Chapter 28 I Perception of Motion, Depth, and Form 549
A 28 Stripes in area 18
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V2 V1
B The Parvocellular and Magnocellular Pathways Feed Into N VISION, AS IN OTHER mental operations, we experi- Two Processing Pathways in Extrastriate Cortex ence the world as a whole. Independent attributes- motion, depth, form, and color-are coordinated Motion Is Analyzed Primarily in the Dorsal Pathway to the I Parietal Cortex into a single visual image. In the two previous chapters Motion Is Represented in the Middle Temporal Area we began to consider how two parallel pathways-the magnocellular and parvocellular pathways, that extend Cells in MT Solve the Aperture Problem from the retina through the lateral geniculate nucleus of Control of Movement Is Selectively Impaired by Lesions the thalamus to the primary visual (striate) cortex- ofMT might produce a coherent visual image. In this chapter Perception of Motion Is Altered by Lesions and we examine how the information from these two path- Microstimulation of MT ways feeds into multiple higher-order centers of visual Depth Vision Depends on Monocular Cues and Binocular processing in the extrastriate cortex. How do these path- Disparity ways contribute to our perception of motion, depth, Monocular Cues Create Far-Field Depth Perception form, and color? The magnocellular (M) and parvocellular (P) path- Stereoscopic Cues Create Near-Field Depth Perception ways feed into two extrastriate cortical pathways: a dor- Figure 28-1 Organization of V1 and V2. B. Connections between V1 and V2. The blobs in V1 connect Information From the Two Eyes Is First Combined in the sal pathway and a ventral pathway. In this chapter we A. Subregions in V1 (area 17) and V2 (area 18). This section primarily to the thin stripes in V2, while the interblobs in V1 Primary Visual Cortex examine, in cell-biological terms, the information pro- from the occipital lobe of a squirrel monkey at the border of ar- connect to interstripes in V2. Layer 48 projects to the thick Random Dot Stereograms Separate Stereopsis From eas 17 and 18 was reacted with cytochrome oxidase. The cy- stripes in V2 and to the middle temporal area (MT). Both thin cessing in each of these pathways. and interstripes project to V4. Thick stripes in V2 also project Object Vision We shall first consider the perception of motion and tochrome oxidase stains the blobs in V1 and the thick and thin stripes in V2. (Courtesy of M. Livingstone.) to MT. Object Vision Depends on the Ventral Pathway to the Inferior depth, mediated in large part by the dorsal pathway to Temporal Lobe the posterior parietal cortex. We then consider the per- Cells in V2 Respond to Both Illusory and Actual ception of contrast and contours, mediated largely by Contours the ventral pathway extending to the inferior temporal The Parvocellular and Magnocellular Pathways dorsal pathway that extends to the posterior parietal Cells in V4 Respond to Form cortex. This pathway also is concerned with the assess- Feed Into Two Processing Pathways cortex. However, the actual relationships are probably Recognition of Faces and Other Complex Forms Depends ment of color, which we will consider in Chapter 29. in Extrastriate Cortex not so exclusive. Upon the Inferior Temporal Cortex Finally, we shall consider the binding problem in the The evidence for separation of function of the dor- Visual Attention Facilitates Coordination Between Separate visual system: how information conveyed in parallel In Chapter 27 we saw that the parallel parvocellular sal and ventral pathways begins in the primary visual, Visual Pathways but separate pathways is brought together into a coher- and magnocellular pathways remain segregated even in or striate, cortex (Vl). Staining for the mitochondrial en- the striate cortex. What happens to these P and M path- zyme cytochrome oxidase reveals a precise and repeat- The Binding Problem in the Visual System ent perception. ways beyond the striate cortex? Early research on these ing pattern of dark, peg-like regions about 0.2 mm in di- An Overall View pathways indicated that the P pathway continues in the ameter called blobs. The blobs are especially prominent ventral cortical pathway that extends to the inferior in the superficial layers 2 and 3, where they are sepa- temporal cortex, and that theM pathway becomes the rated by intervening regions that stain lighter, the in-
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24 24
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E. Chicago: Chicago: evidence evidence and and Scientific Scientific of of W. W. Neurosci Neurosci and and JA. JA. neuronal neuronal posterior posterior in in J J visual visual for for perception perception area area are are The The form, form, of of limited-life- Form Form Organization Organization 16:369--402. 16:369--402. Symp Symp 12:4745-4765. 12:4745-4765. object object 10:363-401. 10:363-401. and and 16:245-263. 16:245-263. Rev Rev 1990. 1990. of of monkey monkey York: York: judgements judgements role role Theory Theory and and cortical cortical in in where where 1967. 1967. with with visual visual Columnar Columnar 11:168-190. 11:168-190. A A motion motion and and processing processing Movshon Movshon stabilization stabilization 1991. 1991. integration integration Perception. Perception. parallel parallel Schneider Schneider WT. WT. New New McClendon McClendon in in 11:454-461. 11:454-461. modularity modularity Annu Annu segmentation segmentation Harbor Harbor JD. JD. mechanisms mechanisms R. R. L, L, 7:3416-3468. 