Biomedical subjects

Eberhard Gwinner

Publications and source records attributed to Eberhard Gwinner.

At least 19 recordsLinked to original sources

PubMed2006

Testosterone and corticosterone during the breeding cycle of equatorial and European stonechats (Saxicola torquata axillaris and S. t. rubicola).

Northern-temperate male birds show seasonal changes in testosterone concentrations with a peak during the breeding season. Many tropical birds express much lower concentrations of testosterone with slight elevations during breeding. Here we describe testosterone and corticosterone concentrations of male stonechats from equatorial Kenya during different substages of breeding and molt. This tropical species has a short breeding season of approximately 3 months. We compare their hormone concentrations to previously published data of males of a northern-temperate relative, the European stonechat, also a seasonal breeder but with a breeding season of approximately 5 months. Equatorial stonechats show a pronounced peak of testosterone during the nest-building and laying stage. During all other stages, testosterone concentrations are low, similar to other year-round territorial tropical bird species. Corticosterone concentrations peak also during the nest-building and laying stage suggesting that this period of maximum female fecundity is a demanding period for the male. Equatorial stonechats have significantly lower concentrations of testosterone than European stonechats during all stages, except during the nest-building and laying stage. During this stage of maximum female fertility, testosterone levels tend to be higher in equatorial than in European stonechats. Our results suggest that equatorial stonechats belong to a group of tropical bird species that are characterized by a short breeding season and a brief high peak of testosterone during the female's fertile period. Such brief, but substantial peaks of testosterone may be common in tropical birds, but they may easily be missed if the exact breeding stage of individual birds is not known.

Animals↗

PubMed2006

Migratory restlessness in an equatorial nonmigratory bird.

The urge of captive birds to migrate manifests itself in seasonally occurring restlessness, termed "Zugunruhe." Key insights into migration and an endogenous basis of behavior are based on Zugunruhe of migrants but have scarcely been tested in nonmigratory birds. We recorded Zugunruhe of African stonechats, small passerine birds that defend year-round territories and have diverged from northern migrants at least 1 million years ago. We demonstrate that Zugunruhe is a regular feature of their endogenous program, one that is precisely timed by photoperiod. These results extend ideas of programs for periodic movement to include nonmigratory birds. Such programs could be activated when movements become necessary, in line with observed fast changes and high flexibility of migration. Attention to Zugunruhe of resident birds promises new insights into diverse and dynamic migration systems and enhances predictions of avian responses to global change.

Animal Migration↗

PubMed2006

Stress and the city: urbanization and its effects on the stress physiology in European blackbirds.

Animals colonizing cities are exposed to many novel and potentially stressful situations. There is evidence that chronic stress can cause deleterious effects. Hence, wild animals would suffer from city life unless they adjusted their stress response to the conditions in a city. Here we show that European Blackbirds born in a city have a lower stress response than their forest conspecifics. We hand-raised urban and forest-living individuals of that species under identical conditions and tested their corticosterone stress response at an age of 5, 8, and 11 months. The results suggest that the difference is genetically determined, although early developmental effects cannot be excluded. Either way, the results support the idea that urbanization creates a shift in coping styles by changing the stress physiology of animals. The reduced stress response could be ubiquitous and, presumably, necessary for all animals that thrive in ecosystems exposed to frequent anthropogenic disturbances, such as those in urban areas.

Animals↗

PubMed2005

Pinealectomy shortens resynchronisation times of house sparrow (Passer domesticus) circadian rhythms.

