Control of metabolism by hypoxia and starvation and the consequences for the pattern of ecdysone secretion in Manduca sexta

When larvae of Manduca sexta pass the critical weight, they initiate an endocrine cascade that leads to molting and metamorphosis. The critical weight coincides with a leveling off of the metabolic rate, and we hypothesize that it is the size at which the metabolic needs of a growing body reach the maximum capacity of the tracheal system to deliver oxygen. We examine two simple predictions about the role of oxygen supply in the regulation of growth. First, that restricting access to oxygen by blocking spiracles should affect growth and metabolism, and second, that placing larvae in a hypoxic atmosphere shortly before they reached the critical weight should prematurely trigger cessation of growth and metamorphosis. When sets of spiracles were blocked, growth rate was reduced, as was the metabolic rate and the body size at metamorphosis. The effect of blocking the posterior-most spiracles was greater than that of blocking the anterior-most ones, suggesting the presence of a required abdominal factor. Contrary to expectations, placing larvae in hypoxia a day before they reach the critical weight delayed the molt significantly, suggesting that hypoxia is not a trigger for the initiation of metamorphosis. Nevertheless, an increase in lactate levels in the second half of the final instar, and the leveling off of metabolic rate, indicate that a metabolic shift occurs at the critical weight. Ecdysone secretion in starved and hypoxic larvae is delayed by about 2 days relative to feeding controls, which explains the delayed timing of the metamorphic molt.