Microelectrode Mapping Methods Research Articles

Altered Spatiotemporal Dynamics of Cortical Activation to Tactile Stimuli in Somatosensory Area 3b and Area 1 of Monkeys after Spinal Cord Injury.

Reactivation of deafferented cortex plays a key role in mediating the recovery of lost functions, although the precise mechanism is not fully understood. This study simultaneously characterized the dynamic spatiotemporal features of tactile responses in areas 3b and 1 before and 6–8 weeks after partial dorsal column lesion (DCL), and examined how the reactivation relates to the recovery of simple hand use in squirrel monkeys. A combination of high spatiotemporal resolution functional intrinsic optical imaging, microelectrode mapping, behavioral assessment, and tracer histology methods were used. Compared with the normal cortex, we found that the responses of deafferented areas 3b and 1 to 3 s of continuous 8 Hz tactile stimulation of a single digit were significantly weaker and more transient. This finding indicates a loss of response to sustained tactile stimuli. The activation area enlarged for areas 3b and 1 in both directions along digit representation (medial–lateral) and across areas (anterior–posterior). All subjects showed behavioral deficits in a food reaching-grasping-retrieving task within the first 5 weeks after DCL, but recovered at the time when optical images were acquired. Summarily, we showed that these populations of cortical neurons responded to peripheral tactile inputs, albeit in significantly altered manners in each area, several weeks after deafferentation. We propose that compromised ascending driven inputs, impaired lateral inhibition, and local integration of input signals may account for the altered spatiotemporal dynamics of the reactivated areas 3b and 1 cortices. Further investigation with large sample sizes is needed to fully characterize the effects of deafferentation on area 1 activation size.

Cortical and thalamic connections of the parietal ventral somatosensory area in marmoset monkeys (Callithrix jacchus).

Microelectrode mapping methods were used to define the parietal ventral somatosensory area (PV) on the upper bank of the lateral sulcus in five marmosets (Callithrix jacchus). In the same animals, neuroanatomical tracers were placed into electrophysiologically identified sites in PV and/or the second somatosensory area (S2). Foci of anterograde and retrograde label were related to electrophysiological maps of cortical areas and cortical and thalamic architecture. The results lead to the following conclusions: (1) Multiunit recordings from cortex on the upper bank of the lateral sulcus demonstrate that PV is somatotopically organized, with the face representation adjoining area 3b and the hindlimb and tail representations away from this border in cortex deep on the upper bank of the lateral sulcus. The forelimb representation is caudal in PV adjacent to the S2 forelimb representation. The body surface representation in PV approximates a mirror image of that in S2; (2) Areas PV and S2 are less myelinated and have less cytochrome oxidase enzyme activity than area 3b; (3) The ventroposterior inferior nucleus (VPI) of the thalamus provides the major somatosensory projections to PV. PV is reciprocally connected with VPI and anterior pulvinar; (4) PV has ipsilateral cortical connections with areas 3a, 3b, 1, and M1 and higher order somatosensory fields, and at least most of these connections are somatotopically matched; and (5) Callosal connections of PV are with S2 and PV of the other cerebral hemisphere. These results further establish PV as one of at least four somatosensory areas of the lateral sulcus of primates.

Organization of somatosensory cortex in monotremes: in search of the prototypical plan.

