The discharge profiles of identified abducens, accessory abducens, and orbicularis oculi motoneurons have been recorded extra- and intracellularly in alert behaving cats during spontaneous, reflexively evoked, and classically conditioned eyelid responses. The movement of the upper lid and the electromyographic activity of the orbicularis oculi muscle also were recorded. Animals were conditioned by short, weak air puffs or 350-ms tones as conditioned stimuli (CS) and long, strong air puffs as unconditioned stimulus (US) using both trace and delayed conditioning paradigms. Motoneurons were identified by antidromic activation from their respective cranial nerves. Orbicularis oculi and accessory abducens motoneurons fired an early, double burst of action potentials (at 4-6 and 10-16 ms) in response to air puffs or to the electrical stimulation of the supraorbital nerve. Orbicularis oculi, but not accessory abducens, motoneurons fired in response to flash and tone presentations. Only 10-15% of recorded abducens motoneurons fired a late, weak burst after air puff, supraorbital nerve, and flash stimulations. Spontaneous fasciculations of the orbicularis oculi muscle and the activity of single orbicularis oculi motoneurons that generated them also were recorded. The activation of orbicularis oculi motoneurons during the acquisition of classically conditioned eyelid responses happened in a gradual, sequential manner. Initially, some putative excitatory synaptic potentials were observed in the time window corresponding to the CS-US interval; by the second to the fourth conditioning session, some isolated action potentials appeared that increased in number until some small movements were noticed in eyelid position traces. No accessory abducens motoneuron fired and no abducens motoneuron modified their discharge rate for conditioned eyelid responses. The firing of orbicularis oculi motoneurons was related linearly to lid velocity during reflex blinks but to lid position during conditioned responses, a fact indicating the different neural origin and coding of both types of motor commands. The power spectra of both reflex and conditioned lid responses showed a dominant peak at approximately 20 Hz. The wavy appearance of both reflex and conditioned eyelid responses was clearly the result of the high phasic activity of orbicularis oculi motor units. Orbicularis oculi motoneuron membrane potentials oscillated at approximately 20 Hz after supraorbital nerve stimulation and during other reflex and conditioned eyelid movements. The oscillation seemed to be the result of both intrinsic (spike afterhyperpolarization lasting approximately 50 ms, and late depolarizations) and extrinsic properties of the motoneuronal pool and of the circuits involved in eye blinks.

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The eyelid motor system represents an excellent experimental model for the study of reflex and learned motor responses. Eyelid responses can be recorded quantitatively with the search coil in a magnetic-field technique. Stimuli able to evoke reflex blinks (air puffs, flashes of light, tones) can also be controlled quantitatively. Eyelid movements can be classified as spontaneous, passive (such as those following eye saccades), reflex, and acquired with classical conditioning procedures. Information is available regarding the firing activity of brainstem motoneuronal pools (abducens, accessory abducens, and facial motoneurons) involved in these types of eyelid response. In particular, facial motoneurons present different encoding properties for the generation of reflex against learned eyelid responses. In cats, accessory abducens motoneurons are involved only in reflex (but not in learned) blinks. The recent description of the complete organization of premotoneuronal pathways related to eyelid motorics opens new experimental possibilities for the study of this particular motor system.

Reflexively evoked and eye‐related eyelid responses were recorded using the search coil in a magnetic field technique in alert cats. The downward phase of a blink was a large (up to 21 deg), fast (up to 2000 deg s−1) eyelid displacement in the closing direction, with an almost fixed rise time duration (15‐20 ms); its maximum velocity was achieved in ≈10 ms. Upward eyelid motion was separated into two phases. The first phase consisted of a fast eyelid displacement, with a short duration (≈30 ms) and a maximum velocity up to 900 deg s−1. The second phase had an exponential‐like form, lasting for 200–400 ms, and a maximum velocity ranging between 30 and 250 deg s−1. Maximum blink velocity in the downward direction was linearly related to maximum velocity of the first upward phase. The first phase in the upward direction was never observed if the eyelid stayed closed for a long period (> 50 ms) or moved slowly in the closing direction before it started to open. In these two cases, the upswing motion of the blink reflex contained only the exponential‐like movement characteristic of the second upward phase, and maximum velocity in the downward direction was not related to that of the eyelid upward displacement. Mean duration of eyelid downward saccades was ≈130 ms, and their peak velocities ranged between 50 and 440 ms. A physiological model is presented explaining the active and passive forces involved in both reflex and saccadic eyelid responses. A second‐order system seems to be appropriate to describe the postulated biomechanical model.

Harmaline's effects on reflex and classically conditioned eyelid responses and on tremor picked up by a coil attached to the back were measured in alert cats. Harmaline at a dose of 10 mg/kg produced skeletal muscle tremogenic effects that lasted 4h. Back movements presented a tremor-like displacement with a frequency peak at 10 Hz, but lid responses oscillated as in controls, at 20 Hz during both reflex and conditioned eyelid movements, with no increase in oscillation amplitude or frequency. The learning curves of harmaline-injected animals remained as in controls, but eyelid conditioned responses showed longer latencies, and smaller amplitude and peak velocity. Reflex and already-learned eyelid responses were not modified by harmaline. These results imply that neuronal control systems for skeletal-motor and facial responses are differentially affected by harmaline.