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

The two basic phenomena that define the problem of visual attention can be illustrated in a simple example. Consider the arrays shown in each panel of Figure 1. In a typical experiment, before the arrays were presented, subjects would be asked to report letters appearing in one color (targets, here black letters), and to disregard letters in the other color (nontargets, here white letters). The array would then be briefly flashed, and the subjects, without any opportunity for eye movements, would give their report. The display mimics our. usual cluttered visual environment: It contains one or more objects that are relevant to current behavior, along with others that are irrelevant. The first basic phenomenon is limited capacity for processing information. At any given time, only a small amount of the information available on the retina can be processed and used in the control of behavior. Subjectively, giving attention to any one target leaves less available for others. In Figure 1, the probability of reporting the target letter N is much lower with two accompa­ nying targets (Figure la) than with none (Figure Ib). The second basic phenomenon is selectivity-the ability to filter out un­ wanted information. Subjectively, one is aware of attended stimuli and largely unaware of unattended ones. Correspondingly, accuracy in identifying an attended stimulus may be independent of the number of nontargets in a display (Figure la vs Ie) (see Bundesen 1990, Duncan 1980).

Neural Mechanisms of Selective Visual Attention

The measurement of meaning

Functional brain imaging in humans has revealed task-specific increases in brain activity that are associated with various mental activities. In the same studies, mysterious, task-independent decreases have also frequently been encountered, especially when the tasks of interest have been compared with a passive state, such as simple fixation or eyes closed. These decreases have raised the possibility that there might be a baseline or resting state of brain function involving a specific set of mental operations. We explore this possibility, including the manner in which we might define a baseline and the implications of such a baseline for our understanding of brain function.

https://www.biac.duke.edu/education/courses/fall04/fmri/readings/week10/2001_NatRevNeuro_Gusnard.pdf

Searching for a baseline: Functional imaging and the resting human brain

The borders of human visual areas V1, V2, VP, V3, and V4 were precisely and noninvasively determined. Functional magnetic resonance images were recorded during phase-encoded retinal stimulation. This volume data set was then sampled with a cortical surface reconstruction, making it possible to calculate the local visual field sign (mirror image versus non-mirror image representation). This method automatically and objectively outlines area borders because adjacent areas often have the opposite field sign. Cortical magnification factor curves for striate and extrastriate cortical areas were determined, which showed that human visual areas have a greater emphasis on the center-of-gaze than their counterparts in monkeys. Retinotopically organized visual areas in humans extend anteriorly to overlap several areas previously shown to be activated by written words.

https://www.cogsci.ucsd.edu/~sereno/papers/HumanRetin95.pdf

Borders of multiple visual areas in humans revealed by functional magnetic resonance imaging

The representation of the visual field in areas V3 and V4 of the macaque was mapped with multiunit electrodes. Twelve Macaca fascicularis were studied in repeated recording sessions while immobilized and anesthetized. V3 is a narrow strip (4-5 mm wide) of myeloarchitectonically distinct cortex located immediately anterior to V2. It contains a systematic representation of the central 35-40 degrees of the contralateral visual field; the representation of the upper quadrant is located ventrally in the hemisphere and that of the lower quadrant, dorsally. There is a small gap between the dorsal (V3d) and ventral (V3v) portions of V3. The representation of the horizontal meridian is adjacent to that in V2 and forms the posterior border of both V3d and V3v. Most or all of the anterior border of V3d consists of the representation of the lower vertical meridian. The entire anterior border of V3v consists of the representation of the upper vertical meridian. V4 is a strip of myeloarchitectonically distinct cortex 5-8 mm wide, immediately anterior to V3. It contains a coarse, but systematic, representation of approximately the central 35-40 degrees of the contralateral visual field. The representation of the upper visual field is located ventrally in the hemisphere. Most of the representation of the lower visual field is located dorsally. The posterior border of V4 corresponds to the representation of the vertical meridian, and the representation of the horizontal meridian is located at or near its anterior border. In both V3 and V4, the representation of the central visual field is magnified relative to that of the periphery. In both areas, the size of receptive fields increases with increasing eccentricity; however, at a given eccentricity, the receptive fields of V4 are larger than those of V3.

https://www.jneurosci.org/content/jneuro/8/6/1831.full.pdf

Visuotopic organization and extent of V3 and V4 of the macaque

The representation of the visual field in the area adjacent to striate cortex was mapped with multiunit electrodes in the macaque. The animals were immobilized and anesthetized and in each animal 30 to 40 electrode penetrations were typically made over several recording sessions. This area, V2, contains a topographically organized representation of the contralateral visual field up to an eccentricity of at least 80 degrees. The representation of the vertical meridian is adjacent to that in striate cortex (V1) and forms the posterior border of V2. The representation of the horizontal meridian in V2 forms the anterior border of V2 and is split so that the representation of the lower visual field is located dorsally and that of the upper field ventrally. As in V1, the representation of the central visual field is magnified relative to that of the periphery. The area of V2 is slightly smaller than that of V1. At a given eccentricity, receptive field size in V2 is larger than in V1. The myeloarchitecture of V2 is distinguishable from that of the surrounding cortex. The location of V2 corresponds, at least approximately, to that of cytoarchitectonic Area OB. V2 is bordered anteriorly by several other areas containing representations of the visual field.

