Studies on the evolution of multiple somatosensory representations in primates: the organization of anterior parietal cortex in the New World Callitrichid, Saguinus

Because members of the New World family, Callithricidae, are generally regarded as the most primitive of monkeys, we studied the organization of somatosensory cortex in the tamarin (Saguinus) in hopes of better understanding differences in the organization of anterior parietal cortex in primates and how these differences relate to phylogeny. In most prosimian primates only one complete representation of cutaneous receptors has been found in the region of primary cortex, S-I, while in all Old and New World monkeys studied to date, two cutaneous representations exist in distinct architectonic fields, areas 3b and 1. In detailed microelectrode mapping studies in anesthetized tamarins, only one complete representation responsive to low-threshold cutaneous stimulation was evident in the S-I region. This topographic representation was in a parietal koniocortical field that architectonically resembles area 3b of other monkeys, and the general somatotopic organization of the field was similar to that of area 3b of other monkeys. Cortex rostral to the single representation was generally unresponsive to somatosensory stimuli, or required more intense stimulation for neural activation. Cortex caudal to the representation, in the region of area 1 of other monkeys, was generally either unresponsive or responded to only high-threshold stimulation, although some recording sites were activated by low-threshold tactile stimulation. The present evidence, together with that from previous studies, suggests that the single, complete body surface representation in Saguinus is homologous to the S-I representation found in some prosimians (Galago, Perodicticus) and the area 3b cutaneous representation found in New World Cebidae (Aotus, Saimiri, and Cebus) and Old World Macaca. Cortex rostral to S-I in Saguinus has the appearance of areas 3a and 4 of other primates. The cortex caudal to S-I in Saguinus, while resembling area 1 in some ways, does not have all of the features of area 1 of other monkeys. In particular, the field was not easily activated by low-threshold cutaneous stimuli, as area 1 is in other monkeys, and therefore a second cutaneous representation of all body parts was not demonstrated. Thus, cortex in the expected location of area 1 of Saguinus was not as responsive as area 1 of other monkeys, and it somewhat resembled the high-threshold fringe zones found caudal to S-I in anesthetized prosimians and some nonprimates. The results raise the possibility that the area 1 cutaneous representation that is characteristic of other New World monkeys and Old World monkeys evolved from a less responsive precursor along the caudal border of S-I in early monkeys.     

Representation of the body surface in somatosensory area I of tree shrews,Tupaia glis

Microelectrode mapping methods were used to determine the organization of the first somatosensory area, S-I, of tree shrews. Tree shrews were chosen for study because of their generalized body form, phylogenetic relationship to primates, and smooth, easily mapped cortex. A systematic representation of the contralateral body surface was found in an architectonically distinct zone identified as somatic koniocortex. Overall features of somatotopic organization were similar to S-I of other mammals, S-I of prosimian primates, and the Area 3b “S-I proper” representation of monkeys. Like Area 3b in monkeys and the somatic koniocortex in galagos, S-I in tree shrews is bordered caudally by cortex also responsive to somatosensory stimuli. Several aspects of S-I organization in tree shrews appear to be primitive and generalized. These include the representation of the trunk with the ventrum at the caudal margin of S-I, the restriction of the glabrous digits of the hand and foot to the rostral half of the representation and pointed rostralward, the representation of an anterior strip of the forelimb lateral to the hand, and a posterior strip of hindlimb medial to the foot representation. As in a number of other mammals, a large portion of S-I in tree shrews is devoted to the head. However, the proportion of S-I activated from the glabrous nose is greater in tree shrews than in any previously studied mammal. We conclude that S-I of tree shrews has both specialized and generalized features, and that the generalized features importantly relate to an understanding of the evolution of the primate somatosensory system.     

