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Philos Trans R Soc Lond B Biol Sci. Jul 29, 1999; 354(1387): 1371–1382.
PMCID: PMC1692626

The clinical and functional measurement of cortical (in)activity in the visual brain, with special reference to the two subdivisions (V4 and V4 alpha) of the human colour centre.


We argue below that, at least in studying the visual brain, the old and simple methods of detailed clinical assessment and perimetric measurement still yield important insights into the organization of the visual brain as a whole, as well as the organization of the individual areas within it. To demonstrate our point, we rely especially on the motion and colour systems, emphasizing in particular how clinical observations predicted an important feature of the organization of the colour centre in the human brain. With the use of data from functional magnetic resonance imaging analysed by statistical parametric mapping and independent component analysis, we show that the colour centre is composed of two subdivisions, V4 and V4 alpha the two together constituting the V4 complex of the human brain. These two subdivisions are intimately linked anatomically and act cooperatively. The new evidence about the architecture of the colour centre might help to explain why the syndrome, cerebral achromatopsia, produced by lesions in it is so variable.

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Selected References

These references are in PubMed. This may not be the complete list of references from this article.
  • Albert ML, Reches A, Silverberg R. Hemianopic colour blindness. J Neurol Neurosurg Psychiatry. 1975 Jun;38(6):546–549. [PMC free article] [PubMed]
  • Bartels A, Zeki S. The theory of multistage integration in the visual brain. Proc Biol Sci. 1998 Dec 7;265(1412):2327–2332. [PMC free article] [PubMed]
  • Beckers G, Zeki S. The consequences of inactivating areas V1 and V5 on visual motion perception. Brain. 1995 Feb;118(Pt 1):49–60. [PubMed]
  • Bell AJ, Sejnowski TJ. An information-maximization approach to blind separation and blind deconvolution. Neural Comput. 1995 Nov;7(6):1129–1159. [PubMed]
  • BORNSTEIN B, KIDRON DP. Prosopagnosia. J Neurol Neurosurg Psychiatry. 1959 May;22(2):124–131. [PMC free article] [PubMed]
  • Büchel C, Coull JT, Friston KJ. The predictive value of changes in effective connectivity for human learning. Science. 1999 Mar 5;283(5407):1538–1541. [PubMed]
  • Corbetta M, Miezin FM, Dobmeyer S, Shulman GL, Petersen SE. Selective and divided attention during visual discriminations of shape, color, and speed: functional anatomy by positron emission tomography. J Neurosci. 1991 Aug;11(8):2383–2402. [PubMed]
  • Cragg BG. The topography of the afferent projections in the circumstriate visual cortex of the monkey studied by the Nauta method. Vision Res. 1969 Jul;9(7):733–747. [PubMed]
  • Crick F, Koch C. Are we aware of neural activity in primary visual cortex? Nature. 1995 May 11;375(6527):121–123. [PubMed]
  • Damasio A, Yamada T, Damasio H, Corbett J, McKee J. Central achromatopsia: behavioral, anatomic, and physiologic aspects. Neurology. 1980 Oct;30(10):1064–1071. [PubMed]
  • DANIEL PM, WHITTERIDGE D. The representation of the visual field on the cerebral cortex in monkeys. J Physiol. 1961 Dec;159:203–221. [PMC free article] [PubMed]
  • DeYoe EA, Van Essen DC. Concurrent processing streams in monkey visual cortex. Trends Neurosci. 1988 May;11(5):219–226. [PubMed]
  • DeYoe EA, Carman GJ, Bandettini P, Glickman S, Wieser J, Cox R, Miller D, Neitz J. Mapping striate and extrastriate visual areas in human cerebral cortex. Proc Natl Acad Sci U S A. 1996 Mar 19;93(6):2382–2386. [PMC free article] [PubMed]
