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Processing of Visual Information within the Retinostriate System

P. O. Bishop

10.1002/cphy.cp010309

Source: Supplement 3: Handbook of Physiology, The Nervous System, Sensory Processes

Originally published: 1984

Published online: January 2011

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 Retinostriate Pathway
2 Retina
2.1 Cat Retina
2.2 Monkey Retina
3 Visual Thalamus
3.1 Cat Dorsal Lateral Geniculate Nucleus (LGN)
3.2 Medial Interlaminar Nucleus
3.3 Perigeniculate Nucleus and Nucleus Reticularis Thalami
3.4 Extrageniculate Visual Thalamus
3.5 Macaque Lateral Geniculate Nucleus
3.6 Macaque Pregeniculate Nucleus
3.7 Extrageniculate Visual Thalamus in Monkey
4 Visual Cortex
4.1 Cat Striate Cortex (Area 17)
4.2 Monkey Striate Cortex
5 Binocular Vision
6 Conclusion
Figure 1. Figure 1.

Human retinostriate system, which consists, in effect, of two mirror‐symmetrical retinostriate systems, one for each cerebral hemisphere. Optic nerve fibers from nasal retina of one eye cross in optic chiasma to join with fibers from temporal retina of other eye to form an optic tract that, in due course, relays through lateral geniculate nucleus before projecting to striate cortex in occipital lobe of one cerebral hemisphere. Vertical through fixation point (center of gaze) separates visual field into two hemifields, left and right. As a result of decussation of optic nerve fibers at chiasma, right visual hemifield is represented in left striate cortex (solid pathway) and vice versa for left hemifield (dotted pathway). Solid arrows: projection of hemifield onto striate cortex, center of gaze being represented posteriorly and horizontal periphery of hemifield anteriorly.
Figure 2. Figure 2.

Responses of cat retinal ganglion cells to introduction and withdrawal of a stationary sinusoidal grating pattern. A: off‐center X‐cell; spatial frequency 0.13 cycles/deg. B: off‐center Y‐cell; spatial frequency 0.16 cycles/deg. Contrast (0.32) was turned off and on at 0.45 Hz, as indicated by square waves (downward deflection, off). Length of zero line, duration of 2 s; ordinate, spikes/s. Right: sketches showing phase angle of pattern, i.e., angular position (in deg) of (cosine) grating relative to midpoint of receptive‐field center (dashed line).

From Enroth‐Cugell and Robson 105
Figure 3. Figure 3.

Golgi‐stained ganglion cells from flat mounts of the cat's retina as seen in plan view. A,B: α‐cells, C‐E: β‐cells; F: γ‐cell. G: δ‐cell. Cells A,C,F were located 1.2 mm from center of area centralis; D,G at 2.9 and 2.8 mm, respectively; B,E at 10.0 mm. All are shown at same magnification: calibration bar (near E) = 100 μm.

From Levick 234 as adapted from Boycott and Wassle 45
Figure 4. Figure 4.

Distribution of cat retinal ganglion cell types as function of horizontal and vertical distance from area centralis. T: temporal; N: nasal; I: inferior; S: superior. Top two curves show total ganglion cell densities as a function of eccentricity. Lower three pairs of curves show distributions of α‐, β‐, and γ‐cell types, either counted directly (α) or based on β‐ and γ‐modes (m) of the Nissl‐stained (n) soma diameter spectra. Relative peak densities of βn‐ and γn‐modes were estimated from horseradish peroxidase filling from superior colliculus. The β‐ and γ‐modes can be regarded as only a guide to class cell densities, not only because of cross‐modal class contamination but also because the γn‐mode contains a class of displaced amacrine cells. Density profiles for all three cell‐diameter modes are very similar in form and resemble that of total neuronal density map. Hence the relative densities of various cell types are much the same at all eccentricities.

From Hughes 184
Figure 5. Figure 5.

A: frequency histogram of response latencies of different types of cat retinal ganglion cells (X,Y,W) following antidromic electrical stimulation from optic chiasm (OX). Y, brisk‐transient; X, brisk‐sustained; W, sluggish and nonconcentric types. Cells were recorded in four preparations, in each case, over a fairly wide range of retinal eccentricities. B: frequency histogram of response latencies of different types of cells in dorsal nucleus of lateral geniculate body (LGNd) of cat following electrical stimulation of retina by an electrode located near center of receptive field of particular LGNd cell. BT, brisk‐transient class; BS, brisk‐sustained. C: frequency histogram of responses of cat LGNd cells following antidromic activation from visual cortex. Height of each column in all three histograms (A,B,C) represents total cell count for a particular latency value.

A: from Rowe and Stone 310; B,C: from Cleland et al. 73
Figure 6. Figure 6.

A: histograms made by recording receptive fields of crossed and uncrossed brisk‐sustained retinal ganglion cells in a succession of narrow vertical strips across visual field of left eye of cat. Retinal azimuth (horizontal eccentricity) inferred from positions of receptive fields on a frontal screen. Antidromic activation of cells (accomplished by stimulating electrodes located in the two optic tracts) determined crossed and uncrossed destinations of axons. Pooled data from 30 cats. B: data in A expressed as percentages of crossed (open circles), and uncrossed (filled circles) ganglion cells. Zero azimuth (dotted line) coincides with center of band of crossed‐uncrossed overlap of brisk‐sustained cells. The two complementary curves have been drawn to illustrate symmetry of transition on either side of line of zero azimuth. Abscissa scale magnified to 5 times that of A. C: same as A for brisk‐transient retinal ganglion cells. Pooled data from 30 cats. D: data in C expressed as percentages of crossed (open circles) and uncrossed (continuous line) ganglion cells. Zero azimuth same as for B. Position where 50% of brisk‐transient cells are crossed lies about 2° into temporal retina from that of brisk‐sustained system.

Adapted from Kirk et al. 207 and Levick 235
Figure 7. Figure 7.

A‐D: location of optic nerve axons and terminals in coronal sections of contralateral hemisphere of Macaca mulatta. Composite drawings based on degeneration patterns revealed by Nauta methods following eye enucleation and from autoradiographs following vitreal injection of tritiated leucine. Brain section numbers (+6.9, etc.) are mm anterior to interaural plane. Small dots, terminal degeneration; large dots and dashes, degenerating fiber bundles; PGN, pregeniculate nucleus with subdivisions 1,2; LGN, dorsal lateral geniculate nucleus; MGN, medial geniculate nucleus; PTN, pretectal nucleus; NAOT, lateral terminal nucleus of accessory optic tract; NO, nucleus olivaris; AOT, posterior accessory optic tract; Br, brachium of superior colliculus; SC, superior colliculus.

From Hendrickson et al. 155
Figure 8. Figure 8.

