Diffusion on a tree with stochastically gated nodes

Many insects exchange respiratory gases cyclically and discontinuously. A typical discontinuous gas exchange cycle (DGC) starts with a closed-spiracle (C) phase, during which little external gas exchange takes place, followed by a fluttering-spiracle (F) phase, which is usually dominated by diffusive oxygen uptake. The DGC is terminated by an open-spiracle (O) phase, during which accumulated CO2 escapes. This review critically examines the applicability of the DGC to insect gas exchange in general, discusses the primary mechanisms of gas exchange in the F and O phases, evaluates the widespread hypothesis that the DGC lowers respiratory water loss rates adaptively, and proposes new hypotheses concerning the evolutionary genesis of the DGC in insects and other tracheate arthropods.

Many adult and diapausing pupal insects exchange respiratory gases discontinuously in a three-phase discontinuous gas exchange cycle (DGC). We summarize the known biophysical characteristics of the DGC and describe current research on the role of convection and diffusion in the DGC, emphasizing control of respiratory water loss. We summarize the main theories for the evolutionary genesis (or, alternatively, nonadaptive genesis) of the DGC: reduction in respiratory water loss (the hygric hypothesis), optimizing gas exchange in hypoxic and hypercapnic environments (the chthonic hypothesis), the hybrid of these two (the chthonic-hygric hypothesis), reducing the toxic properties of oxygen (the oxidative damage hypothesis), the outcome of interactions between O(2) and CO(2) control set points (the emergent property hypothesis), and protection against parasitic invaders (the strolling arthropods hypothesis). We describe specific techniques that are being employed to measure respiratory water loss in the presence or absence of the DGC in an attempt to test the hygric hypothesis, such as the hyperoxic switch and H(2)O/CO(2) regression, and summarize specific areas of the field that are likely to be profitable directions for future research.