The cortical column: A structure without a function

Philosophical Transactions of the Royal Society B: Biological Sciences

Volumn 360, Issue 1456, 2005, Pages 837-862

  Horton, Jonathan G ^a   Adams, Daniel L ^a  

^a UNIVERSITY OF CALIFORNIA   (United States)

Author keywords

Angioscotoma; Barrel; Macaque; Pinwheel; Retinal wave; Spandrel

Indexed keywords

BRAIN; PRIMATE;

MACACA;

EID: 21244487876     PISSN: 09628436     EISSN: None     Source Type: Journal    
DOI: 10.1098/rstb.2005.1623     Document Type: Review

References (220)

1. Adams, D. L. & Horten, J. C. 2002 Shadows cast by retinal blood vessels mapped in primary visual cortex. Science 298, 572-576.
2. Adams, D. L. & Horton, J. C. 2003a Capricious expression of cortical columns in the primate brain. Nat. Neurosci. 6, 113-114.
3. Adams, D. L. & Horton, J. C. 2003b The representation of retinal blood vessels in primate striate cortex. J. Neurosci. 23, 5984-5997.
4. Adams, D. L. & Zeki, S. 2001 Functional organization of macaque V3 for stereoscopic depth. J. Neurophysiol. 86, 2195-2203.
5. Albright, T. D., Desimone, R. & Gross, C. G. 1984 Columnar organization of directionally selective cells in visual area MT of the macaque. J. Neurophysiol. 51, 16-31.
6. Albus, K. 1975 A quantitative study of the projection area of the central and the paracentral visual field in area 17 of the cat. II. The spatial organization of the orientation domain. Exp. Brain Res. 24, 181-202.
7. Amirikian, B. & Georgopoulos, A. P. 2003 Modular organization of directionally tuned cells in the motor cortex: is there a short-range order? Proc. Natl Acad. Sci. USA 100, 12474-12479.
8. Angelucci, A., Levitt, J. B., Walton, E. J., Hupe, J. M., Bullier, J. & Lund, J. S. 2002 Circuits for local and global signal integration in primary visual cortex. J. Neurosci. 22, 8633-8646.
9. Angevine, J. & Sidman, R. 1961 Autoradiographic study of cell migration during histogenesis of cerebral cortex in the mouse. Nature 192, 766-768.
10. Anthony, T. E., Klein, C., Fishell, G. & Heintz, N. 2004 Radial glia serve as neuronal progenitors in all regions of the central nervous system. Neuron 41, 881-890.
11. Bartfeld, E. & Grinvald, A. 1992 Relationships between orientation-preference pinwheels, cytochrome oxidase blobs, and ocular-dominance columns in primate striate cortex. Proc. Natl Acad. Sci. USA 89, 11905-11909.
12. Basole, A., White, L. E. & Fitzpatrick, D. 2003 Mapping multiple features in the population response of visual cortex. Nature 423, 986-990.
13. Bear, M. F. & Rittenhouse, C. D. 1999 Molecular basis for induction of ocular dominance plasticity. J. Neurobiol. 41, 83-91.
14. Blasdel, G. G. 1992a Differential imaging of ocular dominance and orientation selectivity in monkey striate cortex. J. Neurosci. 12, 3115-3138.
15. Blasdel, G. G. 1992b Orientation selectivity, preference, and continuity in monkey striate cortex. J. Neurosci. 12, 3139-3161.
16. Blasdel, G. & Campbell, D. 2001 Functional retinotopy of monkey visual cortex. J. Neurosci. 21, 8286-8301.
17. Blasdel, G. G. & Salama, G. 1986 Voltage-sensitive dyes reveal a modular organization in monkey striate cortex. Nature 321, 579-585.
18. Bonhoeffer, T. & Grinvald, A. 1991 Iso-orientation domains in cat visual cortex are arranged in pinwheel-like patterns. Nature 353, 429-431.
19. Born, R. T. & Tootell, R. B. 1992 Segregation of global and local motion processing in primate middle temporal visual area. Nature 357, 497-499.
20. Bosking, W. H., Zhang, Y., Schofield, B. & Fitzpatrick, D. 1997 Orientation selectivity and the arrangement of horizontal connections in tree shrew striate cortex. J. Neurosci. 17, 2112-2127.
21. Boyd, J. D. & Casagrande, V. A. 1999 Relationships between cytochrome oxidase (CO) blobs in primate primary visual cortex (V1) and the distribution of neurons projecting to the middle temporal area (MT). J. Comp. Neurol. 409, 573-591.
22. Boyd, J. D. & Matsubara, J. A. 1996 Laminar and columnar patterns of geniculocortical projections in the cat: relationship to cytochrome oxidase. J. Comp. Neurol. 365, 659-682.
23. Bugbee, N. M. & Goldman-Rakic, P. S. 1983 Columnar organization of corticocortical projections in squirrel and rhesus monkeys: similarity of column width in species differing in cortical volume. J. Comp. Neurol. 220, 355-364.
24. Bureau, I., Shepherd, G. M. & Svoboda, K. 2004 Precise development of functional and anatomical columns in the neocortex. Neuron 42, 789-801.
25. Buxhoeveden, D. P. & Casanova, M. F. 2002 The mini-column hypothesis in neuroscience. Brain 125, 935-951.
26. Callaway, E. M. 1998 Local circuits in primary visual cortex of the macaque monkey. Annu. Rev. Neurosci. 21, 47-74.
27. Carder, R. K. 1997 Immunocytochemical characterization of AMPA-selective glutamate receptor subunits: laminar and compartmental distribution in macaque striate cortex. J. Neurosci. 17, 3352-3363.