Lateral geniculate nucleus in cat. A: Nissl‐stained parasagittal section through middle of mediolateral extent of dorsal lateral geniculate nucleus (LGNd). A, A1, C, C1‐C3, cell laminae of dorsal lateral geniculate nucleus; PGN, perigeniculate nucleus; OT, optic tract. B: semidiagrammatic coronal section through middle of the anteroposterior extent of LGNd showing laminar distribution of crossed and uncrossed optic nerve fibers. LGNv, ventral lateral geniculate nucleus; MIN, medial interlaminar nucleus; R, reticular nucleus; NP, posterior nucleus; Pul, pulvinar. C: projection map of a parasagittal section through middle of LGNd (lateral 9.2) showing pattern of isoelevation lines; D: projection map of a coronal section through middle of the LGNd (anterior 6.5) showing pattern of isoazimuth and isoelevation lines. Isoelevation values are marked as + or − to distinguish them from isoazimuth lines (unmarked).

C, D: from Sanderson 314
Figure 9. Figure 9.

Spherical polar coordinate systems for defining visual direction relative to visual axis and fixation point (F). A: coordinate system most suitable for mapping projection of visual field onto visual centers in brain. Polar axis passes vertically through nodal point of eye, at right angles to fixation plane. Visual direction expressed by two angles, azimuth (A) and elevation (E). ZV, zero vertical; ZH, zero horizontal (fixation plane). B: coordinate system used for human perimetry. Polar axis coincident with visual axis.
Figure 10. Figure 10.

Semischematic perspective drawing showing planes of isoazimuth and isoelevation in dorsal lateral geniculate nucleus (LGNd) of cat. The two planes intersect along a projection line, which is defined by column of geniculate cells whose receptive fields have a common direction in visual field. The LGNd planes are viewed from a point posterior, medial, and slightly above the nucleus. A, A1, C, main LGNd cell laminae.

Adapted from Bioshop et al. 39
Figure 11. Figure 11.

A: relationship between a projection subunit in dorsal lateral geniculate nucleus (LGNd) and a projection (position) subunit in striate cortex. B: parasagittal section through LGNd showing distribution of cells in a projection subunit, the column in LGNd that contains 90% of all cells whose receptive fields have a common visual direction. Projection line is axis of a projection subunit. Diagram of cross section of a projection column is shown immediately below outline of LGNd. Cortical projection (position) subunit gets its input from a corresponding subunit in LGNd.

Adapted from Sanderson 315
Figure 12. Figure 12.

Major cell and fiber types seen in Golgi preparations of laminae A and A1 of dorsal lateral geniculate nucleus of the cat. Top, lamina A; bottom, lamina A1; dotted lines, region of interlaminar plexus between laminae A and A1. Not all of the elements have been drawn to the same scale. Cells labeled 1, 2, and 3 according to class; RG, retinogeniculate fibers; CG, corticogeniculate fibers.

From Guillery 145
Figure 13. Figure 13.

A: principles of organization of dorsal lateral geniculate nucleus (LGNd). Relay cells (open circles) are shown receiving direct, excitatory retinal inputs as well as inhibitory interneuronal inputs. Interneurons (filled circles) of both types are depicted. An intrageniculate interneuron (right) is placed at same level as relay cells to indicate its position within geniculate laminae. Direct retinal inputs as well as proposed cortical afferent input are shown as being excitatory to such interneurons. A perigeniculate interneuron (left) is placed above the other cells, indicative of its position just above geniculate lamina A. Recurrent collateral and other, presently undefined, inputs are shown exciting such interneurons. B: diagram of LGNd illustrating interneuronal connections and connections from visual cortex and reticular formation on a principal cell (P) and an interneuron (I). RLP (round vesicles, large terminals, pale mitochondria, retinogeniculate axon) and RSD (round vesicles, small terminals, dark mitochondria, corticogeniculate axon) terminals contain round vesicles that are presumed to be excitatory. Intrageniculate axon terminals F1 and F2 contain flat (pleomorphic) vesicles, presumed to be inhibitory. Single lines, axons; broad elements, dendrites (D); narrower elements, presynaptic dendrites, except that their synaptic contacts are enlarged (F2). Bottom center, “triad,” in which optic nerve terminal is presynaptic to both relay cell dendrite and presynaptic dendrite; the latter is also presynaptic to relay cell dendrite.

A: from Dubin and Cleland 103; B: from Burke and Cole 53
Figure 14. Figure 14.

Scatter diagram of antidromic corticogeniculate latency against orthodromic retinogeniculate latency for 115 dorsal lateral geniculate nucleus (LGNd) cells. BT, brisk‐transient cells; BS, brisk‐sustained cells; ST, sluggish‐transient cells; SS, sluggish‐sustained cells; LED, local‐edge‐detector cells.

From Cleland et al. 73
Figure 15. Figure 15.

A: distribution on left (“wrong”) side of zero vertical meridian in visual field of receptive‐field center points of cells recorded in laminae A and A1 in the left dorsal lateral geniculate nucleus (LGNd) of cat. Filled circles, cells with input from contralateral eye; open circles, cells with input from ipsilateral eye. Standard deviation of distribution, 0.76°; standard deviation of error in determination of zero vertical meridian, 0.50°. B: distribution in visual field of the receptive‐field center points of cells recorded in left medial interlaminar nucleus (MIN) in 28 cats. Filled and open circles as in A. Note that cells with their receptive fields in left (“wrong”) hemifield have their input almost exclusively from temporal retina of contralateral eye.

A: from Sanderson and Sherman 318; B: from Sanderson 314
Figure 16. Figure 16.

A‐E: Chartings of transverse sections through cat's lateral posterior‐pulvinar complex illustrate zones of termination of afferent fibers from indicated regions. Chartings based on anterograde autoradiographic experiments. Da, nucleus of Darkschewitsch; Hb, habenular nuclear complex; LP, nucleus lateralis posterior; MGm, medial geniculate body, magnocellular division; MGv, medial geniculate body, ventral division; NIC, interstitial nucleus of Cajal; NP, nucleus posterior of Rioch; Pul, pulvinar; SNr, substantia nigra, pars reticulata; III, oculomotor nerve.

From Berson and Graybiel 21
Figure 17. Figure 17.

A: visuotopic organization of striate recipient zone within feline pulvinar complex. Heavy line, zero vertical meridian; dotted line, zero horizontal; dashed line, peripheral extent of visual field; U and L, upper and lower visual field, respectively. B: nuclei constituting cat's visual thalamus in transverse section showing lateral and posterior nuclear groups and lateral geniculate complex. PI, inferior pulvinar; PM, medial pulvinar; PL, lateral pulvinar; SG, suprageniculate nucleus; LGNd, dorsal lateral geniculate nucleus; LGNv, ventral lateral geniculate nucleus; MIN, medial interlaminar nucleus; NPG, perigeniculate nucleus; OT, optic tract. Nomenclature from Niimi and Kuwahara 271.

