Directional selectivity in the responses of units in cat primary visual cortex to passive eye movement.

The responses of single units in the primary visual cortex (Area 17) of anaesthetized, paralysed cats, to passive movement of the ipsilateral eye were studied. Responses to passive eye movement were found in about one-third of the cortical units isolated. Appropriate control experiments excluded visual, auditory and cutaneous inputs as the source of the effective signal during passive eye movement. The magnitudes of the responses to a number (usually four) of radial directions of passive eye movement were estimated from sets of peristimulus time histograms "interleaved" in time. Units were defined as "radially selective" if the responses to movement along one radius (e.g. vertically upwards) exceeded that along at least one other orthogonal radius (e.g. horizontal-temporal). Of 60 units tested, 53 (88%) were "radially selective" according to this definition. Some of the "radially selective" units showed an additional type of specificity to passive eye movement: (a) Some units responded preferentially to movement along one of the arcs of passive eye movement which were tested (e.g. vertical movement above the equator of the orbit). These units we have called "arc selective". (b) Other units were sensitive to the direction of movement and preferred movement in a particular direction over more than one arc (e.g. horizontal movement towards the temporal side in both nasal and temporal halves of the orbit). These we have called "direction selective". Twenty-one "radially selective" units showed one of these additional properties, nine were arc selective and twelve were direction selective. The implications of these results for the understanding of the function of orbital proprioceptive signals in the cortex are discussed briefly. Responses to passive eye movement were found in all of layers II-VI in Area 17 and the implications of this for the understanding of the pathway by which orbital proprioceptive signals reach the primary visual cortex are discussed. The experiments have shown that many units in cat visual cortex respond to passive eye movement and that most of these units have some specificity for particular radial directions of movement while some have additional specific properties. We believe that these properties of radial, directional and arc sensitivity are likely to be important in understanding the function of the orbital proprioceptive signal which arises during eye movement and they are particularly interesting in relation to the findings of others that this proprioceptive signal appears to be concerned in the normal development of visual properties in the cortex and in the control of visually guided movement in adult cats.