In many birds periodic melatonin secretion by the pineal organ is essential for the high-amplitude self-sustained output of the circadian pacemaker, and thus for the persistence of rhythmicity in 24 h oscillations controlled by it. The elimination of the pineal melatonin rhythm, or a reduction of its amplitude, renders the circadian pacemaker a less self-sustained, often highly damped, oscillatory system. A reduction in the degree of self-sustainment of a rhythm should not only increase its range of entrainment but also shorten the resynchronization times following phase-shifts of the zeitgeber. This hypothesis has not yet been directly tested. We therefore carried out the present study in which house sparrows (Passer domesticus) were subjected to both 6-h advance and 6-h delay phase-shifts of the light-dark cycle before and after the pinealectomy, and the rhythms in locomotion and feeding were recorded. The results indicate that following the delay, but not the advance, phase shift, resynchronization times were significantly shorter after pinealectomy. The dependence of resynchronization times on the presence or absence of the pineal organ is not only of theoretical interest but might also be of functional significance in the natural life of birds. A reduction or elimination of the amplitude of the melatonin secretion rhythm by the pineal organ might be responsible for faster adjustment to changes in zeitgeber conditions in nature.

Animals↗

PubMed2005

Testosterone, reproductive stage, and territorial behavior of male and female European stonechats Saxicola torquata.

We investigated territorial behavior and circulating testosterone (T) levels in a multiple-brooded population of the European stonechat, a socially monogamous passerine bird with biparental care. Between arrival at and departure from the breeding territories, we (1) quantified behavior of both sexes in response to a simulated territorial intrusion (STI) of a male conspecific and (2) measured plasma T concentrations in males and females. Male response scores to a STI and male T concentrations varied with phase, but there was no temporal association between plasma T levels and the intensity of territorial behavior. During both two sexual and two parental phases, at least half of the tested males showed aggressive responses. About 20% of the tested males responded with courtship prior to laying of the first clutch, but none of the males courted during later phases. Age had a positive overall effect on male plasma T. Females also reacted to the STI of a male, but their responses did not vary with breeding phase. Female plasma T varied with phase, being elevated during production of the first but not of the second clutch. As with males, female responses to the STI were not correlated with T levels. Responses of pair partners were positively correlated with each other. We conclude that modulation of male territorial aggression with breeding phase is not regulated by changes of plasma T concentrations. In light of other studies showing reduced male aggression by pharmacological inhibition of cellular actions of T, we propose that T is permissive for male territorial aggression, but does not mediate short-term changes associated with breeding phase. The function of the high female plasma T concentrations during formation of first clutches could be related to the production of eggs with high concentrations of androgens.

Age Factors↗

PubMed2005

The hormonal response of female European Stonechats to a territorial intrusion: the role of the male partner.

In many bird species, the female participates in defending a pair's breeding territory, however, the endocrine control mechanism of female aggressive behavior is largely unknown. The general statement that androgens are involved in the regulation of aggressive behavior is based on studies conducted only in males. Here, we tested whether paired female stonechats show a hormonal response to a simulated male territorial intruder. Since in males of territorial bird species androgen levels usually increase following a male-male encounter, we measured androgen-levels before and after a simulated male intrusion. In addition, we measured estradiol, the main gonadal hormone in females, and corticosterone, a stress hormone. The results show that a male intruder does not affect any of the measured hormones in females. In a second experiment, we also tested whether the endocrine state of the male partner affects the hormonal response of females to a male intruder by comparing the hormonal response of females paired with pharmacologically castrated males and females paired with control males. Females paired with pharmacologically castrated males had lower corticosterone levels both before and after the intrusion than females paired with control males. Additionally, in both groups, female corticosterone levels were increased following a male intrusion. We suggest that the differences found between females paired with pharmacologically castrated males and females paired with control males are due to differences in intra-pair interactions.

Aggression↗

PubMed2005

A separate circadian oscillator controls nocturnal migratory restlessness in the songbird Sylvia borin.