  • Engel SA, Rumelhart DE, Wandell BA, Lee AT, Glover GH, Chichilnisky EJ, Shadlen MN. fMRI of human visual cortex. Nature. 1994 Jun 16;369(6481):525–525. [PubMed]
  • ffytche DH, Guy CN, Zeki S. The parallel visual motion inputs into areas V1 and V5 of human cerebral cortex. Brain. 1995 Dec;118(Pt 6):1375–1394. [PubMed]
  • Friston KJ, Holmes AP, Poline JB, Grasby PJ, Williams SC, Frackowiak RS, Turner R. Analysis of fMRI time-series revisited. Neuroimage. 1995 Mar;2(1):45–53. [PubMed]
  • Girard P, Salin PA, Bullier J. Response selectivity of neurons in area MT of the macaque monkey during reversible inactivation of area V1. J Neurophysiol. 1992 Jun;67(6):1437–1446. [PubMed]
  • Hadjikhani N, Liu AK, Dale AM, Cavanagh P, Tootell RB. Retinotopy and color sensitivity in human visual cortical area V8. Nat Neurosci. 1998 Jul;1(3):235–241. [PubMed]
  • Heywood C, Cowey A. With color in mind. Nat Neurosci. 1998 Jul;1(3):171–173. [PubMed]
  • Holmes G. DISTURBANCES OF VISION BY CEREBRAL LESIONS. Br J Ophthalmol. 1918 Jul;2(7):353–384. [PMC free article] [PubMed]
  • Horton JC, Hoyt WF. Quadrantic visual field defects. A hallmark of lesions in extrastriate (V2/V3) cortex. Brain. 1991 Aug;114(Pt 4):1703–1718. [PubMed]
  • Hubel DH, Livingstone MS. Segregation of form, color, and stereopsis in primate area 18. J Neurosci. 1987 Nov;7(11):3378–3415. [PubMed]
  • Humphrey GK, Goodale MA, Corbetta M, Aglioti S. The McCollough effect reveals orientation discrimination in a case of cortical blindness. Curr Biol. 1995 May 1;5(5):545–551. [PubMed]
  • Jaeger W, Krastel H, Braun S. Cerebrale Achromatopsie (Symptomatik, Verlauf, Differentialdiagnose und Strategie der Untersuchung). Teil II. Klin Monbl Augenheilkd. 1989 Jan;194(1):32–36. [PubMed]
  • Kastner S, De Weerd P, Desimone R, Ungerleider LG. Mechanisms of directed attention in the human extrastriate cortex as revealed by functional MRI. Science. 1998 Oct 2;282(5386):108–111. [PubMed]
  • Kennard C, Lawden M, Morland AB, Ruddock KH. Colour identification and colour constancy are impaired in a patient with incomplete achromatopsia associated with prestriate cortical lesions. Proc Biol Sci. 1995 May 22;260(1358):169–175. [PubMed]
  • Kulikowski JJ, Walsh V, McKeefry D, Butler SR, Carden D. The electrophysiological basis of colour processing in macaques with V4 lesions. Behav Brain Res. 1994 Jan 31;60(1):73–78. [PubMed]
  • Lapresle J, Metreau R, Annabi A. Transient achromatopsia in vertebrobasilar insufficiency. J Neurol. 1977 May 13;215(2):155–158. [PubMed]
  • Lueck CJ, Zeki S, Friston KJ, Deiber MP, Cope P, Cunningham VJ, Lammertsma AA, Kennard C, Frackowiak RS. The colour centre in the cerebral cortex of man. Nature. 1989 Aug 3;340(6232):386–389. [PubMed]
  • McKeefry DJ, Zeki S. The position and topography of the human colour centre as revealed by functional magnetic resonance imaging. Brain. 1997 Dec;120(Pt 12):2229–2242. [PubMed]
  • McKeown MJ, Makeig S, Brown GG, Jung TP, Kindermann SS, Bell AJ, Sejnowski TJ. Analysis of fMRI data by blind separation into independent spatial components. Hum Brain Mapp. 1998;6(3):160–188. [PubMed]
  • Nakamura H, Gattass R, Desimone R, Ungerleider LG. The modular organization of projections from areas V1 and V2 to areas V4 and TEO in macaques. J Neurosci. 1993 Sep;13(9):3681–3691. [PubMed]
  • Ogden JA. Visual object agnosia, prosopagnosia, achromatopsia, loss of visual imagery, and autobiographical amnesia following recovery from cortical blindness: case M.H. Neuropsychologia. 1993 Jun;31(6):571–589. [PubMed]
  • Paulson HL, Galetta SL, Grossman M, Alavi A. Hemiachromatopsia of unilateral occipitotemporal infarcts. Am J Ophthalmol. 1994 Oct 15;118(4):518–523. [PubMed]
  • Pearlman AL, Birch J, Meadows JC. Cerebral color blindness: an acquired defect in hue discrimination. Ann Neurol. 1979 Mar;5(3):253–261. [PubMed]
