Area 3b Research Articles

JEPM October 2018—Abstracts

Traditionally, cytoarchitectonic area 3a of primary somatosensory cortex (SI) has been regarded as a proprioceptive relay to motor cortex. However, neuronal spike-train recordings and optical intrinsic signal imaging, obtained from nonhuman sensorimotor cortex, show that neuronal activity in some of the cortical columns in area 3a can be readily triggered by a C-nociceptor afferent drive. These findings indicate that area 3a is a critical link in cerebral cortical encoding of secondary/slow pain. Also, area 3a contributes to abnormal pain processing in the presence of activity-dependent reversal of gamma-aminobutyric acid A receptor-mediated inhibition. Accordingly, abnormal processing within area 3a may contribute mechanistically to generation of clinical pain conditions.Optical imaging and neurophysiological mapping of area 3a of SI has revealed substantial driving from unmyelinated cutaneous nociceptors, complementing input to areas 3b and 1 of SI from myelinated nociceptors and non-nociceptors. These and related findings force a reconsideration of mechanisms for SI processing of pain.

Suppression of somatosensory stimuli during motor planning may explain levels of balance and mobility after stroke.

The ability to actively suppress, or gate, irrelevant sensory information is required for safe and efficient walking in sensory-rich environments. Both motor attention and motor planning alter somatosensory evoked potentials (SEPs) in healthy adults. This study's aim was to examine the effect of motor attention on processing of irrelevant somatosensory information during plantar flexion motor planning after stroke. Thirteen healthy older adults and 11 individuals with stroke participated. Irrelevant tibial nerve stimulation was delivered while SEPs were recorded over Cz, overlaying the leg portion of the sensorimotor cortex at the vertex of the head. Three conditions were tested in both legs: (1) Rest, (2) Attend To, and (3) Attend Away from the stimulated limb. In conditions 2 and 3, relevant vibration cued voluntary plantar flexion movements of the stimulated (Attend To) or non-stimulated (Attend Away) leg. SEP amplitudes were averaged during motor planning per condition. Individuals with stroke did not show attention-mediated gating of the N40 component associated with irrelevant somatosensory information during motor planning. It may be that dysfunction in pathways connecting to area 3b explains the lack of attention-mediated gating of the N40. Also, attention-mediated gating during motor planning explained significant and unique variance in a measure of community balance and mobility combined with response time. Thus, the ability to gate irrelevant somatosensory information appears important for stepping in both older adults and after stroke. Our data suggest that therapies that direct motor attention could positively impact walking after stroke.

Effect of acceleration of auditory inputs on the primary somatosensory cortex in humans

Cross-modal interaction occurs during the early stages of processing in the sensory cortex; however, its effect on neuronal activity speed remains unclear. We used magnetoencephalography to investigate whether auditory stimulation influences the initial cortical activity in the primary somatosensory cortex. A 25-ms pure tone was randomly presented to the left or right side of healthy volunteers at 1000 ms when electrical pulses were applied to the left or right median nerve at 20 Hz for 1500 ms because we did not observe any cross-modal effect elicited by a single pulse. The latency of N20 m originating from Brodmann’s area 3b was measured for each pulse. The auditory stimulation significantly shortened the N20 m latency at 1050 and 1100 ms. This reduction in N20 m latency was identical for the ipsilateral and contralateral sounds for both latency points. Therefore, somatosensory–auditory interaction, such as input to the area 3b from the thalamus, occurred during the early stages of synaptic transmission. Auditory information that converged on the somatosensory system was considered to have arisen from the early stages of the feedforward pathway. Acceleration of information processing through the cross-modal interaction seemed to be partly due to faster processing in the sensory cortex.