28. Carder, R. K. & Hendry, S. H. 1994 Neuronal characterization, compartmental distribution, and activity-dependent regulation of glutamate immunoreactivity in adult monkey striate cortex. J. Neurosci. 14, 242-262.
29. Casagrande, V. A. & Harting, J. K. 1975 Transneuronal transport of tritiated fucose and proline in the visual pathways of tree shrew Tupaia glis. Brain Res. 96, 367-372.
30. Cases, O., Vitalis, T., Seif, I., De Maeyer, E., Sotelo, C. & Gaspar, P. 1996 Lack of barrels in the somatosensory cortex of monoamine oxidase A-deficient mice: role of a serotonin excess during the critical period. Neuron 16, 297-307.
31. Catania, K. C. & Kaas, J. H. 1995 Organization of the somatosensory cortex of the star-nosed mole. J. Comp. Neurol. 351, 549-567.
32. Celio, M. R., Scharer, L., Morrison, J. H., Norman, A. W. & Bloom, F. E. 1986 Calbindin immunoreactivity alternates with cytochrome c-oxidase-rich zones in some layers of the primate visual cortex. Nature 323, 715-717.
33. Chatterjee, S. & Callaway, E. M. 2003 Parallel colour-opponent pathways to primary visual cortex. Nature 426, 668-671.
34. Chaudhuri, A. & Cynader, M. S. 1993 Activity-dependent expression of the transcription factor Zif268 reveals ocular dominance columns in monkey visual cortex. Brain Res. 605, 349-353.
35. Chow, K. L., Masland, R. H. & Stewart, D. L. 1971 Receptive field characteristics of striate cortical neurons in the rabbit. Brain Res. 33, 337-352.
36. Constantine-Paton, M. & Law, M. I. 1978 Eye-specific termination bands in tecta of three-eyed frogs. Science 202, 639-641.
37. Crair, M. C., Ruthazer, E. S., Gillespie, D. S. & Stryker, M. P. 1997 Ocular dominance peaks at pinwheel center singularities of the orientation map in cat visual cortex. J. Neurophysiol. 77, 3381-3385.
38. Crair, M. C., Horton, J. C., Antonini, A. & Stryker, M. P. 2001 Emergence of ocular dominance columns in cat visual cortex by 2 weeks of age. J. Comp. Neurol. 430, 235-249.
39. Crowley, J. C. & Katz, L. C. 1999 Development of ocular dominance columns in the absence of retinal input. Nat. Neurosci. 2, 1125-1130.
40. Crowley, J. C. & Katz, L. C. 2000 Early development of ocular dominance columns. Science 290, 1321-1324.
41. DeAngelis, G. C. & Newsome, W. T. 1999 Organization of disparity-selective neurons in macaque area MT. J. Neurosci. 19, 1398-1415.
42. DeAngelis, G. C., Ghose, G. M., Ohzawa, I. & Freeman, R. D. 1999 Functional micro-organization of primary visual cortex: receptive field analysis of nearby neurons. J. Neurosci. 19, 4046-4064.
43. DeBruyn, E. J., Casagrande, V. A., Beck, P. D. & Bonds, A. B. 1993 Visual resolution and sensitivity of single cells in the primary visual cortex (V1) of a nocturnal primate (Bush Baby): correlations with cortical layers and cytochrome oxidase patterns. J. Neurophysiol. 69, 3-18.
44. DeYoe, E. A. & Van Essen, D. C. 1985 Segregation of efferent connections and receptive field properties in visual area V2 of the macaque. Nature 317, 58-61.
45. DeYoe, E. A., Felleman, D. J., Van Essen, D. C. & McClendon, E. 1994 Multiple processing streams in occipitotemporal visual cortex. Nature 371, 151-154.
46. Douglas, R. & Martin, K. 2004 Neuronal circuits of the neocortex. Annu. Rev. Neurosci. 27, 419-451.
47. Drager, U. C. 1975 Receptive fields of single cells and topography in mouse visual cortex. J. Comp. Neurol. 160, 269-290.
48. Dufly, K. R. & Livingstone, M. S. 2003 Distribution of non-phosphorylated neurofilament in squirrel monkey V1 is complementary to the pattern of cytochrome-oxidase blobs. Cereb. Cortex 13, 722-727.
49. Dyck, R. H., Chaudhuri, A. & Cynader, M. S. 2003 Experience-dependent regulation of the zincergic innervation of visual cortex in adult monkeys. Cereb. Cortex 13, 1094-1109.
50. Everson, R. M., Prashanth, A. K., Gabbay, M., Knight, B. W., Sirovich, L. & Kaplan, E. 1998 Representation of spatial frequency and orientation in the visual cortex. Proc. Natl Acad. Sci. USA 95, 8334-8338.
51. Favorov, O. V., Diamond, M. E. & Whitsel, B. L. 1987 Evidence for a mosaic representation of the body surface in area 3b of the somatic cortex of cat. Proc. Natl Acad. Sci. USA 84, 6606-6610.
52. Felleman, D. J., Burkhalter, A. & Van Essen, D. C. 1997 Cortical connections of areas V3 and VP of macaque monkey extrastriate visual cortex. J. Comp. Neurol. 379, 21-47.
53. Feller, M. B., Wellis, D. P., Stellwagen, D., Werblin, F. S. & Shatz, C. J. 1996 Requirement for cholinergic synaptic transmission in the propagation of spontaneous retinal waves. Science 112, 1182-1187.
