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

Circuitry and functional aspects of the insular lobe in primates including humans

The association cortices, those regions interposed between the primary sensory and primary motor areas, are thought to mediate a variety of cognitive functions. Among these, the prefrontal cortex has traditionally been considered the least understood and most complex. The only point that seems to marshall agreement among basic neuroscientists is that prefrontal functions and operations, acknowledged to be important in human behavior, are less accessible to analysis than are many other areas of the cortex. One reason for this perception may be the lack of an agreed upon anatomical blueprint that firmly establishes the relationship between prefrontal cortex and other, better understood, parts of the brain. However, this circumstance has changed radically in the last decade, and the structure and functions of the frontal lobes, particularly the prefrontal "silent" portion, which have intrigued investigators since the last century (Ferrier 1886, Bianchi 1895, Franz 1907), have recently again become the subject of intense interest (for reviews, see Damasio 1979, Fuster 1980, Goldman-Rakic 1984, 1987, Milner et al 1985, Mesulam 1986, Ingvar 1983, Stuss & Benson 1984). The availability of solid new findings in experimental animals and human patients and the promise of further discoveries are undoubtedly the basis of this renewed interest. Several extraordinarily influential papers paved the way for the modern era of frontal lobe research. Without a doubt, anatomical studies of pre­ frontal cortex were stimulated by Nauta's 1971 paper on "The Problem of the Frontal Lobe: A Reinterpretation." This review made an indelible impression both for its eloquence and fundamental validity, suggesting as

Topography of cognition: parallel distributed networks in primate association cortex.

Neurons in area 7a of the posterior parietal cortex of monkeys respond to both the retinal location of a visual stimulus and the position of the eyes and by combining these signals represent the spatial location of external objects. A neural network model, programmed using back-propagation learning, can decode this spatial information from area 7a neurons and accounts for their observed response properties.

A back-propagation programmed network that simulates response properties of a subset of posterior parietal neurons.

Previous research in non-human primates has shown that the superior longitudinal fascicle (SLF), a major intrahemispheric fiber tract, is actually composed of four separate components. In humans, only post-mortem investigations have been available to examine the trajectory of this tract. This study evaluates the hypothesis that the four subcomponents observed in non-human primates can also be found in the human brain using in vivo diffusion tensor magnetic resonance imaging (DT-MRI). The results of our study demonstrated that the four subdivisions could indeed be identified and segmented in humans. SLF I is located in the white matter of the superior parietal and superior frontal lobes and extends to the dorsal premotor and dorsolateral prefrontal regions. SLF II occupies the central core of the white matter above the insula. It extends from the angular gyrus to the caudal-lateral prefrontal regions. SLF III is situated in the white matter of the parietal and frontal opercula and extends from the supramarginal gyrus to the ventral premotor and prefrontal regions. The fourth subdivision of the SLF, the arcuate fascicle, stems from the caudal part of the superior temporal gyrus arches around the caudal end of the Sylvian fissure and extends to the lateral prefrontal cortex along with the SLF II fibers. Since DT-MRI allows the precise definition of only the stem portion of each fiber pathway, the origin and termination of the subdivisions of SLF are extrapolated from the available data in experimental material from non-human primates.

Segmentation of Subcomponents within the Superior Longitudinal Fascicle in Humans: A Quantitative, In Vivo, DT-MRI Study

The boundaries of the second somatic sensory cortex (SII) in primates are difficult to define physiologically because cutaneous stimulation activates several regions around SII that do not receive projections from the ventroposterior nucleus of the thalamus. These cortical regions, which include portions of area 7b, the retroinsular (Ri) and postauditory fields (PA), and the granular insula (Ig) are largely buried within the lateral sulcus and most lie posterior to the caudal end of the insula. The differences in somatic activity in these various cortical fields in the unanesthetized cynomolgus monkey became apparent only after the properties of many neighboring neurons could be compared. Receptive fields for area 7b and Ig neurons were generally large (< 10 cm2), with bilateral, moderately defined boundaries; some neurons in area 7b had receptive fields with labile borders as a function of wakefulness. In contrast, receptive fields for Ri neurons were generally (< 10 cm2 and contralateral, with stable, well-defined boundaries. Taken as an ensemble, the neurons in areas neighboring SII exhibited a very crude topography; but at the level of an individual neuron and its neighbor, there was never a pattern of gradual transition in peripheral receptive field locations between one unit and the next, like that seen in SII. In area 7b, this crude map was organized mediolaterally across the inferior parietal lobule and into the upper bank of the lateral sulcus, with the head represented medially and the lower trunk and hindlimb laterally. In Ri-PA, an anteroposterior organization was noted along the fundus of the lateral sulcus with the head represented anterior to the lower trunk and hindlimb. No organization was apparent in Ig. Additional sensitivity to visual stimuli was noted in the more medial aspects of area 7b that were located on the exposed inferior parietal lobule. Sensitivity to auditory stimuli was principally found in PA and occasionally in Ri. The results, especially from area 7b, are discussed with respect to previous notions about the organization of SII.

