Vertical neuronal arrays in the postcentral gyrus signaling active touch: a receptive field study in the conscious monkey

Summary Single neuronal activity was recorded in the crown of the postcentral gyrus (areas 1 and 2) in 5 conscious monkeys. A total of 88 penetrations entered perpendicularly to the cortical surface. Among them, 6 examples are chosen and the receptive field characteristics of constituent neurons were described in some detail. Receptive fields of neurons recorded along a particular penetration were variable in their positions, but the largest receptive field usually covered the others. Neurons with the largest receptive fields were found most frequently in the infragranular layers. Often they included inhibitory receptive fields. The inhibitory receptive fields were arranged side-by-side to the excitatory ones, rather than in a center-surround fashion. The key stimulus common to neurons in a vertical penetration was the contact of an object to the receptive field achieved during animal's active behavior to manipulate the object. We thus designated the largest receptive field as a functional surface. Our results demonstrate that a vertical array of neurons in this cortical region can be regarded as a functional assemblage which deals with a set of information concerning one of various aspects of active touch.Supported by grants from the Japanese Ministry of Education     

Rostrocaudal gradients in the neuronal receptive field complexity in the finger region of the alert monkey's postcentral gyrus

Summary In the primate postcentral gyrus, the cytoarchitectonic characteristics gradually shift from those of koniocortex to more homotypical parakoniocortex along its rostrocaudal axis. To find the physiological correlates of these changes we examined a large body of data accumulated during a series of our experiments with alert monkeys. Along the rostrocaudal axis of the postcentral gyrus, we found a gradual and continuous increase in the number of neurons with converging receptive fields and those in which receptive field positions or submodalities were not determined. Deep or skin submodality neurons were dominant in area 3a or 3b respectively. The proportion of skin submodality neurons decreased gradually from area 3b to the more caudal part of the gyrus. The proportion of deep submodality neurons was almost constant from area 3b to area 2 inclusive; they were not the majority in area 2. The data are consistent with the hierarchical scheme, i.e., within the postcentral gyrus sensory information is processed from the primary sensory receiving stage to the more associative, integrative stages.     

Hierarchical somesthetic processing of tongue inputs in the postcentral somatosensory cortex of conscious macaque monkeys

The representation of the oral structures in the lateralmost part of the postcentral somatosensory cortex in conscious macaque monkeys was studied by recording the activities of single neurons. A total of 104 penetrations were made in the oral regions of six hemispheres in four animals and 2,292 neurons were isolated. The characteristics of the receptive fields (RF) of 1,598 neurons were identified. Of them, 513 neurons (area 3b, 196; area 1, 104; area 2, 213) along 44 penetrations responded to mechanical stimulation of the tongue (tongue neurons). The relative incidence of tongue neurons that had bilateral RFs increased gradually (bilateral integration) on moving caudally from area 3b to area 2. There was also a tendency for the RFs on the tongue to expand in the anteroposterior axis of the tongue (anteroposterior integration). Furthermore, the relative incidence of tongue neurons with composite RFs covering both the tongue and other surrounding oral structures was significantly higher in area 2 than in areas 3b and 1 (interstructural integration). As a result of the bilateral, anteroposterior and interstructural integration, the extent of the RFs of tongue neurons increased progressively from area 3b to area 2. We therefore concluded that hierarchical somatosensory processing, which has been established in the postcentral somatosensory cortex representing other body parts, is also present in the oral representation. We speculate that the hierarchical scheme in the oral representation might be a prerequisite neural process for the oral stereognosis that eventually takes place in the association cortices. Electronic Publication     

Sensory response properties of pyramidal tract neurons in the precentral motor cortex and postcentral gyrus of the rhesus monkey

Summary Pyramidal tract neurons (PTNs) were identified in precentral motor cortex (MI) and in postcentral cortex (PoC) of a monkey trained to pronate and supinate its forearm. PTN responses to passive, ramp-form displacements of the forearm were examined in relation to the size of the neuron (as reflected by its antidromic latency). Larger PTNs tended to exhibit transient responses to passive limb displacement, whereas smaller PTNs more frequently showed sustained responses. These findings suggest that smaller PTNs, that make up the majority of the total PTN population, receive continuous feedback during posture as well as during the dynamic phase of movement.     