  • Rizzo M, Smith V, Pokorny J, Damasio AR. Color perception profiles in central achromatopsia. Neurology. 1993 May;43(5):995–1001. [PubMed]
  • Rockland KS, Saleem KS, Tanaka K. Divergent feedback connections from areas V4 and TEO in the macaque. Vis Neurosci. 1994 May-Jun;11(3):579–600. [PubMed]
  • Rodman HR, Gross CG, Albright TD. Afferent basis of visual response properties in area MT of the macaque. I. Effects of striate cortex removal. J Neurosci. 1989 Jun;9(6):2033–2050. [PubMed]
  • Sereno MI, Dale AM, Reppas JB, Kwong KK, Belliveau JW, Brady TJ, Rosen BR, Tootell RB. Borders of multiple visual areas in humans revealed by functional magnetic resonance imaging. Science. 1995 May 12;268(5212):889–893. [PubMed]
  • Shipp S, Zeki S. The Organization of Connections between Areas V5 and V2 in Macaque Monkey Visual Cortex. Eur J Neurosci. 1989;1(4):333–354. [PubMed]
  • Zeki S, McKeefry DJ, Bartels A, Frackowiak RS. Has a new color area been discovered? Nat Neurosci. 1998 Sep;1(5):335–336. [PubMed]
  • Tootell RB, Dale AM, Sereno MI, Malach R. New images from human visual cortex. Trends Neurosci. 1996 Nov;19(11):481–489. [PubMed]
  • Tootell RB, Mendola JD, Hadjikhani NK, Ledden PJ, Liu AK, Reppas JB, Sereno MI, Dale AM. Functional analysis of V3A and related areas in human visual cortex. J Neurosci. 1997 Sep 15;17(18):7060–7078. [PubMed]
  • Vaina LM. Functional segregation of color and motion processing in the human visual cortex: clinical evidence. Cereb Cortex. 1994 Sep-Oct;4(5):555–572. [PubMed]
  • Essen DC, Zeki SM. The topographic organization of rhesus monkey prestriate cortex. J Physiol. 1978 Apr;277:193–226. [PMC free article] [PubMed]
  • Walsh V, Carden D, Butler SR, Kulikowski JJ. The effects of V4 lesions on the visual abilities of macaques: hue discrimination and colour constancy. Behav Brain Res. 1993 Feb 26;53(1-2):51–62. [PubMed]
  • Young RS, Fishman GA, Chen F. Traumatically acquired color vision defect. Invest Ophthalmol Vis Sci. 1980 May;19(5):545–549. [PubMed]
  • Zeki SM. Representation of central visual fields in prestriate cortex of monkey. Brain Res. 1969 Jul;14(2):271–291. [PubMed]
  • Zeki SM. Uniformity and diversity of structure and function in rhesus monkey prestriate visual cortex. J Physiol. 1978 Apr;277:273–290. [PMC free article] [PubMed]
  • Zeki S. Colour coding in the cerebral cortex: the reaction of cells in monkey visual cortex to wavelengths and colours. Neuroscience. 1983 Aug;9(4):741–765. [PubMed]
  • Zeki S. A century of cerebral achromatopsia. Brain. 1990 Dec;113(Pt 6):1721–1777. [PubMed]
  • Zeki S. Cerebral akinetopsia (visual motion blindness). A review. Brain. 1991 Apr;114(Pt 2):811–824. [PubMed]
  • Zeki S. The mystery of Louis Verrey (1854-1916). Gesnerus. 1993;50(Pt 1-2):96–112. [PubMed]
  • Zeki S, Ffytche DH. The Riddoch syndrome: insights into the neurobiology of conscious vision. Brain. 1998 Jan;121(Pt 1):25–45. [PubMed]
  • Zeki S, Shipp S. The functional logic of cortical connections. Nature. 1988 Sep 22;335(6188):311–317. [PubMed]
  • Zeki S, Shipp S. Modular Connections between Areas V2 and V4 of Macaque Monkey Visual Cortex. Eur J Neurosci. 1989;1(5):494–506. [PubMed]
  • Zeki S, Watson JD, Lueck CJ, Friston KJ, Kennard C, Frackowiak RS. A direct demonstration of functional specialization in human visual cortex. J Neurosci. 1991 Mar;11(3):641–649. [PubMed]
  • Zeki S, McKeefry DJ, Bartels A, Frackowiak RS. Has a new color area been discovered? Nat Neurosci. 1998 Sep;1(5):335–336. [PubMed]
  • Zheng D, LaMantia AS, Purves D. Specialized vascularization of the primate visual cortex. J Neurosci. 1991 Aug;11(8):2622–2629. [PubMed]

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