Arm Representation Research Articles

Deafferentation of neighbouring motor cortex areas does not further enhance saturated practice-dependent plasticity in healthy adults

Objective To assess effects of deafferentation of the arm representation of primary motor cortex (M1) on practice-dependent plasticity in healthy adults. Methods Twelve healthy, right-handed adults (18–48 years, median 20.2 years) performed two consecutive experiments (exp. 1 and exp. 2). Exp. 1 consisted of a motor practice (MP) of repeated ballistic flexion movements of the left thumb. This was followed by exp. 2 consisting of selective anaesthesia of the upper brachial plexus (SPA) to disinhibit the training M1 and a second period of the same MP. Peak acceleration of the trained thumb movement and the motor evoked potential (MEP) amplitude in the flexor pollicis brevis muscle elicited by single-pulse transcranial magnetic stimulation of the training M1 were studied before and after exp. 1 and after exp. 2. Results After exp. 1 all subjects demonstrated an increase of peak acceleration (baseline: 19.23 ± 3.81 m s −2; after exp. 1: 43.28 ± 17.63 m s −2, p = 0.008) and MEP amplitude (from 0.46 ± 0.23 mV to 1.26 ± 0.77 mV, p = 0.03). There was no additional increase of these measures after exp. 2 (44.37 ± 19.56 m s −2, p = 0.78, 1.69 ± 1.21 mV ( p = 0.07)). Conclusions Training of ballistic thumb movements leads to behavioural improvement as well as to an increased excitability of the corresponding M1 representation. These effects do not increase further during deafferentation of the training M1. In contrast to stroke patients [Muellbacher W, Richards C, Ziemann U, Wittenberg G, Weltz D, Boroojerdi B, et al. Improving hand function in chronic stroke. Arch Neurol 2002;59:1278–82], practice-dependent plasticity in healthy subjects cannot be enhanced by deafferentation of neighbouring motor cortex areas. Significance Healthy subjects, in contrast to patients with central motor lesions, are capable of saturating practice-dependent plasticity to a level that cannot be further enhanced by experimental manipulation.

Exploiting redundancy for flexible behavior: Unsupervised learning in a modular sensorimotor control architecture.

Autonomously developing organisms face several challenges when learning reaching movements. First, motor control is learned unsupervised or self-supervised. Second, knowledge of sensorimotor contingencies is acquired in contexts in which action consequences unfold in time. Third, motor redundancies must be resolved. To solve all 3 of these problems, the authors propose a sensorimotor, unsupervised, redundancy-resolving control architecture (SURE_REACH), based on the ideomotor principle. Given a 3-degrees-of-freedom arm in a 2-dimensional environment, SURE_REACH encodes 2 spatial arm representations with neural population codes: a hand end-point coordinate space and an angular arm posture space. A posture memory solves the inverse kinematics problem by associating hand end-point neurons with neurons in posture space. An inverse sensorimotor model associates posture neurons with each other action-dependently. Together, population encoding, redundant posture memory, and the inverse sensorimotor model enable SURE_REACH to learn and represent sensorimotor grounded distance measures and to use dynamic programming to reach goals efficiently. The architecture not only solves the redundancy problem but also increases goal reaching flexibility, accounting for additional task constraints or realizing obstacle avoidance. While the spatial population codes resemble neurophysiological structures, the simulations confirm the flexibility and plausibility of the model by mimicking previously published data in arm-reaching tasks.

Localization of arm representation in the cerebral peduncle of the non‐human primate

