Functional magnetic resonance imaging (fMRI) was used to identify and map the representation of the visual field in seven areas of human cerebral cortex and to identify at least two additional visually responsive regions.The cortical locations of neurons responding to stimulation along the vertical or horizontal visual field meridia were charted on three-dimensional models of the cortex and on unfolded maps of the cortical surface. These maps were used to identify the borders among areas that would be topographically homologous to areas Vl, V2, V3, VP, and parts of V3A and V4 of the macaque monkey. Visually responsive areas homologous to the middle temporal/medial superior temporal area complex and unidentified parietal visual areas were also observed. The topography of the visual areas identified thus far is consistent with the organization in macaque monkeys. However, these and other findings suggest that human and simian cortical organization may begin to differ in extrastriate cortex at, or beyond, V3A and V4.In contrast to our extensive knowledge of cortical organization in nonhuman primates, far less is known about the identity and topography of functional subdivisions in human cerebral cortex. In the macaque monkey cortex, 30 or more distinct visual areas have been tentatively identified (1). In the human brain, a strong case has been made for only two or three areas (Vi, V2, and middle temporal) (2, 4-6), though several additional areas have been proposed (6-11). Brain images produced by functional magnetic resonance imaging (fMRI) show signals that are thought to represent local changes in blood oxygenation. These changes can be elicited by sensory-evoked neural activity though other factors such as alterations in blood volume and proton movement may also contribute [for review, see DeYoe et al. (12)]. We have used this technology to chart several functionally distinct visual areas in the human. Preliminary reports of this work have appeared (13,14). MATERIALS AND METHODSTo map angular positions within the visual field, four male and two female subjects (ages 24-40 years old) viewed a flickering black-and-white-checkered hemifield that rotated slowly about a central fixation point during a fMRI scan. To map visual field eccentricity (distance from the center of gaze), subjects viewed an expanding checkered annulus. In this manner, neurons responding to stimulation at different locations in the visual field were activated at different times during the stimulus sequence. Corresponding differences in the temporal phase of the fMRI response thus identified the retinotopic locationThe publication costs of this article were defrayed in part by page charge payment. This article must therefore be hereby marked "advertisement" in accordance with 18 U.S.C. §1734 solely to indicate this fact. 2382represented by each active site in the brain (12). A similar technique has been described by Engel et al. (15).To enhance activation of extrastriate cortex (12) and to help maintain attention and arousal, the subj...

Here we identify a neural correlate of the ability to precisely direct visual attention to locations other than the center of gaze. Human subjects performed a task requiring shifts of visual attention (but not of gaze) from one location to the next within a dense array of targets and distracters while functional MRI was used to map corresponding displacements of neural activation within visual cortex. The cortical topography of the purely attention-driven activity precisely matched the topography of activity evoked by the cued targets when presented in isolation. Such retinotopic mapping of attention-related activation was found in primary visual cortex, as well as in dorsomedial and ventral occipital visual areas previously implicated in processing the attended target features. These results identify a physiological basis for the effects of spatially directed visual attention.

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V2 is a visual area of the macaque monkey which is at the second level in a recently proposed hierarchy of cortical visual areas. Histochemical staining for cytochrome oxidase (CO) in V2 reveals a pattern of alternate thick and thin CO-rich stripes separated by CO-sparse interstripes. These subregions receive distinct inputs from neurones in CO-rich and CO-sparse zones arrayed within the superficial layers of V1 (refs 4, 5). Are output projections from V2 to higher visual areas also segregated? Using an anatomical double-label paradigm, we have now demonstrated that V2 cells projecting to two of its major target areas, MT and V4 (refs 6, 7), are arranged in stripe-like clusters which are largely segregated from one another and which are closely related to the pattern of CO stripes. Concomitant electrophysiological recordings from V2 indicate that groups of cells having similar receptive field properties are clustered within the subregions defined by these anatomical techniques.

Human listeners can effortlessly categorize a wide range of environmental sounds. Whereas categorizing visual object classes (e.g., faces, tools, houses, etc.) preferentially activates different regions of visually sensitive cortex, it is not known whether the auditory system exhibits a similar organization for different types or categories of complex sounds outside of human speech. Using functional magnetic resonance imaging, we show that hearing and correctly or incorrectly categorizing animal vocalizations (as opposed to handmanipulated tool sounds) preferentially activated middle portions of the left and right superior temporal gyri (mSTG). On average, the vocalization sounds had much greater harmonic and phase-coupling content (acoustically similar to human speech sounds), which may represent some of the signal attributes that preferentially activate the mSTG regions. In contrast, correctly categorized tool sounds (and even animal sounds that were miscategorized as being tool-related sounds) preferentially activated a widespread, predominantly left hemisphere cortical "mirror network." This network directly overlapped substantial portions of motor-related cortices that were independently activated when participants pantomimed tool manipulations with their right (dominant) hand. These data suggest that the recognition processing for some sounds involves a causal reasoning mechanism (a high-level auditory "how" pathway), automatically evoked when attending to hand-manipulated tool sounds, that effectively associates the dynamic motor actions likely to have produced the sound(s).