During migration, birds undergo alternating periods of fasting and re-feeding that are associated with dynamic changes in body mass (m(b)) and in organ size, including that of the digestive tract. After arrival at a migratory stopover site, following a long flight, a bird must restore the tissues of its digestive tract before it can refuel. In the present study we examined how the availability of dietary protein influences refueling of migrating blackcaps (Sylvia atricapilla) during a migratory stopover. We tested the following predictions in blackcaps deprived of food and water for 1-2 days to induce stopover behavior: (1) birds provided with a low-protein diet will gain m(b), lean mass and fat mass, and increase in pectoral muscle size slower than do birds fed a high-protein diet; (2) since stopover time is shorter in spring, birds will gain m(b) and build up fat tissue and lean tissue faster than in autumn; and (3) if low dietary protein limits a bird's ability to gain m(b) and fat reserves, then birds that do not obtain enough protein will initiate migratory restlessness (Zugunruhe) earlier than will birds with adequate dietary protein. These predictions were tested by providing captured migrating blackcaps with semisynthetic isocaloric diets differing only in their protein content. Each day, we measured m(b), and food intake; also lean mass and fat mass were measured using dual energy X-ray absorptiometry. In addition, we monitored nocturnal activity with a video recording system. In both spring and autumn, birds fed diets containing either 3 or 20% protein increased in m(b), lean mass and fat mass at similar rates during the experiment. However, the group receiving 3% protein ate more than did the group receiving 20% protein. In support of our predictions, m(b), lean mass, fat mass, and intake of food all were higher in spring than in autumn. We also found that in spring all birds had higher levels of migratory restlessness, but birds fed 3% protein were less active at night than were birds fed 20% protein, possibly an adaptation conserving energy and protein. We conclude that protein requirements of migrating blackcaps during stopover are lower than expected, and that birds can compensate for low dietary protein by behavioral responses, i.e. hyperphagia and decreased migratory restlessness, that ensure rapid refueling.

Migrating blackcaps (Sylvia atricapilla) were used to test the predictions that (1) the rebuilding of the digestive tract, as reflected by mass-specific consumption of food on the first 2-3 days of a stopover, is faster in birds with access to drinking water than in birds without, and (2) that adipose tissue and pectoral muscles grow faster and to a greater extent in birds with unlimited access to water. We simulated migratory stopover in two experiments. In Experiment I, each of 31 birds was randomly assigned to one of three experimental groups for 6 days. Along with mealworms (∼64% water) ad libitum, Group 1 received drinking water ad libitum; Group 2 had 0.5 h/day access to water; and Group 3 had no access to water. In Experiment II, 30 birds were offered a mixed diet for insectivorous birds (∼33% water) ad libitum for 6 days, while randomly assigned to two groups: (1) Water ad libitum-control; and (2) 30 min access to water twice a day. We measured lean mass and fat mass using dual energy X-ray absorptiometry, as well as body mass (m(b)), pectoral muscle index (PMI), and daily intake of food and water. Mean daily water intake was significantly different among the groups in both experiments. However, the availability of drinking water positively affected the rates of gain of lean and fat mass only in birds fed with the mixed, relatively dry diet. Furthermore, mass-specific daily food intake was affected by the availability of drinking water only in the mixed diet experiment, in which birds with unlimited access to drinking water reached an asymptote, 1 day earlier than birds in the water-restricted group. We suggest that in birds consuming diets with low water content, the lack of sufficient drinking water may result in slower rebuilding of the digestive tract, or may influence biochemical processes in the gut that result in slower growth of tissue. Although blackcaps obtained sufficient water from preformed and metabolic water to renew lost tissues when eating mealworms, given access to water, the birds drank prodigiously. Our results also suggest that if drinking water is unavailable to migrating blackcaps, their choices are restricted to water-rich foods, which may constrain their rate of feeding and thus the rate at which they deposit fat. Consequently, drinking water may have an important influence on birds' migratory strategies with respect to habitat selection, use of energy, and the saving of time.