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At least 19 recordsLinked to original sources

Changes in dry weight of pyramidal cell nuclei in the motor cortex during local motor-food conditioning.

The formation of a local motor-food conditioned reflex in rats, consisting of pressing on a level with the unpreferred paw, in one session led to a decrease in dry weight of the large pyramidal cell nuclei in the area of representation of that paw in the motor cortex. Frequent repetition of the local motor-food conditioned reflex consisting of pressing on the level with the preferred paw was not accompanied by any such changes. It is suggested that changes in dry weight of the large pyramidal cell nuclei in the area of representation of the unpreferred paw are due to involvement of these neurons in the formation of the new motor coordination (the local conditioned reflex).

Animals

Patterns of projections from area 2 of the sensory cortex to area 3a and to the motor cortex in cats.

Peripheral information reaches the motor cortex partly through corticocortical pathways that arise from two functional subdivisions, area 2 and area 3a, of the sensory cortex. These sensory areas are synaptically linked with one another. The patterns of connectivity and the different submodality input that each area receives suggest that they send different efferent signals to the motor cortex. The projections from area 2 to area 3a and to the motor cortex were studied with retrogradely transported fluorescent tracers. The pattern and distribution of neuronal labeling in area 2 was determined following injections of different tracers into the forelimb regions of area 3a and the motor cortex. The results showed that the projections from area 2 to the two target regions were topographically and somatotopically related. Multiple clusters of motor cortex projection neurons were found in area 2, and these clusters overlapped extensively with clusters of area 3a projection neurons. Although cells labeled with one of the dyes were often in close proximity to cells labeled with the other dye, no double-labeled cells were found. Two different laminar patterns were seen for the two populations of neurons. The projection to area 3a originated from cells located in layers II-III and layers V-VI. The projection to the motor cortex originated from cells spread throughout layers II-IV, but predominantly in layer III. Differences in laminar arrangement of the two populations of cells suggest a directional flow of information processing in the sensorimotor cortex.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals

Effects of anodal transcranial direct current stimulation over the right primary motor cortex on a sequential motor finger tapping task in developmental stuttering.

INTRODUCTION: This study investigates the impact of anodal transcranial direct current stimulation (tDCS) on non-speech sequential motor practice in adults who stutter (AWS), compared to non-stuttering controls (ANS). Recent research has explored the effects of tDCS on speech fluency in stuttering. However, its effect on non-speech motor tasks has not yet been studied. METHODS: 20 AWS and 30 ANS right-handed participants were randomly assigned to anodal or sham tDCS conditions, performing a sequential finger tapping task. We targeted over the right primary motor cortex, stimulating at 2 mA for 20 min. Sequence duration and reaction time were analyzed. RESULTS: AWS analysis revealed that the anodal condition had significantly slower reaction times in the second half of the task compared to sham. For sequence durations, AWS in the anodal condition had slower overall sequence durations than the sham condition. However, there were no block-by-block differences in sequence duration. When comparing AWS and ANS, no significant differences were observed for sequence duration. However, there were significant differences in reaction time between AWS and ANS, specifically in earlier blocks. Additionally, there was no significant Group × Condition interaction. DISCUSSION: The findings suggest that anodal stimulation impeded finger sequencing in AWS, showing overall slower sequence durations and a diminishing effect on reaction times in the second half of the experiment, suggesting anodal tDCS may interact uniquely with the neural mechanisms in stuttering. Future studies should explore the effects of anodal tDCS on non-speech motor tasks to gain a broader understanding of its impact on motor control and motor learning.

Humans

Peripheral input pathways projecting to the motor cortex in the cat.

The possibility that the motor cortex receives peripheral input directly from the thalamus was examined using the evoked potential method and the following results were obtained. Potentials in the motor cortex evoked by stimulation of superficial radial (SR) or group II deep radial (DR) nerve were neither abolished nor delayed by ablation of the sensory cortex. Potentials in the motor cortex evoked by stimulation of group II DR nerve were most severely reduced by interruption of the spinocervcial tract. Potentials evoked by stimulation of SR nerve were more severely reduced in the sensory cortex than in the motor cortex by section of the dorsal funiculus or cooling of the cuneate nucleus. The size of evoked potentials in the motor cortex increased rapidly when stimulus intensity to DR nerve exceeded the threshold to group II fibers. The results suggest that some inputs from the SR and group II DR nerves reach the motor cortex without a relay through the sensory cortex.