Thalamic connections of three representations of the body surface in somatosensory cortex of gray squirrels

The anatomical tracer, wheat germ agglutinin, was used to determine the connections of electrophysiologically identified locations in three architectonically distinct representations of the body surface in the somatosensory cortex of gray squirrels. Injections in the first somatosensory area, S-I, revealed reciprocal connections with the ventroposterior nucleus (VP), a portion of the thalamus just dorsomedial to VP, the posterior medial nucleus, Pom, and sometimes the ventroposterior inferior nucleus (VPI). As expected, injections in the representation of the face in S-I resulted in label in ventroposterior medial (VPM), the medial subnucleus of VP, whereas injections in the representation of the body labeled ventroposterior lateral (VPL), the lateral subnucleus of VP. Furthermore, there was evidence from connections that the caudal face and head are represented dorsolaterally in VPM, and the forelimb is represented centrally and medially in VPL. The results also support the conclusion that a representation paralleling that in VP exists in Pom, so that the ventrolateral part of Pom represents the face and the dorsomedial part of Pom is devoted to the body. Because connections with VPI were not consistently revealed, the possibility exists that only some parts or functional modules of S-I are interconnected with VPI. Two separate small representations of the body surface adjoin the caudoventral border of S-I. Both resemble the second somatosensory area, S-II, enough to be identified as S-II in the absence of evidence for the other. We term the more dorsal of the two fields S-II because it was previously defined as S-II in squirrels (Nelson et al., '79), and because it more closely resembles the S-II identified in most other mammals. We refer to the other field as the parietal ventral area, PV (Krubitzer et al, '86). Injections in S-II revealed reciprocal connections with VP, Pom, and a thalamic region lateral and caudal to Pom and dorsal to VP, the posterior lateral nucleus, Pol. Whereas major interconnections between S-II and VPI have been reported for cats, raccoons, and monkeys, no such interconnections were found for S-II in squirrels. The parietal ventral area, PV, was found to have prominent reciprocal interconnections with VP, VPI, and the internal (magnocellular) division of the medial geniculate complex (MGi). The pattern of connections conforms to the established somatotopic organization of VP and suggests a crude parallel somatotopic organization in VPI. Less prominent interconnections were with Pol.(ABSTRACT TRUNCATED AT 400 WORDS)     

The representation of the body surface in S-I of cats

In both cats and monkeys, the traditional region of the first somatosensory area of cortex, S-I, has been described as containing four strip-like architectonic fields, areas 3a, 3b, 1, and 2. In monkeys, a number of recent studies have provided evidence that each of these architectonic fields constitutes a separate representation of the body. Because of the observations in monkeys, we decided to re-examine the S-I region of cats to determine whether the evidence supported the traditional concept of a single representation, or the existence of several representations related to the described architectonic fields. Microelectrode multiunit mapping techniques were used to determine the somatotopic organization of the S-I region of 10 cats. The results indicate that a single representation of the body surface occupies most or all of the traditional S-I region including cortex defined as area "3b," area "1," and much of area "2," but excluding area "3a." Neurons throughout this single representation were activated by cutaneous stimuli, indicating that all parts of S-I receive input from cutaneous receptors. Neurons in area 3a were activated by inputs from deep receptors, as reported by others. Neurons caudal to S-I were activated by cutaneous stimuli and appeared to constitute the additional body surface representations of S-II and possibly S-III. Thus, the significance of the architectonic fields "3b," "1," and "2" is quite different in cats than in monkeys. We propose that most or all of these three fields, as described in cats, constitute the homologue of area 3b in monkeys.     

The somatosensory thalamus of monkeys: cortical connections and a redefinition of nuclei in marmosets.

Thalamic connections of three subdivisions of somatosensory cortex in marmosets were determined by placing wheatgerm agglutinin conjugated with horseradish peroxidase and fluorescent dyes as tracers into electrophysiologically identified sites in S-I (area 3b), S-II, and the parietal ventral area, PV. The relation of the resulting patterns of transported label to the cytoarchitecture and cytochrome oxidase architecture of the thalamus lead to three major conclusions. 1) The region traditionally described as the ventroposterior nucleus (VP) is a composite of VP proper and parts of the ventroposterior inferior nucleus (VPi). Much of the VP region consists of groups of densely stained, closely packed neurons that project to S-I. VPi includes a ventral oval of pale, less densely packed neurons and finger-like protrusions that extend into VP proper and separate clusters of VP neurons related to different body parts. Neurons in both parts of VPi project to S-II rather than S-I. Connection patterns indicate that the proper and the embedded parts of VPi combine to form a body representation paralleling that in VP. 2) VPi also provides the major thalamic input into PV. 3) In architecture, location, and cortical connections, the region traditionally described as the anterior pulvinar (AP) of monkeys resembles the medial posterior nucleus, Pom, of other mammals and we propose that all or most of AP is homologous to Pom. AP caps VP dorsomedially, has neurons that are moderately dense in Nissl staining, and reacts moderately in CO preparations. AP neurons project to S-I, S-II, and PV in somatotopic patterns.     