54. Ferster, D. 1981 A comparison of binocular depth mechanisms in areas 17 and 18 of the cat visual cortex. J. Physiol. (Lond.) 311, 623-655.
55. Fonta, C., Chappert, C. & Imbert, M. 1997 N-Methyl-D-aspartate subunit R1 involvement in the postnatal organization of the primary visual cortex of Callithrix jacchus. J. Comp. Neurol. 386, 260-276.
56. Fonta, C., Chappert, C. & Imbert, M. 2000 Effect of monocular deprivation on NMDAR1 immunostaining in ocular dominance columns of the marmoset Callithrix jacchus. Vis. Neurosci. 17, 345-352.
57. Fujita, I., Tanaka, K., Ito, M. & Cheng, K. 1992 Columns for visual features of objects in monkey inferotemporal cortex. Nature 360, 343-346.
58. Galuske, R. A., Schlote, W., Bratzke, H. & Singer, W. 2000 Interhemispheric asymmetries of the modular structure in human temporal cortex. Science 289, 1946-1949.
59. Gardner, J. C. & Raiten, E. J. 1986 Ocular dominance and disparity-sensitivity: why there are cells in the visual cortex driven unequally by the two eyes. Exp. Brain Res. 64, 505-514.
60. Gegenfurtner, K. R., Kiper, D. C. & Fenstemaker, S. B. 1996 Processing of color, form, and motion in macaque area V2. Vis. Neurosci. 13, 161-172.
61. Gilbert, C. D. 1977 Laminar differences in receptive field properties of cells in cat primary visual cortex. J. Physiol. (Lond.) 268, 391-421.
62. Gilbert, C. D. & Wiesel, T. N. 1989 Columnar specificity of intrinsic horizontal and corticocortical connections in cat visual cortex. J. Neurosci. 9, 2432-2442.
63. Girman, S. V., Sauve, Y. & Lund, R. D. 1999 Receptive field properties of single neurons in rat primary visual cortex. J. Neurophysiol. 82, 301-311.
64. Goldman, P. S. & Nauta, W. J. 1977 Columnar distribution of cortico-cortical fibers in the frontal association, limbic, and motor cortex of the developing rhesus monkey. Brain Res. 122, 393-413.
65. Goto, S. & Singer, W. 1994 Laminar and columnar organization of immunoreactivity for calcineurin, a calcium- and calmodulin-regulated protein phosphatase, in monkey striate cortex. Cereb. Cortex 4, 636-645.
66. Gould, S. J. 1997 The exaptive excellence of spandrels as a term and prototype. Proc. Natl Acad. Sci. USA 94, 10750-10755.
67. Haseltine, E. C., DeBruyn, E. J. & Casagrande, V. A. 1979 Demonstration of ocular dominance columns in Nissl-stained sections of monkey visual cortex following enucleation. Brain Res. 176, 153-158.
68. Hendrickson, A. E., Wilson, J. R. & Ogren, M. P. 1978 The neuroanatomical organization of pathways between the dorsal lateral geniculate nucleus and visual cortex in Old World and New World primates. J. Comp. Neurol. 182, 123-136.
69. Hendrickson, A. E., Hunt, S. P. & Wu, J. Y. 1981 Immunocytochemical localization of glutamic acid decarboxylase in monkey striate cortex. Nature 292, 605-607.
70. Hendry, S. H. & Bhandari, M. A. 1992 Neuronal organization and plasticity in adult monkey visual cortex: immunoreactivity for microtubule-associated protein 2. Vis. Neurosci. 9, 445-459.
71. Hendry, S. H. & Calkins, D. J. 1998 Neuronal chemistry and functional organization in the primate visual system. Trends Neurosci. 21, 344-349.
72. Hendry, S. H. & Yoshioka, T. 1994 A neurochemically distinct third channel in the macaque dorsal lateral geniculate nucleus. Science 264, 575-577.
73. Hendry, S. H., Hockfield, S., Jones, E. G. & McKay, R. 1984 Monoclonal antibody that identifies subsets of neurones in the central visual system of monkey and cat. Nature 307, 267-269.
74. Hendry, S. H., Huntsman, M. M., Viñuela, A., Möhler, H., de Blas, A. L. & Jones, E. G. 1994 GABAA receptor subunit immunoreactivity in primate visual cortex: distribution in macaques and humans and regulation by visual input in adulthood. J. Neurosci. 14, 2383-2401.
75. Hetherington, P. A. & Swindale, N. V. 1999 Receptive field and orientation scatter studied by tetrode recordings in cat area 17. Vis. Neurosci. 16, 637-652.
76. Horton, J. C. 1984 Cytochrome oxidase patches: a new cytoarchitectonic feature of monkey visual cortex. Phil. Trans. R. Soc. B 304, 199-253.
77. Horton, J. C. & Hedley-Whyte, E. T. 1984 Mapping of cytochrome oxidase patches and ocular dominance columns in human visual cortex. Phil. Trans. R. Soc. B 304, 255-272.
78. Horton, J. C. & Hocking, D. R. 1996a Anatomical demonstration of ocular dominance columns in striate cortex of the squirrel monkey. J. Neurosci. 16, 5510-5522.
79. Horton, J. C. & Hocking, D. R. 1996b Intrinsic variability of ocular dominance column periodicity in normal macaque monkeys. J. Neurosci. 16, 7228-7239.