Motor deficit severity and the potential for recovery in patients with brain injury depend on the integrity of descending corticofugal projections. Clinical assessment of these conditions following subtotal brain trauma requires a comprehensive understanding of the anatomical structures involved in the lesion as well as those structures that are spared. To assist in this endeavor, we investigated motor fiber organization in the crus cerebri of the cerebral peduncle (ccCP) in the rhesus monkey. Fibers originating from the arm representations of the primary (M1), supplementary (M2), rostral cingulate (M3), caudal cingulate (M4), dorsolateral pre- (LPMCd) and ventrolateral pre- (LPMCv) motor cortices were studied. The projections from the frontal and cingulate motor cortices formed descending longitudinal bundles that occupied the medial three-fifths of the ccCP at superior and middle levels. Although considerable overlap characterized these corticofugal projections, a general topography was discernable. Fibers from M1 and M4 occupied the central subsector of the ccCP, and fibers from M3 resided medially. The main distribution of LPMCd, LPMCv, and M2 fibers occupied the centromedial region and overlapped extensively. Progressing inferiorly, all fiber bundles in the central and centromedial sectors gradually extended medially, and overlap increased. A common location of fiber passage occurred at the midbrain-pontine isthmus where all of the fiber bundles overlapped. Our findings indicate that the widespread distribution of corticofugal motor projections may account for the favorable levels of motor recovery that accompany subtotal midbrain injury. At superior and mid-levels of the ccCP anteromedial lesions may disrupt projections from M3, whereas anterolateral lesions may disrupt projections from M1 and M4. Fibers from M2, LPMCv, and LPMCd may be compromised to some degree in both situations. The compact and commixed nature of motor fiber organization at inferior levels and the midbrain-pontine isthmus suggests a vulnerable region of passage for comprehensive disruption of frontal and cingulate corticofugal projection fibers.

Interhemispheric Inhibition in Distal and Proximal Arm Representations in the Primary Motor Cortex

Interhemispheric inhibitory interactions (IHI) operate between homologous distal hand representations in primary motor cortex (M1). It is not known whether proximal arm representations exhibit comparable effects on their homologous counterparts. We studied IHI in different arm representations, targeting triceps brachii (TB, n = 13), first dorsal interosseous (FDI, n = 13), and biceps brachii (BB, n = 7) muscles in healthy volunteers. Transcranial magnetic stimulation test stimuli (TS) were delivered to M1 contralateral to the target muscle preceded 10 ms by a conditioning stimulus (CS) to the opposite M1 at 110-150% resting motor threshold (RMT). IHI was calculated as the ratio between motor-evoked potential (MEP) amplitudes in conditioned relative to unconditioned trials. Mean RMTs were 38.9, 46.9, and 46.0% of stimulator output in FDI, TB, and BB muscles, respectively. IHI was 0.45 +/- 0.41 (FDI), 0.78 +/- 0.38 (TB), and 0.52 +/- 0.32 (BB, P < 0.01) when test MEP amplitudes were matched and 0.28 +/- 0.17 (FDI) and 0.85 +/- 0.31 (TB, P < 0.05) when TS intensities expressed as percentage RMT were matched. Significant IHI (P < 0.05) was identified with minimal CS intensities (expressed as percentage stimulator output) in the 30 s for FDI, 60 s for TB, and 40 s for BB. Additionally, a CS of roughly 120% RMT suppressed the test MEP but not a test H-reflex in BB, suggesting IHI observed in BB is likely mediated by a supraspinal mechanism. We conclude that IHI differs between different arm muscle representations, comparable between BB and FDI but lesser for TB. This finding suggests the amount of IHI between different arm representations does not strictly follow a proximal-to-distal gradient, but may be related to the role of each muscle in functional movement synergies.

Amygdala interconnections with the cingulate motor cortex in the rhesus monkey

Amygdala interconnections with the cingulate motor cortices were investigated in the rhesus monkey. Using multiple tracing approaches, we found a robust projection from the lateral basal nucleus of the amygdala to Layers II, IIIa, and V of the rostral cingulate motor cortex (M3). A smaller source of amygdala input arose from the accessory basal, cortical, and lateral nuclei, which targeted only the rostral region of M3. We also found a light projection from the lateral basal nucleus to the same layers of the caudal cingulate motor cortex (M4). Experiments examining this projection to cingulate somatotopy using combined neural tracing strategies and stereology to estimate the total number of terminal-like immunoreactive particles demonstrated that the amygdala projection terminates heavily in the face representation of M3 and moderately in its arm representation. Fewer terminal profiles were found in the leg representation of M3 and the face, arm, and leg representations of M4. Anterograde tracers placed directly into M3 and M4 revealed the amygdala connection to be reciprocal and documented corticofugal projections to the facial nucleus, surrounding pontine reticular formation, and spinal cord. Clinically, such pathways would be in a position to contribute to mediating movements in the face, neck, and upper extremity accompanying medial temporal lobe seizures that have historically characterized this syndrome. Alterations within or disruption of the amygdalo-cingulate projection to the rostral part of M3 may also have an adverse effect on facial expression in patients presenting with neurological or neuropsychiatric abnormalities of medial temporal lobe involvement. Finally, the prominent amygdala projection to the face region of M3 may significantly influence the outcome of higher-order facial expressions associated with social communication and emotional constructs such as fear, anger, happiness, and sadness.