Shape responses in a macaque frontal area connected to posterior parietal cortex

The primate dorsal visual stream processes object shape to guide actions involving an object, but the transmission of shape information beyond posterior parietal cortex remains largely unknown. To clarify the information flow between parietal and frontal cortex, we applied electrical microstimulation during functional Magnetic Resonance Imaging (fMRI) in a shape-selective patch in the posterior part of the Anterior Intraparietal area (pAIP) to chart its connectivity. Subsequently, we recorded single-unit responses to images of objects in the fMRI activation in prefrontal cortex, corresponding to area 45B, elicited by pAIP microstimulation. Neurons in area 45B had properties similar to neurons in pAIP, responding selectively to shape contours and to very small shape fragments measuring less than one deg at exceedingly short latencies. However, contrary to the prevailing view on the hierarchical organization of cortical areas, neurons in area 45B preferred even smaller shape fragments and had smaller receptive fields than neurons in pAIP. These findings provide the first evidence for ultra-fast shape processing in prefrontal cortex, and suggest that the pathway from pAIP to area 45B may not be important for object grasping.

Connectivity of neuronal populations within and between areas of primate somatosensory cortex

Functions of the cerebral cortex emerge via interactions of horizontally distributed neuronal populations within and across areas. However, the connectional underpinning of these interactions is not well understood. The present study explores the circuitry of column-size cortical domains within the hierarchically organized somatosensory cortical areas 3b and 1 using tract tracing and optical intrinsic signal imaging (OIS). The anatomical findings reveal that feedforward connections exhibit high topographic specificity, while intrinsic and feedback connections have a more widespread distribution. Both intrinsic and inter-areal connections are topographically oriented across the finger representations. Compared to area 3b, the low clustering of connections and small cortical magnification factor supports that the circuitry of area 1 scaffolds a sparse functional representation that integrates peripheral information from a large area that is fed back to area 3b. Fast information exchange between areas is ensured by thick axons forming a topographically organized, reciprocal pathway. Moreover, the highest density of projecting neurons and groups of axon arborization patches corresponds well with the size and locations of the functional population response reported by OIS. The findings establish connectional motifs at the mesoscopic level that underpin the functional organization of the cerebral cortex.

Cortical connections of area 2 and posterior parietal area 5 in macaque monkeys.

The overarching goal of the current investigation was to examine the connections of anterior parietal area 2 and the medial portion of posterior parietal area 5 in macaque monkeys; two areas that are part of a network involved reaching and grasping in primates. We injected neuroanatomical tracers into specified locations in each field and directly related labeled cells to histologically identified cortical field boundaries. Labeled cells were counted so that the relative density of projections to areas 2 and 5 from other cortical fields could be determined. Projections to area 2 were restricted and were predominantly from other somatosensory areas of the anterior parietal cortex (areas 1, 3b, and 3a), the second somatosensory area (S2), and from medial and lateral portions of area 5 (5M and 5L respectively). On the other hand, area 5M had very broadly distributed projections from a number of cortical areas including anterior parietal areas, from primary motor cortex (M1), premotor cortex (PM), the supplementary motor area (SMA), cortex on the medial wall, and from posterior parietal areas 5L and 7b. The more restricted pattern of connections of area 2 indicates that it processes somatic inputs locally and provides proprioceptive information to area 5M. 5M, which at least partially overlaps with functionally defined area MIP, receives inputs from somatosensory (predominantly from area 2), posterior parietal and motor cortex, which could provide the substrate for representing multiple coordinate systems necessary for planning ethologically relevant movements, particularly those involving the hand.

Architectonic features and relative locations of primary sensory and related areas of neocortex in mouse lemurs.

Mouse lemurs are the smallest of the living primates, and are members of the understudied radiation of strepsirrhine lemurs of Madagascar. They are thought to closely resemble the ancestral primates that gave rise to present day primates. Here we have used multiple histological and immunochemical methods to identify and characterize sensory areas of neocortex in four brains of adult lemurs obtained from a licensed breeding colony. We describe the laminar features for the primary visual area (V1), the secondary visual area (V2), the middle temporal visual area (MT) and area prostriata, somatosensory areas S1(3b), 3a, and area 1, the primary motor cortex (M1), and the primary auditory cortex (A1). V1 has "blobs" with "nonblob" surrounds, providing further evidence that this type of modular organization might have evolved early in the primate lineage to be retained in all extant primates. The laminar organization of V1 further supports the view that sublayers of layer 3 of primates have been commonly misidentified as sublayers of layer 4. S1 (area 3b) is proportionately wider than the elongated area observed in anthropoid primates, and has disruptions that may distinguish representations of the hand, face, teeth, and tongue. Primary auditory cortex is located in the upper temporal cortex and may include a rostral area, R, in addition to A1. The resulting architectonic maps of cortical areas in mouse lemurs can usefully guide future studies of cortical connectivity and function.

