Contributions of endocrinology to the migration life history of birds.

Migration is a key life cycle stage in nearly 2000 species of birds and is a greatly appreciated phenomenon in both cultural and academic arenas. Despite a long research tradition concerning many aspects of migration, investigations of hormonal contributions to migratory physiology and behavior are more limited and represent a comparatively young research field. We review advances in our understanding of the hormonal mechanisms of migration with particular emphasis on the sub-stages of the migration life history: development, departure, flight and arrival. These sub-stages vary widely in their behavioral, ecological and physiological contexts and, as such, should be given appropriate individual consideration.

Coloration signals the ability to cope with elevated stress hormones: effects of corticosterone on growth of barn owls are associated with melanism.

Stressful situations during development can shape the phenotype for life by provoking a trade-off between development and survival. Stress hormones, mainly glucocorticoids, play an important orchestrating role in this trade-off. Hence, how stress sensitive an animal is critically determines the phenotype and ultimately fitness. In several species, darker eumelanic individuals are less sensitive to stressful conditions than less eumelanic conspecifics, which may be due to the pleiotropic effects of genes affecting both coloration and physiological traits. We experimentally tested whether the degree of melanin-based coloration is associated with the sensitivity to an endocrine response to stressful situations in the barn owl. We artificially administered the mediator of a hormonal stress response, corticosterone, to nestlings to examine the prediction that corticosterone-induced reduction in growth rate is more pronounced in light eumelanic nestlings than in darker nest mates. To examine whether such an effect may be genetically determined, we swapped hatchlings between randomly chosen pairs of nests. We first showed that corticosterone affects growth and, thus, shapes the phenotype. Second, we found that under corticosterone administration, nestlings with large black spots grew better than nestlings with small black spots. As in the barn owl the expression of eumelanin-based coloration is heritable and not sensitive to environmental conditions, it is therefore a reliable, genetically based sign of the ability to cope with an increase in blood corticosterone level.

Regulation of stress response is heritable and functionally linked to melanin-based coloration.

Sexual selection theory posits that ornaments can signal the genetic quality of an individual. Eumelanin-based coloration is such an ornament and can signal the ability to cope with a physiological stress response because the melanocortin system regulates eumelanogenesis as well as physiological stress responses. In the present article, we experimentally investigated whether the stronger stress sensitivity of light than dark eumelanic individuals stems from differential regulation of stress hormones. Our study shows that darker eumelanic barn owl nestlings have a lower corticosterone release after a stressful event, an association, which was also inherited from the mother (but not the father) to the offspring. Additionally, nestlings sired by darker eumelanic mothers more quickly reduced experimentally elevated corticosterone levels. This provides a solution as to how ornamented individuals can be more resistant to various sources of stress than drab conspecifics. Our study suggests that eumelanin-based coloration can be a sexually selected signal of resistance to stressful events.

Postexercise ketosis in night-migrating passerine birds.

This study investigated the postexercise metabolism of six species of free-living, night-migrating passerine birds (European robin, pied flycatcher, wheatear, redstart, blackcap, and garden warbler). The birds were caught during autumn migration out of their nocturnal flight, and their metabolism changed from a fasting, highly active state to a fasting, resting state. Concentrations of six plasma metabolites of the fat, carbohydrate, and protein metabolism were measured during up to 10 h of recovery time. The metabolic changes indicated a biphasic pattern: (a) a quick first response to the reduced energy demands during the first 20 min of recovery, suggested by an increase and subsequent decrease of free fatty acid levels, and (b) subsequently, a postexercise ketosis and a reduction of lipolysis and proteolysis, suggested by high beta-hydroxy-butyrate and low free fatty acid, glycerol, triglyceride, and uric acid levels. This metabolic pattern differs from that of humans and rats, in which ketosis starts immediately postexercise or is absent in trained subjects. Since migrating birds are naturally adapted to endurance exercise, it is hypothesized that the high and long-lasting postexercise ketosis does not evoke physiological problems (such as hypoglycemia) but, by contrast, increases the ability of birds to rely on lipids, to a very high extent, during and after flight and decreases the dependence on glucose and glucogenic amino acids. Differences between species in fat stores and metabolic pattern support this hypothesis.