On the need to revise the arm structure in stylized anthropomorphic phantoms in lateral photon irradiation geometry

Distributions of radiation absorbed dose within human anatomy have been estimated through Monte Carlo radiation transport techniques implemented for two different classes of computational anthropomorphic phantoms: (1) mathematical equation-based stylized phantoms and (2) tomographic image-based voxel phantoms. Voxel phantoms constructed from tomographic images of real human anatomy have been actively developed since the late 1980s to overcome the anatomical approximations necessary with stylized phantoms, which themselves have been utilized since the mid 1960s. However, revisions of stylized phantoms have also been pursued in parallel to the development of voxel phantoms since voxel phantoms (1) are initially restricted to the individual-specific anatomy of the person originally imaged, (2) must be restructured on an organ-by-organ basis to conform to reference individual anatomy and (3) cannot easily represent very fine anatomical structures and tissue layers that are thinner than the voxel dimensions of the overall phantom. Although efforts have been made to improve the anatomic realism of stylized phantoms, most of these efforts have been limited to attempts to alter internal organ structures. Aside from the internal organs, the exterior shapes, and especially the arm structures, of stylized phantoms are also far from realistic descriptions of human anatomy, and may cause dosimetry errors in the calculation of organ-absorbed doses for external irradiation scenarios. The present study was intended to highlight the need to revise the existing arm structure within stylized phantoms by comparing organ doses of stylized adult phantoms with those from three adult voxel phantoms in the lateral photon irradiation geometry. The representative stylized phantom, the adult phantom of the Oak Ridge National Laboratory (ORNL) series and two adult male voxel phantoms, KTMAN-2 and VOXTISS8, were employed for Monte Carlo dose calculation, and data from another voxel phantom, VIP-Man, were obtained from literature sources. The absorbed doses for lungs, oesophagus, liver and kidneys that could be affected by arm structures in the lateral irradiation geometry were obtained for both classes of phantoms in lateral monoenergetic photon irradiation geometries. As expected, those organs in the ORNL phantoms received apparently higher absorbed doses than those in the voxel phantoms. The overestimation is mainly attributed to the relatively poor representation of the arm structure in the ORNL phantom in which the arm bones are embedded within the regions describing the phantom's torso. The results of this study suggest that the overestimation of organ doses, due to unrealistic arm representation, should be taken into account when stylized phantoms are employed for equivalent or effective dose estimates, especially in the case of an irradiation scenario with dominating lateral exposure. For such a reason, the stylized phantom arm structure definition should be revised in order to obtain more realistic evaluations.

Poster 93: Interhemispheric Inhibition in Distal and Proximal Arm Representations in the Primary Motor Cortex

Poster 93: Interhemispheric Inhibition in Distal and Proximal Arm Representations in the Primary Motor Cortex

Visualizing Chromosomes as Transcriptome Correlation Maps: Evidence of Chromosomal Domains Containing Co-expressed Genes—A Study of 130 Invasive Ductal Breast Carcinomas