Bilateral receptive field neurons in the hindlimb region of the postcentral somatosensory cortex in awake macaque monkeys

Single-neuron activities were recorded in the hindlimb region of the primary somatosensory cortex and part of area 5 in awake Japanese monkeys. A total of 1050 units were isolated from five hemispheres of four animals. Receptive fields (RFs) and submodalities were identified for 90% of isolated neurons in areas 3a and 3b. The percentage decreased as the recoding site moved to the more caudal areas. Deep or skin submodality neurons were dominant in area 3a or area 3b, respectively. Deep submodality neurons increased in more caudal areas and were the majority in areas 2 and 5. These observations were consistent with those in the hand and/or digit or arm and/or trunk region. The identified neurons were classified by their RF positions into four types: the foot, leg, foot and leg, or hindlimb and other body parts type. Among 831 identified neurons, 33 neurons had bilateral RFs, 14 had ipsilateral RFs, and the rest (N=784) had contralateral RFs. The relative incidence of neurons with bilateral or ipsilateral RFs among identified neurons was less than 1% in areas 3a, 3b, and 1, and 16% or 25% in areas 2 or 5, respectively. Within areas 2 and 5, the percentage of neurons with bilateral or ipsilateral RFs was significantly smaller in the foot type (5%) than in other RF types (24-57%). RFs of the foot type were on the sole or single toe but never on multiple toes. These observations contrasted with the previous findings that neurons with bilateral RFs were more frequently seen in the hand and/or digit region and that RFs on multiple digit tips were dominant there. The present study thus demonstrated that neurons with bilateral RFs do exist in the hindlimb region. Similarly to the forelimb region, they were found mostly in areas 2 and 5, the caudalmost areas of the postcentral gyrus and hierarchically higher stages in information processing. The relative paucity of neurons with bilateral RFs on the foot, especially those with RFs on multiple toes, may reflect functional differences between the foot and the hand.     

Representation of the visual field in the lateral intraparietal area of macaque monkeys: a quantitative receptive field analysis

The representation of the visual field in the primate lateral intraparietal area (LIP) was examined, using a rapid, computer-driven receptive field (RF) mapping procedure. RF characteristics of single LIP neurons could thus be measured repeatedly under different behavioral conditions. Here we report data obtained using a standard ocular fixation task during which the animals were required to monitor small changes in color of the fixated target. In a first step, statistical analyses were conducted in order to establish the experimental limits of the mapping procedure on 171 LIP neurons recorded from three hemispheres of two macaque monkeys. The characteristics of the receptive fields of LIP neurons were analyzed at the single cell and at the population level. Although for many neurons the assumption of a simple two-dimensional gaussian profile with a central area of maximal excitability at the center and progressively decreasing response strength at the periphery can represent relatively accurately the spatial structure of the RF, about 19% of the cells had a markedly asymmetrical shape. At the population level, we observed, in agreement with prior studies, a systematic relation between RF size and eccentricity. However, we also found a more accentuated overrepresentation of the central visual field than had been previously reported and no marked differences between the upper and lower visual representation of space. This observation correlates with an extension of the definition of LIP from the posterior third of the lateral intraparietal sulcus to most of the middle and posterior thirds. Detailed histological analyses of the recorded hemispheres suggest that there exists, in this newly defined unitary functional cortical area, a coarse but systematic topographical organization in area LIP that supports the distinction between its dorsal and ventral regions, LIPd and LIPv, respectively. Paralleling the physiological data, the central visual field is mostly represented in the middle dorsal region and the visual periphery more ventral and posterior. An anteroposterior gradient from the lower to the upper visual field representations can also be identified. In conclusion, this study provides the basis for a reliable mapping method in awake monkeys and a reference for the organization of the properties of the visual space representation in an area LIP extended with respect to the previously described LIP and showing a relative emphasis of central visual field. Electronic Publication     

Representation of the midline trunk, bilateral arms, and shoulders in the monkey postcentral somatosensory cortex