From Mason 257
Figure 18. Figure 18.

Laminar organization of lateral geniculate nucleus of rhesus monkey (Macaca mulatta) as seen in transverse section showing two systems of laminar nomenclature (numbers and letters). Medial at left; lateral at right. Laminae receiving an input from the contralateral eye are stippled, those with an input from the ipsilateral eye are not. SE, SI, layer S (superficial) external and internal; ME, MI, external and internal magnocellular layers; PE, PI, parvocellular layers.

Adapted from Kaas et al. 191
Figure 19. Figure 19.

Projection of visual field onto macaque lateral geniculate nucleus (LGN). A, B: semischematic view of right visual hemifield (A) and its projection onto dorsal surface of layer 6 of left LGN (B) using spherical polar coordinate system of Fig. 9A. Surface of LGN is distorted to lie in a plane, distortion being mainly in anteroposterior direction. Thin broken lines, azimuths; solid lines, elevations. Scales in degrees, positive values above zero horizontal, negative values below it. Thick broken lines and solid lines, perimeter of right monocular hemifield; they correspond in both drawings. Heavy dotted line in A, outer limit of right binocular hemifield. C: coronal (A = 7.0) and parasagittal (L = 12.0) sections through macaque LGN. Broken lines, isoazimuth; solid lines, isoelevation. Only a few isoazimuth (A) and isoelevation (E) lines are labeled and their angular values in degrees indicated (e.g., A 1, A 8 … and E − 4, E 0 … etc.). Horsley‐Clarke coordinate scales in mm. The coronal section is approximately at middle of anteroposterior extent of nucleus (cf. parasagittal section at L = 12.0) and parasagittal section is just lateral to middle of mediolateral extent (cf. coronal section at A = 7.0).

Adapted from Malpeli and Baker 251
Figure 20. Figure 20.

Pattern of gyri and sulci associated with visual parts of cat cerebral cortex showing plan (left) and medial (right) aspects of left hemisphere.
Figure 21. Figure 21.

Semischematic view of right visual hemifield (A) and its projection onto left cerebral hemisphere of cat (B‐D). Spherical polar coordinate system of Fig. 9A. A‐C: thin solid lines, azimuths (meridians); broken lines, elevations (parallels). ZH, zero horizontal (dashed line); ZV, zero vertical (solid line). Scales are in degrees, positive values above zero horizontal, negative below. Thick solid lines in A, perimeter of right monocular hemifield; dotted line, outer limit of right binocular hemifield. B: perspective drawing showing extent of representation of visual field on exposed dorsal surface of area 17. The 1‐cm bar refers only to B and D. C: topographic representation in area 17 on medial surface of left cerebral hemisphere with cingulate gyrus removed. Upper cross‐hatched area shows portion of cortex normally hidden by cingulate gyrus. The Horsley‐Clarke scale PA (posterior, anterior) is in mm and refers only to drawing C. Dotted line, upper edge of splenial and postsplenial sulci. D: perspective drawing showing various visual areas on exposed outer surface of cerebral cortex. Lettered designations, divisions of lateral suprasylvian cortex according to Tusa et al. 418.

A‐C: adapted from Tusa et al. 376; D: from Sprague et al. 416
Figure 22. Figure 22.

Lamination schemes and nomenclature of laminae of area 17 of the cat and monkey according to various authors.

Adapted from Garey 119
Figure 23. Figure 23.

Diagrams summarizing details of afferent and efferent relationships and intrinsic spinous neuron relays of cat and monkey visual cortex, area 17.

From Lund et al. 246
Figure 24. Figure 24.

A: length‐response curve from a hypercomplex I cell showing 96% end‐zone inhibition. B: length‐response curve from a simple cell showing considerable response variability. Stippled band indicates ± 1 SD of response variability to either side of mean plateau level. Response indicated by upward‐pointing arrow is approximately 28% below response indicated by downward‐pointing arrow. C: short and long bar stimulus orientation tuning curves from a simple cell showing how a length‐response curve recorded with the bar sufficiently away from the optimal (here 25°) can resemble the type of length‐response curve that is obtained from a hypercomplex cell

From Kato, Bishop, and Orban 195
Figure 25. Figure 25.

A: optimal stimulus length plotted as function of end‐zone inhibition, a, Simple cell with the largest response variability shown in Fig. 24B; b, hypercomplex I cell in Fig. 24A. With the exception of cell c, all cells classified as hypercomplex had end‐zone inhibition of about 40% or more. B: distribution of various levels of end‐zone inhibition.

From Kato, Bishop, and Orban 195
Figure 26. Figure 26.

Comparison of responses of a hypercomplex I cell (A) and a complex cell (B) to light and dark bars, both stationary flashing (Aa, Ab, Ba) and moving (Ac, Ad, Bb), as well as to moving light and dark edges (Ae, Bc). Stationary bars (0.1° or 0.5° wide) were flashed at indicated frequencies either on or off (filled and open triangles) or reversed in contrast (filled and open circles), change in contrast being the same in each case (Δc = 0.2). Keys to symbols same for both cells. Hypercomplex I cell has an antisymmetrical response profile. Responses to light bar (open histograms) and to dark bar (filled histograms) were recorded separately and only subsequently combined, regions common to both responses being dotted. Responses for each cell in accurate vertical alignment after correcting for response latency for moving stimuli.

From Kulikowski, Bishop, and Kato 215
Figure 27. Figure 27.

Monocular stimulus orientation tuning curves for each of the two eyes from a binocular simple cell in striate cortex of cat. Curve (R) for right eye has been normalized to same height as the curve for left eye. The abscissa zero is arbitrarily located so that curves are approximately symmetrical about it. HW at HH: half‐width at half‐height of tuning curves of left (L) and right (R) eye.

From Bishop 407
Figure 28. Figure 28.

A: histograms of average responses from a hypercomplex I cell in cat striate cortex to sinusoidal gratings of four different spatial frequencies drifting over receptive field at a constant rate (0.2 Hz) in preferred direction. Response is modulated in synchrony with passage of bars of grating up to highest spatial frequency to which cell responds. B: responses from same cell as in A to a stationary flashing sinusoidal grating (0.5 cycles/deg) placed at different phase positions across receptive field. Grating was flashed at 2 Hz either on and off (a) at each position or reversed in contrast (b), the change in contrast being the same in each case (ΔC = 0.2). In both cases there are two null phases, 180° apart, at which the grating elicits no response.

From Kulikowski and Bishop 213
Figure 29. Figure 29.

