Focal visual attention typically produces enhanced perceptual processi
ng at the psychological level and relatively stronger neural responses
at the physiological level. A longstanding mechanistic question is wh
ether these attentional effects pertain specifically to the attended (
target) object or to the region of space it occupies, We show here tha
t attentional response enhancement in macaque area V4 extends to behav
iorally irrelevant objects in the vicinity of the target object, indic
ating that focal attention has a strong spatial component at the physi
ological level, In addition, we find that spatial attention effects ty
pically show a striking directional asymmetry. The direction of the as
ymmetry varies between cells, so that some cells respond best when att
ention is directed to the left of the stimulus, some when attention is
directed to the right, etc, Thus, attention involves not only enhance
d responses to behavioral targets but also a complex modulation of res
ponses to other stimuli in the surrounding visual space.