Animals

Somatotopic localization in cat motor cortex.

Punctate intracortical stimulation of the motor cortex (areas 4 and 6), with parallel observation of the induced movements, permits description of a fine somatotopic organization of the motor control areas for different parts of the musculature in freely moving adult cats. The results show that movements produced by electrical stimulation of the motor cortex are always single and non-repetitive, regardless of the duration and intensity of the stimulation. These movements are restricted to a very precise part of the musculature, and experiments show that this localization is related to the exact position of the tip of the stimulating electrode in the motor cortex. Other experimental data show that motor responses which disturb the animal's equilibrium are accompanied by postural adjustments. Stimulation of the cerebral cortex permits the definition of a separate motor control area for each part of the cat musculature, with an individual control area for each of the joints of the forelimb. This was not possible for the hindlimb, which is always activated in its entirety. These results establish a new representation of the somatotopic organization of the cat motor cortex. This diagram shows that area 6 controls the more axial parts of the musculature, while area 4 controls the proximal and distal parts of the limb muscles. This map was compared to numerous previous data on the somatotopic organization in the cat motor cortex, especially to the map of Woolsey.

Animals

High-frequency contralesional dorsal premotor cortex and low-frequency contralesional primary motor cortex rTMS in subacute stroke with severe upper limb impairment: comparable motor outcomes and differential regional degree centrality changes.

BACKGROUND: The contralesional dorsal premotor cortex has been proposed as a potential neuromodulatory target for patients with severe upper limb impairment due to subacute ischemic stroke. This proof-of-concept study aimed to compare behavioral outcomes and resting-state neuroimaging findings between high-frequency repetitive transcranial magnetic stimulation (rTMS) over the contralesional dorsal premotor cortex and guideline-supported low-frequency stimulation over the contralesional primary motor cortex. METHODS: In this randomized trial, 46 patients with severe upper limb impairment in the subacute stage after ischemic stroke were randomly assigned to receive either high-frequency rTMS over the contralesional dorsal premotor cortex or low-frequency rTMS over the contralesional primary motor cortex. Low-frequency stimulation over the contralesional primary motor cortex served as an evidence-supported active comparator for poststroke upper limb motor recovery. Stimulation was administered five times per week for two weeks using magnetic resonance imaging-guided neuronavigation. All participants received concurrent standard rehabilitation therapy. The primary outcome was the Fugl-Meyer Assessment for Upper Extremity. Secondary outcomes included the Arm Subscore of the Motricity Index, the Hong Kong version of the Functional Test for the Hemiplegic Upper Extremity, the Modified Barthel Index, and resting-state functional magnetic resonance imaging-derived degree centrality. RESULTS: Both groups showed significant improvements in the primary and secondary behavioral measures (p&#x202f;<&#x202f;0.01), with no significant between-group differences in the magnitude of change (p&#x202f;>&#x202f;0.05). In neuroimaging analyses, patients receiving high-frequency rTMS over the contralesional dorsal premotor cortex showed significantly greater degree centrality changes in the ipsilesional middle occipital gyrus, contralesional medial superior frontal gyrus, and contralesional middle frontal gyrus than those receiving low-frequency rTMS over the contralesional primary motor cortex (p&#x202f;<&#x202f;0.05). Within the high-frequency stimulation group, degree centrality changes in the ipsilesional middle occipital gyrus were positively correlated with improvements in the Fugl-Meyer Assessment for Upper Extremity (r&#x202f;=&#x202f;0.619, false discovery rate-corrected p&#x202f;=&#x202f;0.018). CONCLUSIONS: High-frequency rTMS over the contralesional dorsal premotor cortex produced behavioral improvements comparable to guideline-supported low-frequency rTMS over the contralesional primary motor cortex, without establishing superiority or formal non-inferiority. Exploratory neuroimaging analyses showed greater degree centrality changes in the ipsilesional middle occipital gyrus after high-frequency premotor stimulation, and these changes correlated with upper-limb motor improvement. These findings support further investigation of contralesional dorsal premotor cortex-targeted high-frequency rTMS for severe subacute post-stroke upper limb impairment. REGISTRATION: URL: http://www.chictr.org.cn; Unique identifier: ChiCTR2000038049.

