Erwin Bünning and Tony Lees, two giants of chronobiology, and the problem of time measurement in insect photoperiodism

This paper examines the views of Erwin Bünning and Tony Lees on the mechanism of photoperiodic time measurement, the former advocating a circadian basis for the phenomenon and the latter a non-circadian hourglass-like timer. This difference in opinion led to a protracted split among workers on photoperiodism, some supporting an oscillatory clock and others an 'hourglass', and gave rise to the often stated opinion that the two forms of time measurement were mutually exclusive. This paper, however, suggests that both oscillatory and hourglass-like properties are to be seen in insect photoperiodism. Furthermore, the differences between the two apparently conflicting models may be resolved if, following Bünning, 'hourglasses' are regarded as damping circadian oscillators, with the more self-sustained (clearly oscillatory) and more highly damped (hourglass-like) responses being parts of a continuous series. Since circadian rhythmicity is an all-pervading and fundamental aspect of insect biology, currently opening up to genetic and molecular analysis, recognition of the basic similarity of a wide range of insect photoperiodic timers may help to unravel the biochemical nature of the mechanism(s) involved.