Histograms of average responses from three simple cells (A, B, C) in cat striate cortex to moving bars (a) and edges (b) showing predictability of these responses from inverse Fourier transformation (continuous lines) of respective contrast sensitivity tuning curves (c). Shapes of these tuning curves were chosen to produce best fit to data points, thereby testing predictability of responses to bars and edges (essential for proper classification of simple cells). However, fitted tuning curves are not greatly different from ideal Gaussian functions for tuning curves with medium and narrow bandwidths (<1.5 octaves). Cell A is fitted best by an antisymmetrical profile (sinusoidal Fourier transform), whereas cells B and C are a pair fitted with symmetrical and antisymmetrical profiles, respectively. Contrast (C) = 0.2.

From Kulikowski and Bishop 212
Figure 30. Figure 30.

Tangential microelectrode penetrations through left postlateral gyrus in cat. A, B, C from one experiment; D from another. Receptive fields (A) and receptive‐field centers (B) in lower right hemifield are from cells recorded along microelectrode track shown in coronal histological sections through postlateral gyrus (C). Line in B, mean receptive‐field position (drawn by hand); AC, projection of area centralis. D: centers of two series of receptive fields, one close to area centralis, the other farther away, recorded from cells along two further tangential electrode penetrations.

Adapted by Bishop 28 from Albus 4
Figure 31. Figure 31.

Spatial organization of orientation domain. A: plan view of an extended surface of postlateral gyrus in cat. Striate cells were recorded along three tangential electrode tracks indicated by arrows from the scale at top. The 43 cell positions (filled circles) are projected onto surface and their preferred stimulus orientation in each case indicated by a line. Area (orientation matrix) selected is 1 mm long in anteroposterior direction and 0.5 mm wide in mediolateral direction. Gaps between experimentally recorded cells have been filled in with hypothetical neurons located at regular spacings of 50 μm, each preferred orientation being set 10° different from that of preceding cell. The interpolation procedure started at the second experimentally recorded neuron (star) and proceeded always toward next experimentally recorded cell from left to right and from top to bottom. Squares labeled 1 and 2 are arbitrarily selected regions, one with a side length of 400 μm, the other of 300 μm, in which full range, or nearly the full range, of preferred orientations is represented, although not to an equal degree. B: positions of cells having same preferred orientation and not more than 50 μm apart as shown in A are connected by continuous lines. Broken lines interconnect cells between 50 and 100 μm apart. Orientation represented by each isoorientation line is indicated at one or both ends of line.

From Albus 5
Figure 32. Figure 32.

Pattern of sulci associated with, or surrounding, visual parts of left hemisphere of macaque monkey cerebral cortex. A: lateral; B: medial. The following sulci are labeled: central, intraparietal, superior temporal, lunate, lateral calcarine, inferior occipital, parietooccipital, medial calcarine, collateral, and callosomarginal (cingulate). Lateral fissure and corpus callosum are also labeled.
Figure 33. Figure 33.

Topography of striate cortex in the macaque monkey. A: posterolateral view of brain showing area 17‐area 18 border (broken line) and extent of roof of buried calcarine sulcus (dotted line). Three oblique lines indicate levels of parasagittal sections shown at c, d, and e. B: same view, with representation of zero horizontal (0) and of the parallels 1°, 3°, and 6° above (+) and below (−) zero horizontal. Fovea is represented laterally where zero horizontal meets the area 17‐area 18 border (zero vertical meridian). C‐E: parasagittal sections (medial to lateral) to show extent of area 17 (black). Three levels of cortex lie in parallel planes: 1, operculum; 2, roof of calcarine sulcus; 3, leaves joining roof to stem. Stem of sulcus is oriented perpendicular to other levels. Dotted white lines, approximate planes of tangential sections used for reconstructed patterns of ocular dominance columns in Fig. 38B and Fig. 39. LS, lunate sulcus; CS, calcarine sulcus.

From LeVay et al. 228
Figure 34. Figure 34.

A: Nissl‐stained section of area 17 of Macaca mulatta from the perimacular area of occipital operculum. Average depth in this area from pia to white matter in frozen sections is 1,700 μm; average thickness for lamina 1 = 100 μm, lamina 2 + 3 = 650 μm, lamina 4A = 70 μm, lamina 4B = 150 μm, lamina 4Cα = 140 μm, lamina 4Cβ = 140 μm, lamina 5 = 210 μm, lamina 6 = 240 μm. B: Golgi rapid preparation of perimacular area 17. Large pyramidal cells of Meynert can be seen in lamina 6.

From Lund 244
Figure 35. Figure 35.

Retinal, lateral geniculate (LGN), and striate cortical magnification factors as a function of visual eccentricity in macaque monkey. The three curves have been normalized with respect to their ordinate values at 0° eccentricity.

Data for retinal curve from Rolls and Cowey 305; for LGN curve from Malpeli and Baker 251; for striate cortex from Daniel and Whitteridge 82
Figure 36. Figure 36.

Graph of average receptive‐field size (crosses) and magnification−1 in deg/mm (circles) against eccentricity for 5 striate cortical locations. Points for 4°, 8°, 18°, and 22° were from one monkey; point for 1° from a second. Receptive‐field size was determined by averaging fields at each eccentricity, estimating size from (length × width)0.5.

From Hubel and Wiesel 178
Figure 37. Figure 37.

Graph of preferred stimulus orientation of striate cells vs. electrode track distance for an oblique penetration restricted to layers 2 and 3 (inset) in monkey cortex. Filled circles, cells dominated by right eye; open circles, by left eye. Several reversals in direction of rotation occur, with two very long, almost linear, sequences followed by two short ones. Right eye was dominant until almost end of the sequence.

From Hubel and Wiesel 179
Figure 38. Figure 38.

A: reconstructed pattern of orientation columns in macaque striate cortex viewed face‐on. Reconstruction made from deoxyglucose autoradiographs by cutting tangential sections through exposed part of left occipital cortex and mounting the parts of each section that passed through layer 6. B: reconstructed pattern of ocular‐dominance columns in the same region as A, made from autoradiographs of [3H]proline sections following injection of right eye; dark‐field photographs. Both reconstructions A and B based on the same series of tangential sections, every third section being set aside for transneuronal labeling with tritium. For B, parts of each autoradiograph that passed through layer 4C were cut and mounted.

From Hubel et al. 182
Figure 39. Figure 39.

Reconstruction of ocular‐dominance columns over whole of exposed part of striate cortex in right occipital lobe of a macaque monkey, as shown also in Fig. 33. Reconstruction prepared from set of serial sections roughly tangential to exposed surface of lobe and stained by reduced silver method of Liesegang 228. In diagram, every other column has been inked in, dark stripes corresponding to one eye and the light stripes to the other. F, representation of fovea on cortex; broken line VFV′, area 17‐ area 18 border, representing vertical midline; ZH, zero horizontal drawn by eye along path of confluence of stripes as they stream in from VF and V′F. Line VV′ at medial edge of lobe indicates where cortex bends over abruptly to continue as a buried fold. Line approximates the 8° isoazimuth.