Humans

Early and late lower limb motor evoked potentials elicited by transcranial magnetic motor cortex stimulation.

Transcranial magnetic motor cortex stimulation can elicit a series of responses recorded with different latencies from relaxed muscles of the lower limbs. In 7 healthy subjects, ranging in age from 16 to 62 years, stimulation was delivered by a 9 cm coil centered over Cz with the subject in the supine position. Surface polyelectromyography was used to record motor evoked potentials (MEPs) from the quadriceps (QD), hamstrings (HS), tibialis anterior (TA) and triceps surae (TS) muscles bilaterally. Three characteristic responses were identified in each muscle group on the basis of amplitude and latency criteria, identified by latencies: the direct oligosynaptic response MEP30 appeared with a latency of 24.3 msec in the QD, 26.3 msec in the HS, 30.5 msec in the TA and 31.3 msec in the TS; MEP70 with latencies of 64 msec in the QD, 59 msec in the HS, 79 msec in the TA and 72 msec in the TS; MEP120 with latencies of 115 msec in the QD, 126 msec in the HS, 117 msec in the TA and 124 msec in the TS. These 3 responses have distinct latencies, amplitudes and durations. MEP70 appears to be the result of activation of long descending tracts which end on spinal interneuronal circuits. As MEP120 has different features, it may have a different mechanism.

Adolescent

Anatomical analysis of ventrolateral thalamic input to primate motor cortex.

1. The origin and topographical organization of input to the arm area of the primate motor cortex from the ventrolateral thalamus were examined using the method of retrograde transport of horseradish peroxidase (HRP). 2. A thin, continuous slab of labeled neurons was found in the ventrolateral thalamus followingmultiple injections of HRP into the arm area of the motor cortex. The slab of labeled neurons was flanked, medially and laterally, by groups of unlabeled neurons. 3. The origin of ventrolateral thalamic input was more extensive than previously thought. Labeled neurons were found from A10.0 to A6.0 and occurred in three ventolateral thalamic subdivisions: ventralis lateralis pars oralis (VLo), ventralis lateralis pars caudalis (VLc), and ventralis posterior lateralis pars oralis (VPLo). For simplicity this region containing labeled neurons has been termed the ventrolateral thalamic (VL) arm area. 4. Injections of HRP into the somatic sensory cortex indicated that the thalamic regions which project to the somatic sensory cortex are separate from the VL arm area. 5. The distribution of labeled neurons following single injections of HRP into different regions of the motor cortex arm area indicated that the VL arm area is topographically organized, particularly its caudal part. Ventral regions of the VL arm area were labeled following HRP injections into motor cortex regions adjacent to the central sulcus where the representation of largely distal musculature is localized. Dorsal regions of the VL arm area were labeled following HRP injections into motor cortex regions more rostral to the central sulcus where the representation of more proximal musculature is localized. 6. A larger region of the VL arm area was labeled following HRP injections adjacent to the central sulcus than following the more rostral motor cortex injections. This suggests that, like the arm area of the motor cortex, more of the VL arm area is allotted to the representation of distal than proximal musculature. 7. Following very small cortical HRP injections, isolated labeled thalamic neurons were diffusely scattered throughout a 3-mm rostrocaudal extent of the VL arm area. In addition, a small focal cluster of labeled thalamic neurons was also seen. The labeled cluster was limited to 0.5 mm rostrocaudally and 300 mum in width. The focal distribution of labeled thalamic neurons suggests that aspects of a point to point organization may exist in the connection between VL and the motor cortex arm area.

Afferent Pathways

Afferent-efferent linkages in motor cortex for single forelimb muscles.

