The Corticocuneate Pathway in the Cat: Relations among Terminal Distribution Patterns,Cytoarchitecture, and Single Neuron Functional Properties

A combined anatomical and physiological strategy was used to investigate the organization of the corticocuneate pathway in the cat. The distribution of the corticocuneate projection was mapped by means of the anterograde horseradish peroxidase (HRP) labeling technique and correlated with the nuclear cytoarchitecture in Nissl and Golgi material, the distribution of retrogradely labeled relay cells after HRP injections in the ventrobasal complex of the thalamus, and the topographic organization derived from single-and multiunit recordings in the decerebrate, unanesthetized cat. This approach provided details about the arrangement of the corticocuneate pathway that were not available from previous studies with anterograde degeneration methods.

On the basis of cytoarchitectonic and connectional features, a number of subdivisions are identified in the cuneate nucleus, each of which is associated with characteristic functional properties. In agreement with previous studies, it is found that a large portion of the cuneate nucleus, the middle dorsal part (MCd), is exclusively devoted to the representation of cutaneous receptive fields on the digits. This “core” region contains more thalamic projecting neurons than any other subdivision of the cuneate nucleus. A topographic arrangement also exists in the subdivisions of the rostral cuneate and of the nuclear region ventral to MCd, although in these, receptive fields are larger and predominantly, but not exclusively, related to deep receptors and involve the arm, shoulder, and trunk.

Observations on corticocuneate projections were based on injections, mainly focused on functional subdivisions of the primary somatosensory cortex (SI) as described by McKenna et al (1981). Although cortical projections are mainly to cuneate regions other than its core, a significant proportion of fibers from the region of SI where the digits are represented (particularly area 3b) do project to the MCd region of the cuneate nucleus. Similarly, nuclear areas associated with receptive fields on the arm and trunk are labeled after injection in SI arm and trunk regions, respectively. Thus, a close topographic relationship appears to exist between the somatosensory cortex and cuneate regions related to the same body representation, although nuclear regions in which receptive fields on the neck area are represented receive very sparse or no detectable cortical projections even when the injection of the tracer involves the entire sensorimotor cortex. The topographic arrangement of SI projections upon the cuneate nucleus suggests that a similar pattern exists in both structures with regard to the relative representations of distal versus proximal and deep versus cutaneous receptive fields (e.g., “core” vs. “shell” organization), and that cuneate regions preferentially related to either of these classes of receptive fields receive direct connections from the corresponding regions in SI.

A comparison of the results from cats with tracer injections in areas 4 and 3b reveals that the projections from the former is denser than that arising from the latter and that their territories of termination largely overlap in the ventral portions of the cuneate nucleus. However, cortical projections to MCd may be derived from the somatosensory cortex with no contribution from area 4. The demonstration of the relative selectivity of cortical projections from different cytoarchitectonic and functional cortical areas to cuneate regions identified here provides a structural basis for the elucidation of the physiological and behavioral observations, particularly on cortical modulation of somatosensory transmission during movements.     

Experience-induced plasticity of cutaneous maps in the primary somatosensory cortex of adult monkeys and rats