Adapted from Hubel and Freeman 166
Figure 40. Figure 40.

Comparison of patterns of orientation and ocular‐dominance columns in same area of striate cortex in same monkey. Orientation columns from Fig. 38A have been traced as thick lines, left‐eye ocular‐dominance columns from Fig. 38B as thin lines. Average widths of the hypercolumns are 770 μm for ocular dominance, 570 μm for orientation. Angles of intersection of two sets of columns show a distribution not obviously different from that expected for any two randomly superimposed sets of lines.

From Hubel et al. 182


Figure 1.

Human retinostriate system, which consists, in effect, of two mirror‐symmetrical retinostriate systems, one for each cerebral hemisphere. Optic nerve fibers from nasal retina of one eye cross in optic chiasma to join with fibers from temporal retina of other eye to form an optic tract that, in due course, relays through lateral geniculate nucleus before projecting to striate cortex in occipital lobe of one cerebral hemisphere. Vertical through fixation point (center of gaze) separates visual field into two hemifields, left and right. As a result of decussation of optic nerve fibers at chiasma, right visual hemifield is represented in left striate cortex (solid pathway) and vice versa for left hemifield (dotted pathway). Solid arrows: projection of hemifield onto striate cortex, center of gaze being represented posteriorly and horizontal periphery of hemifield anteriorly.



Figure 2.

Responses of cat retinal ganglion cells to introduction and withdrawal of a stationary sinusoidal grating pattern. A: off‐center X‐cell; spatial frequency 0.13 cycles/deg. B: off‐center Y‐cell; spatial frequency 0.16 cycles/deg. Contrast (0.32) was turned off and on at 0.45 Hz, as indicated by square waves (downward deflection, off). Length of zero line, duration of 2 s; ordinate, spikes/s. Right: sketches showing phase angle of pattern, i.e., angular position (in deg) of (cosine) grating relative to midpoint of receptive‐field center (dashed line).

From Enroth‐Cugell and Robson 105


Figure 3.

Golgi‐stained ganglion cells from flat mounts of the cat's retina as seen in plan view. A,B: α‐cells, C‐E: β‐cells; F: γ‐cell. G: δ‐cell. Cells A,C,F were located 1.2 mm from center of area centralis; D,G at 2.9 and 2.8 mm, respectively; B,E at 10.0 mm. All are shown at same magnification: calibration bar (near E) = 100 μm.

From Levick 234 as adapted from Boycott and Wassle 45


Figure 4.

Distribution of cat retinal ganglion cell types as function of horizontal and vertical distance from area centralis. T: temporal; N: nasal; I: inferior; S: superior. Top two curves show total ganglion cell densities as a function of eccentricity. Lower three pairs of curves show distributions of α‐, β‐, and γ‐cell types, either counted directly (α) or based on β‐ and γ‐modes (m) of the Nissl‐stained (n) soma diameter spectra. Relative peak densities of βn‐ and γn‐modes were estimated from horseradish peroxidase filling from superior colliculus. The β‐ and γ‐modes can be regarded as only a guide to class cell densities, not only because of cross‐modal class contamination but also because the γn‐mode contains a class of displaced amacrine cells. Density profiles for all three cell‐diameter modes are very similar in form and resemble that of total neuronal density map. Hence the relative densities of various cell types are much the same at all eccentricities.

From Hughes 184


Figure 5.

A: frequency histogram of response latencies of different types of cat retinal ganglion cells (X,Y,W) following antidromic electrical stimulation from optic chiasm (OX). Y, brisk‐transient; X, brisk‐sustained; W, sluggish and nonconcentric types. Cells were recorded in four preparations, in each case, over a fairly wide range of retinal eccentricities. B: frequency histogram of response latencies of different types of cells in dorsal nucleus of lateral geniculate body (LGNd) of cat following electrical stimulation of retina by an electrode located near center of receptive field of particular LGNd cell. BT, brisk‐transient class; BS, brisk‐sustained. C: frequency histogram of responses of cat LGNd cells following antidromic activation from visual cortex. Height of each column in all three histograms (A,B,C) represents total cell count for a particular latency value.

A: from Rowe and Stone 310; B,C: from Cleland et al. 73


Figure 6.

A: histograms made by recording receptive fields of crossed and uncrossed brisk‐sustained retinal ganglion cells in a succession of narrow vertical strips across visual field of left eye of cat. Retinal azimuth (horizontal eccentricity) inferred from positions of receptive fields on a frontal screen. Antidromic activation of cells (accomplished by stimulating electrodes located in the two optic tracts) determined crossed and uncrossed destinations of axons. Pooled data from 30 cats. B: data in A expressed as percentages of crossed (open circles), and uncrossed (filled circles) ganglion cells. Zero azimuth (dotted line) coincides with center of band of crossed‐uncrossed overlap of brisk‐sustained cells. The two complementary curves have been drawn to illustrate symmetry of transition on either side of line of zero azimuth. Abscissa scale magnified to 5 times that of A. C: same as A for brisk‐transient retinal ganglion cells. Pooled data from 30 cats. D: data in C expressed as percentages of crossed (open circles) and uncrossed (continuous line) ganglion cells. Zero azimuth same as for B. Position where 50% of brisk‐transient cells are crossed lies about 2° into temporal retina from that of brisk‐sustained system.

Adapted from Kirk et al. 207 and Levick 235


Figure 7.

A‐D: location of optic nerve axons and terminals in coronal sections of contralateral hemisphere of Macaca mulatta. Composite drawings based on degeneration patterns revealed by Nauta methods following eye enucleation and from autoradiographs following vitreal injection of tritiated leucine. Brain section numbers (+6.9, etc.) are mm anterior to interaural plane. Small dots, terminal degeneration; large dots and dashes, degenerating fiber bundles; PGN, pregeniculate nucleus with subdivisions 1,2; LGN, dorsal lateral geniculate nucleus; MGN, medial geniculate nucleus; PTN, pretectal nucleus; NAOT, lateral terminal nucleus of accessory optic tract; NO, nucleus olivaris; AOT, posterior accessory optic tract; Br, brachium of superior colliculus; SC, superior colliculus.

From Hendrickson et al. 155


Figure 8.

Lateral geniculate nucleus in cat. A: Nissl‐stained parasagittal section through middle of mediolateral extent of dorsal lateral geniculate nucleus (LGNd). A, A1, C, C1‐C3, cell laminae of dorsal lateral geniculate nucleus; PGN, perigeniculate nucleus; OT, optic tract. B: semidiagrammatic coronal section through middle of the anteroposterior extent of LGNd showing laminar distribution of crossed and uncrossed optic nerve fibers. LGNv, ventral lateral geniculate nucleus; MIN, medial interlaminar nucleus; R, reticular nucleus; NP, posterior nucleus; Pul, pulvinar. C: projection map of a parasagittal section through middle of LGNd (lateral 9.2) showing pattern of isoelevation lines; D: projection map of a coronal section through middle of the LGNd (anterior 6.5) showing pattern of isoazimuth and isoelevation lines. Isoelevation values are marked as + or − to distinguish them from isoazimuth lines (unmarked).