Completion of the working draft of the human genome has made it possible to analyze the expression of genes according to their position on the chromosomes. Here, we used a transcriptome data analysis approach involving for each gene the calculation of the correlation between its expression profile and those of its neighbors. We used the U133 Affymetrix transcriptome data set for a series of 130 invasive ductal breast carcinomas to construct chromosomal maps of gene expression correlation (transcriptome correlation map). This highlighted nonrandom clusters of genes along the genome with correlated expression in tumors. Some of the gene clusters identified by this method probably arose because of genetic alterations, as most of the chromosomes with the highest percentage of correlated genes (1q, 8p, 8q, 16p, 16q, 17q, and 20q) were also the most frequent sites of genomic alterations in breast cancer. Our analysis showed that several known breast tumor amplicons (at 8p11-p12, 11q13, and 17q12) are located within clusters of genes with correlated expression. Using hierarchical clustering on samples and a Treeview representation of whole chromosome arms, we observed a higher-order organization of correlated genes, sometimes involving very large chromosomal domains that could extend to a whole chromosome arm. Transcription correlation maps are a new way of visualizing transcriptome data. They will help to identify new genes involved in tumor progression and new mechanisms of gene regulation in tumors.

Cortical Representation of Bimanual Movements

It is well established that the discharge of neurons in primate motor cortex is tuned to the movement direction of the contralateral arm. Interestingly, it has been found that these neurons exhibit a directional tuning to the ipsilateral arm as well and that the preferred directions to both arms tend to be similar. A recent study showed that motor cortex cells are also directionally selective to bimanual movements, but the relationship between the bimanual and unimanual representations remains unclear. To address this issue, we analyzed the responses of motor cortical neurons recorded from two macaque monkeys during unimanual and bimanual reaching movements. We decomposed the bimanual movement representation into contralateral and ipsilateral directionally tuned components. Our major finding is that the movement of the contralateral arm modifies the tuning of the cells to the ipsilateral arm such that: (1) the offset and modulation depth of the tuning are suppressed; and (2) the preferred directions are randomly shifted. Both these effects eliminate the correlation between the contralateral and ipsilateral representations during bimanual movements. We suggest that the modification of the ipsilateral arm representation is caused by the recruitment of local inhibition that conveys callosal inputs during bimanual movements. This hypothesis is supported by the analysis of a model of two motor cortical networks, coupled with sparse random interhemispheric projections that reproduce the main features observed in the data. Finally, we show that the modification of the ipsilateral arm representation reduces the interference between the movements of both arms.

Stimulation-Induced Within-Representation and Across-Representation Plasticity in Human Motor Cortex

The human motor cortex contains a dynamic and distributed network of motor representations. Formation, maintenance, and modification of these representations is an activity-driven process. Repeated stimulation of one representation results in increased motor output from this representation, a process referred to as "within-representation plasticity." We showed previously that within-representation plasticity of the upper arm representation occurs when repetitive transcranial magnetic stimulation (rTMS) is delivered to that representation during transient ischemic nerve block (INB) of the contralateral hand. INB reduces inhibition in the motor cortex and thus lowers the threshold for stimulation-induced plasticity. Here we studied the effects of rTMS delivered to nearby body part representations on the motor output from the upper arm representation. Six healthy subjects underwent INB-alone (control), or INB plus 30 min of focal 0.1 Hz rTMS of either the face, hand, arm, leg, or overlap arm/hand representations in motor cortex. INB-alone and rTMS of the leg representation resulted in only a short-lasting (< 20 min) increase in motor output from the arm representation, as measured by motor evoked potentials in the biceps. rTMS of arm and arm/hand representations induced a prolonged (> 60 min) within-representation increase. In contrast, rTMS of face or hand representations canceled the short-lasting increase and even led to a long-lasting decrease of motor output from the arm representation. Therefore, rTMS of the experimentally disinhibited motor cortex induces within-representation increase, and across-representation decrease of motor cortical output. This bidirectional plasticity might be used for purposeful modulation of human cortical function.

Stimulation-induced within-representation and across-representation plasticity in human motor cortex.