In a first study, the representations of skin surfaces of the hand in the primary somatosensory cortex, area 3b, were reconstructed in owl monkeys and squirrel monkeys trained to pick up food pellets from small, shallow wells, a task which required skilled use of the digits. Training sessions included limited manual exercise over a total period of a few hours of practice. From an early clumsy performance in which many retrieval attempts were required for each successful pellet retrieval, the monkeys exhibited a gradual improvement. Typically, the animals used various combinations of digits before developing a successful retrieval strategy. As the behavior came to be stereotyped, monkeys consistently engaged surfaces of the distal phalanges of one or two digits in the palpation and capture of food pellets from the smallest wells. Microelectrode mapping of the hand surfaces revealed that the glabrous skin of the fingertips predominantly involved in the dexterity task was represented over topographically expanded cortical sectors. Furthermore, cutaneous receptive fields which covered the most frequently stimulated digital tip surfaces were less than half as large as were those representing the corresponding surfaces of control digits. In a second series of experiments, Long-Evans rats were assigned to environments promoting differential tactile experience (standard, enriched, and impoverished) for 80 to 115 days from the time of weaning. A fourth group of young adult rat experienced a severe restriction of forepaw exploratory movement for either 7 or 15 days. Cortical maps derived in the primary somatosensory cortex showed that environmental enrichment induced a substantial enlargement of the cutaneous forepaw representation, and improved its spatial resolution (smaller glabrous receptive fields). In contrast, tactile impoverishment resulted in a degradation of the forepaw representation that was characterized by larger cutaneous receptive fields and the emergence of non-cutaneous responses. Cortical maps derived in the hemispheres contralateral to the immobilized forelimb exhibited a severe decrease of about 50% in the overall areal extent of the cutaneous representation of the forepaw, which resulted from the invasion of topographically organized cortical zones of non-cutaneous responses, and numerous discontinuities in the representation of contiguous skin territories. The size and the spatial arrangement of the cutaneous receptive fields were not significantly modified by the immobilization of the contralateral forelimb. Similar results were obtained regardless of whether the forelimb restriction lasted 7 or 15 days. These two studies corroborate the view that representational constructs are permanently reshaped by novel experiences through dynamic competitive processes. These studies also support the notion that subject-environment interactions play a crucial role in the maintenance of basic organizational features of somatosensory representations.     

Ablations of areas 3b (SI proper) and 3a of somatosensory cortex in marmosets deactivate the second and parietal ventral somatosensory areas

Partial ablations of specific parts of cortical areas 3b (SI proper) and 3a in marmosets were found to render somatotopically equivalent parts of two other cortical somatosensory fields, the second somatosensory area (SII) and the parietal ventral area (PV), unresponsive to peripheral stimulation. Microelectrode recordings in anesthetized marmosets first established the responsiveness and locations of the representations of body parts, including the hand in areas 3a and 3b, SII, and in some cases PV. The hand representations in areas 3a and 3b were then removed by aspiration. Immediately afterwards, additional recordings established that regions of SII and PV that formerly represented the hand were no longer responsive to cutaneous stimulation of the hand (or any other skin surface). Other parts of these fields, representing parts of the body other than the hand, remained responsive to stimulation of the previously effective receptive fields. We conclude that SII and PV depend on inputs (either direct or indirect) from areas 3a and 3b for their activation.     

Multiple Representations of the Body and Input-Output Relationships in the Agranular and Granular Cortex of the Chronic Awake Guinea Pig

The organization of somatosensory input and the input-output relationships in regions of the agranular frontal cortex (AGr) and granular parietal cortex (Gr) were examined in the chronic awake guinea pig, using the combined technique of single-unit recording and intracortical microstimulation (ICMS). AGr, which was cytoarchitectonically subdivided into medial (AGrm) and lateral (AGrl) parts, also can be characterized on a functional basis. AGrl contains the head, forelimb, and most hindlimb representations; only a small number of hindlimb neurons are confined in AGrm. Different distributions of submodalities exist in AGr and Gr: AGr receives predominantly deep input (with the exception of the vibrissa region, which receives cutaneous input), whereas neurons of Gr respond almost exclusively to cutaneous input. The cutaneous or deep receptive field (RF) of each neuron was determined by natural peripheral stimulation. All studied neurons were activated by small RFs, with the exception of lip, nose, pinna, and limb units of lateral Gr (Grl), for which the RFs were larger.

Microelectrode mapping experiments revealed the existence of three spatially separate, incomplete body maps in which somatosensory and motor representations overlap. One body map, with limbs medially and head rostrolaterally, is contained in AGr. A second map, comparable to the first somatosensory cortex (SI) of other mammals, is found in Gr, with hindlimb, trunk, forelimb, and head representations in an orderly mediolateral sequence. An unresponsive zone separates the head area from the forelimb region. A third map, with the forelimb rostrally and the hindlimb caudally, lies adjacent and lateral to the SI head area. This limb representation, which is characterized by an upright and small size compared to that found in SI, can be considered to be part of the second somatosensory cortex (SII). A distinct head representation was not recognized as properly belonging to SII, but the evidence that neurons of the SI head region respond to stimulation of large RFs located in lips, nose, and pinna leads us to hypothesize that the SII face area overlaps that of SI to some extent, or, alternatively, that the two areas are strictly contiguous and the limits are ambiguous, making them difficult to distinguish.