80. Horton, J. C. & Hocking, D. R. 1998 Monocular core zones and binocular border strips in primate striate cortex revealed by the contrasting effects of enucleation, eyelid suture, and retinal laser lesions on cytochrome oxidase activity. J. Neurosci. 18, 5433-5455.
81. Horton, J. C. & Hubel, D. H. 1981 Regular patchy distribution of cytochrome oxidase staining in primary visual cortex of macaque monkey. Nature 292, 762-764.
82. Horton, J. C., Hocking, D. R. & Adams, D. L. 2000 Rapid identification of ocular dominance columns in macaques using cytochrome oxidase, Zif268, and dark-field microscopy. Vis. Neurosci. 17, 495-508.
83. Hubel, D. H. 1975 An autoradiographic study of the retino-cortical projections in the tree shrew (Tupaia glis). Brain Res. 96, 41-50.
84. Hubel, D. H. 1982 Exploration of the primary visual cortex, 1955-78. Nature 299, 515-524.
85. Hubel, D. H. & Livingstone, M. S. 1987 Segregation of form, color, and stereopsis in primate area 18. J. Neurosci. 7, 3378-3415.
86. Hubel, D. H. & Wiesel, T. N. 1962 Receptive fields, binocular interaction and functional architecture in the cat's visual cortex. J. Physiol. (Lond.) 160, 106-154.
87. Hubel, D. H. & Wiesel, T. N. 1963 Shape and arrangement of columns in cat's striate cortex. J. Physiol. (Lond.) 165, 559-568.
88. Hubel, D. H. & Wiesel, T. N. 1968 Receptive fields and functional architecture of monkey striate cortex. J. Physiol. (Lond.) 195, 215-243.
89. Hubel, D. H. & Wiesel, T. N. 1969 Anatomical demonstration of columns in the monkey striate cortex. Nature 221, 747-750.
90. Hubel, D. H. & Wiesel, T. N. 1972 Laminar and columnar distribution of geniculo-cortical fibers in the macaque monkey. J. Comp. Neurol. 146, 421-450.
91. Hubel, D. H. & Wiesel, T. N. 1974a Sequence regularity and geometry of orientation columns in the monkey striate cortex. J. Comp. Neurol. 158, 267-293.
92. Hubel, D. H. & Wiesel, T. N. 1974b Uniformity of monkey striate cortex: a parallel relationship between field size, scatter, and magnification factor. J. Comp. Neurol. 158, 295-305.
93. Hubel, D. H. & Wiesel, T. N. 1977 The Ferrier lecture: functional architecture of macaque monkey visual cortex. Proc. R. Soc. B 198, 1-59.
94. Hubel, D. H. & Wiesel, T. N. 1979 Brain mechanisms of vision. Sci. Am. 241, 150-162.
95. Hubel, D. H., LeVay, S. & Wiesel, T. N. 1975 Mode of termination of retinotectal fibers in macaque monkey: an autoradiographic study. Brain Res. 96, 25-40.
96. Hubel, D. H., Wiesel, T. N. & LeVay, S. 1976 Functional architecture of area 17 in normal and monocularly deprived macaque monkeys. Cold Spring Harb. Symp. Quant. Biol. 40, 581-589.
97. Hübener, M., Shoham, D., Grinvald, A. & Bonhoeffer, T. 1997 Spatial relationships among three columnar systems in cat area 17. J. Neurosci. 17, 9270-9284.
98. Huberman, A. D., Wang, G. Y., Liets, L. C., Collins, O. A., Chapman, B. & Chalupa, L. M. 2003 Eye-specific retinogeniculate segregation independent of normal neuronal activity. Science 300, 994-998.
99. Humphrey, A. L., Skeen, L. C. & Norton, T. T. 1980 Topographic organization of the orientation column system in the striate cortex of the tree shrew (Tupaia glis). II. Deoxyglucose mapping. J. Comp. Neurol. 192, 549-566.
100. Imig, T. J. & Adrian, H. O. 1977 Binaural columns in the primary field (A1) of cat auditory cortex. Brain Res. 138, 241-257.
101. Imig, T. J. & Reale, R. A. 1981 Ipsilateral corticocortical projections related to binaural columns in cat primary auditory cortex. J. Comp. Neurol. 203, 1-14.
102. Issa, N. P., Trepel, C. & Stryker, M. P. 2000 Spatial frequency maps in cat visual cortex. J. Neurosci. 20, 8504-8514.
103. Jacob, F. 1977 Evolution and tinkering. Science 196, 1161-1166.
104. Jain, N., Catania, K. C. & Kaas, J. H. 1998 A histologically visible representation of the fingers and palm in primate area 3b and its immutability following long-term deafferentations. Cereb. Cortex 8, 227-236.
105. Jones, E. G., Burton, H. & Porter, R. 1975 Commissural and cortico-cortical "columns" in the somatic sensory cortex of primates. Science 190, 572-574.
106. Kaas, J. H., Krubitzer, L. A. & Johanson, K. L. 1989 Cortical connections of areas 17 (V-I) and 18 (V-II) of squirrels. J. Comp. Neurol. 281, 426-446.
107. Kaschube, M., Wolf, F., Geisel, T. & Löwel, S. 2002 Genetic influence on quantitative features of neocortical architecture. J. Neurosci. 22, 7206-7217.
108. Kaschube, M., Wolf, F., Puhlmann, M., Rathjen, S., Schmidt, K. F., Geisel, T. & Lowel, S. 2003 The pattern of ocular dominance columns in cat primary visual cortex: intra- and interindividual variability of column spacing and its dependence on genetic background. Eur. J. Neurosci. 18, 3251-3266.
