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Journal of Neuroscience, Vol 4, 309-356, Copyright © 1984 by Society for Neuroscience
Anatomy and physiology of a color system in the primate visual cortex
MS Livingstone and DH Hubel
Staining for the mitochondrial enzyme cytochrome oxidase reveals an array
of dense regions (blobs) in the primate primary visual cortex. They are
most obvious in the upper layers, 2 and 3, but can also be seen in layers
4B, 5, and 6, in register with the blobs in layers 2 and 3. We compared
cells inside and outside blobs in macaque and squirrel monkeys, looking at
their physiological responses and anatomical connections. Cells within
blobs did not show orientation selectivity, whereas cells between blobs
were highly orientation selective. Receptive fields of blob cells had
circular symmetry and were of three main types, Broad-Band Center-Surround,
Red-Green Double-Opponent, and Yellow-Blue Double-Opponent. Double-Opponent
cells responded poorly or not at all to white light in any form, or to
diffuse light at any wavelength. In contrast to blob cells, none of the
cells recorded in layer 4C beta were Double-Opponent: like the majority of
cells in the parvocellular geniculate layers, they were either Broad-Band
or Color- Opponent Center-Surround, e.g., red-on-center green-off-surround.
To our surprise cells in layer 4C alpha were orientation selective. In
tangential penetrations throughout layers 2 and 3, optium orientation, when
plotted against electrode position, formed long, regular, usually linear
sequences, which were interrupted but not perturbed by the blobs. Staining
area 18 for cytochrome oxidase reveals a series of alternating wide and
narrow dense stripes, separated by paler interstripes. After small
injections of horseradish peroxidase into area 18, we saw a precise set of
connections from the blobs in area 17 to thin stripes in area 18, and from
the interblob regions in area 17 to interstripes in area 18. Specific
reciprocal connections also ran from thin stripes to blobs and from
interstripes to interblobs. We have not yet determined the area 17
connections to thick stripes in area 18. In addition, within area 18 there
are stripe-to-stripe and interstripe- to-interstripe intrinsic connections.
These results suggest that a system involved in the processing of color
information, especially color-spatial interactions, runs parallel to and
separate from the orientation-specific system. Color, encoded in three
coordinates by the major blob cell types, red-green, yellow-blue, and
black-white, can be transformed into the three coordinates, red, green, and
blue, of the Retinex algorithm of Land.
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