In recent years, many new cortical areas have been identified in the macaque monkey. The number of identified connections between areas has increased even more dramatically. We report here on (1) a summary of the layout of cortical areas associated with vision and with other modalities, (2) a computerized database for storing and representing large amounts of information on connectivity patterns, and (3) the application of these data to the analysis of hierarchical organization of the cerebral cortex. Our analysis concentrates on the visual system, which includes 25 neocortical areas that are predominantly or exclusively visual in function, plus an additional 7 areas that we regard as visual-association areas on the basis of their extensive visual inputs. A total of 305 connections among these 32 visual and visual-association areas have been reported. This represents 31% of the possible number of pathways if each area were connected with all others. The actual degree of connectivity is likely to be closer to 40%. The great majority of pathways involve reciprocal connections between areas. There are also extensive connections with cortical areas outside the visual system proper, including the somatosensory cortex, as well as neocortical, transitional, and archicortical regions in the temporal and frontal lobes. In the somatosensory/motor system, there are 62 identified pathways linking 13 cortical areas, suggesting an overall connectivity of about 40%. Based on the laminar patterns of connections between areas, we propose a hierarchy of visual areas and of somatosensory/motor areas that is more comprehensive than those suggested in other recent studies. The current version of the visual hierarchy includes 10 levels of cortical processing. Altogether, it contains 14 levels if one includes the retina and lateral geniculate nucleus at the bottom as well as the entorhinal cortex and hippocampus at the top. Within this hierarchy, there are multiple, intertwined processing streams, which, at a low level, are related to the compartmental organization of areas V1 and V2 and, at a high level, are related to the distinction between processing centers in the temporal and parietal lobes. However, there are some pathways and relationships (about 10% of the total) whose descriptions do not fit cleanly into this hierarchical scheme for one reason or another. In most instances, though, it is unclear whether these represent genuine exceptions to a strict hierarchy rather than inaccuracies or uncertainities in the reported assignment.

/pdf/distributed-hierarchical-processing-in-the-primate-cerebral-1dvblm6d82.pdf

Distributed Hierarchical Processing in the Primate Cerebral Cortex

A theory of edge detection is presented. The analysis proceeds in two parts. (1) Intensity changes, which occur in a natural image over a wide range of scales, are detected separately at different scales. An appropriate filter for this purpose at a given scale is found to be the second derivative of a Gaussian, and it is shown that, provided some simple conditions are satisfied, these primary filters need not be orientation-dependent. Thus, intensity changes at a given scale are best detected by finding the zero values of delta 2G(x,y)*I(x,y) for image I, where G(x,y) is a two-dimensional Gaussian distribution and delta 2 is the Laplacian. The intensity changes thus discovered in each of the channels are then represented by oriented primitives called zero-crossing segments, and evidence is given that this representation is complete. (2) Intensity changes in images arise from surface discontinuities or from reflectance or illumination boundaries, and these all have the property that they are spatially. Because of this, the zero-crossing segments from the different channels are not independent, and rules are deduced for combining them into a description of the image. This description is called the raw primal sketch. The theory explains several basic psychophysical findings, and the operation of forming oriented zero-crossing segments from the output of centre-surround delta 2G filters acting on the image forms the basis for a physiological model of simple cells (see Marr & Ullman 1979).

http://www.hms.harvard.edu/bss/neuro/bornlab/qmbc/beta/day4/marr-hildreth-edge-prsl1980.pdf

Theory of Edge Detection

Many current neurophysiological, psychophysical, and psychological approaches to vision rest on the idea that when we see, the brain produces an internal representation of the world. The activation of this internal representation is assumed to give rise to the experience of seeing. The problem with this kind of approach is that it leaves unexplained how the existence of such a detailed internal representation might produce visual consciousness. An alternative proposal is made here. We propose that seeing is a way of acting. It is a particular way of exploring the environment. Activity in internal representations does not generate the experience of seeing. The outside world serves as its own, external, representation. The experience of seeing occurs when the organism masters what we call the governing laws of sensorimotor contingency. The advantage of this approach is that it provides a natural and principled way of accounting for visual consciousness, and for the differences in the perceived quality of sensory experience in the different sensory modalities. Several lines of empirical evidence are brought forward in support of the theory, in particular: evidence from experiments in sensorimotor adaptation, visual \"filling in,\" visual stability despite eye movements, change blindness, sensory substitution, and color perception.