109. Katz, L. C. & Crowley, J. C. 2002 Development of cortical circuits: lessons from ocular dominance columns. Nat. Rev. Neurosci. 3, 34-42.
110. Katz, L. C. & Shatz, C. J. 1996 Synaptic activity and the construction of cortical circuits. Science 274, 1133-1138.
111. Kiper, D. C., Fenstemaker, S. B. & Gegenfurtner, K. R. 1997 Chromatic properties of neurons in macaque area V2. Vis. Neurosci. 14, 1061-1072.
112. Krubitzer, L. A. & Kaas, J. H. 1990 Cortical connections of MT in four species of primates: areal, modular, and retinotopic patterns. Vis. Neurosci. 5, 165-204.
113. Lachica, E. A., Beck, P. D. & Casagrande, V. A. 1992 Parallel pathways in macaque monkey striate cortex: anatomically defined columns in layer III. Proc. Natl Acad. Sci. USA 89, 3566-3570.
114. Land, P. W., Buffer, S. A., Jr & Yaskosky, J. D. 1995 Barreloids in adult rat thalamus: three-dimensional architecture and relationship to somatosensory cortical barrels. J. Comp. Neurol. 355, 573-588.
115. Landisman, C. E. & Ts'o, D. Y. 2002 Color processing in macaque striate cortex: relationships to ocular dominance, cytochrome oxidase, and orientation. J. Neurophysiol. 87, 3126-3137.
116. Lennie, P., Krauskopf, J. & Sclar, G. 1990 Chromatic mechanisms in striate cortex of macaque. J. Neurosci. 10, 649-669.
117. LeVay, S. & Nelson, S. B. 1991 Columnar organization of the visual cortex. In The neural basis of visual function (ed. A. G. Leventhal), pp. 266-315. Boston: CRC Press.
118. LeVay, S. & Voigt, T. 1988 Ocular dominance and disparity coding in cat visual cortex. Vis. Neurosci. 1, 395-414.
119. LeVay, S., Hubel, D. H. & Wiesel, T. N. 1975 The pattern of ocular dominance columns in macaque visual cortex revealed by a reduced silver stain. J. Comp. Neurol. 159, 559-575.
120. LeVay, S., Stryker, M. P. & Shatz, C. J. 1978 Ocular dominance columns and their development in layer IV of the cat's visual cortex: a quantitative study. J. Comp. Neurol. 179, 223-244.
121. LeVay, S., Connolly, M., Houde, J. & Van Essen, D. C. 1985 The complete pattern of ocular dominance stripes in the striate cortex and visual field of the macaque monkey. J. Neurosci. 5, 486-501.
122. Leventhal, A. G., Thompson, K. G., Du, D., Zhou, Y. & Ault, S. J. 1995 Concomitant sensitivity to orientation, direction, and color of cells in layers 2, 3 and 4 of monkey striate cortex. J. Neurosci. 15, 1808-1818.
123. Levitt, J. B., Kiper, D. C. & Movshon, J. A. 1994 Receptive fields and functional architecture of macaque V2. J. Neurophysiol. 71, 2517-2542.
124. Levitt, J. B., Yoshioka, T. & Lund, J. S. 1995 Connections between the pulvinar complex and cytochrome oxidase-defined compartments in visual area V2 of macaque monkey. Exp. Brain Res. 104, 419-430.
125. Lia, B. & Olavarria, J. F. 1996 The distribution of corticotectal projection neurons correlates with the interblob compartment in macaque striate cortex. Vis. Neurosci. 13, 461-466.
126. Livingstone, M. S. 1996 Ocular dominance columns in New World monkeys. J. Neurosci. 16, 2086-2096.
127. Livingstone, M. S. & Hubel, D. H. 1984a Anatomy and physiology of a color system in the primate visual cortex. J. Neurosci. 4, 309-356.
128. Livingstone, M. S. & Hubel, D. H. 1984b Specificity of intrinsic connections in primate primary visual cortex. J. Neurosci. 4, 2830-2835.
129. Livingstone, M. S., Nori, S., Freeman, D. C. & Hubel, D. H. 1995 Stereopsis and binocularity in the squirrel monkey. Vision Res. 35, 345-354.
130. Lorento De Nó, R. 1922 La corteza cerebral del raton. Trab. Lab. Invest. Biol. (Madrid) 20, 41-78.
131. Lorento De Nó, R. 1949 Cerebral cortex: architecture, intracortical connections, motor projections. In Physiology of The nervous system (ed. J. Fulton), pp. 288-330. New York: Oxford University Press.
132. Lübke, J., Egger, V., Sakmann, B. & Feldmeyer, D. 2000 Columnar organization of dendrites and axons of single and synaptically coupled excitatory spiny neurons in layer 4 of the rat barrel cortex. J. Neurosci. 20, 5300-5311.
133. Lund, J. S., Angelucci, A. & Bressloff, P. C. 2003 Anatomical substrates for functional columns in macaque monkey primary visual cortex. Cereb. Cortex 13, 15-24.
134. Malach, R., Amir, Y., Harel, M. & Grinvald, A. 1993 Relationship between intrinsic connections and functional architecture revealed by optical imaging and in vivo targeted biocytin injections in primate striate cortex. Proc. Natl Acad. Sci. USA 90, 10469-10473.
135. Malach, R., Schirman, T. D., Harel, M., Tootell, R. B. & Malonek, D. 1997 Organization of intrinsic connections in owl monkey area MT. Cereb. Cortex 7, 386-393.
