Spiracle activity in moth pupae--the role of oxygen and carbon dioxide revisited

After decades of intensive research, the actual mechanism behind discontinuous gas exchange in insects has not been fully understood. One open question concerns the actual way (closed, flutter, and open) of how spiracles respond to tracheal gas concentrations. As the results of a classic paper [Burkett, B.N., Schneiderman, H.A., 1974. Roles of oxygen and carbon dioxide in the control of spiracular function in cecropia pupae. Biological Bulletin 147, 274-293] allow ambiguous interpretation, we thus reexamined the behavior of the spiracles in response to fixed, controlled endotracheal gas concentrations. The tracheal system of diapausing pupae of Attacus atlas (Saturniidae, Lepidoptera) was flushed with gas mixtures varying in P(O(2)) and P(CO(2)) while the behavior of the spiracles was monitored using changes in the pressure signal. This novel pressure based technique proved to be superior to classic visual observation of single spiracles. A two-dimensional map of the spiracle behavior in response to endotracheal P(O(2)) and P(CO(2)) was established. Typically, it contained two distinct regions only, corresponding to "closed" and "open" spiracles. A separate "flutter" region was missing. Because fluttering is commonly observed in moth pupae, we suggest that the intermittent spiracle opening during a flutter phase is an effect of non-steady-state conditions within the tracheal system. For low P(CO(2)) the minimum P(O(2)) resulting in open spiracles was linearly dependent upon P(CO(2)). Above a threshold of 1-1.5 kPa CO(2) the spiracles were open irrespective of P(O(2)). We propose a hypothetical spiracular control model, which is simple and explains the time course of endotracheal partial pressures during all phases of discontinuous gas exchange.