A sensorimotor account of vision and visual consciousness-Authors' Response-Acting out our sensory experience

Many current neurophysiological, psychophysical, and psychological approaches to vision rest on the idea that when we see, the brain produces an internal representation of the world. The activation of this internal representation is assumed to give rise to the experience of seeing. The problem with this kind of approach is that it leaves unexplained how the existence of such a detailed internal representation might produce visual consciousness. An alternative proposal is made here. We propose that seeing is a way of acting. It is a particular way of exploring the environment. Activity in internal representations does not generate the experience of seeing. The outside world serves as its own, external, representation. The experience of seeing occurs when the organism masters what we call the governing laws of sensorimotor contingency. The advantage of this approach is that it provides a natural and principled way of accounting for visual consciousness, and for the differences in the perceived quality of sensory experience in the different sensory modalities. Several lines of empirical evidence are brought forward in support of the theory, in particular: evidence from experiments in sensorimotor adaptation, visual “filling in,” visual stability despite eye movements, change blindness, sensory substitution, and color perception.

/pdf/a-sensorimotor-account-of-vision-and-visual-consciousness-1o8kf5szfs.pdf

A sensorimotor account of vision and visual consciousness

Neuronal Synchrony: A Versatile Code for the Definition of Relations?

We have studied the mechanism of contour perception by recording from neurons in the visual cortex of alert rhesus monkeys. In order to assess the relationship between neural signals and perception, we compared the responses to edges and lines with the responses to patterns in which human observers perceive a contour where no line or edge is given (anomalous contour), such as the border between gratings of thin lines offset by half a cycle. With only one exception out of 60, orientation-selective neurons in area V1 did not signal the anomalous contour. Many neurons failed to respond to this stimulus at all, others responded according to the orientation of the grating lines. In area V2, 45 of 103 neurons (44%) signaled the orientation of the anomalous contour. Sixteen did so without signaling the orientation of the inducing lines. Some responded better to anomalous contours than to the optimum bars or edges. Preferred orientations and widths of tuning for anomalous contour and bar or edge were found to be highly correlated, but not identical, in each neuron. Similar to perception, the neuronal responses depended on a minimum number of lines inducing the contour, but not so much on line spacing, and tended to be weaker when the lines were oblique rather than orthogonal to the border. With oblique lines, the orientations signaled were biased towards the orientation orthogonal to the lines, as in the Zollner illusion. We conclude that contours may be defined first at the level of V2. While the unresponsiveness of neurons in V1 to this type of anomalous contour is in agreement with linear filter predictions, the responses of V2 neurons need to be explained. We assume that they sum the signals of 2 parallel paths, one that defines edges and lines and another that defines anomalous contours by pooling signals from end-stopped receptive fields oriented mainly orthogonal to the contour.

https://www.jneurosci.org/content/jneuro/9/5/1731.full.pdf

Mechanisms of contour perception in monkey visual cortex. I. Lines of pattern discontinuity

We have studied the mechanism of contour perception by recording from neurons in the visual cortex of alert rhesus monkeys. We used stimuli in which human observers perceive anomalous contours: A moving pair of notches in 2 bright rectangles mimicked an overlaying dark bar. For control, the notches were closed by thin lines so that the anomalous contours disappeared or half of the figure was blanked, with a similar effect. Orientation-selective neurons were studied. With the receptive fields centered in the gap, 23 of 72 (32%) neurons tested in area V2 responded to the moving “bar” even though the stimulus spared their response fields, and when the notches were closed, their responses were reduced or abolished. Likewise, when half of the figure was removed, the neurons usually failed to respond. Neurons with receptive fields within 4 degrees of the fovea signaled anomalous contours bridging gaps of 1 degree-3.5 degrees. The anomalous-contour responses were compared quantitatively with response field profiles and length-summation curves and found to exceed the predictions by linear-summation and summation- to-threshold models. Summation models also fail to explain the effect of closing lines which add only negligible amounts of light. In V1, only one of 26 neurons tested showed comparable responses, and only with a narrow gap. The others responded only when the stimulus invaded the response field and did not show the effect of closing lines, or failed to respond at all. The contour responses in V2, the nonadditivity, and the effect of closure can be explained by the previously proposed model (Peterhans et al., 1986), assuming that the corners excite end-stopped fields orthogonal to the contour whose signals are pooled in the contour neurons.

https://www.jneurosci.org/content/jneuro/9/5/1749.full.pdf

Mechanisms of contour perception in monkey visual cortex. II. Contours bridging gaps

We studied the relation between anatomical structure and functional properties of cells in area V2 of the macaque. Visual function was assessed in the alert animal during fixation of gaze. Recording sites were reconstructed with respect to cortical lamination and the cytochrome oxidase pattern. We measured orientation and direction selectivity, end-stopping, sensitivity to binocular disparity and ocular dominance, and determined more complex functions like sensitivity to anomalous contours and lines defined by coherent motion. Orientation selectivity was found in all parts of area V2, with high frequencies in the pale and thick stripes of the cytochrome oxidase pattern, and with lower frequency in the thin stripes. Representations of anomalous contours were found in the pale and thick stripes with similar frequencies, but generally not in the thin stripes, which have been thought to process colour. Lines defined by coherent motion were most frequently represented in the thick stripes; they were less frequent in the pale stripes, and (as with anomalous contours) were not found in the thin stripes. Sensitivity to binocular disparity was found in all types of stripes, but more frequently in the thick stripes, where the exclusively binocular neurons were also concentrated. By contrast, no segregation was found for direction selectivity and end-stopping. All neuronal properties were distributed evenly across cortical laminae. We conclude that mechanisms for figure-ground segregation involve the pale and the thick stripes of the cytochrome oxidase pattern, perhaps with greater emphasis on 'shape from motion' and 'stereoscopic depth' in the thick stripes, while more elementary neuronal properties are distributed almost evenly across the stripe pattern.