136. Maldonado, P. E., Godecke, I., Gray, C. M. & Bonhoeffer, T. 1997 Orientation selectivity in pinwheel centers in cat striate cortex. Science 276, 1551-1555.
137. Marin, O. & Rubenstein, J. L. 2003 Cell migration in the forebrain. Annu. Rev. Neurosci. 26, 441-483.
138. Martinez-Millán, L. & Holländer, H. 1975 Cortico-cortical projections from striate cortex of the squirrel monkey (Saimiri sciureus). A radioautographic study. Brain Res. 83, 405-417.
139. Meister, M., Wong, R. O. L., Baylor, D. A. & Shatz, C. J. 1991 Synchronous bursts of action potentials in ganglion cells of the developing mammalian retina. Science 252, 939-943.
140. Miller, K. D., Keller, J. B. & Stryker, M. P. 1989 Ocular dominance column development: analysis and simulation. Science 245, 605-615.
141. Mooney, R., Penn, A. A., Gallego, R. & Shatz, C. J. 1996 Thalamic relay of spontaneous retinal activity prior to vision. Neuron 17, 863-874.
142. Mountcastle, V. B. 1957 Modality and topographic properties of cat's somatic sensory cortex. J. Neurophysiol 20, 408-434.
143. Mountcastle, V. 1978 An organizing principle for cerebral function: the unit module and the distributed system. In The mindful brain (ed. G. M. Edelman & V. B. Mountcastle), pp. 7-50. Massachusetts: MIT Press.
144. Mountcastle, V. B. 1997 The columnar organization of the neocortex. Brain 120, 701-722.
145. Mountcastle, V. B., Berman, A. L. & Davies, P. W. 1955 Topographic organization and modality representation in first somatic area of cat's cerebral cortex by method of single unit analysis. Am. J. Physiol. 183, 464.
146. Mountcastle, V., Davies, P. & Berman, A. 1957 Response properties of neurons of cat's somatic sensory cortex to peripheral stimuli. J. Neurophysiol. 20, 374-407.
147. Murphy, E. H. & Berman, N. 1979 The rabbit and the cat: a comparison of some features of response properties of single cells in the primary visual cortex. J. Comp. Neurol. 188, 401-427.
148. Murphy, K. M., Jones, D. G. & Van Sluyters, R. C. 1995 Cytochrome-oxidase blobs in cat primary visual cortex. J. Neurosci. 15, 4196-4208.
149. Nelson, S. 2002 Cortical microcircuits: diverse or canonical. Neuron 36, 19-27.
150. Noctor, S. C., Flint, A. C., Weissman, T. A., Dammerman, R. S. & Kriegstein, A. R. 2001 Neurons derived from radial glial cells establish radial units in neocortex. Nature 409, 714-720.
151. Ohki, K., Chung, S., Ch'ng, Y. H., Kara, P. & Reid, R. C. 2005 Functional imaging with cellular resolution reveals precise micro-architecture in visual cortex. Nature 433, 597-603.
152. O'Keefe, L. P., Levitt, J. B., Kiper, D. C., Shapley, R. M. & Movshon, J. A. 1998 Functional organization of owl monkey lateral geniculate nucleus and visual cortex. J. Neurophysiol. 80, 594-609.
153. Olavarria, J. F. & Van Essen, D. C. 1997 The global pattern of cytochrome oxidase stripes in visual area V2 of the macaque monkey. Cereb. Cortex 7, 395-404.
154. O'Rourke, N. A., Sullivan, D. P., Kaznowski, C. E., Jacobs, A. A. & McConnell, S. K. 1995 Tangential migration of neurons in the developing cerebral cortex. Development 121, 2165-2176.
155. Peterhans, E. & von der Heydt, R. 1993 Functional organization of area V2 in the alert macaque. Eur. J. Neurosci. 5, 509-524.
156. Poggio, G. F. & Fischer, B. 1977 Binocular interaction and depth sensitivity in striate and prestriate cortex of behaving rhesus monkey. J. Neurophysiol. 40, 1392-1405.
157. Poggio, G. F., Gonzalez, F. & Krause, F. 1988 Stereoscopic mechanisms in monkey visual cortex: binocular correlation and disparity selectivity. J. Neurosci. 8, 4531-4550.
158. Polimeni, J. R., Granquist-Fraser, D., Wood, R. J. & Schwartz, E. L. 2005 Physical limits to spatial resolution of optical recording: clarifying the spatial structure of cortical hypercolumns. Proc. Natl Acad. Sci. USA 102, 4158-4163.
159. Powell, T. P. S. & Mountcastle, V. B. 1959 Some aspects of the functional organization of the cortex of the postcentral gyrus of the monkey: a correlation of findings obtained in a single unit analysis with cytoarchitecture. Bull. Johns Hopkins Hosp. 105, 133-162.
160. Purves, D., Riddle, D. R. & LaMantia, A. S. 1992 Iterated patterns of brain circuitry (or how the cortex gets its spots). Trends Neurosci. 15, 362-368.
161. Rakic, P. 1971 Guidance of neurons migrating to the fetal monkey neocortex. Brain Res. 33, 471-476.
162. Rakic, P. 1977 Prenatal development of the visual system in rhesus monkey. Phil. Trans. R. Soc. B 278, 245-260.
163. Rakic, P. 1988 Specification of cerebral cortical areas. Science 241, 170-176.
164. Rakic, P. 1995 Radial versus tangential migration of neuronal clones in the developing cerebral cortex. Proc. Natl Acad. Sci. USA 92, 11323-11327.
