Temperature-induced pupil movements in insect superposition eyes

In this paper, we describe the hitherto largely overlooked effect of temperature on the pupil of insect compound eyes. In the turnip moth Agrotis segetum and in two other nocturnal insects with superposition eyes, the lacewing Euroleon nostras and the codling moth Cydia pomonella, the pupil not only opens and closes with changes in the ambient light level, as expected, but also with changes in temperature in the absence of light. In complete darkness, the pupil of A. segetum responds over a wide range of temperatures, with the pupillary pigments migrating to a light-adapted position when the animal is exposed to either low or high temperatures. At temperatures between 21.0 and 22.7 C, the pigments migrate to the fully dark-adapted position, resulting in an open pupil and maximal eye glow. Pupil closure at high temperatures shows two distinct thresholds: the first at 23.8+/-0.7 C and a second some degrees higher at 25.7+/-1.2 C (means +/- s.d., N=10). Temperatures exceeding the first threshold (the activation temperature, T(a)) initiate a closure of the pupil that is completed when the temperature exceeds the second threshold (the closure temperature, T(c)), which causes rapid and complete migration of pigment to the light-adapted position. All temperatures above T(a) affect the pupil, but only temperatures exceeding T(c) result in complete closure. Temperatures between T(a) and T(c) cause a slow, partial and rather unpredictable closure. The lacewing and the codling moth both show very similar responses to those of A. segetum, suggesting that this response to temperature is widespread in superposition eyes. The possibility that the ambient temperature could be used to pre-adapt the eye to different light intensities is discussed.