When confined to a cage, migratory songbirds exhibit nocturnal migratory restlessness (also called Zugunruhe) during the spring and autumn migratory periods, even though these birds are exclusively diurnal during the remainder of the year. Zugunruhe, which has been demonstrated to be under the direct control of a circannual timer, is characterized by a stereotypic "wing-whirring" behavior while the bird is perched. To elucidate the role played by the circadian system in the regulation of Zugunruhe, the authors studied the activity of garden warblers (Sylvia borin), long-distance nocturnal migrants, under skeleton photoperiods of different lengths and under constant dim light. In 11.5D:1L:10.5D:1L skeleton photoperiods, the authors found that Zugunruhe free-ran in a substantial proportion of birds, while their normal daily activities (e.g., feeding and preening) remained synchronized to 24 h. Some birds expressing Zugunruhe under constant dim light continued to show 2 distinct bouts of activity: one corresponding to daily activities, the other to wing-whirring. In some cases, these 2 bouts crossed while free-running with different periods. Birds expressing Zugunruhe also had significantly longer free-running periods than birds that did not. The study data suggest that the seasonal appearance of Zugunruhe is the result of the interactions of at least 2 circadian oscillators and that it is the phase relationship of these 2 oscillators that determines when nocturnal migratory restlessness is expressed. Furthermore, these data are consistent with the previously proposed internal coincidence hypothesis as a model for the ontogeny of circannual rhythms.

Animal Migration↗

PubMed2005

Flexible seasonal timing and migratory behavior: results from stonechat breeding programs.

Rigid schedules of long-distance migrants could be among candidate traits for adaptive migratory syndromes. This prediction was tested on stonechats, passerines that differ widely in migratory behavior and seasonal schedules. Stonechats in Europe are short-distance migrants and multiclutched, whereas African residents and Siberian long-distance migrants usually raise single broods. In captivity, all subspecies displayed endogenous cycles of reproductive development and molt. The subspecies differed in time afforded to life cycle stages. Under conducive aviary conditions, African stonechats were multibrooded, whereas Siberian stonechats did not add clutches. This difference in flexibility was exclusively related to the length of breeding windows. Stonechats also differed in premigratory preparations. Postjuvenile molt started early in Siberian stonechats, but in European and African stonechats, depended strongly on hatching date. In contrast, all subspecies shortened molt duration at the same rate when hatched from late broods. Plasticity of Zugunruhe timing was identical in Siberian and European subspecies and nearly compensated for hatching late. The stonechat data suggest a refined understanding of temporal plasticity in long-distance migrants. Overall, plasticity was not reduced, but was differently organized. Apparently rigid migrant schedules were related to short breeding cycles and inflexible molt onset. Short windows for breeding and juvenile development could provide safety measures for timely departure. Once molt was initiated, temporal plasticity of long-distance migrants matched that of less migratory conspecifics. In addition to adjusting endogenous programs, stonechats differed in implementing them in the field. Modifying the conditions under which programs are expressed may be an efficient way to enhance seasonal plasticity.

Africa↗

PubMed2005

Melatonin and nocturnal migration.

Many species of diurnal birds migrate nocturnally. Here, a series of studies of the blackcap (Sylvia atricapilla) on the relationship between nocturnal restlessness and melatonin, a hormone that in birds modulates day-night rhythms, are reviewed. Migratory populations from Sweden and Kenya were compared with resident populations from Cape Verde. In blackcaps of migratory populations, night levels of melatonin were lower during the migratory period, when birds showed nocturnal activity, than before and after this period, when birds did not show nocturnal activity. On the contrary, the occurrence of periodic or irregular phases of nocturnal activity in some nonmigratory birds from Cape Verde was not accompanied by a reduction in melatonin levels. In a second series of experiments, it was studied whether melatonin levels change when nocturnally active blackcaps are experimentally transferred from a migratory to a nonmigratory state. A long migratory flight and a refueling stopover were simulated by depriving birds of food for 2 days, subsequently readministering food. The experiments were done in autumn with birds collected in Sweden, and repeated in spring with birds collected in Kenya. In autumn, there was a suppression of nocturnal activity and an increase in melatonin in the night following food reintroduction. In spring, the effects were qualitatively similar, but their extent depended on the amount of body fat reserves. Taken together, the studies demonstrate the existence of a functional relationship between melatonin and nocturnal restlessness and of seasonal differences in the response of the migratory program to food availability.