Corticocortical projections to area 1 in squirrel monkeys (Saimiri sciureus).

Cortical area 1 is a non-primary somatosensory area in the primate anterior parietal cortex that is critical to tactile discrimination. The corticocortical projections to area 1 in squirrel monkeys were determined by placing multiple injections of anatomical tracers into separate body part representations defined by multiunit microelectrode mapping in area 1. The pattern of labeled cells in the cortex indicated that area 1 has strong intrinsic connections within each body part representation and has inputs from somatotopically matched regions of areas 3b, 3a, 2 and 5. Somatosensory areas in the lateral sulcus, including the second somatosensory area (S2), the parietal ventral area (PV), and the presumptive parietal rostral (PR) and ventral somatosensory (VS) areas, also project to area 1. Topographically organized projections to area 1 also came from the primary motor cortex (M1), the dorsal and ventral premotor areas (PMd and PMv), and the supplementary motor area (SMA). Labeled cells were also found in cingulate motor and sensory areas on the medial wall of the hemisphere. Previous studies revealed a similar pattern of projections to area 1 in Old World macaque monkeys, suggesting a pattern of cortical inputs to area 1 that is common across anthropoid primates.

Discrete Modules and Mesoscale Functional Circuits for Thermal Nociception within Primate S1 Cortex.

This study addresses one long-standing question of whether functional separations are preserved for somatosensory modalities of touch, heat, and cold nociception within primate primary somatosensory (S1) cortex. This information is critical for understanding how the nature of pain is represented in the primate brain. Using a combination of submillimeter-resolution fMRI and microelectrode local field potential (LFP) and spike recordings, we identified spatially segregated cortical zones for processing touch and nociceptive heat and cold stimuli in somatotopically appropriate areas 3a, 3b, 1, and 2 of S1 in male monkeys. The distances between zones were comparable (∼3.4 mm) across stimulus modalities (heat, cold, and tactile), indicating the existence of uniform, modality-specific modules. Stimulus-evoked LFP maps validated the fMRI maps in areas 3b and 1. Isolation of heat and cold nociceptive neurons from the fMRI zones confirmed the validity of using fMRI to probe nociceptive regions and circuits. Resting-state fMRI analysis revealed distinct intrinsic functional circuits among functionally related zones. We discovered distinct modular structures and networks for thermal nociception within S1 cortex, a finding that has significant implications for studying chronic pain syndromes and guiding the selection of neuromodulation targets for chronic pain management.SIGNIFICANCE STATEMENT Primate S1 subregions contain discrete heat and cold nociceptive modules. Modules with the same properties exhibit strong functional connection. Nociceptive fMRI response coincides with LFP and spike activities of nociceptive neurons. Functional separation of heat and cold pain is retained within primate S1 cortex.

Visually-Driven Maps in Area 3b.