Physiological and anatomical evidence for a discontinuous representation of the trunk in S-I of tree shrews

Microelectrode mapping methods revealed that the representation of the body surface in the first somatosensory area of cortex, S-I, of the tree shrew is unique in that only the ventral trunk was found in the usual location of the trunk representation in cortex of the dorsolateral surface of the cerebral hemisphere. Instead, the dorsal trunk was found as an extension of the representation of the posterior leg in cortex on the medial wall. The separation of the representation of the trunk occurs along a line that is counter to the orientation of the dorsal root dermatomes, so that S-I of the tree shrew clearly cannot be characterized as a serial representation of dermatomes. Anatomical studies of connections support the conclusion that the representation of the trunk is split in S-I. Both the representation of the dorsal trunk on the medial wall of the cerebral hemisphere and S-I of the dorsolateral surface were found to project to S-II when horseradish peroxidase was injected into S-II.     

The organization and connections of somatosensory cortex in marmosets

Microelectrode mapping methods were used to define and describe 3 representations of the body surface in somatosensory cortex of marmosets: S-I proper or area 3b of anterior parietal cortex, S-II, and the parietal ventral area (PV) of the upper bank of the lateral sulcus. In the same animals, injections of anatomical tracers were placed into electrophysiologically determined sites in area 3b or S-II. Mapping results and patterns of connections were later related to architectonic fields that were delimited in sections cut parallel to the surface of manually flattened cortex and stained for myelin. There were several major results. (1) Recordings from area 3b revealed a characteristic somatotopic organization of foot to face in a mediolateral sequence as previously reported in other members of the marmoset family (Carlson et al., 1986). (2) Multiple injections of WGA-HRP in area 3b demonstrated dense, patchy interconnections with ipsilateral S-II, PV, area 3a, and area 1, less dense interconnections with primary motor cortex (M-I), the supplementary motor area (SMA), limbic cortex of the medial wall (L), and rostrolateral parietal cortex of the lateral sulcus (PR), and callosal connections with areas 3b, S-II, and PV. Injections of 3 different tracers into the representation of 3 body regions in area 3b indicated that the connections with areas 3a, 3b, 1, S-II, and PV are topographically organized. (3) Recordings from cortex on the upper bank of the lateral sulcus demonstrated a somatotopic representation of the body surface that matches that of S-II of other mammals. S-II immediately adjoined areas 3b along the dorsal lip of the lateral sulcus. The face representation in S-II was adjacent to the face representation in 3b while the trunk, hindlimb, and forelimb were represented in a caudorostral sequence deeper in the sulcus. (4) Injections in S-II revealed ipsilateral connections with areas 3a, 3b, 1, a presumptive area 2, PV, PR, M-I, SMA, limbic cortex, the frontal eye fields, and the frontal ventral visual area. Dense callosal connections were with S-II and PV. (5) The recordings also revealed a systematic representation just rostral to S-II that has not been previously described in primates.(ABSTRACT TRUNCATED AT 400 WORDS)     