 Single neuronal activities were recorded in the arm/trunk region of the postcentral gyrus in awake Japanese monkeys. A total of 1608 units were isolated from four hemispheres of two animals, and receptive fields (RFs) and submodalities were identified in 1162 units. Deep or skin submodality neurons were dominant in area 3a or area 3b, respectively. The deep/skin ratio increased as the recording site moved from area 3b to the more caudal areas. In areas 3a and 3b, neuronal RFs were almost exclusively on either the arm or trunk. In areas 2 and 5, neurons with RFs on the trunk decreased and those with RFs on the hand or covering more than one body part, etc. increased. We found a total of 107 neurons with bilateral RFs and 56 with ipsilateral RFs, while the rest (n=999) were with contralateral RFs. Bilateral or ipsilateral neurons of skin submodality (n=37) were found in areas 1, 2, and 5. Twenty six (70%) had RFs on the trunk and/or occiput, five on the forelimb, and the rest (n=6) on both the trunk and forelimb (the combined type). Among 33 skin bilateral neurons, 90% (n=30) had RFs across the midline. Bilateral or ipsilateral neurons responding to joint manipulation (n=104) were found in areas 2 and 5. Most of them were activated by manipulation of the shoulder and/or elbow (the proximal type, n=72, 69%). There were 25 neurons of the combined type (both the proximal and distal joints were effective, 24%). Bilateral or ipsilateral neurons of deep-others submodality (n=20) were found in areas 1, 2, and 5. The forelimb type (n=12, 60%) was dominant in this category. The combined-type neurons in both the skin- and joint-manipulation categories were found only or mostly in area 5. These results indicate the presence of hierarchical processing for bilateral as well as contralateral information within the arm/trunk region of the postcentral gyrus. Received: 12 January 1998 / Accepted: 19 May 1998     

Sprouting of remaining substance P-immunoreactive fibers in the monkey dentate gyrus following denervation from its substance P-containing hypothalamic afferents

This study analyzed the response of intrinsic substance P-immunoreactive fibers in the monkey dentate gyrus to disruption of the supramammillohippocampal projection. This projection normally forms a thin plexus of large, substance P-immunoreactive terminals in the innermost portion of the dentate molecular layer and establishes exclusively asymmetric synapses with dendritic shafts and spines of dentate neurons. Conversely, substance P-containing terminals have never been observed in synaptic contact with granule cell bodies. Ten days after ipsilateral fimbria-fornix transection, the prominent band of large immunostained axons in the inner molecular layer of the ipsilateral fascia dentata disappeared. Four and five weeks following transection, however, some small, substance P-containing terminals were observed in the innermost portion of the dentate molecular layer and the granule cell layer. These terminals established exclusively symmetric synapses with the somata and proximal dendritic shafts of granule cells. These results suggest that, following transection of the hypothalamo-hippocampal fiber tract, presumptive intrinsic substance P-containing axons are capable of sprouting into the granule cell layer and the former termination field of the hypothalamic fibers. The symmetric synapses established with granule cell bodies and their proximal dendrites might indicate a shift from an extrinsic excitation to an intrinsic inhibition of granule cells following disruption of substance P-containing hypothalamic afferents.     

Sprouting of remaining substance P-immunoreactive fibers in the monkey dentate gyrus following denervation from its substance P-containing hypothalamic afférents

This study analyzed the response of intrinsic substance P-immunoreactive fibers in the monkey dentate gyrus to disruption of the supramammillo-hippocampal projection. This projection normally forms a thin plexus of large, substance P-immunoreac tive terminals in the innermost portion of the dentate molecular layer and establishes exclusively asymmetric synapses with dendritic shafts and spines of dentate neurons. Conversely, substance P-containing terminals have never been observed in synaptic contact with granule cell bodies. Ten days after ipsilateral fimbria-fornix transection, the prominent band of large immunostained axons in the inner molecular layer of the ipsilateral fascia dentata disappeared. Four and five weeks following transection, however, some small, substance P-containing terminals were observed in the innermost portion of the dentate molecular layer and the granule cell layer. These terminals established exclusively symmetric synapses with the somata and proximal dendritic shafts of granule cells. These results suggest that, following transection of the hypothalamo-hippocampal fiber tract, presumptive intrinsic substance P-containing axons are capable of sprouting into the granule cell layer and the former termination field of the hypothalamic fibers. The symmetric synapses established with granule cell bodies and their proximal dendrites might indicate a shift from an extrinsic excitation to an intrinsic inhibition of granule cells following disruption of substance P-containing hypothalamic afferents.     