Regulation of protein breakdown and adrenocortical response to stress in birds during migratory flight.

During long-term fasting at rest, protein utilization is maintained at low levels until it increases at a threshold adiposity. This study examines 1) whether such a shift in energy substrate use also occurs during endurance exercise while fasting, 2) the role of corticosterone, and 3) the adrenocortical response to an acute stressor. Ten species of migrating birds caught after an endurance flight over at least 500 km were examined. Plasma uric acid and corticosterone levels were low in birds with fat stores >5% of body mass and high in birds with smaller fat stores. Corticosterone levels were very high in birds with no visible fat stores and emaciated breast muscles. Corticosterone levels increased with handling time only in birds with large fat stores. These findings suggest that 1) migrating birds with appreciable fat stores are not stressed by endurance flight, 2) a metabolic shift (increased protein breakdown), regulated by an endocrine shift (medium corticosterone levels), occurs at a threshold adiposity, as observed in birds at rest, 3) adrenocortical response to an acute stressor is inhibited after this shift, and 4) an adrenocortical response typical for an emergency situation (high corticosterone levels) is only reached when muscle protein is dangerously low.

Diurnal rhythms in neurotransmitter receptor binding and choline acetyltransferase activity: different patterns in two rat lines of Wistar origin.

Diurnal cycles for 8 ligand receptor pairs and choline acetyltransferase (ChAT) activity in 3 brain regions differed markedly in two rat lines, both of Wistar origin. Statistically significant differences between diurnal cycles in the two rat lines were found in the following parameters: 24 h means in 6 of 11 measurements, magnitude of cycle amplitudes, and phase position in 6 of 11 measurements, up to complete reversal of the acrophases in the case of ChAT activity in hippocampus. The importance of these findings--such major differences in two closely related rat lines--is obvious in any attempt to compare receptor-binding studies per se between laboratories using the same strain but not line, in studies of receptor rhythm characteristics, and in particular, for analysing the effects of brain-reactive drugs. While there are some reports on strain-dependency of cyclic functions, we are not aware that line-dependency has previously been described.

Circadian variations of neurotransmitter binding in three age groups of rats.

Binding in 14 ligand membrane receptor pairs and choline acetyltransferase activity were studied at 4-hour intervals during a 24-hour cycle (12:12 light:dark). Cortex, hippocampus, cerebellum, striatum and brainstem were prepared from 3-, 12- and 24-month-old male rats. Cholinergic, alpha- and beta-adrenergic, dopaminergic, serotonergic, gabaergic and opiate binding were determined. In the aged animals, the times of the binding maxima are no longer locked to the same time of the light:dark cycle, and the cycle phases themselves are shifted in comparison to those of the young and adult group. Cycle amplitudes were smallest in the 12-month-old rats, increasing significantly in the 24-month group. The age changes in the overall (24-hour) means fall into 4 different patterns, none of which shows a linear age-related decrease. This points out the great importance of including an adult group (12 months) in all ageing studies on small mammals.

Studies on neurotransmitter binding in senile dementia. Comparison of Alzheimer's and mixed vascular-Alzheimer's dementias.

Binding of 3H-labeled agonists and antagonists to muscarinic-cholinergic, alpha- and beta-adrenergic, dopaminergic, serotoninergic, and opiate receptors was studied in four regions of the neocortex and in hippocampus, thalamus, putamen, and caudatus in autoptic material from patients with senile dementia of Alzheimer type and of mixed vascular-Alzheimer pathogenesis. Different patterns of changes in ligand binding were found for the two groups. Some of these changes were quantitatively correlated with the histological scores of plaques and neurofibrillary tangles.