Interhemispheric connections of cortical sensory areas in tree shrews

Interhemispheric connections were studied in tree shrews (Tupaia belangeri) after multiple injections of horseradish peroxidase or horseradish peroxidase conjugated to wheat germ agglutinin into the cortex of one cerebral hemisphere. After an appropriate survival period, the areal pattern of connections was revealed by flattening the other hemisphere, cutting sections parallel to the cortical surface, and staining with tetramethylbenzidine. Architectonic boundaries were identified by using sections stained for myelinated fibers. Labeled cells and axon terminations formed largely overlapping distributions that covaried in density, although labeled cells appeared to be more evenly distributed than labeled terminations. Connections were concentrated along the border of area 17 (V-I) with area 18 (V-II). However, connections also extended as far as 2 mm into area 17 to include cortex representing parts of the visual field 10° or more from the zero vertical meridian. Clusters of dense connections spanned the width of area 18, where they alternated with regions of fewer connections. These clusters roughly corresponded in location to regions with heavier myelination. In the visually responsive temporal cortex, connections were also unevenly distributed. The organization of most of this cortex is not understood, but one subdivision, the temporal dorsal area (TD), has been identified on the basis of reciprocal connections with area 17. The central part of the TD had few interhemispheric connections, while most of the outer border had dense connections. The auditory cortex had dense and patchy connections throughout. The pattern in the primary somatosensory cortex (S-I) varied according to the representation of body parts, so that the cortex related to the forepaw had sparse connections, while connections were dense but uneven over much of the representation of the face, nose, and mouth. A focus of connections was found at the border of the forepaw and face representations, where the myelination of S-I cortex is interrupted. Dense, uneven connections also characterized the second somatosensory area, S-II. The motor cortex was densely connected, with only slightly fewer terminations rostral to the forepaw region of S-I. Other parts of frontal cortex had dense connections, The distribution of cortical connections varied with depth for at least some areas, so that clusters of cells and terminations were found in supragranular layers in S-I, S-II, and TD, while infragranular labeled cells were more evenly distributed. The results indicate that interhemispheric connections in tree shrews are widely distributed and include large portions of primary sensory fields, and that the primary somatic and visual areas have more interhemispheric connections than their homologues in higher primates. The local unevenness of the connections suggests that functions are unevenly distributed within cortical areas. Because visual and somatic areas representing the contralateral visual hemifield or body surface receive callosal inputs, many of these connections are not reflected in the excitatory receptive fields of cortical neurons.     

Connections of the ventral granular frontal cortex of macaques with perisylvian premotor and somatosensory areas: anatomical evidence for somatic representation in primate frontal association cortex

In macaque monkeys with injections of tritiated amino acids or horseradish peroxidase in the ventrolateral granular frontal cortex, we observed extensive anterograde and retrograde labeling of the premotor and somatosensory cortex in and around the lateral sulcus. Comparable labeling was not present with large and small control injections of the dorsal granular cortex. Cytoarchitectonic evaluation of the perisylvian cortex in the three cases examined in detail indicated that labeled areas included the ventral premotor cortex (area 6V); the precentral opercular and orbitofrontal opercular areas (PrCO and OFO); the second somatosensory area (S-II); the opercular cortex immediately anterior to S-II, possibly corresponding to area 2 of the S-I complex; and the central part of the insular cortex, including portions of the granular and dysgranular insular fields (Ig, Idg). Labeling was particularly dense and extensive in areas 6V, S-II, and OFO. Lighter labeling was also present in the rostral inferior parietal lobule (areas 7b and POa). The distribution of label within perisylvian areas was not uniform: certain parts were heavily labeled, while other parts were lightly labeled or unlabeled. Comparison of label distribution with published accounts of the somatotopy of these areas indicates that forelimb and orofacial representations were selectively labeled. Further, our results, taken together with other recent anatomical findings (e.g., Matelli et al.: Journal of Comparative Neurology 251:281-298, 1987; Barbas and Pandya: Journal of Comparative Neurology 256:211-228, 1987) suggest strongly that there is a network of interconnected forelimb and orofacial representations in macaque cortex, involving the ventral granular frontal cortex, area 6V, OFO, opercular area 2, S-II, the central insula, and area 7b. Each injection of frontal cortex which labeled the perisylvian somatic cortex involved the cortex of the ventral rim of the principal sulcus (PSvr). The cortex surrounding the PSvr does not stand out as a distinct area in Nissl-stained material. However, examination of myelin-stained sections prepared from uninjected hemispheres with the Gallyas technique revealed the existence of a distinct zone centered on the PSvr. This myeloarchitectonic area, which we term area 46vr, is more heavily myelinated than the ventral bank and fundus of the principal sulcus (area 46v) but is less heavily myelinated than the ventral (inferior) convexity (area 12). Involvement of area 46vr in our injections was probably responsible for the strong labeling observed in perisylvian somatic areas.(ABSTRACT TRUNCATED AT 400 WORDS)     

Representation of the body surface in somatic koniocortex in the prosimian Galago

Microelectrode mapping methods were used to determine the organization of somatosensory cortex in galagos, a prosimian primate. A systematic representation of the controlateral body surface was found within somatic koniocortex with an organization comparable to that of Area 3b of parietal cortex of monkeys (S-I proper) and primary somatosensory cortex (S-I) of other mammals. Limited studies of the response properties of single neurons within the representation revealed further similarities with the 3b field of monkeys. We conclude that somatic koniocortex of galagos and Area 3b of monkeys are homologous, and suggest the term S-I proper for the representation in both prosimians and monkeys. Differences in the details of S-I proper in galagos and monkeys suggest a more primitive organization in galagos.     