The present investigation was designed to determine the number and internal organization of somatosensory fields in monotremes. Microelectrode mapping methods were used in conjunction with cytochrome oxidase and myelin staining to reveal subdivisions and topography of somatosensory cortex in the platypus and the short-billed echidna. The neocortices of both monotremes were found to contain four representations of the body surface. A large area that contained neurons predominantly responsive to cutaneous stimulation of the contralateral body surface was identified as the primary somatosensory area (SI). Although the overall organization of SI was similar in both mammals, the platypus had a relatively larger representation of the bill. Furthermore, some of the neurons in the bill representation of SI were also responsive to low amplitude electrical stimulation. These neurons were spatially segregated from neurons responsive to pure mechanosensory stimulation. Another somatosensory field (R) was identified immediately rostral to SI. The topographic organization of R was similar to that found in SI; however, neurons in R responded most often to light pressure and taps to peripheral body parts. Neurons in cortex rostral to R were responsive to manipulation of joints and hard taps to the body. We termed this field the manipulation field (M). The mediolateral sequence of representation in M was similar to that of both SI and R, but was topographically less precise. Another somatosensory field, caudal to SI, was adjacent to SI laterally at the representation of the face, but medially was separated from SI by auditory cortex. Its position relative to SI and auditory cortex, and its topographic organization led us to hypothesize that this caudal field may be homologous to the parietal ventral area (PV) as described in other mammals. The evidence for the existence of four separate representations in somatosensory cortex in the two species of monotremes indicates that cortical organization is more complex in these mammals than was previously thought. Because the two monotreme families have been separate for at least 55 million years (Richardson, B.J. [1987] Aust. Mammal. 11:71-73), the present results suggest either that the original differentiation of fields occurred very early in mammalian evolution or that the potential for differentiation of somatosensory cortex into multiple fields is highly constrained in evolution, so that both species arrived at the same solution independently.

Electrophysiological examination of the representation of the face in the suprasylvian gyrus of the ferret: a correlative study with cytoarchitecture.

Using high-resolution microelectrode mapping methods, we explored the organization of the face representation within the primary somatosensory cortex of ferrets, finding evidence for at least two and probably four representations of the face distributed consecutively from anterior to posterior along the long axis of the suprasylvian gyrus. Examination of the cytoarchitecture (Rice et al., this issue) revealed that these four areas corresponded to four different cytoarchitectonic fields within the crown of the suprasylvian gyrus. The two central, most completely defined representations were oriented so that the dorsal cutaneous surfaces of the face were represented on the lateral side of the gyrus, while the perioral and ventral surfaces were represented on the medial side. The rostral-to-caudal organization within these two representations was reversed; the glabrous rhinaria were represented at the opposite ends of the maps, and penetrations progressively further away from the cortex serving the rhinaria encountered neurons activated by sites progressively more caudal on the face. Receptive fields obtained more rostrally on the gyrus suggested another reversal, implying a third representation. A small area with large receptive fields near the caudal and medial border of the two central maps suggested the presence of a fourth representation. Since the projections of adjacent skin surfaces overlapped considerably, cortical sites serving a particular cutaneous surface were illustrated as enclosed areas that overlapped the territories of other, adjacent representations. The results of this study and of others suggest a need for a re-evaluation of the hypothesis establishing a homology between the representation found in area 3b of primates and that of the primary somatosensory area in nonprimates.

Subdivisions and connections of auditory cortex in owl monkeys

The organization and connections of auditory cortex in owl monkeys, Aotus trivirgatus, were investigated by combining microelectrode mapping methods with studies of architecture and connections in the same animals. In most experiments, portions of auditory cortex were first explored with microelectrodes, neurons were characterized as responsive or not to auditory stimuli, and best frequencies were determined whenever possible. Most recordings were in cortex previously designated as primary (A-I) and rostral (R) auditory fields (Imig et al. J Comp Neurol 171:111, '77) and in a newly defined rostrotemporal field (RT) located rostral to R. Injections of wheat germ agglutinin-horseradish peroxidase (WGA-HRP) and fluorescent tracers were placed in electrophysiologically identified locations of A-I, R, and RT; the posterolateral (PL) and anterolateral (AL) divisions of a narrow belt of auditory cortex lateral and adjacent to A-I and R; cortex of the superior temporal gyrus lateral and rostrolateral to PL and AL; and regions of prefrontal cortex that receive inputs from auditory cortex. There were several major findings: 1. Best frequencies were most clearly determined for neurons within a densely myelinated strip of cortex on the lower bank and lip of the lateral sulcus. We divided this strip into three fields, A-I, R, and RT, although an alternative interpretation that A-I and R are parts of a single field remains tenable. In some cases, isofrequency contours appeared to continue uninterrupted across fields A-I and R, with lower frequencies represented laterally and higher frequencies represented deeper in the sulcus. In other cases, there was a tendency for high frequencies to be represented caudally and medially, and low frequencies laterally in A-I and rostrally in R, with partial discontinuity in the isofrequency contours. A reversal of the tonotopic gradient appeared in RT with a common low-frequency representation at the caudal border with R, and progressively higher frequencies encountered rostrally. Of the three fields, A-I appears slightly more myelinated than R, and RT slightly less than R. The distinctiveness of the three fields is further demonstrated by the patterns of connections. In particular, A-I and RT are both interconnected with R, but not with each other. Connections between A-I and R are between tonotopically matched locations. 2. A narrow 2-3 mm wide band of cortex lateral to A-I, R, and RT was also responsive to auditory stimuli, but typically neurons were more difficult to activate, and best frequencies were more difficult to determine. No distinctions in myeloarchitecture or CO activity were obvious.(ABSTRACT TRUNCATED AT 400 WORDS)