Magnification factor and receptive field size in foveal striate cortex of the monkey

Summary Receptive field size and magnification have been studied in striate cortex of awake, behaving rhesus monkeys at visual eccentricities in the range of 5–160 min. The major findings that emerge are (1) magnification in the foveola achieves values in the range of 30 mm/deg, (2) mean field size is not proportional to inverse magnification in contrast with previous reports, and (3) the product, magnification X aggregate field size, is greater in central vision than in peripheral vision. Thus, a point of light projected onto foveal retina is seen by larger numbers of striate cortical cells than a point of light projected onto peripheral retina.Implications of these findings for visual localization and two-point discrimination are discussed.Dedicated to Hermann RahnSupported by NIH grants EY02349 and 5 T32 EY07019     

Are extent and force independent movement parameters? Preparation- and movement-related neuronal activity in the monkey cortex

Movement extent and movement force can be independently controlled in motor performance. Therefore, independent representations of extent and force should exist in the central nervous system (CNS). To test this hypothesis, microelectrode recordings were made in sensorimotor cortex of monkeys trained to perform visually cued wrist flexion movements of two extents, against two levels of frictional resistance. An initial preparatory signal (PS) provided complete, partial or no information about extent and/or force of the movement, which had to be performed in response to a second, response signal (RS). The activity of 511 neurons of the primary motor cortex (MI), the premotor cortex (PM), the postcentral cortex (PC), and the posterior parietal cortex (PA) was recorded in two monkeys. Both reaction time (RT) and neuronal data suggest that there exist independent, neuronal mechanisms responsible for the programming of either parameter. On the one hand, partial information about either movement parameter shortened RT when compared with the condition of no prior information. On the other hand, there were, among others, two discrete populations of neurons, one related only to extent, the other only to force. Preparatory changes in activity related to either movement parameter were mainly located in the frontal cortex, especially in the PM. After occurrence of the RS, the percentage of selective changes in activity increased and tended to extend to the parietal cortex. In particular during the movement, force-related changes in activity have been encountered in PA. Furthermore, we conducted trial-by-trial correlation analyses between RT and preparatory neuronal activity for all conditions of prior information. The mean correlation coefficient was significantly higher in the condition of information about movement extent than of information about movement force and it was significantly higher in MI/ PM than in PC/PA.     

Efferent connections of the cingulate gyrus in the rhesus monkey

Summary Efferent cortical connections of the cingulate gyrus are investigated in rhesus monkey using autoradiographic technique. The results indicate that the rostralmost part of the cingulate gyrus (area 32) sends projections to the lateral prefrontal and midorbitofrontal cortex and to the rostral portion of the superior temporal gyrus. In contrast, the other two major subdivisions of the cingulate gyrus, areas 24 and 23, have widespread connections within the cortex. Area 24, for example, projects to the pre-motor region (areas 6 and 8), the fronto-orbital cortex (area 12), the rostral part of the inferior parietal lobule, the anterior insular cortex, the perirhinal area and the laterobasal nucleus of amygdala. Area 23, likewise, sends its connections to the dorsal prefrontal cortex (areas 9 and 10), the rostral orbital cortex (area 11), the parieto-temporal cortex (posterior part of the inferior parietal lobule and the superior temporal sulcus), the parahippocampal gyrus (areas TH and TF), the retrosplenial region and the presubiculum. It seems that the connections of the rostralmost part of the cingulate gyrus resemble the efferent cortical connectional patterns described for lateral prefrontal and orbito-frontal cortex, whereas the projections of areas 24 and 23 are directed to the neocortical, the paralimbic and the limbic areas.This study was in part supported by NIH Grant NS09211 and V.A. Research Project No. 6901Preliminary results of this investigation were presented in abstract form (Pandya et al. 1979)     

Finger movements during reach-to-grasp in the monkey: amplitude scaling of a temporal synergy

To reduce the complexity of controlling hand-shaping, recent evidence suggests that the central nervous system uses synergies. In this study, two Rhesus monkeys reached-to-grasp 15 objects, varying in geometric properties, at five grasp force levels. Hand kinematics were recorded using a video-based tracking system. Individual finger movements were described as vectors varying in length and angle. Inflection points (i.e., stereotypic minima/maxima in the temporal profile of each finger vector) exhibited a temporal synchrony for individual fingers and in the coupling across fingers. Inflection point amplitudes varied significantly across objects grasped, scaling linearly with the object grasp dimension. Thus, differences in the vectors as a function of the objects were in the relative scaling of the vector parameters over time rather than a change in the temporal structure. Mahalanobis distance analysis of the inflection points confirmed that changes in inflection point amplitude as a function of objects were greater than changes in timing. Inflection points were independent of the grasp force, consistent with the observation that reach-to-grasp kinematics and grasp force are controlled independently. In summary, the shaping of the hand during reach-to-grasp involves scaling the amplitude of highly stereotypic temporal movements of the fingers.     