1. In locally anesthetized cats, extracellular recordings were made from single neurons in the lateral cruciate gyrus of cerebral cortex. These neurons responded to natural activation of stretch receptors in single, contralateral, forelimb wrist muscles, typically with phasic excitation. Low-velocity stretches, which activate primary endings of muscle spindles, excited one set of neurons at a mean latency of 11 ms; high-velocity stretches, which principally activate Golgi tendon organs and/or secondary spindle endings, excited a second set at 18 ms. The cortical neurons showing threshold responses to low-velocity stretches were found exclusively within restricted columns, 0.5-2.0 mm in diameter, which were spatially separate for each muscle. Neurons exhibiting threshold responses to high-velocity stretches were present in high density within the same columns and were also distributed, although more sparsely, outside the columns. 2. These afferent columns were located in cytoarchitectonic area 4gamma, and were shown by intracortical microstimulation to coincide with the efferent columns for contraction of the same muscle from which in input rose. Discrete afferent columns were also found for single muscles in the peridimple region of sensory cortex (area 3a), spatially separate from the columns in motor cortex. The excitation of the columns in motor cortex by these inputs from muscle was independent of that in sensory cortex. 3. The role of the cerebellum in controlling these feedback systems to motor cortex was investigated by selective cooling of interpositus and dentate nucleus, respectively. Cooling of interpositus markedly reduced transmission in the high-threshold system; cooling of dentate had a similar effect on the low-threshold system. 4. The latency, threshold, and cooling data indicated that the low-threshold system to motor cortex utilizes extracerebellar pathways including medial lemniscus and is facilitated by dentate nucleus. The high-threshold system involves a transcerebellar pathway including interpositus nucleus. Both systems transmit velocity-related information, with each showing different and complementary sensitivity and dynamic range. 5. The results are discussed with reference to the cortical load-compensation mechanism postulated by Phillips (37-38).

Animals

Magnetoelectrical stimulation of motor cortex in children with motor disturbances.

Transcranial magnetoelectrical stimulation (TMS) is now widely used as a diagnostic tool in adults. In this study we report our experiences with this technique in children with central motor disturbances. We used a Cadwell MES10 magnetoelectrical stimulator with a maximal magnetic field of 2 tesla. The stimulation procedure followed a standardized protocol, with the patients being as relaxed as possible in order to avoid contamination of parameters with different preinnervational levels. Stimulation data were compared to a data base obtained in 58 normal children. The first group of patients consisted of 20 children aged from 7 months to 16 years with hemiparesis of different etiologies. Neuroimaging data were correlated with the results of magnetoelectrical stimulation. In 13 patients a pathological pattern of TMS could be detected, and in 7 of these a corresponding lesion of the cortico-spinal tract was found in CT or MRI scans. In 7 children TMS was normal, in spite of a clear-cut lesion of the cortico-spinal tract in CT or MRI scans in 4 of them. The second group of patients consisted of 16 children with extrapyramidal disease, mostly of hereditary origin, such as DOPA-responsive dystonia or benign hereditary chorea. TMS showed a normal response pattern in this group. We discuss problems and possible pitfalls in TMS in childhood in evaluating the diagnostic value of TMS. At the moment the diagnostic usefulness of TMS in children with motor disturbances appears limited and calls for careful interpretation.

Adolescent

Receptive fields of thalamic neurons projecting to the motor cortex in the cat.

The locations and receptive fields of thalamic neurons projecting to the motor cortex were examined and the following results were obtained. (1) Neurons located at the border area between nucleus ventralis lateralis (VL) and nucleus ventralis posterolateralis (VPL) could be activated antidromically from the motor cortex. (2) These neurons received topographically organized somesthetic inputs arising from skin and deep receptors. (3) The receptive fields of neurons in the small area of the motor cortex where these thalamic neurons projected could be examined in 8 instances. In 6 instances, the cortical neurons and the thalamic projection neurons were activated by exactly the same stimuli in the periphery. (4) Removal of the sensory cortex did not significantly change the characteristics of afferent inputs from the periphery to the motor cortex. (5) It is concluded that the motor cortex receives somesthetic inputs directly from the thalamus. The functional role of these inputs was discussed in relation to the known cortical reflexes.

Animals

Effect of elaboration of a local motor-food conditioned reflex and reconditioning its effector component on the protein content in neurons of the motor cortex of rats.

Elaboration in rats of a local motor-food conditioned reflex in the form of pressing a lever by a preferred foot was accompanied by an increase of the protein content in interneurons in the absence of a change of this index in large and medium pyramidal neurons in the V layer of the motor cortex in the projection of the preferred foot in comparison with the symmetric point of the opposite hemisphere. Reconditioning of the conditioned reflex, i.e., its accomplishment by the nonpreferred foot, was accompanied by a decrease of the protein content in large pyramidal neurons of the V layer of the motor cortex in the absence of changes in the dry mass of interneurons in the projection of the nonpreferred foot in comparison with the projection of the preferred foot. Pyramidal neurons of the III layer reacted by an insignificant change in the protein content both in the case of elaborating the conditioned reflex and reconditioning of its effector component. It is suggested that the pyramidal neurons of the III layer, large pyramidal neurons and interneurons of the V layer of the motor cortex are involved in metabolic provision of elaboration of the local motor-food conditioned reflex and in the formation of new motor coordination at different stages of these processes.