165. Rakic, P. 2003 Developmental and evolutionary adaptations of cortical radial glia. Cereb. Cortex 13, 541-549.
166. Rathjen, S., Schmidt, K. E. & Lowel, S. 2002 Two-dimensional analysis of the spacing of ocular dominance columns in normally raised and strabismic kittens. Exp. Brain Res. 145, 158-165.
167. Read, J. C. & Cumming, B. G. 2004 Ocular dominance predicts neither strength nor class of disparity selectivity with random-dot stimuli in primate V1. J. Neurophysiol. 91, 1271-1281.
168. Read, H. L., Winer, J. A. & Schreiner, C. E. 2001 Modular organization of intrinsic connections associated with spectral tuning in cat auditory cortex. Proc. Natl Acad. Sci. USA 98, 8042-8047.
169. Reid, C. B., Liang, I. & Walsh, C. 1995 Systematic widespread clonal organization in cerebral cortex. Neuron 15, 299-310.
170. Reser, D. H., Fishman, Y. I., Arezzo, J. C. & Steinschneider, M. 2000 Binaural interactions in primary auditory cortex of the awake macaque. Cereb. Cortex 10, 574-584.
171. Ringach, D. L., Shapley, R. M. & Hawken, M. J. 2002 Orientation selectivity in macaque V1: diversity and laminar dependence. J. Neurosci. 22, 5639-5651.
172. Rockel, A. J., Hiorns, R. W. & Powell, T. P. 1980 The basic uniformity in structure of the neocortex. Brain 103, 221-244.
173. Rockland, K. S. & Lund, J. S. 1983 Intrinsic laminar lattice connections in primate visual cortex. J. Comp. Neurol. 216, 303-318.
174. Roe, A. W. & Ts'o, D. Y. 1995 Visual topography in primate V2: multiple representation across functional stripes. J. Neurosci. 15, 3689-3715.
175. Rosenthal, F., Woodbury, J. W. & Patton, H. D. 1966 Dipole characteristics of pyramidal cell activity in cat postcruciate cortex. J. Neurophysiol. 29, 612-625.
176. Sandell, J. H. 1986 NADPH diaphorase histochemistry in the macaque striate cortex. J. Comp. Neurol. 251, 388-397.
177. Shaw, C., Yinon, U. & Auerbach, E. 1975 Receptive fields and response properties of neurons in the rat visual cortex. Vision Res. 15, 203-208.
178. Shipp, S. & Zeki, S. 1985 Segregation of pathways leading from area V2 to areas V4 and V5 of macaque monkey visual cortex. Nature 315, 322-325.
179. Shipp, S. & Zeki, S. 1989 The organization of connections between areas V5 and V2 in macaque monkey visual cortex. Eur. J. Neurosci. 1, 333-354.
180. Shipp, S. & Zeki, S. 2002 The functional organization of area V2. I. Specialization across stripes and layers. Vis. Neurosci. 19, 187-210.
181. Sincich, L. C. & Blasdel, G. G. 2001 Oriented axon projections in primary visual cortex of the monkey. J. Neurosci. 21, 4416-4426.
182. Sincich, L. C. & Horton, J. C. 2002a Divided by cytochrome oxidase: a map of the projections from V1 to V2 in macaques. Science 295, 1734-1737.
183. Sincich, L. C. & Horton, J. C. 2002b Pale cytochrome oxidase stripes in V2 receive the richest projection from macaque striate cortex. J. Comp. Neurol. 447, 18-33.
184. Sincich, L. C. & Horton, J. C. 2003a An albino-like decussation error in the optic chiasm revealed by anomalous ocular dominance columns. Vis. Neurosci. 19, 541-545.
185. Sincich, L. C. & Horton, J. C. 2003b Independent projection streams from macaque striate cortex to the second visual area and middle temporal area. J. Neurosci. 23, 5684-5692.
186. Sincich, L. C., Adams, D. L. & Horton, J. C. 2003 Complete flatmounting of the macaque cerebral cortex. Vis. Neurosci. 20, 663-686.
187. Slimp, J. C. & Towe, A. L. 1990 Spatial distribution of modalities and receptive fields in sensorimotor cortex of awake cats. Exp. Neurol. 107, 78-96.
188. Sperry, R. W., Miner, N. & Myers, R. E. 1955 Visual pattern perception following subpial slicing and tantalum wire implantations in the visual cortex. J. Comp. Physiol. Psychol. 48, 50-58.
189. Stosiek, C., Garaschuk, O., Holthoff, K. & Konnerth, A. 2003 In vivo two-photon calcium imaging of neuronal networks. Proc. Natl Acad. Sci. USA 100, 7319-7324.
190. Stryker, M. P. & Harris, W. A. 1986 Binocular impulse blockade prevents the formation of ocular dominance columns in cat visual cortex. J. Neurosci. 6, 2117-2133.
191. Swindale, N. V. 1990 Is the cerebral cortex modular? Trends Neurosci. 13, 487-492.
192. Swindale, N. V., Shoham, D., Grinvald, A., Bonhoeffer, T. & Hubener, M. 2000 Visual cortex maps are optimized for uniform coverage. Nat. Neurosci. 3, 822-826.
193. Tamamaki, N., Nakamura, K., Okamoto, K. & Kaneko, T. 2001 Radial glia is a progenitor of neocortical neurons in the developing cerebral cortex. Neurosci. Res. 41, 51-60.