The input-output relationships were based on the results of RF mapping and ICMS in the same electrode penetration. The intrinsic specific interconnections of cortical neurons whose afferent input and motor output is related to identical body regions show a considerable degree of refinement. The input-output correspondence is especially pronounced for neurons with small RFs. This study confirms and extends similar data recently reported for other rodents.     

Motor organization of the lateral cerebellar nucleus of the rat

The motor organization of the nucleus lateralis (NL) of the cerebellum of the rat was investigated by studying the motor effects following the electrical microstimulation. The movements evoked by the NL stimulation concerned prevalently the forelimb and the head segments. The movements of the hindlimb segments were evoked in only few cases. The NL is organized as a mosaic of zones without, or at least very little overlap. The various body segments are differently represented in the NL. Some of them are once represented (single representations). In other cases, the same movements were evoked by different NL regions (multiple representations). Finally, in a last lot of cases, various representations concerned the same body regions but from each representation a different type of movement was evoked (specific representations, i.e. displacement of an individual digit and flexion of all digits together). The topographical distribution of the representations in the NL cytological regions (magnicellularis, NLm; dorsolateral hump, DLH; subnucleus lateralis parvocellularis, slp) suggests the idea that each of them may be concerned in a specific motor activity: the NLm would control the position of the body, or of part of it, in the space; the DLH would be concerned in the oral (prevalently) and in the forelimb motor activity; the slp would be concerned in the exploration of the environment as well as in skilled movements of the distalmost forelimb segments.     

Somatotopic organization and cortical projections of the ventrobasal complex of the flying fox: an "inverted" wing representation in the thalamus

The present study investigates the somatotopic representation in the somatosensory thalamus of a megachiropteran bat. Using standard microelectrode mapping techniques, representational maps were generated for the ventrobasal (Vb) and posterior (Po) thalamic complexes of the Grey-headed flying fox. Anatomical tracing from neocortical injections provided additional data confirming the somatotopy found physiologically. A full representation of the body surface innervated by the trigeminal and spinal nerves was found. However, in contrast with other mammals, the representations of the forelimb and adjacent thoracic trunk within the thalamus were inverted. This means that the distal portions of the wing membrane and the tips of the digits were represented dorsally in Vb, and the thoracic trunk was represented ventrally. In Po the digit tips were represented in the ventral most portion and the thoracic trunk in the dorsal portion of the nucleus.These results are discussed in relation to similarities of megachiropteran somatosensory thalamic nuclei to those of other mammalian species and in relation to the formation of thalamic somatotopic maps and fiber sorting.     

Somatotopic organization and cortical projections of the ventrobasal complex of the flying fox: an "inverted" wing representation in the thalamus

The present study investigates the somatotopic representation in the somatosensory thalamus of a megachiropteran bat. Using standard microelectrode mapping techniques, representational maps were generated for the ventrobasal (Vb) and posterior (Po) thalamic complexes of the Grey-headed flying fox. Anatomical tracing from neocortical injections provided additional data confirming the somatotopy found physiologically. A full representation of the body surface innervated by the trigeminal and spinal nerves was found. However, in contrast with other mammals, the representations of the forelimb and adjacent thoracic trunk within the thalamus were inverted. This means that the distal portions of the wing membrane and the tips of the digits were represented dorsally in Vb, and the thoracic trunk was represented ventrally. In Po the digit tips were represented in the ventral most portion and the thoracic trunk in the dorsal portion of the nucleus. These results are discussed in relation to similarities of megachiropteran somatosensory thalamic nuclei to those of other mammalian species and in relation to the formation of thalamic somatotopic maps and fiber sorting.     