Animals

Localization of multiple dopamine receptor subtype mRNAs in human and monkey motor cortex and striatum.

Dopamine plays a critical role in motor and cognitive function through actions mediated by specific receptors, multiple subtypes of which have recently been identified. The distribution of mRNAs encoding D1, D2 and D5 receptors in the motor cortex of humans and in the motor cortex and striatum of macaque monkeys was examined using in situ hybridization. In motor cortices from both primate species, hybridization to each receptor probe resulted in numerous labeled cells throughout layers II-VI. In contrast to neocortex, in monkey striatum only the D1 and D2 receptor probes showed significant hybridization. Thus, not only does primate neocortex possess a broader representation of the dopamine receptor subtype mRNAs examined in comparison with striatum, but the unexpected presence and widespread distribution of D2 and D5 receptor mRNAs in cortex suggests that, along with D1 receptors, D2 and D5 receptors play a crucial role in the dopaminergic modulation of cognition and motor behavior, and in dopamine dysfunction associated with neuropsychiatric disorders.

Aged

Synaptic proliferation in the motor cortex of adult cats after long-term thalamic stimulation.

1. One of the hypotheses for information storage in the CNS postulates the induction of structural changes in synaptic circuits. This postulate predicts that behavioral experiences produce changes in neural activity that subsequently induce synaptogenesis in the mature CNS. Available data indicate that the establishment of engrams for novel motor acts may involve alterations of synaptic interactions within the primary motor cortex. The present study examines the hypothesis that patterns of synaptic circuitry and of synaptic activation are rearranged after enhanced neural activity in pathways projecting to the motor cortex. 2. Electrodes implanted in the ventroposterolateral (VPL) nucleus of the thalamus were used for long-term stimulation (20 microA, 4 days) of afferents to the motor cortex in freely behaving, adult cats. This stimulation primarily affected corticocortical inputs from the somatosensory cortex (area 2) to area 4 gamma of the motor cortex. Electron microscopy and stereological procedures were used to compare the numerical density (Nv) of various types of synapses in layers II/III of the stimulated (experimental) motor cortex with the Nv of the corresponding synapses in the contralateral (control) hemisphere. 3. Long-term stimulation produced a significant increase (25.6%) in synaptic Nv in experimental motor cortex. This increase was due primarily to an increase in the Nv of asymmetrical synapses with dendritic spines. The numbers of symmetrical synapses, and of asymmetrical synapses with dendritic shafts, were not affected by long-term stimulation. 4. Synaptic active zones [calculated by measuring the lengths of postsynaptic densities (PSDs)] were significantly longer in experimental motor cortex. Lengthening of PSDs occurred selectively in asymmetrical synapses with dendritic shafts (28% increase). 5. The Nv of synapses having perforations in their PSDs (perforated synapses) was significantly higher in experimental hemispheres. Also increased was the incidence of synapse-associated polyribosomes, which are most commonly found at the base of dendritic spines. An increase in the number of perforated synapses and of polyribosomes are both morphological hallmarks of synaptogenesis. 6. The percentages of synapses having different curvatures (i.e., presynaptically concave, convex, or flat) were similar in experimental and in control motor cortex.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals

[Magnetic stimulation of the motor cortex in healthy persons].

In 49 healthy volunteers the motor cortex was stimulated transcranially by the intensive short magnetic field and the responses of the m. abductor digiti minimi (m. ADM) in 20 volunteers were recorded and of the m. tibialis anterior (m. TA) in 29 volunteers in relaxation and in mild contraction. In all volunteers by the electric stimulation of the ulnar nerve in the region of the wrist and the peroneal nerve in the region of the fibular capitulum there have been measured the F wave latency, distal latency of M responses and the conduction time of the peripheral motor neuron has been calculated. The difference between the latency of m. ADM and n. TA responses and the conduction time of the peripheral motor nerve represented the central motor conduction time from the motor cortex to the motor neurons of the anterior horns of the spinal cord (C8 and L4 segments). The normative values of the absolute latencies of the muscle responses and the central motor conduction time in the relaxed and contracted muscle are calculated as well as the normatives of the normal differences in latencies and the central motor conduction time between the left and right side of the same subject.