Somatosensory cortex of the neonatal pig: II. Topographic organization of the secondary somatosensory cortex (SII)

Multiunit microelectrode recording techniques were used to delineate the somatotopic organization of the secondary somatosensory cortex (SII) of the neonatal pig. Barbiturate anesthetized piglets ranging in age from 7 days preterm to 2 months postpartum were used. The SII area, located lateral to the rostral and middle suprasylvian sulci, was found to contain a complete somatotopic representation of the contralateral body surface with a significant proportion of bilateral input for all body regions except the forehoof and forelimb. The SII forelimb and hindlimb representations were found to possess a "striplike" orientation in a rostral to caudal sequence, and the trunk representation was located posterolateral to the hindlimb representation, giving SII an inverted appearance. Two apparently separate face representations were delineated; one posterolateral to the projection from the trunk and the other anterior to the forehoof region. Unlike SI, which possesses a disproportionately large representation of the rostrum, SII has no specialized representation of the rostrum. The overall organization of SII supports the contention that this cortical region provides a more generalized representation of the entire body surface than does SI.

Somatosensory cortex of the neonatal pig: I. Topographic organization of the primary somatosensory cortex (SI)

The cerebral cortex of adult mammals contains several somatotopic representations of the body surface. Although the organization of the various somatosensory cortices of numerous species of adult animals has been elucidated, data on the somatosensory representations of fetal and neonatal animals are limited. As part of an investigation into the perinatal development of the somatosensory cortices, it was necessary to delineate the organization of the somatosensory cortices of the perinatal pig. This study presents the topographical organization of the primary somatosensory cortex (SI) of the perinatal pig. Multiunit microelectrode mapping methods were used to produce topographic maps of SI from barbiturate anesthetized pigs ranging in age from 7 days preterm to 2 months postpartum. It was demonstrated that the overall organization of this region of cortex was similar to that of other mammals: a somatotopic projection of predominantly the contralateral body surface was delineated in which the hindlimb is represented medially and the face laterally across the cortex. A disproportionately enlarged rostrum representation was mapped in detail, and multiple representations of the rostrum, face, and mouth were found. Several of these representations exhibited bilateral and ipsilateral input. The SI trunk and hindlimb representations were located on the medial wall of the hemisphere; these representations were small but their presence refutes speculation that ungulates do not have a complete body representation in SI.

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)

Microelectrode maps, myeloarchitecture, and cortical connections of three somatotopically organized representations of the body surface in the parietal cortex of squirrels.

Microelectrode mapping methods and anatomical procedures were combined in the same animals to reveal the cortical connections of three architectonically distinct representations of the body surface in the somatosensory cortex of grey squirrels. In individual experiments, microelectrode multiunit recordings were used to determine the somatotopic organization of regions of the cortex and to identify sites for injections of the anatomical tracer, wheat germ agglutinin conjugated to horseradish peroxidase. After the brains were perfused, the cortex was separated from the brainstem, flattened, and cut parallel to the flattened surface to facilitate comparisons of areal connection patterns, physiological data, and architectonic subdivisions. Recordings in the primary (S-I) and secondary (S-II) somatosensory fields confirmed earlier descriptions of the somatotopic organization of these fields (Sur et al.: J. Comp. Neurol. 179:425-450, '78; Nelson et al.: J. Comp. Neurol. 184:473-490, '79). In addition, recordings in the cortex caudal to S-I and ventral to S-II revealed a third representation of the body surface in parietal cortex, the parietal ventral area (PV). Neurons in PV were responsive to light tactile stimulation of skin and hairs. Multiple unit receptive fields of neurons in PV were larger than those for neurons in S-I but similar in size to those for neurons in S-II. PV represented the contralateral body surface in a somatotopic manner that can be roughly characterized as an inverted "homunculus" with the limbs directed medially, the trunk located ventrally, and the face congruent with the representations of the upper lip and nose in S-I. Neurons in some electrode penetrations in PV were also responsive to auditory clicks. Microlesions placed at physiologically determined borders allowed all three somatic representations to be related to myeloarchitectonically defined fields. S-I was architectonically distinct as a densely myelinated region. Within S-I, a lightly myelinated oval of the cortex between the representation of the hand and face, the "unresponsive zone" (Sur et al.: J. Comp. Neurol. 179:425-450, '78), was an easily recognized landmark. S-II and PV corresponded to less densely myelinated fields. Other subdivisions such as motor cortex, primary auditory cortex, and visual areas 17 and 18 were distinguished. Connections were revealed by placing injections within S-I, S-II, or PV.(ABSTRACT TRUNCATED AT 400 WORDS)

