Various bird species regularly fast in connection with breeding, migration, or drastic climatic conditions. The metabolic response of penguins and domestic geese to fasting has been studied in detail. These large birds, in contrast to small species, do not become torpid when they are fasting. Nevertheless, they reduce their rate of energy expenditure by decreasing both resting metabolic rate and locomotor activity. From changes in the loss of body mass, the fast of penguins and geese has been divided into three phases: I, the loss of body mass decreases; II, it remains at a minimum level; and III, it increases. These phases reflect metabolic adjustments. Phase I is a rapid phase of adaptation, marked by a decrease in protein catabolism and mobilization of lipids. Phase II is a phase of economy, during which more than 90% of the energy expenditure derives from lipids, while protein catabolism remains at a minimum level. In phase III there still are lipid reserves and this phase is reversible; it is, however, critical because proteins are no longer spared. Data in the literature suggest that these three phases may also be used to describe how a wide variety of wild and domestic birds adapt to fasting.

During long-term fasting in birds and mammals, protein utilization initially decreases (phase I), is thereafter maintained at a low value (phase II), and then further increases (phase III). To delineate hormonal and biochemical changes responsible for these modifications, the effect of food deprivation for 50 days was studied in 6 male king penguins captured at the beginning of their natural breeding fast. During phase II, both rate of mass loss and plasma uric acid concentration remained at low levels, whereas plasma beta-hydroxybutyrate concentration increased. In phase III there was by contrast a 2.5-fold increase in the rate of mass loss, an eightfold increase in plasma uric acid, and an 80% drop in plasma beta-hydroxybutyrate. Plasma corticosterone was low and steady in phase II and increased three times in phase III. During the overall fast, there were no significant variations in plasma insulin, but there was a fourfold increase in plasma glucagon and a decrease in plasma thyroxine and triiodothyronine. These findings suggest that protein sparing (phase II) requires low levels of corticosterone, insulin, and thyroid hormones, whereas the further increase in protein utilization (phase III) is due to an increase in plasma corticosterone. The high plasma glucagon concentration in phase III is presumably responsible for a transient increase in plasma glucose observed at this stage; such increase in glucagon could enhance gluconeogenesis from amino acids.

Mammals and birds adapt to prolonged fasting by mobilizing fat stores and minimizing protein loss. This strategy ends with an increase in protein utilization associated with behavioural changes promoting food foraging. Using the Zucker rat as a model, we have investigated the effect of severe obesity on this pattern of protein loss during long-term fasting. Two interactions between the initial adiposity and protein utilization were found. First, protein conservation was more effective in obese than in lean rats: fatty rats had a three times lower daily nitrogen excretion and proportion of energy expenditure deriving from proteins, and a lower daily protein loss in various muscles. This phase of protein sparing is moreover nine times longer in the fatty rats. Second, obese animals did not show the late increase in nitrogen excretion that occurred in their lean littermates. Total body protein loss during starvation was larger in fatty rats (57% versus 29%) and, accordingly, total protein loss was greater in their muscles. At the end of the experiment, lean and obese rats had lost 98% and 82%, respectively, of their initial lipid reserves, and fatty rats still had an obese body composition. These results support the hypothesis that in severely obese humans and animals a lethal cumulative protein loss is reached long before the exhaustion of fat stores, while the phase of protein conservation is still continuing. In contrast, in lean rats, survival of fasting seems to depend on the availability of lipid fuels.(ABSTRACT TRUNCATED AT 250 WORDS)

The body mass of male emperor penguins is approximately 38 kg at the beginning of the 4-mo winter fast connected with breeding, and it is an estimated approximately 18 kg in leanest birds at time of spontaneous refeeding. For a 38- to 18-kg range, we investigated the changes in the rate of body mass loss, body composition, and plasma concentrations of uric acid and urea. After the first few days (phase I) a steady state (phase II) was reached in the proportions of the energy derived from proteins and lipids with proteins accounting for a constant 4%, and the remaining 96% being from lipids. The same proportions were maintained until body mass had decreased to 24 kg. Below this value the proportion of energy derived from proteins increased progressively (phase III), being 14 times higher at 18 kg than during phase II. Rate of body mass loss and plasma uric acid and urea concentrations closely reflected the changes in protein utilization: being at a low and steady value throughout phase II and increasing during phase III. Emperor penguins also fast during the spring, but for periods of only 2-3 wk. We found a 2.5 times higher value for rate of body mass loss, uric acid, and urea during spring phase II, suggesting lower effectiveness in protein sparing at that time. It may be attributed to the lower initial lipid reserves of spring birds. Would these findings be generalized to the wide variety of birds and mammals that spontaneously fast under natural conditions?(ABSTRACT TRUNCATED AT 250 WORDS)

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