Somatotopic organization of the lateral sulcus of owl monkeys: area 3b, S-II, and a ventral somatosensory area

Multiunit microelectrode recordings and injections of horseradish peroxidase (HRP) were used to reveal neuron response properties, somatotopic organization, and interconnections of somatosensory cortex in the lateral sulcus (sylvian fissure) of New World owl monkeys. There were a number of main findings. 1) Representations of the face and head in areas 3b, 1, and S-II are found on the upper bank of the lateral sulcus. Most of the mouth and lip representations of area 3b were found in a rostral extension along the lip of the lateral sulcus. Adjacent cortex deeper in the lateral sulcus represented the nose, eye, ear, and scalp. 2) S-II was located on the upper bank of the lateral sulcus and extended past the fundus onto the deepest part of the lower bank. The face was represented most superficially in the sulcus, with the hand, foot, and trunk located in a rostrocaudal sequence deeper in the sulcus. The orientation of S-II is "erect," with the limbs pointing away from area 3b. 3) Neurons in S-II were activated by light tactile stimulation of the contralateral body surface. Receptive fields were several times larger than for area 3b neurons. 4) A 1-2-mm strip of cortex separating the face and hand representations in S-II was consistently responsive to the stimulation of deep receptors but was unresponsive to light cutaneous stimulation. 5) Injections of horseradish peroxidase in the electrophysiologically identified hand or foot representations of area 3b revealed somatotopically matched interconnections with mapped hand and foot representations in S-II. 6) A systematic representation of the body, termed the "ventral somatic" area, VS, was found extending laterally from S-II on the lower bank of the lateral sulcus. Within VS, the hand and foot were represented deep in the sulcus along the hand and foot regions of S-II, and the face was lateral near the ventral lip of the sulcus. 7) Neurons at most recording sites in the VS region were activated by contralateral cutaneous stimuli. However, a few sites had neurons with bilateral receptive fields. Receptive field sizes were comparable to those in S-II. In addition, neurons in islands of cortex in the VS region had properties that suggested that they were activated by pacinian receptors, while other regions were difficult to activate by light tactile stimuli but responded to stimuli that would activate deep receptors. 8) A few recording sites caudal to S-II on the upper bank of the lateral sulcus were responsive to somatic stimuli.(ABSTRACT TRUNCATED AT 400 WORDS)     

Corticocortical neurons projecting to the medial and lateral banks of the middle suprasylvian sulcus in the cat: An experimental study with the horseradish peroxidase method

Corticocortical afferents to both cortical walls of the cat middle suprasylvian sulcus (MSs area) were investigated by means of retrograde axonal transport of horseradish peroxidase (HRP). The visual cortex (V-I and V–II) projects to the medial wall of the MSs, the projection from V–II being heavier. The auditory cortex (A-I, A-II, and Ep), including cortical walls of the dorsal part of the anterior and posterior ectosylvian sulci, sends fibers to the lateral wall of the MSs. Connections from the first auditory area (A-I) are heavier than from the second (A-II). In the rostral part of the MSs, both the medial and lateral walls receive fibers from the somatosensory (S-I and S-II) cortex. A larger number of association fibers appear to arise from S-II than S-I. Although the MSs as a whole apparently receives various kinds of sensory inputs, there seems to be a parcellation of the MSs area such that the areas receiving cortical association fibers from the visual, auditory, or somatosensory cortical areas also receive thalamic projections from those parts of the thalamus receiving sensory connections of the same modality. The cells of origin of the association fibers were mostly pyramidal, the majority located in layer III (e.g., 80% in the visual cortex and 74% in the auditory cortex), some in layer V, and a few in other layers. Most (76–79%) of the labeled cell bodies were of 15–20 μm diameter. Smaller (8–15 μm) and larger (20–26 μm) cells constituted less than 15% in each case. The mean diameters were 17.0 ± 2.8 μm (SD) in the visual cortex and 17.7 ± 3.2 μm (SD) in the auditory cortex.     

