.

Age Factors↗

Extraocular control of photorefractoriness in American tree sparrows (Spizella arborea).

Photorefractoriness, a reversible state of unresponsiveness to daylengths of gonadostimulatory duration, terminates seasonal breeding in many photoperiodic species of birds. Whether the eyes are components of the mechanism that triggers photorefractoriness is an important, but heretofore unresolved, question. Although a role for extraocular photoreception in the mechanism of photoinduced gonadal growth is well documented, the eyes may be important in the mechanism of photorefractoriness if, as some evidence suggests, they are gonadoinhibitory. With American tree sparrows (Spizella arborea), I here confirm that the absence of eyes does not impede photoinduced testicular growth and establish that an extraocular mechanism mediates the transition from photosensitivity to photorefractoriness: Tree sparrows blinded by bilateral ocular enucleation, when photosensitivity to long days or by miniature self-powered lights implanted atop the skull, showed marked testicular growth and then, as evidenced by spontaneous testicular regression, became photorefractory, as did sighted controls.

Animals↗

Effect of withdrawing long days from male American tree sparrows (Spizella arborea): implications for understanding thyroid-dependent programming of seasonal reproduction and postnuptial molt.

In previous studies, we withdrew thyroid hormones by thyroidectomy before, at, or after the onset of photostimulation and showed that male American tree sparrows (Spizella arborea) are programmed for seasonal reproduction and postnuptial molt by or before Week 3 on long days. In this corollary study, we withdrew long days before or after the control circuits had been programmed. After 1 day to 4 wk on long days, groups of thyroid-intact males were returned to short days until Week 7, when they were moved to constant light and evaluated for photosensitivity or photorefractoriness and postnuptial molt. Long-day controls held 7 wk on long days showed robust testicular growth through Week 6 and then spontaneous testicular regression. Testes of short-day controls and of males photostimulated for 1 day remained small. In all other groups, photostimulation induced testicular growth, which gave way to regression during exposure to short days. Long-day controls tested photorefractory at Week 7 and initiated molt by Week 10. All other groups tested photosensitive and did not molt. Our demonstration that long days are required for expression of seasonal reproduction and postnuptial molt in thyroid-intact male tree sparrows previously programmed for these events suggests that long days create a milieu that is permissive for expression.

Animals↗

Estimating the time to extinction in an island population of song sparrows.

We estimated and modelled how uncertainties in stochastic population dynamics and biases in parameter estimates affect the accuracy of the projections of a small island population of song sparrows which was enumerated every spring for 24 years. The estimate of the density regulation in a theta-logistic model (theta = 1.09 suggests that the dynamics are nearly logistic, with specific growth rate r1 = 0.99 and carrying capacity K = 41.54. The song sparrow population was strongly influenced by demographic (ŝigma2(d) = 0.66) and environmental (ŝigma2(d) = 0.41) stochasticity. Bootstrap replicates of the different parameters revealed that the uncertainties in the estimates of the specific growth rate r1 and the density regulation theta were larger than the uncertainties in the environmental variance sigma2(e) and the carrying capacity K. We introduce the concept of the population prediction interval (PPI), which is a stochastic interval which includes the unknown population size with probability (1 - alpha). The width of the PPI increased rapidly with time because of uncertainties in the estimates of density regulation as well as demographic and environmental variance in the stochastic population dynamics. Accepting a 10% probability of extinction within 100 years, neglecting uncertainties in the parameters will lead to a 33% overestimation of the time it takes for the extinction barrier (population size X = 1) to be included into the PPI. This study shows that ignoring uncertainties in population dynamics produces a substantial underestimation of the extinction risk.

Animals↗

Testosterone influences basal metabolic rate in male house sparrows: a new cost of dominance signalling?

Sexually selected signals of individual dominance have profound effects on access to resources, mate choice and gene flow. However, why such signals should honestly reflect individual quality is poorly understood. Many such signals are known to develop under the influence of testosterone. We conducted an experiment in male house sparrows in which testosterone was manipulated independently during two periods: before the onset of the breeding season and prior to the autumn moult. We then measured the effects of these manipulations on basal metabolic rate and on the size of the chest bib, a sexually selected signal. The results demonstrate that testosterone simultaneously affects both signal development and basal metabolic rate in the house sparrow (Passer domesticus). This evidence, therefore, supports a novel conclusion: that testosterone-dependent signals act as honest indicators of male quality possibly because only high-quality individuals can sustain the energetic costs associated with signal development.

Animals↗

Immune activity elevates energy expenditure of house sparrows: a link between direct and indirect costs?