Africa, Western↗

PubMed2004

Differences in the timing of reproduction between urban and forest European blackbirds (Turdus merula): result of phenotypic flexibility or genetic differences?

Species which have settled in urban environments are exposed to different conditions from their wild conspecifics. A previous comparative study of an urban and a forest-living European blackbird population had revealed a three weeks earlier onset of gonadal growth in urban individuals. These physiological adjustments are either the result of genetic differences that have evolved during the urbanization process, or of phenotypic flexibility resulting from the bird's exposure to the different environmental conditions of town or forest. To identify which of these two mechanisms causes the differences in reproductive timing, hand-reared birds originating from the urban and the forest populations were kept in identical conditions. The substantial differences in the timing of reproduction between urban and forest birds known from the field did not persist under laboratory conditions, indicating that temporal differences in reproductive timing between these two populations are mainly a result of phenotypic flexibility. Nevertheless, urban males initiated plasma luteinizing hormone (LH) secretion and testicular development earlier than forest males in their first reproductive season. Moreover, plasma LH concentration and follicle size declined earlier in urban females than in forest females, suggesting that genetic differences are also involved and might contribute to the variations in the timing of reproduction in the wild.

Animals↗

PubMed2004

Simulation of migratory flight and stopover affects night levels of melatonin in a nocturnal migrant.

Several species of diurnal birds are nocturnal migrants. The activation of nocturnal activity requires major physiological changes, which are essentially unknown. Previous work has shown that during migratory periods nocturnal migrants have reduced night-time levels of melatonin. Since this hormone is involved in the modulation of day-night rhythms, it is a good candidate regulator of nocturnal migratory activity. We studied whether melatonin levels change when nocturnally active blackcaps (Sylvia atricapilla) are experimentally transferred from a migratory to a non-migratory state. We simulated a long migratory flight by depriving birds of food for 2 days, and a refuelling stopover by subsequently re-administering food. Such a regimen is known to induce a reduction in migratory restlessness ('Zugunruhe') in the night following food reintroduction. The experiments were performed in both autumn and spring using blackcaps taken from their breeding grounds (Sweden) and their wintering areas (Kenya). In autumn, the food regimen induced a suppression of Zugunruhe and an increase in melatonin in the night following food reintroduction. In spring, the effects of the treatment were qualitatively similar but their extent depended on the amount of body-fat reserves. This work shows that the reduction of night-time melatonin during migratory periods is functionally related to nocturnal migration, and that fat reserves influence the response of the migratory programme to food deprivation.

Adipose Tissue↗

PubMed2004

Excremental androgen metabolite concentrations and gonad sizes in temperate zone vs. tropical Stonechats (Saxicola torquata ssp.).

Many songbirds in the temperate zones have comparatively short breeding seasons and are migratory. This often implies high breeding synchrony and competition for food, space, and mates. Tropical birds, in contrast, tend to be resident and react more flexibly to a more variable onset of environmental conditions conducive for breeding. Therefore, it can be expected that androgen metabolite concentrations in breeding males vary among birds from different latitudes. Studies on tropical birds, however, have revealed inconsistent results suggesting that additional factors not directly related to latitude, modify hormone levels. Furthermore, there is a potential for a phylogenetic bias when comparing birds of different taxonomic groups. In the present study, we compared androgen metabolite concentrations among closely related taxa of hand-raised male stonechats originating from tropical and temperate-zone populations in Kenya (0 degrees latitude), Kazakhstan (52 degrees N), and Ireland (52 degrees N), which differ partly in latitude of their provenance but also in the length of their breeding season (birds from Kazakhstan vs. Ireland). Hand-raised birds were housed in their natural photoperiod under otherwise identical environmental conditions. Androgen levels were determined from excrements to reduce disturbance. All three groups demonstrated a seasonal cycle of androgen metabolite concentrations concomitant with testis growth. Peak androgen metabolite concentrations were significantly lower in Kenyan stonechats compared to Kazakh stonechats. Irish birds had intermediate concentrations. Differences between Kazakh and Kenyan stonechats correlate with latitude, but data from the Irish population suggest the involvement of factors not directly related to latitude.

