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Cerebral Cortex Advance Access originally published online on June 4, 2007
Cerebral Cortex 2008 18(3):477-499; doi:10.1093/cercor/bhm061
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Published by Oxford University Press 2007.
This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/2.0/uk/) which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.

Cortical Connections of Area V4 in the Macaque

Leslie G. Ungerleider1, Thelma W. Galkin2, Robert Desimone2,3 and Ricardo Gattass4

1 Laboratory of Brain and Cognition, 2 Laboratory of Neuropsychology, National Institute of Mental Health, National Institutes of Health, Department of Health and Human Services, Bethesda, MD 20892 USA, 3 McGovern Institute, MIT, Cambridge, MA, USA, 4 Instituto de Biofísica Carlos Chagas Filho, UFRJ, Rio de Janeiro RJ 21941-941, Brazil

Address correspondence to Dr Leslie G. Ungerleider, Laboratory of Brain and Cognition, NIMH, NIH, Building 10, Room 4C104, Bethesda, MD 20892-1366, USA. Email: ungerlel{at}mail.nih.gov.

To determine the locus, full extent, and topographic organization of cortical connections of area V4 (visual area 4), we injected anterograde and retrograde tracers under electrophysiological guidance into 21 sites in 9 macaques. Injection sites included representations ranging from central to far peripheral eccentricities in the upper and lower fields. Our results indicated that all parts of V4 are connected with occipital areas V2 (visual area 2), V3 (visual area 3), and V3A (visual complex V3, part A), superior temporal areas V4t (V4 transition zone), MT (medial temporal area), and FST (fundus of the superior temporal sulcus [STS] area), inferior temporal areas TEO (cytoarchitectonic area TEO in posterior inferior temporal cortex) and TE (cytoarchitectonic area TE in anterior temporal cortex), and the frontal eye field (FEF). By contrast, mainly peripheral field representations of V4 are connected with occipitoparietal areas DP (dorsal prelunate area), VIP (ventral intraparietal area), LIP (lateral intraparietal area), PIP (posterior intraparietal area), parieto-occipital area, and MST (medial STS area), and parahippocampal area TF (cytoarchitectonic area TF on the parahippocampal gyrus). Based on the distribution of labeled cells and terminals, projections from V4 to V2 and V3 are feedback, those to V3A, V4t, MT, DP, VIP, PIP, and FEF are the intermediate type, and those to FST, MST, LIP, TEO, TE, and TF are feedforward. Peripheral field projections from V4 to parietal areas could provide a direct route for rapid activation of circuits serving spatial vision and spatial attention. By contrast, the predominance of central field projections from V4 to inferior temporal areas is consistent with the need for detailed form analysis for object vision.

Key Words: extrastriate cortex • inferior temporal cortex • object recognition • primates • visual system


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