Integration of Sensory Quanta in Cuneate Nucleus Neurons In Vivo

Discriminative touch relies on afferent information carried to the central nervous system by action potentials (spikes) in ensembles of primary afferents bundled in peripheral nerves. These sensory quanta are first processed by the cuneate nucleus before the afferent information is transmitted to brain networks serving specific perceptual and sensorimotor functions. Here we report data on the integration of primary afferent synaptic inputs obtained with in vivo whole cell patch clamp recordings from the neurons of this nucleus. We find that the synaptic integration in individual cuneate neurons is dominated by 4–8 primary afferent inputs with large synaptic weights. In a simulation we show that the arrangement with a low number of primary afferent inputs can maximize transfer over the cuneate nucleus of information encoded in the spatiotemporal patterns of spikes generated when a human fingertip contact objects. Hence, the observed distributions of synaptic weights support high fidelity transfer of signals from ensembles of tactile afferents. Various anatomical estimates suggest that a cuneate neuron may receive hundreds of primary afferents rather than 4–8. Therefore, we discuss the possibility that adaptation of synaptic weight distribution, possibly involving silent synapses, may function to maximize information transfer in somatosensory pathways.     

Somatosensory areas S2 and PV project to the superior colliculus of a prosimian primate,Galago garnetti

As part of an effort to describe the connections of the somatosensory system in Galago garnetti, a small prosimian primate, injections of tracers into cortex revealed that two somatosensory areas, the second somatosensory area (S2) and the parietal ventral somatosensory area (PV), project densely to the ipsilateral superior colliculus, while the primary somatosensory area (S1 or area 3b) does not. The three cortical areas were defined in microelectrode mapping experiments and recordings were used to identify appropriate injection sites in the same cases. Injections of wheat germ agglutinin conjugated with horseradish peroxidase (WGA-HRP) were placed in S1 in different mediolateral locations representing body regions from toes to face in five galagos, and none of these injections labeled projections to the superior colliculus. In contrast, each of the two injections in the face representation of S2 in two galagos and three injections in face and forelimb representations of PV in three galagos produced dense patches of labeled terminations and axons in the intermediate gray (layer IV) over the full extent of the superior colliculus. The results suggest that the higher-order somatosensory areas, PV and S2, are directly involved in the visuomotor functions of the superior colliculus in prosimian primates, while S1 is not. The somatosensory inputs appear to be too widespread to contribute to a detailed somatotopic representation in the superior colliculus, but they may be a source of somatosensory modulation of retinotopically guided oculomotor instructions.     

Somatosensory areas S2 and PV project to the superior colliculus of a prosimian primate, Galago garnetti

As part of an effort to describe the connections of the somatosensory system in Galago garnetti, a small prosimian primate, injections of tracers into cortex revealed that two somatosensory areas, the second somatosensory area (S2) and the parietal ventral somatosensory area (PV), project densely to the ipsilateral superior colliculus, while the primary somatosensory area (S1 or area 3b) does not. The three cortical areas were defined in microelectrode mapping experiments and recordings were used to identify appropriate injection sites in the same cases. Injections of wheat germ agglutinin conjugated with horseradish peroxidase (WGA-HRP) were placed in S1 in different mediolateral locations representing body regions from toes to face in five galagos, and none of these injections labeled projections to the superior colliculus. In contrast, each of the two injections in the face representation of S2 in two galagos and three injections in face and forelimb representations of PV in three galagos produced dense patches of labeled terminations and axons in the intermediate gray (layer IV) over the full extent of the superior colliculus. The results suggest that the higher-order somatosensory areas, PV and S2, are directly involved in the visuomotor functions of the superior colliculus in prosimian primates, while S1 is not. The somatosensory inputs appear to be too widespread to contribute to a detailed somatotopic representation in the superior colliculus, but they may be a source of somatosensory modulation of retinotopically guided oculomotor instructions.     