Organization of somatosensory receptive fields in cortical areas 7b, retroinsula, postauditory and granular insula of M. fascicularis.

The second somatosensory area (SII) of awake, untrained cynomolgus monkeys was surveyed with recordings from nearly 1,000 single neurons. A detailed somatotopographic organization could be demonstrated in SII because the majority of these neurons had contralateral, moderate to well-defined receptive fields of < 10 cm2, and because neighboring neurons possessed receptive fields that were only slightly displaced from one another. Different body regions were represented in successive anterior to posterior strips that were oriented across the parietal operculum with an anterolateral to posteromedial slant. Neurons with trigeminal receptive fields were found in the anterior portion of SII; these neurons were the only ones in SII with predominantly bilateral receptive fields (r.f.'s.). Neurons with digit or hand r.f'.s form the largest component of the map, and were located posterior to those with face r.f.s. Most of these neurons had only contralateral activation. The hand and digit region was followed in turn by the arm, the upper and lower trunk, and the hindlimb regions. Although the overall SII orientation was along an anterior-posterior gradient, recordings at individual coronal planes often demonstrated isolated sequences of receptive fields that exhibited a medial-lateral progression. The principle example of this latter gradient was seen in the forelimb region where digits one through five were represented in an overlapping sequence across the parietal operculum. Except for portions of the digit representation, neighboring sequences of neurons in SII do not form a precise topologic map of the body that is comparable to the somatotopic maps observed in areas 3b and 1. The present findings contrast with previous physiological studies of SII in the primate. These discrepancies are discussed in relation to methodological differences and in terms of distinctions used to define the boundaries of SII.

Somatotopographic organization in the second somatosensory area of M. fascicularis.

Somatic response properties were determined for over 1,300 neurons isolated within and near the lateral sulci of unanesthetized and unparalyzed cynomolgus monkeys. Somatic stimuli unequivocally activated the majority of units studied in SII (93%) and in cortical fields surrounding SII: area 7b (65%), the retroinsular field (74%), and the granular insula (76%). No activation other than somatic was seen for SII neurons, and noxious somatic stimulation was rarely required. The SII units almost always responded in a rapidly adapting manner to hair or skin stimulation, but not both; however, the submodality distribution seen in SII varied as a function of peripheral receptor locations. Two small zones within SII contained neurons that responded only if the animal actively interacted with the stimulus. In contrast, one-half of the sample of neurons from area 7b unequivocally responded only to somatic stimulation. Although many neurons in the lateral parts of area 7b were vigorously activated by innocuous tactile stimulation, others demonstrated little association with an identifiable somatic submodality, had sluggish responses, required complex, noxious, visual or other non-somatic stimuli for activation, and had labile response properties and receptive fields. Indeed, the responses of some area 7b neurons suggested a possible relationship with the animal's attention towards or anticipation of a noxious or a novel somatic stimulus. Neurons within the retroinsular cortex (Ri), which receives projections from the posterior nucleus (PO), primarily responded to light tactile stimulation of rapidly adapting skin receptors; less than 3% responded to moderate or high threshold mechanical stimulation. The sensitivity to tactile stimulation in Ri closely resembled the responses of SII neurons. Neurons in the granular insula (Ig) often responded to gentle hair deflection within receptive fields covering large areas of the body. Ig and area 7b were the principle loci within the lateral sulcus that contained neurons responding to noxious stimulation. Owing to the great similarity in the somatic response properties within these areas in the awake and unparalyzed animal, the designation of cortical areas could only be made after correlating the recording sites with connectional and cytoarchitectonic analyses in the same animal. Consequently, previous physiological studies may have attributed to SII some of the response characteristics of neurons in neighboring areas.

Somatic submodality distribution within the second somatosensory (SII), 7b, retroinsular, postauditory, and granular insular cortical areas of M. fascicularis.

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Organization of somatosensory receptive fields in cortical areas 7b, retroinsula, postauditory and granular insula of M. fascicularis.

Somatotopographic organization in the second somatosensory area of M. fascicularis.

Somatic submodality distribution within the second somatosensory (SII), 7b, retroinsular, postauditory, and granular insular cortical areas of M. fascicularis.