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Laboratorio de Neurociencia, Facultad de Biología, Universidad de Sevilla, 41012-Sevilla, Spain
Domingo, José A., Agnes Gruart, and José M. Delgado-García. Quantal organization of reflex and conditioned eyelid responses. J. Neurophysiol. 78: 2518-2530, 1997. Upper lid movements and the electromyographic activity of the orbicularis oculi muscle were recorded in behaving cats during spontaneous and experimentally evoked reflex blinks, and conditioned eyelid responses. Reflex blinks evoked by the presentation of air puffs, flashes, or tones consisted of a fast downward lid movement followed by late, small downward waves, recurring at 
50-ms intervals. The latency, maximum amplitude, peak velocity, and number of late waves depended on the modality, intensity, and duration of the evoking stimulus. The power spectra of acceleration records indicated a dominant frequency of 20 Hz for air puff-evoked blinks. Flashes and tones usually evoked small and easily fatigable reflex responses of lower dominant frequencies (14-17 and 9-11 Hz, respectively). A basic 20-Hz oscillation was also noticed during lid fixation, and ramplike lid displacements evoked by optokinetic stimuli. Five classical conditioning paradigms were used to analyze the frequency-domain properties of conditioned eyelid responses. These learned lid movements differed in latency, maximum amplitude, and profile smoothness depending on the modality (air puff, tone), intensity (weak, strong), and presentation site (ipsi-, contralateral to the unconditioned stimulus) of the conditioned stimulus. It was found that the characteristic ramplike profile of a conditioned response was not smooth, but appeared to be formed by a succession of small waves at a dominant frequency of 20 Hz. The amplitude (and number) of the constituting waves depended on the characteristics of the conditioned stimulus and on the time interval until unconditioned stimulus presentation. Thus conditioned responses seemed to be formed from lid displacements of 2-6° in amplitude and 50 ms in duration, which increased in number throughout conditioning sessions, until a complete (i.e., lid closing) conditioned response was reached. It is suggested that a 20-Hz oscillator underlies the generation of reflex and conditioned eyelid responses. The oscillator is susceptible to being neurally modulated to modify the velocity of a given quantum of movement, and the total duration of the lid response. Learned eyelid movements are probably the result of a successively longer release of the oscillator as a function of the temporal-spatial needs of the motor response.
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