Barrels and septa: separate circuits in rat barrels field cortex.

The neural circuitry within sensory cortex determines its functional properties, and different solutions have evolved for integrating the activity that arises from an array of sensory inputs to cortex. In rodent, circumscribed receptors, such as whiskers, are represented in somatic sensory (S-I) cortex in islands of cells in layer IV called "barrels" surrounded by narrow channels that separate barrels called "septa." These two cortical domains were previously shown to receive sensory inputs through parallel subcortical pathways. Here, by using small biocytin injections, we demonstrate that distinct intrinsic and corticocortical circuitries arise from barrel and septal columns. The intracortical S-I projections originating from barrel columns are rather short-ranged, terminating for the most part within the far boundaries of the most immediate neighboring barrel columns, whereas corticocortical projections reach the second somatosensory (S-II) cortex. In contrast, the intrinsic projections arising from septal columns extend two to three barrels' distance along the row of whisker representation, producing terminals preferentially in other septal columns. Septal corticocortical projections terminate in the dysgranular cortex anterior to E-row barrels and in the posteromedial parietal cortex in addition to S-II. Whereas layer IV barrels are largely isolated from lateral connections, septa are the main conduits of intracortical projections arising from neighboring barrel and septal columns. These results indicate that the two subcortical pathways from whiskers to cortex continue as two distinct partially segregated pathways in cortex.     

Relative contributions of SII and area 5 to tactile discrimination in monkeys

Rhesus monkeys with ablations of either the second somatosensory cortex (SII) or of the superior parietal lobule (area 5) were tested on a battery of tactile discrimination tasks in order to help determine which of these areas might constitute part of a postulated cortico-limbic tactile processing pathway. Monkeys with ablations of SII were severely impaired on both texture and shape discrimination learning and had markedly elevated size and roughness discrimination thresholds relative to control animals. By contrast, monkeys with area 5 lesions were impaired only on roughness thresholds, and these were elevated only moderately. Although more severe tactile deficits following lesions of area 5 have been reported previously, they were found in the present study only when the area 5 removals were extended slightly rostrally, in a third operated group, to include the posteromedial part of the hand representation of area 2. These results are consistent with the suggestion that SII, but not area 5, is a critical station in a tactile processing pathway that proceeds from the primary somatosensory cortex (SI) to the limbic structures of the temporal lobe through links in SII and the insular cortex.     

Thalamocortical connections of anterior and posterior parietal cortical areas in New World titi monkeys

We examined the thalamocortical connections of electrophysiologically identified locations in the hand and forelimb representations in areas 3b, 1, and 5 in the New World titi monkeys (Callicebus moloch), and of area 7b/AIP. Labeled cells and terminals in the thalamus resulting from the injections were related to architectonic boundaries. As in previous studies in primates, the hand representation of area 3b has dense, restricted projections predominantly from the lateral division of the ventral posterior nucleus (VPl). Projections to area 1 were highly convergent from several thalamic nuclei including the ventral lateral nucleus (VL), anterior pulvinar (PA), VPl, and the superior division of the ventral posterior nucleus (VPs). In cortex immediately caudal to area 1, what we term area 5, thalamocortical connections were also highly convergent and predominantly from nuclei of the thalamus associated with motor, visual, or somatic processing such as VL, the medial pulvinar (PM), and PA, respectively; with moderate projections from VP, central lateral nucleus (CL), lateral posterior nucleus (LP), and VPs. Finally, thalamocortical connections of area 7b/AIP were from a range of nuclei including PA, PM, LP/LD, VL, CL, PL, and CM. The current data support two conclusions drawn from previous studies in titi monkeys and other primates. First, cortex caudal to area 1 in New World monkeys is more like area 5 than area 2. Second, the presence of thalamic input to area 5 from both motor nuclei and somatosensory nuclei of the thalamus, suggests that area 5 could be considered a highly specialized sensorimotor area.     