Extent of the ipsilateral representation in the ventral posterior medial nucleus of the monkey thalamus

Summary Single and multiunit mapping was used to determine the extent of the representation of ipsilateral structures in the ventral posterior medial (VPM) nucleus of the thalamus in cynomolgus monkeys. The extent of the VPM occupied by terminations of afferent fibers arising in the ipsilateral principal trigeminal nucleus was also determined by anterograde transport of horseradish peroxidase. Both methods indicate that most of the medial half of VPM is occupied by the ipsilateral representation. This is much larger than previously suspected. Units in the medial half of VPM have small, well localized receptive fields on the ipsilateral side of the lower lip, tongue and palate, in the ipsilateral cheek pouch and on the ipsilateral teeth. The representation is largest for the ipsilateral side of the tongue and the cheek pouch. Most units in the lateral half of VPM have small, contralateral receptive fields. Few units in VPM have bilateral receptive fields. VPM is clearly distinguishable by cytochrome oxidase (CO) staining. Anteroposteriorly elongated, CO-positive aggreations correspond to elongated aggregations of units with the same or closely similar receptive fields, especially in the medial, ipsilateral representation.Abbreviations CL Central lateral nucleus - CM Centre médian nucleus - DCN Dorsal cochlear nucleus - DIT Dorsal ipsilateral trigeminal tract - IO Inferior olivary nuclei - ML Medial lemniscus - MV Motor trigeminal nucleus - PRV Principal sensory trigeminal nucleus - SO Superior olivary nuclei - SPV Spinal trigeminal nucleus - Ves Vestibular nuclei - VMb Basal ventral medial nucleus - VPI Ventral posterior inferior nucleus - VPL Ventral posterior lateral nucleus - VPM Ventral posterior medial nucleus - IV Trochlear nucleus - VI Abducens nucleus     

Neurons with radial receptive fields in monkey area V4A: evidence of a subdivision of prelunate gyrus based on neuronal response properties

In recordings from two awake, behaving macaque monkeys we found that neurons in the crown of the prelunate gyrus differed in their responsiveness to simple visual stimuli. Neurons in the posterior part of the gyrus (area V4) responded strongly to stationary or moving bars, while neurons in the anterior part (area V4A) responded only weakly to such stimuli. Most receptive fields in area V4A were elongated with long axes oriented radially towards the fovea. These neurons were sensitive to radial movements, especially to sudden shifts of real 3D objects. The border between areas V4 and V4A coincided with the representation of the horizontal meridian. Area V4A extended into the posterior bank of the superior temporal sulcus, where its border corresponded to the representation of the vertical meridian. The sequence of the representations of the horizontal and vertical meridians over the prelunate gyrus suggests the existence of another area between V4A and V4t.     

A quantitative study of chromatic organisation and receptive fields of cells in the lateral geniculate body of the rhesus monkey

Summary The responses of neurones in the lateral geniculate nucleus (LGN) were investigated in anaesthetised rhesus monkeys. A new classification for cells in the parvocellular layers (PCL) is proposed, based on their spectral response curve and their response to white stimuli: (A) narrow-band, short wavelength (NS) excited cells, activity suppressed by white stimuli; (B) wide-band, short-wavelength (WS) excited cells, excited by white stimuli; (C) wide-band, long-wavelength (WL) excited cells, (D) narrow-band, long-wavelength (NL) excited cells, activity suppressed by white stimuli; (E) light suppressed (LI) cells, activity suppressed by all wavelengths, usually with some concealed excitatory input at extreme short or long wavelengths. Responses to moving bars and to spots of various diameters (area response curves) were determined for various wavelengths. It was found that the receptive fields from which wavelength-dependent excitatory or suppressive effects could be elicited are concentrically superimposed. The spectral responsiveness of the excitatory inputs to individual cell types corresponds to the absorption curves of single cones (S-, M- or L-cone for NS, WS and WL cells respectively), the spectral distribution of the suppressive mechanisms of all cells was panchromatic and approximately fitted to a sum of all cones. The excitatory input to NL-cells cannot be related to any of the known cone absorption curves, and a simple (L-M) subtraction model is questioned. Neurones in the magnocellular layers (MCL) can be divided into on- and off-centre cells as in the cat's LGN and give qualitatively similar responses over the whole spectrum. In contrast to the tonic responses of PCL cells, MCL cells respond phasically to chromatic and white flashed spots, even with the smallest stimuli. Implications of these findings for colour processing in the LGN are discussed.     

