Energetic bottlenecks and other design constraints in avian annual cycles.

The flexible phenotypes of birds and mammals often appear to represent adjustments to alleviate some energetic bottleneck or another. By increasing the size of the organs involved in digestion and assimilation of nutrients (gut and liver), an individual bird can increase its ability to process nutrients, for example to quickly store fuel for onward flight. Similarly, an increase in the exercise organs (pectoral muscles and heart) enables a bird to increase its metabolic power for sustained flight or for thermoregulation. Reflecting the stationary cost of organ maintenance, changes in the size of any part of the "metabolic machinery" will be reflected in Basal Metabolic Rate (BMR) unless changes in metabolic intensity also occur. Energetic bottlenecks appear to be set by the marginal value of organ size increases relative to particular peak requirements (including safety factors). These points are elaborated using the studies on long-distance migrating shorebirds, especially red knots Calidris canutus. Red knots encounter energy expenditure levels similar to experimentally determined ceiling levels of ca. 5 times BMR in other birds and mammals, both during the breeding season on High Arctic tundra (probably mainly a function of costs of thermoregulation) and during winter in temperate coastal wetlands (a function of the high costs of processing mollusks, prey poor in nutrients but rich in shell material and salt water). During migration, red knots phenotypically alternate between a "fueling [life-cycle] stage" and a "flight stage." Fueling red knots in tropical areas may encounter heat load problems whilst still on the ground, but high flight altitudes during migratory flights seem to take care of overheating and unacceptably high rates of evaporative water loss. The allocation principles for the flexible phenotypes of red knots and other birds, the costs of their organ flexibility and the ways in which they "organize" all the fast phenotypic changes, are yet to be discovered.