Circadian rhythms and the evolution of photoperiodic timing in insects

This review discusses possible evolutionary trends in insect photoperiodism, mainly from a chronobiological perspective. A crucial step was the forging of a link between the hormones regulating diapause and the systems of biological rhythms, circadian or circannual, which have independently evolved in eukaryotes to synchronize physiology and behaviour to the daily cycles of light and darkness. In many of these responses a central feature is that the circadian system resets to a constant phase at the beginning of the subjective night, and then 'measures' the duration of the next scotophase. In 'external coincidence', one version of such a clock, light now has a dual role. First, it serves to entrain the circadian system to the stream of pulses making up the light/dark cycle and, second, it regulates the nondiapause/diapause switch in development by illuminating/not illuminating a specific light sensitive phase falling at the end of the critical night length. Important work by A. D. Lees on the aphid Megoura viciae using so-called 'night interruption experiments' demonstrates that pulses falling early in the night lead to long-day effects that are reversible by a subsequent dark period longer than the critical night length and also show maximal sensitivity in the blue-green range of the spectrum. Pulses falling in the latter half of the night, however, produce long-day effects that are irreversible by a subsequent long-night and show a spectral sensitivity extending into the red. With movement to higher latitudes, insects develop genetic clines in various parameters, including critical night length, the number of long-night cycles needed for diapause induction, the strength of the response, and the 'depth' or intensity of the diapause thus induced. Evidence for these and other types of photoperiodic response suggests that they provided strong selective advantages for insect survival.