The effects of long-standing limb loss on anatomical reorganization of the somatosensory afferents in the brainstem and spinal cord

We examined the terminations of sensory afferents in the brainstem and spinal cord of squirrel monkeys and prosimian galagos 4-8 years after a therapeutic forelimb or hindlimb amputation within 2 months of birth. In each animal, the distributions of labeled sensory afferent terminations from remaining body parts proximal to the limb stump were much more extensive than in normal animals. These sprouted afferents extended into the portions of the dorsal horn of the spinal cord as well as the cuneate and external cuneate nuclei of the brainstem (forelimb amputees) or spinal Clarke's column (hindlimb amputee) related to the amputated limb. Such reorganization in sensory afferents along with reorganization of the motor efferents to muscles (Wu and Kaas, J Neurosci 19: 7679-7697, 1999, Neuron 28: 967-978, 2000) may provide a basis for mislocated phantom sensations of missing forelimb movements accompanying actual shoulder movements during cortical stimulation or movement imagery in patients with amputations.     

The effects of long-standing limb loss on anatomical reorganization of the somatosensory afferents in the brainstem and spinal cord

We examined the terminations of sensory afferents in the brainstem and spinal cord of squirrel monkeys and prosimian galagos 4-8 years after a therapeutic forelimb or hindlimb amputation within 2 months of birth. In each animal, the distributions of labeled sensory afferent terminations from remaining body parts proximal to the limb stump were much more extensive than in normal animals. These sprouted afferents extended into the portions of the dorsal horn of the spinal cord as well as the cuneate and external cuneate nuclei of the brainstem (forelimb amputees) or spinal Clarke's column (hindlimb amputee) related to the amputated limb. Such reorganization in sensory afferents along with reorganization of the motor efferents to muscles (Wu and Kaas, J Neurosci 19 : 7679-7697, 1999, Neuron 28 : 967-978, 2000) may provide a basis for mislocated phantom sensations of missing forelimb movements accompanying actual shoulder movements during cortical stimulation or movement imagery in patients with amputations.     

Mechanisms of pain arising from articular tissues

This paper reviews the peripheral and central neural mechanisms underlying pain from articular tissues innervated by spinal and trigeminal afferents. The paper especially addresses trigeminal mechanisms related to pain from the temporomandibular joint and its associated craniofacial musculature. Recent studies have shown the existence of articular nociceptive primary afferents that project to the spinal cord dorsal horn and trigeminal brainstem complex. A particular feature of most neurones receiving these deep nociceptive afferent inputs is their responsivity also to cutaneous nociceptive afferent inputs. This suggests the involvement of these neurones not only in the detection of acute articular pain, but also in the hyperalgesia and poor localization, spread, and referral of pain that characterize many painful conditions of joints and other deep structures. While only limited information is available on related higher brain centre mechanisms, convergence and interaction between cutaneous and deep afferent inputs also seem to be a characteristic of somatosensory neurones in the thalamus and somatosensory cerebral cortex. Muscle and autonomic reflexes may be induced by such deep noxious stimuli, but the functional significance of some of these effects (e.g., in relation to clinical concepts of myofascial dysfunction) requires further study in more appropriate functional settings.     

Somatosensory Event-related Potentials from Orofacial Skin Stretch Stimulation

Cortical processing associated with orofacial somatosensory function in speech has received limited experimental attention due to the difficulty of providing precise and controlled stimulation. This article introduces a technique for recording somatosensory event-related potentials (ERP) that uses a novel mechanical stimulation method involving skin deformation using a robotic device. Controlled deformation of the facial skin is used to modulate kinesthetic inputs through excitation of cutaneous mechanoreceptors. By combining somatosensory stimulation with electroencephalographic recording, somatosensory evoked responses can be successfully measured at the level of the cortex. Somatosensory stimulation can be combined with the stimulation of other sensory modalities to assess multisensory interactions. For speech, orofacial stimulation is combined with speech sound stimulation to assess the contribution of multi-sensory processing including the effects of timing differences. The ability to precisely control orofacial somatosensory stimulation during speech perception and speech production with ERP recording is an important tool that provides new insight into the neural organization and neural representations for speech.     