Functional Organization of Area V2 in the Alert Macaque

To study the visual processing of periodic and aperiodic patterns, we have analyzed neuronal responses in areas V1 and V2 of the visual cortex of alert monkeys during behaviorally induced fixation of gaze. Receptive field eccentricities ranged between 0.5 degrees and 4 degrees. We found cells that responded vigorously to gratings, but weakly or not all to bars and edges. In some cells the aperiodic stimuli even reduced the activity below the spontaneous level. The distribution of a bar-grating response index indicated a discrete population of "grating cells" characterized by more than 10-fold superiority of gratings. We estimated that these cells have a frequency of 4% in V1 and 1.6% in V2, and that about 4 million grafting cells of V1 subserve the central 4 degrees of vision. The converse, cells that responded to isolated bars but not to gratings of any periodicity, was also observed. The grating cells of V1 were mostly (23 of 26) found in layers 2, 3, and 4B. They preferred spatial frequencies between 2.6 and 19 cycles/degree (median, 9.3), with tuning widths at half-amplitude between 0.4 and 1.4 octaves (median, 1.0). Their tunings were narrower, and their preferred frequencies higher, than those of other cells on average. Grating cells were also narrowly tuned for orientation. Those of V2 were similarly selective. The responses of grating cells depended critically on the number of cycles of the gratings. With square waves of optimum periodicity responses required a minimum of 2-6 grating cycles and leveled off at 4-14 (median, 7.5). The corresponding receptive field widths were 0.34-2.4 degrees (median, 0.78 degrees) for V1 and 0.72-2.4 degrees (median, 1.4 degrees) for V2. Grating cells typically gave unmodulated responses to drifting gratings, were unselective for direction of motion, and were strongly activated also by stationary gratings. Half of those of V1 were monocular, the others binocular, some showing strong binocular facilitation and disparity sensitivity. Length summation was usually monotonic, but strong end-inhibition was also observed. In contrast to other cells, grating cells were not activated by harmonic components. Spatial-frequency response curves for sine-wave, square-wave, and line gratings were similar. Square-wave gratings of one-third the preferred frequency failed to excite the cells, while the isolated 3f component (f = the fundamental of the square wave) of these gratings evoked strong responses. In spite of the nonlinear features, grating cells had low contrast thresholds.(ABSTRACT TRUNCATED AT 400 WORDS)

https://www.jneurosci.org/content/jneuro/12/4/1416.full.pdf

Periodic-pattern-selective cells in monkey visual cortex.

1.

Receptive field correspondence and binocular interaction were studied in the cat striate cortex using fundus photography for correction of residual eye drifts.




2.

The scatter of incongruities (frequently called receptive field disparities) was found to be equal in horizontal and vertical direction (S.D. 0.5 °, N = 96). Incongruity was not correlated with field orientation.




3.

Using binocular stimulation, disparity tuning and binocular facilitation were analysed quantitatively in 46 cells. Seven of these were disparity specific, i.e. showed marked facilitation critically dependent on disparity.




4.

With monocular stimulation, three of the seven disparity specific cells could be driven only from one eye; one gave weak responses to both left and right monocular stimulation, and one no monocular responses at all.




5.

The disparity specific cells preferred vertical or oblique orientations, and vertical disparity was less critical than horizontal disparity. The disparity tuning curves, plotted with reference to the experimentally determined horopter, showed peaks within 0.5 ° crossed and 0.2 ° uncrossed disparity. The smallest half-widths at half-amplitude were 0.25 °. Some of the curves were asymmetric, or step-like, with a steep slope at zero disparity.




6.

It is concluded that stereoscopic depth may be signalled by disparity specific cells in area 17. These cells could account for stereoscopic acuity of a few min arc, if one assumes that depth is encoded in the amount of activity of single units, or in the graded imbalance of activity of antagonistic units.




7.

It is argued that the distribution of incongruities does not reflect stereoscopic function because the disparity specific cells often cannot be plotted in both eyes, and most of the units that can be plotted are not disparity selective.

R. von der Heydt

Papers

Mechanisms of contour perception in monkey visual cortex. I. Lines of pattern discontinuity

Mechanisms of contour perception in monkey visual cortex. II. Contours bridging gaps

Functional Organization of Area V2 in the Alert Macaque

Periodic-pattern-selective cells in monkey visual cortex.

Disparity sensitivity and receptive field incongruity of units in the cat striate cortex.