The organization and connections of somatosensory cortex in primates

The use of microelectrode mapping methods, in conjunction with studies of cortical architecture, patterns of connections, and the response properties of single neurons, has led to a greatly increased understanding of the functional organization of somatosensory cortex in mammals. All mammals appear to have the traditionally defined first and second somatosensory representations, SI and SII, but at least some mammals have additional body representations as well. Furthermore, the area traditionally regarded as “SI” in monkeys actually includes four functionally distinct representations. Within “SI,” four striplike architectonic fields, areas 3a, 3b, 1, and 2 have long been recognized. It is now clear that areas 3b and 1 form separate parallel and largely mirror‐image representations of the body surface, each with major inputs from a single representation of the body surface in the ventroposterior nucleus (VP) of the thalamus. VP appears to relay information from both slowly and rapidly adapting peripheral neurons to area 3b, while information from rapidly adapting neurons is relayed to area 1. Neurons with properties that could reflect Pacinian receptor influences are found in area 1, but not area 3b. The anatomical pathways mediating Pacinian‐like responses are unknown. Area 3a contains a representation of largely deep receptors, probably muscle spindle afferents, relayed from part of the thalamus dorsal to VP. Area 2 forms a representation of both deep and cutaneous body receptors. The somatosensory cortex receives input from other regions as well, including the anterior pulvinar and ventroposterior inferior nucleus of the thalamus. Other portions of the somatosensory cortex are less well understood.

Representations of the body surface in areas 3b and 1 of postcentral parietal cortex of cebus monkeys

The somatotopic organization of postcentral parietal cortex was determined with microelectrode mapping methods in a New World monkey, Cebus albifrons. As in previous studies in macaque, squirrel and owl monkeys, two separate representations of the body surface were found in regions corresponding to the architectonic fields 3b and 1. The two representations were roughly mirror-images of each other, with receptive field locations matched for recording sites along the common border. As in other monkeys, the glabrous digit tips of the hand and foot pointed rostrally in the Area 3b representation and caudally in the Area 1 representation. Both representations proceeded in parallel from the tail on the medial wall of the cerebral hemisphere to the teeth and tongue in lateral cortex along the Sylvian fissure. Compared with the other monkeys, the tail of the cebus monkey, which is prehensile, was represented in a very large region of cortex in Areas 3b and 1. Like its close relative, the squirrel monkey, the representation of the trunk and parts of the limbs were reversed in orientation in both Area 3b and Area 1 in cebus monkeys as compared to owl and macaque monkeys. The reversals of organization for some but not all parts of the representations in cebus and squirrel monkeys suggest that one line of New World monkeys acquired a unique but functionally adequate pattern of somatotopic organization for the two adjoining fields.

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.

Connections of the ventroposterior nucleus of the thalamus with the body surface representations in cortical areas 3b and 1 of the cynomolgus macaque, (Macaca fascicularis).

