GENERAL METHODSSubjects Male albino rats, descendants of the Sprague-Dawley strain and weighing 250-350 g, were Ss. Individual Ss, 40 in all, were sometimes used in more than one experiment. Miller, 1967), the rat has not been used in studies of eyelid responses. The size of this species and its reluctance to accept restraint have made the measurement of eyelid responses difficult. But it seemed that if this difficulty could be overcome, all of those attributes that make the rat such a good laboratory S would make it most valuable to a long-term research program requiring many animals and experiments of many kinds. The difficulties have not been overcome completely, but the results below are encouraging in this respect and bear on several problems in the study of eyelid responses. Eyelid responses in the rat were recorded as electromyographic activity (EMG) or as movements detected by a photoelectric technique. Spontaneous blinks, startle responses to acoustic stimuli, unconditioned responses to air puffs, and conditioned responses based on the corneal reflex were recorded under a variety of conditions. Special attention was given to topographical and temporal characteristics of the responses. Some relationships between the EMG and eyelid movements are described, as well as changes in eyelid responses related to habituation, conditioning, and various stimulus manipulations. In addition to the substantive findings, the study suggests that the rat may yet become a useful S in the study of eyelid conditioning.One attractive feature of eyelid reflexes that has gone almost unnoticed is a relative simplicity that might well recommend them for analysis at the neuronal level. Eyelid responses are accomplished primarily by two small muscles, orbicularis oculi and levator palpebrae superioris, and they are to a considerable degree independent of other behavior. Although clearly influenced by eye and head movements, they seem less entangled in most postural and movement activities than responses of the skeletal musculature mediated by the spinal cord. Woody and his associates (Engle & Woody, 1972;Woody, 1970; Woody & Brozek, 1969a, b, c; Woody, Vassilevsky, & Engel, 1970; Woody & Yarowsky, 1972), with their studies of the glabella reflex in cats, have been the first to recognize these advantages of eyelid reflexes and the possibilities they afford in electrophysiological inquiries into the neural substrate of conditioning. The experiments on the rat described in this paper were undertaken as part of a program initiated with the same intent, to use eyelid reflexes to study neural events underlying conditioned responses.Except for a few early conditioning studies (Biel & Wickens, 1941; Hughes & Schlosberg, 1938)
Measurement of EyelidResponsesThe eyelid electromyogram (EMG) was recorded from two stainless steel Teflon-insulated stranded wires, 125 /Jm in diam, implanted subcutaneously in the upper lid. The EMG was filtered to eliminate potential changes below 10 Hz. It was also integrated, following full-wave rectificat...