7:3416-3468. Neurosci Neurosci Neurosci Neurosci Neurosci Neurosci 1984. 1984. Neurosci Neurosci Depth, Depth, WT, WT, History, History, 1996. 1996. temporal temporal J J cortex: cortex: occipitotemporal occipitotemporal comparison comparison for for DC, DC, Psychophysical Psychophysical discrimination. discrimination. cells cells How How Visual Visual visual visual cortex cortex perception perception a a feature feature Its Its perceptual perceptual assessed assessed Vision. Vision. Rev Rev in in stereopsis stereopsis Neurosci Neurosci Rev Rev Rev Rev CG. CG. Residual Residual FA. FA. Image Image perception: perception: J J Spring Spring Neural Neural Cyclopean Cyclopean of of the the Neurosci Neurosci 1987. 1987. Essen Essen 1993. 1993. 1987. 1987. Newsome Newsome and and Thomas Thomas Motion, Motion, 346:174-177. 346:174-177. J J depth depth rapid rapid of of Pettigrew Pettigrew Neurosci Neurosci inferior inferior Desimone Desimone Annu Annu Visual Visual 1991. 1991. temporal temporal Murray. Murray. J J Annu Annu Annu Annu Gross Gross of of for for hypothesis. hypothesis. 1993. 1993. Newsome Newsome selective selective Press. Press. 26:1587-1600. 26:1587-1600. Miles Miles motion: motion: 1986. 1986. Cold Cold M, M, JH. JH. visual visual C, C, Van Van patient, patient, DH. DH. J. J. WT. WT. the the streams streams 3:193-204. 3:193-204. R, R, KH, KH, the the 19:109-139. 19:109-139. LG, LG, performance. performance. implications implications Brain, Brain, of of Stereoscope, Stereoscope, influences influences Res Res JD. JD. in in GS, GS, John John TD. TD. DJ, DJ, 1995. 1995. Nature Nature MN, MN, to to stimuli. stimuli. macaques. macaques. 5:129-149. 5:129-149. Zihl Zihl depth. depth. cortex. cortex. visual visual Inferior Inferior Neurophysiol51:16-31. Neurophysiol51:16-31. Inferotemporal Inferotemporal The The binocular binocular Eye, Eye, in in Hubel Hubel 371:151-154. 371:151-154. memory. memory. Mechanisms Mechanisms Chicago Chicago Cortex Cortex J J channels channels dot dot cues cues of of RF, RF, CM. CM. 380:342-345. 380:342-345. Foundations Foundations Perception Perception Britten Britten pathways? pathways? of of and and Maunsell Maunsell control control Vision Vision of of I I Wessinger Wessinger Newsome Newsome Neurosci Neurosci Library. Library. correlation correlation Desimone Desimone inputs inputs Blakemore Blakemore MS, MS, 1993. 1993. Masson Masson directionally directionally processing processing 1856. 1856. London: London: 1988. 1988. Shadlen Shadlen Albright Albright 1996. 1996. 1995. 1995. Pettigrew Pettigrew Felleman Felleman cortex cortex mechanisms. mechanisms. 28 28 meets meets direction. direction. processing: processing: R, R, Hess Hess Y. Y. Neurosci Neurosci Cereb Cereb Gray Gray 193:327-342. 193:327-342. Ungerleider Ungerleider CD, CD, JH, JH, Nature Nature 1971. 1971. of of depth depth C, C, WH, WH, D. D. Rev Rev TD, TD, visual visual K. K. Nature Nature "motion-blind" "motion-blind" HB, HB, random random GF. GF. extrastriate extrastriate vision. vision. GR, GR, analysis analysis PO, PO, 55:963-971. 55:963-971. KH, KH, DH. DH. separate separate B. B. W, W, psychophysical psychophysical JS, JS, EA, EA, macaque. macaque. CL, CL, visual visual a a Cogn Cogn American American for for University University key key ception ception movement, movement, mate mate cortex. cortex. microstimulation microstimulation 18:555-586. 18:555-586. temporal temporal motion motion Annu Annu motion motion J J of of zation zation (Land) (Land) the the in in time time parietal parietal mechanism mechanism Chapter Chapter ular ular and and struction. struction. scopic scopic cortical cortical The The Bioi Bioi eyes. eyes. Attentional Attentional Multiple Multiple cortex. cortex. Hubel Hubel Livingstone Livingstone Miyashita Miyashita Merigan Merigan Julesz Julesz Poggio Poggio Maunsell Maunsell Salzman Salzman References References Singer Singer Stoner Stoner Tanaka Tanaka Baizer Baizer Albright Albright Baker Baker Brewster Brewster Barlow Barlow Bishop Bishop Busettini Busettini Britten Britten Desimone Desimone DeYoe DeYoe is is in in of of in in an an be be re- re- be- the the are are be- the the ab- cor- cor- and and con- both both edge edge Neu- Like- addi- is is path- to to visual visual Multi- of of the the J J Cortex Cortex still still at at parallel parallel be be takes takes talk talk Wurtz Wurtz In In an an process- when when Neurosci Neurosci steps steps more more Kandel Kandel (which (which cue cue on on character- pattern pattern pathways. pathways. pathways. pathways. the the of of tend tend anatomical anatomical hierarchical hierarchical 1997. 1997. parietal parietal H. H. R. R. remembered remembered parietal parietal must must Rev Rev Responses Responses Thus, Thus, Cereb Cereb J. J. these these temporal temporal only only the the field. field. brain brain the the cross cross early early only only later, later, these these the the processing processing temporal temporal should should effect effect scene scene pathway). pathway). the the of of pathways. pathways. Ann Ann the the the the Eric Eric by by the the the the inactivation inactivation Xing Xing areas areas 1994. 1994. motion motion of of But But stimulus stimulus from from the the form, form, that that level, level, in in Robert Robert posterior posterior exclusive exclusive cortex. cortex. more more pathways pathways inferior inferior separately separately LGN LGN visual visual of of respond respond Distributed Distributed the the and and the the DC, DC, JH. 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