The somatotopic pattern of the connections of the ventroposterior nucleus of the thalamus with the representations of the body surface in somatic cortical Areas 3b and 1 were investigated in macaque monkeys using the anatomical tracers horseradish peroxidase (HRP), and 3H-proline. These tracers, in combination or separately, were injected into one or more cortical sites, after the skin surface capable of activating neurons at each site had been defined with microelectrode mapping methods. These injection sites were later related to the overall organizations of the two cutaneous representations as determined in more extensive mapping experiments (Nelson et al, '80). Finally, zones of anterograde and retrograde label in the thalamus were located with reference to architectonically defined subnuclei in the ventroposterior nucleus (VP). The results led to the following conclusions. (1) Zones of cells in VP projecting to a given location in either Area 3b or 1 also receive reciprocal projections from the cortical target. (2) A given cortical location typically relates to a disc-shaped or lamellar region in VP with the disc extending rostrocaudally and dorsoventrally, being most narrow in the mediolateral dimension. (3) The representations of the same body surface in Area 1 and Area 3b relate to the same region of VP. However, since fewer cells were generally labeled after Area 1 injections, less dense terminations are suggested from VP to Area 1 than to Area 3b. (4) VP is the major or exclusive relay of somatosensory information to the two cutaneous representations from the thalamus. Labeled neurons or zones of terminations were found outside VP in other parts of the somatosensory thalamus only when injection sites extended into cortex outside Areas 3b and 1. (5) The face, hand, limbs, and foot are represented in a mediolateral sequence of subnuclei within VP. A dorsal capping subnucleus is apparently largely devoted to the axial trunk and perhaps other hairy skin surfaces such as some parts of the proximal surfaces of the limbs. (6) While anterior and posterior surfaces of the hind limb are widely separated by the glabrous foot in the cortical representations, they appear to be represented in adjacent zones in the thalamus.

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 iun 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.

Somatic sensory cortex (SmI) of the prosimian primate Galago crassicaudatus: Organization of mechanoreceptive input from the hand in relation to cytoarchitecture

Mechanoreceptive input from the hand to the somatic sensory cortex (SmI) of the prosimian primate Galago crassicaudatus was examined with microelectrode mapping methods. In anesthetized animals, low threshold cutaneous input from the hand projects to SmI cortex in a single, complete, somatotopically organized pattern. Within this single pattern, cells with receptive fields on the glabrous skin of the palm, digits and digit tips are located in the rostral half, and cells with RFs on the hairy skin of the dorsal hand and digits are located in the caudal half of the hand areas. The cutaneous hand area is coextensive with the densely granular architectonic region of SmI. Studies of single cells in this region of awake galagos reveal the same pattern of cutaneous input and, in addition, demonstrate the presence of cells responding to joint movement not detected in anesthetized animals. Cells responsive to joint movement are arranged in vertically oriented columns located adjacent to cutaneous columns with receptive fields on the same part of the hand. In anesthetized animals, cells rostral to the granular region, in an area typified by increasing numbers of pyramidal cells in layer V and decreasing numbers of granular cells in upper layers, respond to high threshold stimulation of large areas of the hand. The few cells isolated in this area in awake animals respond to either active or passive hand movements. In such animals, cells caudal to the granular region, in an area characterized as agranular and alaminar cortex, respond to either passive stimulation of single or multiple joints or to active hand movements. These results, together with similar findings in a related prosimian, Nycticebus coucang, emphasize the generality of a single cutaneous hand area in SmI of prosimian species. The demonstration of multiple hand areas corresponding to multiple cytoarchitectonic subdivisions in SmI of Old and New World simians illustrates the increased degree of SmI differentiation from the prosimian to the simian grade of organization. The present results further suggest that determination of the homologues of multiple areas or subdivisions within and surrounding SmI in primates will require comparisons of somatotopy, submodality, sulcal patterns, cytoarchitecture, and connectivity in representative members of prosimian and simian families.

The organization of the second somatosensory area (SmII) of the grey squirrel

Microelectrode mapping methods were used to determine the organization of the second somatosensory representation, SmII, in grey squirrels. A systematic representation of the contralateral body surface was found in lateral parietal cortex adjoining the first somatosensory representation, SmI (Sur et al., '78a). The representation of the body in SmII was found to be much less distorted than in SmI. Under our recording conditions, almost all recording sites were activated from strictly contralateral body locations. The most important finding was that the basic orientation of the body representation in SmII is "erect" rather than "inverted." This orientation allows SmII and SmI to be adjoined along a common border representing the top of the head and face. This type of border has been called congruent (Allman and Kaas, '75; Kaas, '77), and it may have significance in the development of sensory representations.