Visual properties and spatial distribution of neurones in the visual association area on the prelunate gyrus of the awake monkey

Summary We have analysed, in the awake monkey (Macaca sylvana) the functional properties of 489 neurones in the prelunate visual area (PVA, largely corresponding to V4). PVA has a coarse retinotopic organization with the lower quadrant of the visual field represented along the prelunate gyrus. The visual periphery is located medio-dorsally, the central visual field laterally near (and within?) the inferior occipital sulcus and the upper quadrant latero-ventrally. The vertical meridian runs caudally within the lunate sulcus, the horizontal meridian crosses the prelunate gyrus and continues into the superior temporal sulcus. Receptive field diameters of neurones vary between 1° and 10° with increase towards the visual periphery, but are strictly confined to the contralateral visual field. 28% of the neurones showed spectral sensitivity. About half of these cells had strong spectral opponency, the other half showed only weak opponency with broader spectral response curves. 11 cells (2%) showed striking centre/surround interactions with inhibition, disinhibition or occlusion of the two mechanisms, and different spectral response ranges of the centre and the surround, respectively. 43% of the prelunate cells were responsive to various spatial features without spectral sensitivity. We distinguished on- and off-center cells (2%), direction and movement sensitive cells (10%) and cells sensitive to gratings of parallel lines within a limited range of orientations (about 10%). A special group were cells which responded strongly to stimuli which contained many contrasts (textures without specific orientations and without regular spatial arrangements) (9%). Many of these cells were specifically responsive to variations of the internal structure of such stimuli. 3% of the cells were strongly activated in connection with behaviour: 11 neurones discharged strongly when the monkey looked attentively at a human face or when he responded with facial expressions to a threatening expression of a person. Photographs of faces were not effective. Some neurones (1%) were activated in connection with eye movement. These neurones were found in the lateral part of the prelunate gyrus. Neurones with spectral or non-spectral properties were clustered within small, irregularly shaped patches of 1–4 mm diameter. It is concluded that the prelunate visual cortex, which we consider as part of area 19, is not just a colour area, but represents various features of the visual environment (including colour, luminance, movement, texture and behavioral significance), and relates them — through its subcortical and cortical outputs — to behaviour. The various visual cortical areas may be seen as a cooperative of several connections between visual input and behaviour output rather than as links in a hierarchical chain of perceptual and cognitive representations.     

Mechanical response properties of ventroposterior medial thalamic neurons in the alert monkey

Summary The properties of trigeminal cutaneous thalamic neurons were explored in alert cynomolgous monkeys to determine receptive field and response characteristics. Two monkeys received juice reward for sitting quietly while an investigator probed the monkey's face with mechanical stimuli. Extracellular single unit recordings were made from the ventroposterior medial thalamic nucleus (VPM), and mechanical response properties were evaluated for each cell having an extraoral cutaneous receptive field. Of 89 cells examined, 90% responded best to innocuous tactile stimulation, and were classified as low threshold. The other 10%, classified as wide dynamic range, showed a graded response to increasingly intense stimuli, with a maximum discharge to noxious pinch. Of the low threshold neurons, most exhibited excitatory responses, with about half being rapidly adapting and the others slowly adapting. The spontaneous activity of 11% of the low threshold neurons was inhibited by stimulation of the neuron's receptive field. There was no systematic difference in receptive field size for the various types of neurons, but the receptive fields of wide dynamic range cells were smaller than those previously observed in trigeminothalamic neurons of the medullary dorsal horn. The wide dynamic range and inhibitory low threshold neurons were located primarily in the caudal third of VPM, while the excitatory low threshold neurons were located throughout. In summary, response characteristics of VPM neurons show more diversity in the alert monkey than has been reported in paralyzed and/or anesthetized animals.