C, D: from Sanderson 314


Figure 9.

Spherical polar coordinate systems for defining visual direction relative to visual axis and fixation point (F). A: coordinate system most suitable for mapping projection of visual field onto visual centers in brain. Polar axis passes vertically through nodal point of eye, at right angles to fixation plane. Visual direction expressed by two angles, azimuth (A) and elevation (E). ZV, zero vertical; ZH, zero horizontal (fixation plane). B: coordinate system used for human perimetry. Polar axis coincident with visual axis.



Figure 10.

Semischematic perspective drawing showing planes of isoazimuth and isoelevation in dorsal lateral geniculate nucleus (LGNd) of cat. The two planes intersect along a projection line, which is defined by column of geniculate cells whose receptive fields have a common direction in visual field. The LGNd planes are viewed from a point posterior, medial, and slightly above the nucleus. A, A1, C, main LGNd cell laminae.

Adapted from Bioshop et al. 39


Figure 11.

A: relationship between a projection subunit in dorsal lateral geniculate nucleus (LGNd) and a projection (position) subunit in striate cortex. B: parasagittal section through LGNd showing distribution of cells in a projection subunit, the column in LGNd that contains 90% of all cells whose receptive fields have a common visual direction. Projection line is axis of a projection subunit. Diagram of cross section of a projection column is shown immediately below outline of LGNd. Cortical projection (position) subunit gets its input from a corresponding subunit in LGNd.

Adapted from Sanderson 315


Figure 12.

Major cell and fiber types seen in Golgi preparations of laminae A and A1 of dorsal lateral geniculate nucleus of the cat. Top, lamina A; bottom, lamina A1; dotted lines, region of interlaminar plexus between laminae A and A1. Not all of the elements have been drawn to the same scale. Cells labeled 1, 2, and 3 according to class; RG, retinogeniculate fibers; CG, corticogeniculate fibers.

From Guillery 145


Figure 13.

A: principles of organization of dorsal lateral geniculate nucleus (LGNd). Relay cells (open circles) are shown receiving direct, excitatory retinal inputs as well as inhibitory interneuronal inputs. Interneurons (filled circles) of both types are depicted. An intrageniculate interneuron (right) is placed at same level as relay cells to indicate its position within geniculate laminae. Direct retinal inputs as well as proposed cortical afferent input are shown as being excitatory to such interneurons. A perigeniculate interneuron (left) is placed above the other cells, indicative of its position just above geniculate lamina A. Recurrent collateral and other, presently undefined, inputs are shown exciting such interneurons. B: diagram of LGNd illustrating interneuronal connections and connections from visual cortex and reticular formation on a principal cell (P) and an interneuron (I). RLP (round vesicles, large terminals, pale mitochondria, retinogeniculate axon) and RSD (round vesicles, small terminals, dark mitochondria, corticogeniculate axon) terminals contain round vesicles that are presumed to be excitatory. Intrageniculate axon terminals F1 and F2 contain flat (pleomorphic) vesicles, presumed to be inhibitory. Single lines, axons; broad elements, dendrites (D); narrower elements, presynaptic dendrites, except that their synaptic contacts are enlarged (F2). Bottom center, “triad,” in which optic nerve terminal is presynaptic to both relay cell dendrite and presynaptic dendrite; the latter is also presynaptic to relay cell dendrite.

A: from Dubin and Cleland 103; B: from Burke and Cole 53


Figure 14.

Scatter diagram of antidromic corticogeniculate latency against orthodromic retinogeniculate latency for 115 dorsal lateral geniculate nucleus (LGNd) cells. BT, brisk‐transient cells; BS, brisk‐sustained cells; ST, sluggish‐transient cells; SS, sluggish‐sustained cells; LED, local‐edge‐detector cells.

From Cleland et al. 73


Figure 15.

A: distribution on left (“wrong”) side of zero vertical meridian in visual field of receptive‐field center points of cells recorded in laminae A and A1 in the left dorsal lateral geniculate nucleus (LGNd) of cat. Filled circles, cells with input from contralateral eye; open circles, cells with input from ipsilateral eye. Standard deviation of distribution, 0.76°; standard deviation of error in determination of zero vertical meridian, 0.50°. B: distribution in visual field of the receptive‐field center points of cells recorded in left medial interlaminar nucleus (MIN) in 28 cats. Filled and open circles as in A. Note that cells with their receptive fields in left (“wrong”) hemifield have their input almost exclusively from temporal retina of contralateral eye.

A: from Sanderson and Sherman 318; B: from Sanderson 314


Figure 16.

A‐E: Chartings of transverse sections through cat's lateral posterior‐pulvinar complex illustrate zones of termination of afferent fibers from indicated regions. Chartings based on anterograde autoradiographic experiments. Da, nucleus of Darkschewitsch; Hb, habenular nuclear complex; LP, nucleus lateralis posterior; MGm, medial geniculate body, magnocellular division; MGv, medial geniculate body, ventral division; NIC, interstitial nucleus of Cajal; NP, nucleus posterior of Rioch; Pul, pulvinar; SNr, substantia nigra, pars reticulata; III, oculomotor nerve.

From Berson and Graybiel 21


Figure 17.

A: visuotopic organization of striate recipient zone within feline pulvinar complex. Heavy line, zero vertical meridian; dotted line, zero horizontal; dashed line, peripheral extent of visual field; U and L, upper and lower visual field, respectively. B: nuclei constituting cat's visual thalamus in transverse section showing lateral and posterior nuclear groups and lateral geniculate complex. PI, inferior pulvinar; PM, medial pulvinar; PL, lateral pulvinar; SG, suprageniculate nucleus; LGNd, dorsal lateral geniculate nucleus; LGNv, ventral lateral geniculate nucleus; MIN, medial interlaminar nucleus; NPG, perigeniculate nucleus; OT, optic tract. Nomenclature from Niimi and Kuwahara 271.

From Mason 257


Figure 18.

Laminar organization of lateral geniculate nucleus of rhesus monkey (Macaca mulatta) as seen in transverse section showing two systems of laminar nomenclature (numbers and letters). Medial at left; lateral at right. Laminae receiving an input from the contralateral eye are stippled, those with an input from the ipsilateral eye are not. SE, SI, layer S (superficial) external and internal; ME, MI, external and internal magnocellular layers; PE, PI, parvocellular layers.

Adapted from Kaas et al. 191


Figure 19.