194. Tamura, H., Sato, H., Katsuyama, N., Hata, Y. & Tsumoto, T. 1996 Less segregated processing of visual information in V2 than in V1 of the monkey visual cortex. Eur. J. Neurosci. 8, 300-309.
195. Tiao, Y. C. & Blakemore, C. 1976 Functional organization in the visual cortex of the golden hamster. J. Comp. Neurol. 168, 459-481.
196. Tootell, R. B. H., Silverman, M. S., De Valois, R. L. & Jacobs, G. H. 1983 Functional organization of the second cortical visual area in primates.
197. Tootell, R. B., Hamilton, S. L. & Silverman, M. S. 1985 Topography of cytochrome oxidase activity in owl monkey cortex. J. Neurosci. 5, 2786-2800.
198. Ts'o, D. Y. & Gilbert, C. D. 1988 The organization of chromatic and spatial interactions in the primate striate cortex. J. Neurosci. 8, 1712-1727.
199. Ts'o, D. Y., Roe, A. W. & Gilbert, C. D. 2001a A hierarchy of the functional organization for color, form and disparity in primate visual area V2. Vision Res. 41, 1333-1349.
200. Ts'o, D. Y., Roe, A. W. & Gilbert, C. D. 2001b A hierarchy of the functional organization for color, form and disparity in primate visual area V2. Vision Res. 41, 1333-1349.
201. Van Hooser, S. D., Heimel, J. A. F., Chung, S., Nelson, S. B. & Toth, L. J. 2005 Orientation selectivity without orientation maps in visual cortex of a highly visual mammal. J. Neurosci. 25, 19-28.
202. Walsh, C. & Cepko, C. L. 1988 Clonally related cortical cells show several migration patterns. Science 241, 1342-1345.
203. Walsh, C. & Cepko, C. L. 1992 Widespread dispersion of neuronal clones across functional regions of the cerebral cortex. Science 255, 434-440.
204. Wang, G., Tanifuji, M. & Tanaka, K. 1998 Functional architecture in monkey inferotemporal cortex revealed by in vivo optical imaging. Neurosci. Res. 32, 33-46.
205. Weliky, M., Bosking, W. H. & Fitzpatrick, D. 1996 A systemic map of direction preference in primary visual cortex. Nature 379, 725-728.
206. Welker, E., Armstrong-James, M., Bronchti, G., Ourednik, W., Gheorghita-Baechler, F., Dubois, R., Guernsey, D. L., Van der Loos, H. & Neumann, P. E. 1996 Altered sensory processing in the somatosensory cortex of the mouse mutant barrelless. Science 271, 1864-1867.
207. Wiesenfeld, Z. & Kornel, E. E. 1975 Receptive fields of single cells in the visual cortex of the hooded rat. Brain Res. 94, 401-412.
208. Wong-Riley, M. T. T. 1979 Columnar cortico-cortical interconnections within the visual system of the squirrel and macaque monkeys. Brain Res. 162, 201-217.
209. Wong-Riley, M. T. 1989 Cytochrome oxidase: an endogenous metabolic marker for neuronal activity. Trends Neurosci. 12, 94-101.
210. Wong-Riley, M. & Carroll, E. W. 1984 Effect of impulse blockage on cytochrome oxidase activity in monkey visual system. Nature 307, 262-264.
211. Wong-Riley, M., Anderson, B., Liebl, W. & Huang, Z. 1998 Neurochemical organization of the macaque striate cortex: correlation of cytochrome oxidase with Na + K+ ATPase, NADPH-diaphorase, nitric oxide synthase, and N-Methyl-D-aspartate receptor subunit 1. Neuroscience 83, 1025-1045.
212. Woolsey, T. A. & Van Der Loos, H. 1970 The structural organization of layer IV in the somatosensory region (S1) of mouse cerebral cortex: the description of a cortical field composed of cytoarchitectonic units. Brain Res. 17, 205-242.
213. Woolsey, T. A., Welker, C. & Schwartz, R. H. 1975 Comparative anatomical studies of the SmL face cortex with special reference to the occurrence of "barrels" in layer IV. J. Comp. Neurol. 164, 79-94.
214. Xiao, Y., Wang, Y. & Felleman, D. J. 2003 A spatially organized representation of colour in macaque cortical area V2. Nature 421, 535-539.
215. Xu, X., Bosking, W., Sary, G., Stefansic, J., Shima, D. & Casagrande, V. 2004 Functional organization of visual cortex in the owl monkey. J. Neurosci. 24, 6237-6247.
216. Yabuta, N. H. & Callaway, E. M. 1998 Functional streams and local connections of layer 4C neurons in primary visual cortex of the macaque monkey. J. Neurosci. 18, 9489-9499.
217. Yoshioka, T. & Dow, B. M. 1996 Color, orientation and cytochrome oxidase reactivity in areas V1, V2 and V4 of macaque monkey visual cortex. Behav. Brain Res. 76, 71-88.
218. Yoshioka, T., Levitt, J. B. & Lund, J. S. 1994 Independence and merger of thalamocortical channels within macaque monkey primary visual cortex: anatomy of interlaminar projections. Vis. Neurosci. 11, 467-489.
219. Yoshioka, T., Blasdel, G. G., Levitt, J. B. & Lund, J. S. 1996 Relation between patterns of intrinsic lateral connectivity, ocular dominance, and cytochrome oxidase-reactive regions in macaque monkey striate cortex. Cereb. Cortex 6, 297-310.
220. Zeki, S. & Shipp, S. 1989 Modular connections between areas V2 and V4 of macaque monkey visual cortex. Eur. J. Neurosci. 1, 494-506.