The second somatosensory region of the cat thalamus

Recording focal evoked potentials showed that the second somatosensory region of the cat thalamus is located in the posterolateral part of the ventral posteromedial nucleus. The zone of representation of the hindlimb lies somewhat laterally, posteriorly, and superiorly to the zone of the forelimb. The zone of representation of the mouth lies more medially still. Latent periods of responses arising in both regions of the thalamus to cutaneous stimulation of the contralateral limbs are about equal. Neurons of the second somatosensory region of the thalamus send projections to cortical area S2. It can be concluded from the results that formation of the structure of double representation of the somatosensory systems in the cat is completed at the thalamic level.     

Incorporation of neurons of transplants of embryonal rat neocortex in the achievement of sensory function of the cerebral cortex of the recipient

The barrelfield of the adult rats was removed by suction and embryonic tissue of the somatosensory neocortex was transplanted into the cavity. Spontaneous and evoked activity of the grafted neurones was investigated extracellularly 2-3 months after the grafting. The light microscopy of the grafts revealed the presence of normal neuronal cells, but their distribution was diffuse, and they were not organized into barrels as in intact neocortex. The background activity of grafted neurones depended upon the level of the recipient's anaesthesia. The response types of the grafted neurones to vibrissae deflection and to tactile stimulation of the host body surfaces, their latencies and lability did not differ from such of the intact somatosensory cortex, but the receptive fields of the grafted neurones were larger. There was also substantial convergence of inputs from other surfaces upon the grafted neurones. The effectiveness of stimulation of the various skin areas was determined by the proximity of their neocortical representations to the graft.     

The organization and connections of somatosensory cortex in the brush-tailed possum (Trichosurus vulpecula): evidence for multiple, topographically organized and interconnected representations in an Australian marsupial

Microelectrode mapping techniques were used to determine the organization of somatosensory cortex in the Australian brush-tailed possum (Trichosurus vulpecula). The results of electrophysiological mapping were combined with data on the cyto- and myeloarchitecture, and patterns of corticocortical connections, using sections cut tangential to the pial surface. We found evidence for three topographically organized representations of the body surface that were coextensive with architectonic subdivisions. A large, discontinuous cutaneous representation in anterior parietal cortex was termed the primary somatosensory area (SI). Lateral to SI we found evidence for two further areas, the second somatosensory area (SII) and the parietal ventral area (PV). While neurones in all of these areas were responsive to cutaneous stimulation, those of SI were non-habituating, whereas those in SII and PV often habituated to the stimuli. Moreover, neuronal receptive fields in SII and PV were, in general, larger than those in SI. Neurones in cortex adjacent to the rostral and caudal boundaries of SI, including cortex that interdigitated between the discontinuous SI head and body representations, required stimulation of deep receptors in the periphery to elicit responses. Within the region of cortex containing neurones responsive to stimulation of deep receptors, body parts were represented in a mediolateral progression. Injections of anatomical tracers placed in electrophysiologically identified locations in SI revealed ipsilateral connections with other parts of SI, as well as cortex rostral to, caudal to, and interdigitating between, SI. Injections in SI also resulted in labelling in PV, SII, motor cortex, posterior parietal cortex and perirhinal cortex. The patterns of contralateral projections reflected those of ipsilateral projections, although they were relatively less dense. The present findings support recent observations in other marsupials in which multiple representations of the body surface were described, and suggest that multiple interconnected sensory representations may be a common feature of cortical organization and function in marsupials.     

Barrelettes without Barrels in the American Water Shrew

Water shrews (Sorex palustris) depend heavily on their elaborate whiskers to navigate their environment and locate prey. They have small eyes and ears with correspondingly small optic and auditory nerves. Previous investigations have shown that water shrew neocortex is dominated by large representations of the whiskers in primary and secondary somatosensory cortex (S1 and S2). Flattened sections of juvenile cortex processed for cytochrome oxidase revealed clear borders of the whisker pad representation in S1, but no cortical barrels. We were therefore surprised to discover prominent barrelettes in brainstem of juvenile water shrews in the present investigation. These distinctive modules were found in the principal trigeminal nucleus (PrV), and in two of the three spinal trigeminal subnuclei (interpolaris – SpVi and caudalis – SpVc). Analysis of the shrew''s whisker pad revealed the likely relationship between whiskers and barrelettes. Barrelettes persisted in adult water shrew PrV, but barrels were also absent from adult cortex. Thus in contrast to mice and rats, which have obvious barrels in primary somatosensory cortex and less clear barrelettes in the principal nucleus, water shrews have clear barrelettes in the brainstem and no barrels in the neocortex. These results highlight the diverse ways that similar mechanoreceptors can be represented in the central nervous systems of different species.     