Projection of visual field onto macaque lateral geniculate nucleus (LGN). A, B: semischematic view of right visual hemifield (A) and its projection onto dorsal surface of layer 6 of left LGN (B) using spherical polar coordinate system of Fig. 9A. Surface of LGN is distorted to lie in a plane, distortion being mainly in anteroposterior direction. Thin broken lines, azimuths; solid lines, elevations. Scales in degrees, positive values above zero horizontal, negative values below it. Thick broken lines and solid lines, perimeter of right monocular hemifield; they correspond in both drawings. Heavy dotted line in A, outer limit of right binocular hemifield. C: coronal (A = 7.0) and parasagittal (L = 12.0) sections through macaque LGN. Broken lines, isoazimuth; solid lines, isoelevation. Only a few isoazimuth (A) and isoelevation (E) lines are labeled and their angular values in degrees indicated (e.g., A 1, A 8 … and E − 4, E 0 … etc.). Horsley‐Clarke coordinate scales in mm. The coronal section is approximately at middle of anteroposterior extent of nucleus (cf. parasagittal section at L = 12.0) and parasagittal section is just lateral to middle of mediolateral extent (cf. coronal section at A = 7.0).

Adapted from Malpeli and Baker 251


Figure 20.

Pattern of gyri and sulci associated with visual parts of cat cerebral cortex showing plan (left) and medial (right) aspects of left hemisphere.



Figure 21.

Semischematic view of right visual hemifield (A) and its projection onto left cerebral hemisphere of cat (B‐D). Spherical polar coordinate system of Fig. 9A. A‐C: thin solid lines, azimuths (meridians); broken lines, elevations (parallels). ZH, zero horizontal (dashed line); ZV, zero vertical (solid line). Scales are in degrees, positive values above zero horizontal, negative below. Thick solid lines in A, perimeter of right monocular hemifield; dotted line, outer limit of right binocular hemifield. B: perspective drawing showing extent of representation of visual field on exposed dorsal surface of area 17. The 1‐cm bar refers only to B and D. C: topographic representation in area 17 on medial surface of left cerebral hemisphere with cingulate gyrus removed. Upper cross‐hatched area shows portion of cortex normally hidden by cingulate gyrus. The Horsley‐Clarke scale PA (posterior, anterior) is in mm and refers only to drawing C. Dotted line, upper edge of splenial and postsplenial sulci. D: perspective drawing showing various visual areas on exposed outer surface of cerebral cortex. Lettered designations, divisions of lateral suprasylvian cortex according to Tusa et al. 418.

A‐C: adapted from Tusa et al. 376; D: from Sprague et al. 416


Figure 22.

Lamination schemes and nomenclature of laminae of area 17 of the cat and monkey according to various authors.

Adapted from Garey 119


Figure 23.

Diagrams summarizing details of afferent and efferent relationships and intrinsic spinous neuron relays of cat and monkey visual cortex, area 17.

From Lund et al. 246


Figure 24.

A: length‐response curve from a hypercomplex I cell showing 96% end‐zone inhibition. B: length‐response curve from a simple cell showing considerable response variability. Stippled band indicates ± 1 SD of response variability to either side of mean plateau level. Response indicated by upward‐pointing arrow is approximately 28% below response indicated by downward‐pointing arrow. C: short and long bar stimulus orientation tuning curves from a simple cell showing how a length‐response curve recorded with the bar sufficiently away from the optimal (here 25°) can resemble the type of length‐response curve that is obtained from a hypercomplex cell

From Kato, Bishop, and Orban 195


Figure 25.

A: optimal stimulus length plotted as function of end‐zone inhibition, a, Simple cell with the largest response variability shown in Fig. 24B; b, hypercomplex I cell in Fig. 24A. With the exception of cell c, all cells classified as hypercomplex had end‐zone inhibition of about 40% or more. B: distribution of various levels of end‐zone inhibition.

From Kato, Bishop, and Orban 195


Figure 26.

Comparison of responses of a hypercomplex I cell (A) and a complex cell (B) to light and dark bars, both stationary flashing (Aa, Ab, Ba) and moving (Ac, Ad, Bb), as well as to moving light and dark edges (Ae, Bc). Stationary bars (0.1° or 0.5° wide) were flashed at indicated frequencies either on or off (filled and open triangles) or reversed in contrast (filled and open circles), change in contrast being the same in each case (Δc = 0.2). Keys to symbols same for both cells. Hypercomplex I cell has an antisymmetrical response profile. Responses to light bar (open histograms) and to dark bar (filled histograms) were recorded separately and only subsequently combined, regions common to both responses being dotted. Responses for each cell in accurate vertical alignment after correcting for response latency for moving stimuli.

From Kulikowski, Bishop, and Kato 215


Figure 27.

Monocular stimulus orientation tuning curves for each of the two eyes from a binocular simple cell in striate cortex of cat. Curve (R) for right eye has been normalized to same height as the curve for left eye. The abscissa zero is arbitrarily located so that curves are approximately symmetrical about it. HW at HH: half‐width at half‐height of tuning curves of left (L) and right (R) eye.

From Bishop 407


Figure 28.

A: histograms of average responses from a hypercomplex I cell in cat striate cortex to sinusoidal gratings of four different spatial frequencies drifting over receptive field at a constant rate (0.2 Hz) in preferred direction. Response is modulated in synchrony with passage of bars of grating up to highest spatial frequency to which cell responds. B: responses from same cell as in A to a stationary flashing sinusoidal grating (0.5 cycles/deg) placed at different phase positions across receptive field. Grating was flashed at 2 Hz either on and off (a) at each position or reversed in contrast (b), the change in contrast being the same in each case (ΔC = 0.2). In both cases there are two null phases, 180° apart, at which the grating elicits no response.

From Kulikowski and Bishop 213


Figure 29.

Histograms of average responses from three simple cells (A, B, C) in cat striate cortex to moving bars (a) and edges (b) showing predictability of these responses from inverse Fourier transformation (continuous lines) of respective contrast sensitivity tuning curves (c). Shapes of these tuning curves were chosen to produce best fit to data points, thereby testing predictability of responses to bars and edges (essential for proper classification of simple cells). However, fitted tuning curves are not greatly different from ideal Gaussian functions for tuning curves with medium and narrow bandwidths (<1.5 octaves). Cell A is fitted best by an antisymmetrical profile (sinusoidal Fourier transform), whereas cells B and C are a pair fitted with symmetrical and antisymmetrical profiles, respectively. Contrast (C) = 0.2.

From Kulikowski and Bishop 212


Figure 30.