/pdf/visual-topography-of-v2-in-the-macaque-3rvwzfzkbt.pdf

Visual Topography of V2 in the Macaque

We have examined the origin and topography of cortical projections to area PO, an extrastriate visual area located in the parieto-occipital sulcus of the macaque. Distinguishable retrograde fluorescent tracers were injected into area PO at separate retinotopic loci identified by single-neuron recording. The results indicate that area PO receives retinotopically organized inputs from visual areas V1, V2, V3, V4, and MT. In each of these areas the projection to PO arises from the representation of the periphery of the visual field. This finding is consistent with neurophysiological data indicating that the representation of the periphery is emphasized in PO. Additional projections arise from area MST, the frontal eye fields, and several divisions of parietal cortex, including four zones within the intraparietal sulcus and a region on the medial dorsal surface of the hemisphere (MDP). On the basis of the laminar distribution of labeled cells we conclude that area PO receives an ascending input from V1, V2, and V3 and receives descending or lateral inputs from all other areas. Thus, area PO is at approximately the same level in the hierarchy of visual areas as areas V4 and MT. Area PO is connected both directly and indirectly, via MT and MST, to parietal cortex. Within parietal cortex, area PO is linked to particular regions of the intraparietal sulcus including VIP and LIP and two newly recognized zones termed here MIP and PIP. The wealth of connections with parietal cortex suggests that area PO provides a relatively direct route over which information concerning the visual field periphery can be transmitted from striate and prestriate cortex to parietal cortex. In contrast, area PO has few links with areas projecting to inferior temporal cortex. The pattern of connections revealed in this study is consistent with the view that area PO is primarily involved in visuospatial functioning.

Topographical organization of cortical afferents to extrastriate visual area PO in the macaque: a dual tracer study.

SUMMARY AND CONCLUSIONS 1 The representation of the visual field in the striate projection zone in the posterior portion of the superior temporal sulcus of the macaque (MT) was mapped with multiunit electrodes The animals were immobilized and anesthetized and in each animal 25-35 electrode penetrations were typically made over several recording sessions 2 MT contains a representation of virtually the entire contralateral visual field The representation of the vertical meridian forms its ventrolateral border and lies near the bottom of the lower bank of the superior temporal sulcus (STS) The representation of the horizontal meridian runs across the floor of STS The upper field is located ventral and anterior and the lower field dorsal and posterior The medial border lies at the junction of the floor of STS and its upper bank 3 MT is similar to striate cortex in being a first-order transformation of the visual field In both areas, receptive-field size and cortical magnification increase with eccentricity MT is much smaller than striate cortex and has much larger receptive fields at a given eccentricity and a cruder topography 4 The results further support the suggestion that MT in the macaque is homologous to visual area MT in New World primates

Visual topography of striate projection zone (MT) in posterior superior temporal sulcus of the macaque.

To determine the locus, full extent, and topographic organization of cortical connections of area V4 (visual area 4), we injected anterograde and retrograde tracers under electrophysiological guidance into 21 sites in 9 macaques. Injection sites included representations ranging from central to far peripheral eccentricities in the upper and lower fields. Our results indicated that all parts of V4 are connected with occipital areas V2 (visual area 2), V3 (visual area 3), and V3A (visual complex V3, part A), superior temporal areas V4t (V4 transition zone), MT (medial temporal area), and FST (fundus of the superior temporal sulcus [STS] area), inferior temporal areas TEO(cytoarchitectonicareaTEOinposteriorinferiortemporalcortex) and TE (cytoarchitectonic area TE in anterior temporal cortex), and the frontal eye field (FEF). By contrast, mainly peripheral field representations of V4 are connected with occipitoparietal areas DP (dorsalprelunatearea),VIP(ventralintraparietalarea),LIP(lateralintraparietal area), PIP (posterior intraparietal area), parieto-occipital area, and MST (medial STS area), and parahippocampal area TF (cytoarchitectonic area TF on the parahippocampal gyrus). Based on the distribution of labeled cells and terminals, projections from V4 to V2 and V3 are feedback, those to V3A, V4t, MT, DP, VIP, PIP, and FEF aretheintermediatetype,andthosetoFST,MST,LIP,TEO,TE,andTF are feedforward. Peripheral field projections from V4 to parietal areas could provide a direct route for rapid activation of circuits serving spatial vision and spatial attention. By contrast, the predominance of central field projections from V4 to inferior temporal areasisconsistent withtheneedfordetailedformanalysisforobject vision.

Ricardo Gattass

Papers

Visuotopic organization and extent of V3 and V4 of the macaque

Visual Topography of V2 in the Macaque

Topographical organization of cortical afferents to extrastriate visual area PO in the macaque: a dual tracer study.

Visual topography of striate projection zone (MT) in posterior superior temporal sulcus of the macaque.

Cortical Connections of Area V4 in the Macaque