The representation of the body surface in somatosensory area I of the grey squirrel

Microelectrode mapping methods were used to determine the organization of primary somatosensory cortex, SmI, in grey squirrels. A systematic representation of the contralateral body surface was found within somatic konicortex. This primary representation differs from maps of SmI in other mammals in at least two significant ways. The first way in which SmI of squirrels differs from the organization reported for other mammals is that SmI of squirrels contains a double representation of the hand and parts of the forearm. The glabrous skin of the digits is represented twice in a mirror image fashion joined at the finger tips. The hairy skin of the digits, wrist, and parts of the forearm are also represented twice, once on each side of the joined representations of the glabrous skin. A second unique feature of SmI of squirrels is that there is a small region of cortex completely surrounded by SmI that was unresponsive to light cutaneous stimuli under our recording conditions. This unresponsive zone is easily identified in brain sections by architectonic features that deviate from sensory koniocortex and approach motor cortex. A third significant finding was that the back is rostral to the belly in the representation of the trunk in SmI of squirrels. This is the reverse of the orientation reported elsewhere for SmI of mammals, but corresponds to the orientation of the trunk representation in Area 3b of owl monkeys (Kaas et al., '78; Merzenich et al., '78). This similarity supports an earlier contention that the representation of the body in Area 3b of primates is the homolog of SmI in other mammals (Merzenich et al., '78).

Microelectrode mapping of modality-specific somatic sensory cerebral neocortex in slow loris.

Microelectrode mapping methods and strategies were developed to map fine details of the somatotopic organization of somatic sensory cortex (SI) in slow loris. In-depth, multiple-unit recording, and threshold natural stimulation of skin and hari revealed three modality-specific cortical areas. The centrally located one was activated by extremely light touch and the somatopic organization of peripheral projections to it was mapped in detail. As many as 25 punctures/mm2 were made in delineating these details. The microelectrode mapping method used revealed smaller receptive fields, lesser degrees of overlap, a more differentiated pattern of peripheral projections, and a more precise correlation of electrophysiological results with underlying cytoarchitectonic details than did the usual macroelectrode surface recording method. These results and the correlations found between different physiologically defined areas and cortical sulci are discussed as they pertain to methodological advances in studies of cerebral localization of function.

Somatic Sensory Cortex of Llama <i>(Lama glama)</i>

The somatic sensory cortex (SI and SII) was mapped in llamas using microelectrode mapping methods developed earlier in a study of SI of the slow loris. Projections to SI from the llama's prehensile browsing lips were differentially enlarged when compared to those reported for sheep. In llama, SII was reversed in its mediolatreal pattern from that reported for SII in most other mammals. Fissural landmarks reliably demarcated different projections within SI, between SI and SII and between SI or SII and other surrounding nonsensory areas. The use of microelectrode mapping methods in different mammals to determine gyral and fissural homologies is discussed.

Representation of the visual field in striate and adjoining cortex of the owl monkey (Aotus trivirgatus)

Summary Microelectrode mapping methods were used to determine the representation of the visual field in striate cortex and adjacent cortex in the owl monkey. As in other primates, the center of gaze is represented in dorsolateral striate cortex. Most of the lower quadrant of the visual field is represented on the upper bank of the calcarine sulcusm while more central portions of the lower quadrant are represented on the medial wall and dorsal surface. The representation of the upper quadrant extends from the dorsolateral surface the tentorial surface and across the lower bank of the calcarine sulcus. The representation of the temporal periphery of the upper quadrant was found on the upper bank of the calcarine sulcus. Almost half of striate cortex is devoted to the central 20° of the visual field and 90% of the border corresponds to the zero vertical meridian or lilne of decussation. Recordings from cortex adjoining striate cortex indicate that the second visual area, V II, borders the complete vertical meridian of striate cortex. The small representation of the temporal periphery in striate cortex is bordered by cortez that was nonresponsive to visual stimuli. The responsive and nonresponsive bordering areas were found to relate to two histologically distinct types of cortex, areas 18 and ‘pro-striata’ respectively.