Tangential microelectrode penetrations through left postlateral gyrus in cat. A, B, C from one experiment; D from another. Receptive fields (A) and receptive‐field centers (B) in lower right hemifield are from cells recorded along microelectrode track shown in coronal histological sections through postlateral gyrus (C). Line in B, mean receptive‐field position (drawn by hand); AC, projection of area centralis. D: centers of two series of receptive fields, one close to area centralis, the other farther away, recorded from cells along two further tangential electrode penetrations.

Adapted by Bishop 28 from Albus 4


Figure 31.

Spatial organization of orientation domain. A: plan view of an extended surface of postlateral gyrus in cat. Striate cells were recorded along three tangential electrode tracks indicated by arrows from the scale at top. The 43 cell positions (filled circles) are projected onto surface and their preferred stimulus orientation in each case indicated by a line. Area (orientation matrix) selected is 1 mm long in anteroposterior direction and 0.5 mm wide in mediolateral direction. Gaps between experimentally recorded cells have been filled in with hypothetical neurons located at regular spacings of 50 μm, each preferred orientation being set 10° different from that of preceding cell. The interpolation procedure started at the second experimentally recorded neuron (star) and proceeded always toward next experimentally recorded cell from left to right and from top to bottom. Squares labeled 1 and 2 are arbitrarily selected regions, one with a side length of 400 μm, the other of 300 μm, in which full range, or nearly the full range, of preferred orientations is represented, although not to an equal degree. B: positions of cells having same preferred orientation and not more than 50 μm apart as shown in A are connected by continuous lines. Broken lines interconnect cells between 50 and 100 μm apart. Orientation represented by each isoorientation line is indicated at one or both ends of line.

From Albus 5


Figure 32.

Pattern of sulci associated with, or surrounding, visual parts of left hemisphere of macaque monkey cerebral cortex. A: lateral; B: medial. The following sulci are labeled: central, intraparietal, superior temporal, lunate, lateral calcarine, inferior occipital, parietooccipital, medial calcarine, collateral, and callosomarginal (cingulate). Lateral fissure and corpus callosum are also labeled.



Figure 33.

Topography of striate cortex in the macaque monkey. A: posterolateral view of brain showing area 17‐area 18 border (broken line) and extent of roof of buried calcarine sulcus (dotted line). Three oblique lines indicate levels of parasagittal sections shown at c, d, and e. B: same view, with representation of zero horizontal (0) and of the parallels 1°, 3°, and 6° above (+) and below (−) zero horizontal. Fovea is represented laterally where zero horizontal meets the area 17‐area 18 border (zero vertical meridian). C‐E: parasagittal sections (medial to lateral) to show extent of area 17 (black). Three levels of cortex lie in parallel planes: 1, operculum; 2, roof of calcarine sulcus; 3, leaves joining roof to stem. Stem of sulcus is oriented perpendicular to other levels. Dotted white lines, approximate planes of tangential sections used for reconstructed patterns of ocular dominance columns in Fig. 38B and Fig. 39. LS, lunate sulcus; CS, calcarine sulcus.

From LeVay et al. 228


Figure 34.

A: Nissl‐stained section of area 17 of Macaca mulatta from the perimacular area of occipital operculum. Average depth in this area from pia to white matter in frozen sections is 1,700 μm; average thickness for lamina 1 = 100 μm, lamina 2 + 3 = 650 μm, lamina 4A = 70 μm, lamina 4B = 150 μm, lamina 4Cα = 140 μm, lamina 4Cβ = 140 μm, lamina 5 = 210 μm, lamina 6 = 240 μm. B: Golgi rapid preparation of perimacular area 17. Large pyramidal cells of Meynert can be seen in lamina 6.

From Lund 244


Figure 35.

Retinal, lateral geniculate (LGN), and striate cortical magnification factors as a function of visual eccentricity in macaque monkey. The three curves have been normalized with respect to their ordinate values at 0° eccentricity.

Data for retinal curve from Rolls and Cowey 305; for LGN curve from Malpeli and Baker 251; for striate cortex from Daniel and Whitteridge 82


Figure 36.

Graph of average receptive‐field size (crosses) and magnification−1 in deg/mm (circles) against eccentricity for 5 striate cortical locations. Points for 4°, 8°, 18°, and 22° were from one monkey; point for 1° from a second. Receptive‐field size was determined by averaging fields at each eccentricity, estimating size from (length × width)0.5.

From Hubel and Wiesel 178


Figure 37.

Graph of preferred stimulus orientation of striate cells vs. electrode track distance for an oblique penetration restricted to layers 2 and 3 (inset) in monkey cortex. Filled circles, cells dominated by right eye; open circles, by left eye. Several reversals in direction of rotation occur, with two very long, almost linear, sequences followed by two short ones. Right eye was dominant until almost end of the sequence.

From Hubel and Wiesel 179


Figure 38.

A: reconstructed pattern of orientation columns in macaque striate cortex viewed face‐on. Reconstruction made from deoxyglucose autoradiographs by cutting tangential sections through exposed part of left occipital cortex and mounting the parts of each section that passed through layer 6. B: reconstructed pattern of ocular‐dominance columns in the same region as A, made from autoradiographs of [3H]proline sections following injection of right eye; dark‐field photographs. Both reconstructions A and B based on the same series of tangential sections, every third section being set aside for transneuronal labeling with tritium. For B, parts of each autoradiograph that passed through layer 4C were cut and mounted.

From Hubel et al. 182


Figure 39.

Reconstruction of ocular‐dominance columns over whole of exposed part of striate cortex in right occipital lobe of a macaque monkey, as shown also in Fig. 33. Reconstruction prepared from set of serial sections roughly tangential to exposed surface of lobe and stained by reduced silver method of Liesegang 228. In diagram, every other column has been inked in, dark stripes corresponding to one eye and the light stripes to the other. F, representation of fovea on cortex; broken line VFV′, area 17‐ area 18 border, representing vertical midline; ZH, zero horizontal drawn by eye along path of confluence of stripes as they stream in from VF and V′F. Line VV′ at medial edge of lobe indicates where cortex bends over abruptly to continue as a buried fold. Line approximates the 8° isoazimuth.

Adapted from Hubel and Freeman 166


Figure 40.

Comparison of patterns of orientation and ocular‐dominance columns in same area of striate cortex in same monkey. Orientation columns from Fig. 38A have been traced as thick lines, left‐eye ocular‐dominance columns from Fig. 38B as thin lines. Average widths of the hypercolumns are 770 μm for ocular dominance, 570 μm for orientation. Angles of intersection of two sets of columns show a distribution not obviously different from that expected for any two randomly superimposed sets of lines.

From Hubel et al. 182
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Article Title: Processing of Visual Information within the Retinostriate System
 

How to Cite

P. O. Bishop. Processing of Visual Information within the Retinostriate System. Compr Physiol 2011, Supplement 3: Handbook of Physiology, The Nervous System, Sensory Processes: 341-424. First published in print 1984. doi: 10.1002/cphy.cp010309