Sensory perception relies on the precise neuronal encoding of modality-specific environmental features in primary sensory cortices. Some studies have reported the penetration of signals from other modalities even into early sensory areas. So far, no comprehensive account of maps induced by "foreign sources" exists. We addressed this question using surface-based topographic mapping techniques applied to ultra-high resolution fMRI neuroimaging data, measured in female participants. We show that fine-grained finger maps in human primary somatosensory cortex, area 3b, are somatotopically activated not only during tactile mechanical stimulation, but also when viewing the same fingers being touched. Visually-induced maps were weak in amplitude, but overlapped with the stronger tactile maps tangential to the cortical sheet when finger touches were observed in both first- and third-person perspectives. However, visually-induced maps did not overlap tactile maps when the observed fingers were only approached by an object but not actually touched. Our data provide evidence that "foreign source maps" in early sensory cortices are present in the healthy human brain, that their arrangement is precise, and that their induction is feature-selective. The computations required to generate such specific responses suggest that counterflow (feedback) processing may be much more spatially specific than has been often assumed.SIGNIFICANCE STATEMENT Using ultra-high field fMRI, we provide empirical evidence that viewing touches activates topographically aligned single finger maps in human primary somatosensory cortical area 3b. This shows that "foreign source maps" in early sensory cortices are topographic, precise, and feature-selective in healthy human participants with intact sensory pathways.

Chapter 2 - The evolution of parietal cortex in primates

Many of the adaptive changes in the functional organization of parietal cortex of humans emerged in past in the early primates as they depended on visually guided forelimb use to grasp branches and food. Currently, human, apes and some monkeys have four well-defined subdivisions of anterior parietal cortex, areas 3a, 3b, 1 and 2 of Brodmann. In some of the smaller monkeys, and in stepsirrine primates (galagos, lemurs, and lorises), especially areas 1 and 2 are less developed, and the existence of an area 2 is questionable. In galagos, area 3b, the homologue of S1 in other mammals, has a more primitive somatotopy, is less devoted to representing the hand, and information from facial whiskers is more important. Humans and other primates also have more somatosensory areas in lateral parietal cortex than most mammals. While the regions of the second somatosensory area, S2 is divided into S2 and the parietal ventral area, PV in most mammals, primates have the additional caudal ad rostral ventral somatosensory areas, VSc and VSr. Posterior parietal cortex is another region of posterior cortex that has changed greatly from non-primate ancestors in having a more caudal half that is heavily devoted to further processing visual information for guiding different actions, such as running, reaching, looking, and grasping. All primates have at least 8 small subdivisions or domains in PPC, and have matching domains in premotor and motor cortex. In humans, domains for speech and tool use appear to have been added, and other parts of PPC have expanded. In addition, parts of PPC are differently specialized in the right and left cerebral hemispheres of humans more than in other primates.

Chapter 4 - Somatosensory maps

Somatosensory areas containing topographic maps of the body surface are a major feature of parietal cortex. In primates, parietal cortex contains four somatosensory areas, each with its own map, with the primary cutaneous map in area 3b. Rodents have at least three parietal somatosensory areas. Maps are not isomorphic to the body surface, but magnify behaviorally important skin regions, which include the hands and face in primates, and the whiskers in rodents. Within each map, intracortical circuits process tactile information, mediate spatial integration, and support active sensation. Maps may also contain fine-scale representations of touch submodalities, or direction of tactile motion. Functional representations are more overlapping than suggested by textbook depictions of map topography. The whisker map in rodent somatosensory cortex is a canonic system for studying cortical microcircuits, sensory coding, and map plasticity. Somatosensory maps are plastic throughout life in response to altered use or injury. This chapter reviews basic principles and recent findings in primate, human, and rodent somatosensory maps.

Efferents of anterior cingulate areas 24a and 24b and midcingulate areas 24a' and 24b' in the mouse.