Fingers

Responses of neurones in motor cortex and in area 3A to controlled stretches of forelimb muscles in cebus monkeys.

1. The experiments were designed to investigate the effects of longitudinal muscle displacements on neurones of the motor cortex of anaesthetized Cebus monkeys and thus test the hypothesis that signals from muscle spindles may modify motor cortical output. The effects of sinusoidal stretching of the extensor digitorum communis (EDC) at frequencies varying from 6 to 300 Hz and of step and rhomboidal stretches were studied in neurones of the motor cortex. For comparison, neurones of the primary receiving area for low-threshold muscle afferents, cortical area 3a, were also included in this study. Neurones of the motor cortex were subdivided into corticospinal (PT) neurones and non-corticospinal (non-PT) neurones. 2. Threshold stretch amplitudes were clearly higher for neurones of area 4 (PT and non-PT) than for 3a neurones. However, a conspicuous fall in threshold stretch amplitude was observed for all three neurone populations when the frequency of sinusoidal stretching was increased (highest frequency: 300 Hz). A small number of non-PT and PT neurones responded to vibration amplitudes of less than 100 mum and some of these low-threshold cells of area 4 also responded to rhomboidal stretches of 8 mm/sec ramp velocity and 80 mum plateau amplitude. Increasing the stretch amplitude to twice threshold nearly doubled the output magnitude in all three cell types. Neurones of area 3a and non-PT neurones of area 4 had similar latencies, and these were significantly shorter than the latencies of PT neurones tested with trains of high frequency vibration. Dynamic response patterns were observed in all three cell types, but most frequently in 3a neurones. 3. It is concluded that, in Cebus monkeys, signals from both primary and secondary muscle spindle endings from forelimb muscles reach the motor cortex. Under the present experimental conditions, the input from the primaries to the motor cortex was effective only if these spindle receptors were driven maximally by vibratory stimuli. The particularly low probability of stretch-evoked discharges of cortico-spinal neurones in the anaesthetized preparation may be explained by a low gain in transmission from input to output cells of the motor cortex.

Animals

Postnatal development of visually evoked activity within motor cortex of cat.

1. The development of visually evoked activity within motor cortex was studied in chloralose-anesthetized kittens of 3, 4, and 5 mo of age and adult cats. 2. The slow-wave response from motor cortex of the adult cat is a triphasic negative-positive-negative wave. In 3-mo-old kittens the response is only a long-duration negative wave, while at 4 mo the positive and late negative waves emerge, and at 5 mo of age the adult form of the response is present. 3. Single neurons in the adult respond to the light during the positive component of the slow-wave response. When only the negative wave is present in kittens of 3 and 4 mo of age, neurons do not respond to the photic stimulus. At 5 mo of age the correlation between unit activity and the positive component of the response is identical to the adult. 4. The laminar analysis of the positive component of the response suggests that it originates relatively superficially in cortex and migrates down toward the deeper layers. Its correlation with unit activity indicates that the positive component represents excitatory postsynaptic potentials. 5. These results are consistent with an excitatory drive on neurons within motor cortex undergoing development change until the 5th mo of life.

Aging

Thalamic projections to the feline motor cortex studied with horseradish peroxidase.

Cells in the thalamus projecting to the distal forelimb regions of the motor cortex, area 3a, and the sensory cortex were identified with horseradish peroxidase (HRP). These cortical areas were defined by mapping evoked potentials from superficial and deep radial nerve stimulation. Following injections into the motor cortex, the labelled cells were distributed throughout a large region (greater than 2 mm wide) in the ventral portion of the ventral lateral nucleus (VL). The border area between VL and the ventral posterolateral nucleus (VPL) also contained labelled cells. The region containing labelled cells following injections into area 3a was in the rostral, dorsal VPL, and overlapped with the region containing cells labelled by injections into the motor cortex. Cells labelled by injections into the sensory cortex were near the center of VPL distinctly separated from those labelled after injections into either area 3a or the motor cortex.

Animals