The activation of an immune response is beneficial for organisms but may also have costs that affect fitness. Documented immune costs include those associated with acquisition of special nutrients, as well as immunopathology or autoimmunity. Here, we test whether an experimental induction of the immune system with a non-pathological stimulant can elevate energy turnover in passerine birds. We injected phytohaemagglutinin (PHA), a commonly used mitogen that activates the cell-mediated immune response, into the wing web of house sparrows, Passer domesticus. We then examined energetic costs resulting from this immune activity and related those costs to other physiological activities. We found that PHA injection significantly elevated resting metabolic rate (RMR) of challenged sparrows relative to saline controls. We calculated the total cost of this immune activity to be ca. 4.20 kJ per day (29% RMR), which is equivalent to the cost of production of half of an egg (8.23 kJ egg(-1)) in this species. We suggest that immune activity in wild passerines increases energy expenditure, which in turn may influence important life-history characteristics such as clutch size, timing of breeding or the scheduling of moult.

Animals↗

Synergistic effects of food and predators on annual reproductive success in song sparrows.

The behaviour literature is full of studies showing that animals in every taxon balance the probability of acquiring food with the risk of being preyed upon. While interactions between food and predators clearly operate at an individual scale, population-scale studies have tended to focus on only one factor at a time. Consequently, interactive (or 'synergistic') effects of food and predators on whole populations have only twice before been experimentally demonstrated in mammals. We conducted a 2 x 2 experiment to examine the joint effects of food supply and predator pressure on the annual reproductive success of song sparrows (Melospiza melodia). Our results show that these two factors do not operate in an additive way, but instead have a synergistic effect on reproduction. Relative to controls, sparrows reared 1.1 more young when food was added and 1.3 more when predator pressure was low. When these treatments were combined 4.0 extra young were produced, almost twice as many as expected from an additive model. These results are a cause for optimism for avian conservation because they demonstrate that remedial actions, aimed at simultaneously augmenting food and reducing predators, can produce dramatic increases in reproductive success.

Animals↗

Melatonin: effects on the circadian locomotor rhythm of sparrows.

The continuous administration of low levels of melatonin via intraperitoneally placed Silastic capsules either (i) shortened the free-running period of activity or (ii) induced continuous activity in house sparrows (Passer domesticus) maintained in constant darkness. After the melatonin-filled capsules were removed, the period of the circadian rhythm of activity lengthened in rhythmic birds and normal rhythmicity was restored in continuously active birds. The results suggest that melatonin is involved in the physiological control of circadian rhythmicity in sparrows.

Animals↗

Pineal function: the biological clock in the sparrow?

The pineal organ of the house sparrow, Passer domesticus, is essential for persistence of the circadian locomotor rhythm in constant conditions. Upon removal of the pineal body, activity becomes arrhythmic. However, pinealectomy does not abolish the rhythm of locomotor activity in birds exposed to light-dark cycles. Pinealectomized birds are entrained by light cycles in much the same manner as are normal birds. Our data demonstrate that the pineal organ is a crucial component of the endogenous time-measuring system of the sparrow.

Animals↗

Temporal synergism of corticosterone and prolactin controlling gonadal growth in sparrows.

Gonadal growth was controlled in two avian species by corticosterone and prolactin injected daily at various times. Testicular growth was induced in photorefractory house sparrows (Passer domesticus) kept in continuous light by prolactin injected 4 or 8 hours after administration of corticosterone. Other temporal patterns were ineffective. Gonadal growth was also stimulated in photosensitive white-throated sparrows (Zonotrichia albicollis) kept in continuous light by prolactin injected 12 hours after administration of corticosterone. Daily injections of prolactin 8 hours after injection of corticosterone inhibited gonadal growth. The seasonal cycle of reproductive photorefractoriness and photosensitivity is controlled by a changing relation between the daily rhythms of plasma concentrations of corticosterone and prolactin.

Adipose Tissue↗

Pineal function in sparrows: circadian rhythms and body temperature.

Deep body temperature of the house sparrow, Passer domesticus, was monitored continuously by radio telemetry. Pinealectomy abolished the normal circadian rhythm of body temperature in constant darkness, and significantly altered the amplitude of body temperature rhythms entrained to light cycles. The body temperature minima of pinealectomized birds never fell as low as those of unoperated birds regardless of the light conditions; the temperature maxima of both normal and pinealectomized birds were higher in light than in darkness. In sparrows the pineal organ is essential to the normal function of the biological clock controlling both activity and body temperature rhythms and may be directly involved in thermoregulation.

Animals↗