The anterior cingulate cortex (ACC), constituted by areas 25, 32, 24a and 24b in rodents, plays a major role in cognition, emotion and pain. In a previous study, we described the afferents of areas 24a and 24b and those of areas 24a' and 24b' of midcingulate cortex (MCC) in mice and highlighted some density differences among cingulate inputs (Fillinger et al., Brain Struct Funct 222:1509-1532, 2017). To complete this connectome, we analyzed here the efferents of ACC and MCC by injecting anterograde tracers in areas 24a/24b of ACC and 24a'/24b' of MCC. Our results reveal a common projections pattern from both ACC and MCC, targeting the cortical mantle (intracingulate, retrosplenial and parietal associative cortex), the non-cortical basal forebrain, (dorsal striatum, septum, claustrum, basolateral amygdala), the hypothalamus (anterior, lateral, posterior), the thalamus (anterior, laterodorsal, ventral, mediodorsal, midline and intralaminar nuclei), the brainstem (periaqueductal gray, superior colliculus, pontomesencephalic reticular formation, pontine nuclei, tegmental nuclei) and the spinal cord. In addition to an overall denser ACC projection pattern compared to MCC, our analysis revealed clear differences in the density and topography of efferents between ACC and MCC, as well as between dorsal (24b/24b') and ventral (24a/24a') areas, suggesting a common functionality of these two cingulate regions supplemented by specific roles of each area. These results provide a detailed analysis of the efferents of the mouse areas 24a/24b and 24a'/24b' and achieve the description of the cingulate connectome, which bring the anatomical basis necessary to address the roles of ACC and MCC in mice.

Representations of Fine Digit Movements in Posterior and Anterior Parietal Cortex Revealed Using Long-Train Intracortical Microstimulation in Macaque Monkeys.

The current investigation in macaque monkeys utilized long-train intracortical microstimulation to determine the extent of cortex from which movements could be evoked. Not only were movements evoked from motor areas (PMC and M1), but they were also evoked from posterior parietal (5, 7a, 7b) and anterior parietal areas (3b, 1, 2). Large representations of digit movements involving only the index finger (D2) and thumb (D1), were elicited from areas 1, 2, 7b, and M1. Other movements evoked from these regions were similar to ethologically relevant movements that have been described in other primates. These include combined forelimb and mouth movements and full hand grasps. However, many other movements were much more complex and could not be categorized into any of the previously described ethological categories. Movements involving specific digits, which mimic precision grips, are unique to macaques and have not been described in New World or prosimian primates. We propose that these multiple and expanded motor representations of the digits co-evolved with the emergence of the opposable thumb and alterations in grip type in some anthropoid lineages.

Posterior parietal cortex estimates the relationship between object and body location during locomotion.

We test the hypothesis that the posterior parietal cortex (PPC) contributes to the control of visually guided locomotor gait modifications by constructing an estimation of object location relative to body state, and in particular the changing gap between them. To test this hypothesis, we recorded neuronal activity from areas 5b and 7 of the PPC of cats walking on a treadmill and stepping over a moving obstacle whose speed of advance was varied (slowed or accelerated with respect to the speed of the cat). We found distinct populations of neurons in the PPC, primarily in area 5b, that signaled distance- or time-to-contact with the obstacle, regardless of which limb was the first to step over the obstacle. We propose that these cells are involved in a sensorimotor transformation whereby information on the location of an obstacle with respect to the body is used to initiate the gait modification.

Correlated Disruption of Resting-State fMRI, LFP, and Spike Connectivity between Area 3b and S2 following Spinal Cord Injury in Monkeys

This study aims to understand how functional connectivity (FC) between areas 3b and S2 alters following input deprivation and the neuronal basis of disrupted FC of resting-state fMRI signals. We combined submillimeter fMRI with microelectrode recordings to localize the deafferented digit regions in areas 3b and S2 by mapping tactile stimulus-evoked fMRI activations before and after cervical dorsal column lesion in each male monkey. An average afferent disruption of 97% significantly reduced fMRI, local field potential (LFP), and spike responses to stimuli in both areas. Analysis of resting-state fMRI signal correlation, LFP coherence, and spike cross-correlation revealed significantly reduced functional connectivity between deafferented areas 3b and S2. The degrees of reductions in stimulus responsiveness and FC after deafferentation differed across fMRI, LFP, and spiking signals. The reduction of FC was much weaker than that of stimulus-evoked responses. Whereas the largest stimulus-evoked signal drop (∼80%) was observed in LFP signals, the greatest FC reduction was detected in the spiking activity (∼30%). fMRI signals showed mild reductions in stimulus responsiveness (∼25%) and FC (∼20%). The overall deafferentation-induced changes were quite similar in areas 3b and S2 across signals. Here we demonstrated that FC strength between areas 3b and S2 was much weakened by dorsal column lesion, and stimulus response reduction and FC disruption in fMRI covary with those of LFP and spiking signals in deafferented areas 3b and S2. These findings have important implications for fMRI studies aiming to probe FC alterations in pathological conditions involving deafferentation in humans.SIGNIFICANCE STATEMENT By directly comparing fMRI, local field potential, and spike signals in both tactile stimulation and resting states before and after severe disruption of dorsal column afferent, we demonstrated that reduction in fMRI responses to stimuli is accompanied by weakened resting-state fMRI functional connectivity (FC) in input-deprived and reorganized digit regions in area 3b of the S1 and S2. Concurrent reductions in local field potential and spike FC validated the use of resting-state fMRI signals for probing neural intrinsic FC alterations in pathological deafferented cortex, and indicated that disrupted FC between mesoscale functionally highly related regions may contribute to the behavioral impairments.

