Parietal lobe mechanisms for directed visual attention

Abstract

1. Experiments were made on the cortex of the inferior parietal lobule in 10 hemispheres of six alert, behaving monkeys. The electrical signs of the impulse discharges of single cortical cells were recorded as the monkeys executed tasks requiring them to fixate stationary visual targets, track those which moved slowly, and to make saccadic movements to foveate those which suddenly jumped from one locus to another within the field of view. A total of 907 neurons of area 7 were identified in terms of their physiological properties, particularly the correlation of their activity with the oculomotor components of these behavioral acts of directed visual attention; 480 of these were located by cytoarchitectural layer. Most identifiable cells of area 7 are visuomotor neurons, in a special and conditional sense. Their discharge frequencies increase before and during those steady fixations and movements of the eyes which secure and maintain foveation of objects, but only if the visual targets engaged are linked by a strong motivational drive; in our experiments, one between thirst and the light whose dimming the animal has learned to detect for liquid reward. We have identified and studied three major classes of neurons in area 7. 2. The visual fixation neurons (57%) accelerate discharge synchronously with fixation of a visual object the animal desires. The incremented discharge continues until reward, but then declines abruptly even when there is no immediate shift of the line of gaze. Fixation neurons are relatively inactive during those casual fixations by which the animal insepcts the surrounding environment. Mist fixation neurons subtend gaze fields limited to one quadrant or half of the total gaze field. The sum of the gaze fields of the fixation neurons in one hemisphere is weighted moderately toward the contralateral side. Fixation cells also discharge during slow pursuit movements in any direction so long as the movement stays within the gaze field of the neuron under study. About 40% of fixation cells are suppressed before and during saccadic movements of the eyes to a new target within the gaze field of the fixation cell. Those suppressed are located preferentially in layer V of the cortex. Suppression is maximal for saccades directed contralaterally to the hemisphere under study. 3. Visual tracking neurons are active during oculomotor pursuit of slowly moving visual objects, not during steady fixations. They show a marked directional but no laterality relation, and are suppressed before and during a visually evoked saccade superimposed on the smooth pursuit movement. The rate of discharge is a flat function of tracking speed so that these cells do not appear to emit signals which specify the speed of smooth pursuit movements. 4. The saccade neurons are active before and during visually evoked saccadic movements of the eyes but not before spontaneous saccades, no matter whether made in light or near darkness. The discharge of saccade neurons leads the eye movement by as much as 150 ms (mean, 73 ms)...