Circadian and circannual programmes in avian migration

In migratory birds, endogenous daily (circadian) and annual (circannual) rhythms serve as biological clocks that provide the major basis for their temporal orientation. Circannual rhythms are responsible for the initiation of migration both in autumn and spring. This function of timing migrations is particularly important for birds that spend the winter close to the equator where the environment is too constant or irregular to provide accurate timing cues. In addition, circannual rhythms produce programmes that determine both the temporal and the spatial course of migration. In Sylvia warblers, the time programmes controlling autumn migration are organized in a species- or population-specific manner. It has been proposed that, in first-year migrants, the time programme for autumn migration plays a major role in determining migratory distance, thus providing the vector component in a mechanism of vector navigation. It is not yet clear, however, whether this programme does indeed determine migratory distance or whether it only provides the temporal framework within which other factors determine how far a bird flies. Evidence against the first alternative comes from findings indicating that migratory activity can be drastically modified by a constellation of rather specific, but highly relevant, factors and that the resulting changes in migratory activity are not compensated by subsequent increases or decreases of migratory activity. In normally day-active but nocturnally migrating birds, circannual signals cause alterations in the circadian system leading to the development of nocturnal activity. Although the nature of these signals is unknown, there is evidence that changes in the diurnal pattern of melatonin secretion by the pineal gland are associated with, and possibly causally involved in, the waxing and waning of nocturnal activity. These changes in the melatonin pattern presumably also affect general synchronization properties of the circadian system to Zeitgebers in such a way that circadian rhythms adjust faster to new conditions after long transmeridian flights.