Altered Expression of Reorganized Inputs as They Ascend From the Cuneate Nucleus to Cortical Area 3b in Monkeys With Long-Term Spinal Cord Injuries.

Chronic deafferentations in adult mammals result in reorganization of the brain. Lesions of the dorsal columns of the spinal cord at cervical levels in monkeys result in expansion of the intact chin inputs into the deafferented hand representation in area 3b, second somatosensory (S2) and parietal ventral (PV) areas of the somatosensory cortex, ventroposterior lateral nucleus (VPL) of the thalamus, and cuneate nucleus of the brainstem. Here, we describe the extent and nature of reorganization of the cuneate and gracile nuclei of adult macaque monkeys with chronic unilateral lesions of the dorsal columns, and compare it with the reorganization of area 3b in the same monkeys. In both, area 3b and the cuneate nucleus chin inputs expand to reactivate the deafferented neurons. However, unlike area 3b, neurons in the cuneate nucleus also acquire receptive fields on the shoulder, neck, and occiput. A comparison with the previously published results shows that reorganization in the cuneate nucleus is similar to that in VPL. Thus, the emergent topography following deafferentations by spinal cord injuries undergoes transformation as the reorganized inputs ascend from subcortical nuclei to area 3b. The results help us understand mechanisms of the brain plasticity following spinal cord injuries.

P 23 fMRI-mapping of somatosensory finger representation in complex regional pain syndrome

Objective Somatosensory deficits are often reported in patients with complex regional pain syndrome (CRPS). In particular reduced spatial tactile resolution is a typical clinical finding in CRPS patients. The detailed underlying mechanism for development and preservation of these deficits is not yet fully understood. However, a systematic review of studies on pain representation and cerebral representation showed a smaller representation of the hand in the primary somatosensory cortex (S1) contralateral to the affected upper limb ( Di Pietro et al., 2013 ). Associations between S1 hand representation size, pattern of pain and clinical deficits in patients with CRPS are suspected, but studies using high resolution fMRI are rare. Methods In a group of 13 patients diagnosed with CRPS type 1 (CSS: 13.1 ± 9.8; age 54 ± 10 years; handedness: 90 ± 11) and unilateral affection of the upper limb, the thumb (D1) and pinky (D5) representation in Brodmann area (BA) 3b of S1 was determined with our new high resolution fMRI protocol ( Pfannmoller et al., 2016 ). The surface distance between D1 and D5 was used as a measure for the size of the hand representation. In order to investigate the impact of the hand representation size on behavior measures, associations with pain intensity (VAS pain intensity: 4.0 ± 2.5), pinch grip performance (Roeder), and two-point-discrimination (TPD D1) as a measure for the spatial tactile resolution were analyzed. Results Overall, the affected and non-affected hand differed in motor function (Roeder: t (12) = 2.36; p = .036) and tactile resolution (e.g. TPD: t (12) = 2.65; p = .021). Patients with higher D1-D5 distance showed better TPD ( r = −.62; p = .024). We found no association between D1-D5 distance and patient’s pain intensity ( r = −.39; n.s.). Our optimized evaluation revealed no difference between D1-D5 distances of the affected and non-affected side. Without quality optimization the non-affected hand (20.9 ± 4.3 mm) was larger than the affected hand (16.2 ± 3.5 mm) and this difference was highly significant ( t (10) = 4.85; p = .001). However, the non-optimized distances showed no associations to the behavioral measures. Conclusion Our data on an association of D1-D5 S1-distance with spatial tactile resolution are in good agreement with other studies( Pleger et al., 2006 ). The only difference to the existing literature is the similarity of the representation size of non-affected and affected hand. Since our optimized evaluation decreases the statistical significance of the results we assume that this finding is due to our sample size and that the difference between the hand representations may be recovered if more patients are included.