Ablations of areas 3a and 3b of monkey somatosensory cortex abolish cutaneous responsivity in area 1

Cortex traditionally referred to as S-I in monkeys is a composite of 4 separate and complete representations of the contralateral body surface, one in each of the 4 cytoarchitectonic fields, areas 3a, 3b, 1, and 2. We investigated the significance of interconnections between these architectonic areas by assessing the immediate effects of ablations of parts of areas 3a and 3b on the responsivity of neurons in area 1. Ablations of specific parts of the hand representations in areas 3a and 3b immediately deactivated the corresponding part of the hand representation in area 1. We conclude that the processing of somesthetic inputs across anterior parietal cortex is predominantly hierarchical.     

Somatotopic organization of the second somatosensory cortical area after lesions of the primary somatosensory area in infant and adult cats

The somatotopic organization of the second somatosensory cortical area (SII) and receptive fields of multineuron responses to cutaneous stimulation were studied in cats 6–16 months after lesions of the forelimb representation in the primary somatosensory area (SI) at 4 days, 4 weeks of age or in adults. No change was detected in SII. The results contrast with findings of alterations in SII of macaque monkeys following similar ablations of SI.     

Corticocortical connections of area 2 of somatosensory cortex in macaque monkeys: a correlative anatomical and electrophysiological study

The cortical connections of electrophysiologically identified locations in the body representations in somatosensory cortex of macaque monkeys were investigated after injections of horseradish peroxidase, wheat germ agglutinin (WGA) conjugated with horseradish peroxidase, tritiated WGA, or tritiated proline. After extensive microelectrode mapping of portions of the body representations in areas 3b, 1, 2, and 5 and careful determinations of electrophysiological borders between areas, restricted injections of tracers were placed, usually into the representation of the hand in area 2. Other injections were placed in the foot representation in area 2 or in area 1, in the wrist representation in area 1, and in the forearm and wrist representation in area 5. Connection patterns were related to the physiological mapping results and to cortical cytoarchitecture. Injections confined to a lateral portion of area 2 representing the glabrous digits of the hand revealed reciprocal connections with the digit representations in areas 1 and 3b. Projections to area 2 were largely from layer III neurons in both of these fields, and return projections terminated largely in supragranular layers. Other inputs were from layer III cells in one or more separate locations in area 5 and in one or more closely spaced foci in the expected location of S-II in the lateral sulcus. These connections were also reciprocal with terminations apparent in layers IV and III. A few neurons in area 4 were labeled in some of these cases. Results were similar after an injection in the foot representation in area 2 with the differences that infragranular neurons, in addition to supragranular neurons, formed a substantial part of the projection to area 2, terminations as well as projections were noted from area 4, interconnections were found more rostrally in area 6, and a dense focus of label was apparent in the dorsal bank of cingulate sulcus in the apparent location of the supplementary motor area. Injections in the foot representation in area 1 revealed dense layer IV terminations in the foot representation in area 2, as well as connections with area 3b, the S-II region, and areas 5 and 7. The injection in the wrist representation in area 1 resulted in dense terminations in the portion of area 5 responsive to the distal forearm and hand, sparser connections with a lateral location in part of area 2 related to the hand, and interconnections with 3b and S-II.(ABSTRACT TRUNCATED AT 400 WORDS)     

Sensory and motor representation in the cerebral cortex of the three-toed sloth (Bradypus tridactylus)

Cortical sensory receiving areas were studied in 32 specimens of the three-toed sloth,Bradypus tridactylus, using the evoked response technique and barbiturate anesthesia.

Somatotopic organization in the somesthetic first area (S-I) was shown to be similar to that reported in higher mammals. The area devoted to the representation of the forelimb was considerably larger than that for the remaining body parts.

The second somatosensory area (S-II), showing bilateral representation without a precise topographical organization, was identified in the rostral portion of the ectosylvian gyrus. Visual and auditory projections, occupying restricted areas, were found along the caudal banks of the ectosylvian fissure.

Electrically excitable cortical motor area was explored in 8 animals anesthetized with diallyl barbituric acid in urethane. The motor representation of the various body parts was shown to coincide with the sensory projections, demonstrating the existence in a Eutherian mammal of a sensorimotor amalgam, identical to that described in marsupials.

These findings suggest that sloths display a primitive pattern of neural organization, and that the sensory motor amalgam is a generalized form of cortical organization of primitive Therian mammals.