A Comparison of the Ipsilateral Cortical Projections to the Dorsal and Ventral Subdivisions of the Macaque Premotor Cortex

The cortical connections of the dorsal (PMd) and ventral (PMv) subdivisions of the premotor area (PM, lateral area 6) were studied in four monkeys (Macaca fascicularis) through the use of retrograde tracers. In two animals, tracer was injected ventral to the arcuate sulcus (PMv), in a region from which forelimb movements could be elicited by intracortical microstimulation (ICMS). Tracer injections dorsal to the arcuate sulcus (PMd) were made in two locations. In one animal, tracer was injected caudal to the genu of the arcuate sulcus (in caudal PMd [cPMd], where ICMS was effective in eliciting forelimb movements); in another animal, it was injected rostral to the genu of the arcuate sulcus (in rostral PMd [rPMd], where ICMS was ineffective in eliciting movements). Retrogradely labeled neurons were counted in the ipsilateral hemisphere and located in cytoarchitectonically identified areas of the frontal and parietal lobes. Although both PMv and PMd were found to receive inputs from other motor areas, the prefrontal cortex, and the parietal cortex, there were differences in the topography and the relative strength of projections from these areas.

There were few common inputs to PMv and PMd; only the supplementary eye fields projected to all three areas studied. Interconnections within PMd or PMv appeared to link hindlimb and forelimb representations, and forelimb and face representations; however, connections between PMd and PMv were sparse. Areas cPMd and PMv were found to receive inputs from other motor areas—the primary motor area, the supplementary motor area, and the cingulate motor area—but the topography and strength of projections from these areas varied. Area rPMd was found to receive sparse inputs, if any, from these motor areas. The frontal eye field (area 8a) was found to project to PMv and rPMd, and area 46 was labeled substantially only from rPMd. Parietal projections to PMv were found to originate from a variety of somatosensory and visual areas, including the second somatosensory cortex and related areas in the parietal operculum of the lateral sulcus, as well as areas 5, 7a, and 7b, and the anterior intraparietal area. By contrast, projections to cPMd arose only from area 5. Visual areas 7m and the medial intraparietal area were labeled from rPMd. Relatively more parietal neurons were labeled after tracer injections in PMv than in PMd. Thus, PMv and PMd appear to be parts of separate, parallel networks for movement control.     

Face or hand,not both: perceptual correlates of reafferentation in a former amputee

The topography of the somatosensory maps of our body can be largely shaped by alterations of peripheral sensory inputs. Following hand amputation, the hand cortical territory becomes responsive to facial cutaneous stimulation. Amputation-induced remapping, however, reverses after transplantation, as the grafted hand (re)gains its sensorimotor representation. Here, we investigate hand tactile perception in a former amputee by touching either grafted hand singly or in combination with another body part. The results showed that tactile sensitivity recovered rapidly, being remarkably good 5 months after transplant. In the right grafted hand, however, the newly acquired somatosensory awareness was strikingly hampered when the ipsilateral face was touched simultaneously, i.e., right face perception extinguished right hand perception. Ipsilateral face-hand extinction was present in the formerly dominant right hand 5 months after transplant and eventually disappeared 6 months afterwards. Control conditions' results showed that right hand tactile awareness was not extinguished either by contralateral left face and left hand stimulation or ipsilateral stimulation of the arm, which is bodily close to, but cortically far from, the hand. We suggest that ipsilateral face-hand extinction is a perceptual counterpart of the remapping that occurs after allograft and eyewitnesses the inherently competitive nature of sensory representations.