Hemispheric Dominance for Stereognosis in a Patient With an Infarct of the Left Postcentral Sensory Hand Area

The concept of left hemispheric dominance for praxis, speech, and language has been one of the pillars of neurology since the mid-19th century. In 1906, Hermann Oppenheim reported a patient with bilateral stereoagnosia (astereognosis) caused by a left parietal lobe tumor and proposed that the left hemisphere was also dominant for stereognosis. Surprisingly, few cases of bilateral stereoagnosia caused by a unilateral cerebral lesion have been documented in the literature since then. Here we report a 75-year-old right-handed man who developed bilateral stereoagnosia after suffering a small infarct in the crown of the left postcentral gyrus. He could not recognize objects with either hand, but retained the ability to localize stimuli applied to the palm of his left (ipsilesional) hand. He was severely disabled in ordinary activities requiring the use of his hands. The lesion corresponded to Brodmann area 1, where probabilistic anatomic, functional, and electrophysiologic studies have located one of the multiple somatosensory representations of the hand. The lesion was in a strategic position to interrupt both the processing of afferent tactile information issuing from the primary somatosensory cortex (areas 3a and 3b) and the forward higher-order processing in area 2, the secondary sensory cortex, and the contralateral area 1. The lesion also deprived the motor hand area of its afferent regulation from the sensory hand area (grasping), while leaving intact the visuomotor projections from the occipital cortex (reaching). Our patient supports Oppenheim's proposal that the left postcentral gyrus of some individuals is dominant for stereognosis.

Neural encoding of saltatory pneumotactile velocity in human glabrous hand.

Neurons in the somatosensory cortex are exquisitely sensitive to mechanical stimulation of the skin surface. The location, velocity, direction, and adaptation of tactile stimuli on the skin’s surface are discriminable features of somatosensory processing, however the representation and processing of dynamic tactile arrays in the human somatosensory cortex are poorly understood. The principal aim of this study was to map the relation between dynamic saltatory pneumatic stimuli at discrete traverse velocities on the glabrous hand and the resultant pattern of evoked BOLD response in the human brain. Moreover, we hypothesized that the hand representation in contralateral Brodmann Area (BA) 3b would show a significant dependence on stimulus velocity. Saltatory pneumatic pulses (60 ms duration, 9.5 ms rise/fall) were repetitively sequenced through a 7-channel TAC-Cell array at traverse velocities of 5, 25, and 65 cm/s on the glabrous hand initiated at the tips of D2 (index finger) and D3 (middle finger) and sequenced towards the D1 (thumb). The resulting hemodynamic response was sampled during 3 functional MRI scans (BOLD) in 20 neurotypical right-handed adults at 3T. Results from each subject were inserted to the one-way ANOVA within-subjects and one sample t-test to evaluate the group main effect of all three velocities stimuli and each of three different velocities, respectively. The stimulus evoked BOLD response revealed a dynamic representation of saltatory pneumotactile stimulus velocity in a network consisting of the contralateral primary hand somatosensory cortex (BA3b), associated primary motor cortex (BA4), posterior insula, and ipsilateral deep cerebellum. The spatial extent of this network was greatest at the 5 and 25 cm/s pneumotactile stimulus velocities.