Adaptation, Physiological↗

PubMed2004

Convergent evolution of strigiform and caprimulgiform dark-activity is supported by phylogenetic analysis using the arylalkylamine N-acetyltransferase (Aanat) gene.

Alternative hypotheses propose the sister order of owls (Strigiformes) to be either day-active raptors (Falconiformes) or dark-active nightjars and allies (Caprimulgiformes). In an effort to identify molecular characters distinguishing between these hypotheses we examined a gene, arylalkylamine N-acetyltransferase (Aanat), potentially associated with the evolution of avian dark-activity. Partial Aanat coding sequences, and two introns, were obtained from the genomic DNA of 16 species: Strigiformes (four species), Falconiformes (four species), Caprimulgiformes (five species), with outgroups: Ciconiiformes (one species), Passeriformes (one species), and Apterygiformes (one species). Phylogenetic trees derived from aligned, evolutionarily conserved Aanat regions did not consistently recover clades corresponding to orders Strigiformes and Falconiformes but did place a caprimulgiform clade more distant from the strigiform and falconiform species than the latter two groups are to each other. This finding was supported by spectral analysis. The taxonomic distribution of seven intronic indels is consistent with the Aanat derived phylogenetic trees and supports conventional family-level groupings within both Strigiformes and Caprimulgiformes. The phylogenetic analyses also indicate that Caprimulgiformes is a polyphyletic grouping. In conclusion the data support, but do not conclusively prove, the proposal that Falconiformes is the sister order to Strigiformes and therefore, that the dark-activity characteristic of Strigiformes and Caprimulgiformes arose by convergent evolution.

Animals↗

PubMed2003

Slow pace of life in tropical sedentary birds: a common-garden experiment on four stonechat populations from different latitudes.

It has been hypothesized that organisms living at different latitudes or in different environments adjust their metabolic activity to the prevailing conditions. However, do differences in energy turnover simply represent a phenotypic adaptation to the local environment, or are they genetically based? To test this, we obtained nestling stonechats (Saxicola torquata) from equatorial Kenya (0 degrees N), Ireland (51.5 degrees N), Austria (47.5 degrees N) and Kazakhstan (51.5 degrees N). Birds were hand-raised and kept in Andechs, Germany. We measured their resting metabolic rates (RMR) and locomotor activity at an age of ca. 14 months (July) and 20 months (January), when birds went through postnuptial moult (July), and neither moulted nor exhibited enlarged gonads or migratory activity (January). RMR was generally higher during moult, but differed among populations: RMR was lowest in the resident Kenyan birds, higher in mostly sedentary Irish birds, and highest in migratory Austrian and Kazakhstan birds. Thus our data demonstrate that even in birds kept from early life under common-garden conditions, the 'pace of life', as indicated by metabolic turnover, is lower in sedentary tropical than in north-temperate migratory individuals of the same species. Such intrinsically low energy expenditure in sedentary tropical birds may have important implications for slow development, delayed senescence and high longevity in many tropical organisms.

Acclimatization↗

PubMed2003

Long-term memory for a life on the move.

Evidence is accumulating that cognitive abilities are shaped by the specific ecological conditions to which animals are exposed. Long-distance migratory birds may provide a striking example of this. Field observations have shown that, at least in some species, a substantial proportion of individuals return to the same breeding, wintering, and stopover sites in successive years. This observation suggests that migrants have evolved special cognitive abilities that enable them to accomplish these feats. Here we show that memory of a particular feeding site persisted for at least 12 months in a long-distance migrant, whereas a closely related nonmigrant could remember such a site for only 2 weeks. Thus, it seems that the migratory lifestyle has influenced the learning and memorizing capacities of migratory birds. These results build a bridge between field observations suggesting special memorization feats of migratory birds and previous neuroanatomical results from the same two species indicating an increase in relative hippocampal size from the first to the second year of life in the migrant but not in the nonmigrant.

Animals↗

PubMed2003

Circannual rhythms in birds.

In many birds, reproduction, molt, migration and other seasonal activities are controlled by endogenous circannual rhythmicity. Under constant conditions, this rhythm persists for many cycles with a period deviating from 12 months. Whether or not the rhythm is expressed depends on day length (photoperiod), which thus represents an important permissive factor in the process of rhythm generation. In nature, circannual rhythms are usually synchronized by the seasonal changes in photoperiod. However, equatorial birds may use daytime light intensity, which changes with the annual cycle of dry and rainy seasons, as a synchronizing zeitgeber. Photoperiod also modulates the rate of progress of the successive phases of the rhythmicity, such that an optimal adjustment to the annual environmental cycle is guaranteed. Populations of a given species may differ in their responsiveness to photoperiod in a manner that can be described as 'adaptive population-specific reaction norms'. In young migratory songbirds a circannual program determines changes in migratory direction and, at least partly, the time course and distance of migration. This circannual mechanism is replaced or supplemented in older birds by mechanisms formed on the basis of learning and memory. In general, circannual rhythms are intimately involved in the seasonal organization of a bird's behavior, providing the substrate onto which seasonal environmental factors act.

Animal Migration↗

PubMed2003

Comparative analysis of avian BMAL1 and CLOCK protein sequences: a search for features associated with owl nocturnal behaviour.

Animals differ widely in the phasing of their daily rhythms with respect to daily environmental rhythms. While birds are predominantly day-active, nocturnal activity is a characteristic feature of the order Strigiformes (owls). To study the evolution of owl night-activity cDNA sequences encoding the circadian core oscillator (CCO) proteins BMAL1 and CLOCK were obtained from barn owl (Tyto alba). The predicted proteins showed high sequence identity with their Galliform homologues (BMAL1: 99%; CLOCK: 95.6%). A computer-predicted chicken BMAL1 casein kinase-1 phosphorylation site is absent from T. alba BMAL1, but also absent from homologues of other six bird species (5 orders) (night-active (n=2), day-active (n=4)) indicating no evolutionary association with night activity. Sequence differences between T. alba and Galliform CLOCK frequently involved serine and threonine residues suggesting potential differences in their phosphorylation. The length of a poly-glutamine string in the CLOCK C-terminus varied between and within 25 species (6 orders) examined, however, no discernible feature distinguishing day and night active species was found. No differences were found between day (n=5) and night (n=7)-active species (12 species, 6 orders) in a region of the PER2 protein implicated in altered rhythm phasing in humans. In conclusion the avian CCO components examined showed strong evolutionary conservation. Molecular evolution associated with owl night-activity may have involved alterations in the CCO relationship with 'output' genes rather than in the molecular structure of the CCO itself.

Amino Acid Sequence↗

PubMed2002

Seasonal differences in the hormonal control of territorial aggression in free-living European stonechats.

In birds, territorial aggression during the breeding season is regulated by testosterone (T). However, many bird species also express aggressive behavior during the nonbreeding season, when plasma levels of T are low. It has been suggested that during this period estrogens might play a major role in regulating territorial aggression. In the present study we compared the effects of simultaneous blockage of androgenic and estrogenic actions on territorial aggression during the breeding and nonbreeding seasons in free-living male European stonechats (Saxicola torquata rubicola). European stonechats are of particular interest since they establish territories and form pairs during both the breeding and the nonbreeding seasons. Thus territorial aggression and its endocrine control can be compared between reproductive and non-reproductive contexts. Inhibition of androgenic and estrogenic actions by simultaneous application of Flutamide and ATD reduced territorial aggression during the breeding season, but not during the nonbreeding season. Our results show that androgens and/or estrogens are involved in the endocrine control of territorial aggression in stonechats only in a reproductive context, but not in a non-reproductive one.

Aggression↗