The human motor cortex contains a dynamic and distributed network of motor representations. Formation, maintenance, and modification of these representations is an activity-driven process. Repeated stimulation of one representation results in increased motor output from this representation, a process referred to as "within-representation plasticity." We showed previously that within-representation plasticity of the upper arm representation occurs when repetitive transcranial magnetic stimulation (rTMS) is delivered to that representation during transient ischemic nerve block (INB) of the contralateral hand. INB reduces inhibition in the motor cortex and thus lowers the threshold for stimulation-induced plasticity. Here we studied the effects of rTMS delivered to nearby body part representations on the motor output from the upper arm representation. Six healthy subjects underwent INB-alone (control), or INB plus 30 min of focal 0.1 Hz rTMS of either the face, hand, arm, leg, or overlap arm/hand representations in motor cortex. INB-alone and rTMS of the leg representation resulted in only a short-lasting (< 20 min) increase in motor output from the arm representation, as measured by motor evoked potentials in the biceps. rTMS of arm and arm/hand representations induced a prolonged (> 60 min) within-representation increase. In contrast, rTMS of face or hand representations canceled the short-lasting increase and even led to a long-lasting decrease of motor output from the arm representation. Therefore, rTMS of the experimentally disinhibited motor cortex induces within-representation increase, and across-representation decrease of motor cortical output. This bidirectional plasticity might be used for purposeful modulation of human cortical function.

Distribution of eye- and arm-movement-related neuronal activity in the SEF and in the SMA and Pre-SMA of monkeys.

We analyzed neuronal activity in the supplementary eye field (SEF), supplementary motor area (SMA), and presupplementary motor area (pre-SMA) during the performance of three motor tasks: capturing a visual target with a saccade, reaching one arm to a target while gazing at a visual fixation point, or capturing a target with a saccade and arm-reach together. Our data demonstrated that each area was involved in controlling the arm and eye movements in a different manner. Saccade-related neurons were found mainly in the SEF. In contrast, arm-movement-related neurons were found primarily in the SMA and pre-SMA. In addition, we found that the activity of both arm-movement- and saccade-related neurons differed depending on the presence or absence of an accompanying saccade or arm movement. Such context dependency was found in all three areas. We also discovered that activity preceding eye or arm movement alone, and eye and arm movement combined, appeared more often in the pre-SMA and SEF, suggesting their involvement in effector-independent aspects of motor behavior. Subsequent analysis revealed that the laterality of arm representation differed in the three areas: it was predominantly contralateral in the SMA but largely bilateral in the pre-SMA and SEF.

Localization of arm representation in the corona radiata and internal capsule in the non-human primate.

Localization of the corticofugal projection in the corona radiata (CR) and internal capsule (IC) can assist in evaluating a patient's residual motor capacity following subtotal brain damage and predicting their potential for functional restitution. To advance our understanding of the organization of the corticofugal projection in this critical brain region, we studied the trajectories of the projection arising from six different cortical arm representations in rhesus monkeys. They included the arm representation of the primary (M1), ventral lateral pre- (LPMCv), dorsolateral pre- (LPMCd), supplementary (M2), rostral cingulate (M3) and caudal cingulate (M4) motor cortices. In the CR, each pathway was segregated as medial motor area fibres arched over the caudate and lateral motor area fibres arched over the putamen. In the IC, the individual corticofugal pathways were found to be widespread, topographically organized and partially overlapping. At superior levels of the IC, the corticofugal projection from the arm representation of M3 coursed through the middle and posterior portion of the anterior limb (ICa). The projection from M2 passed through the posterior portion of the ICa and the genu (ICg). The projection from LPMCv travelled through the genu and anterior portion of the posterior limb (ICp). The projection from LPMCd occupied the anterior portion of the ICp. The projection from M4 descended through the mid-portion of the ICp. Fibres from M1 also travelled in the ICp, positioned immediately posterior to the M4 projection. As each fibre system progressed inferiorly within the IC, all fibres shifted posteriorly to occupy the ICp. Within the ICp, the projections from M3, M2, LPMCv, LPMCd, M4 and M1 maintained their anterior to posterior orientation, respectively. M2, LPMCd and LPMCv fibres overlapped extensively, as did fibres from M4 and M1. Our data suggest that CR and superior capsular lesions may correlate with more favourable levels of functional recovery due to the widespread nature of arm representation. In contrast, the extensive overlap and comparatively condensed organization of arm representation at inferior capsular levels suggest that lesions seated inferiorly are likely to correlate with poorer levels of recovery of upper limb movement. Based on the relative density of corticospinal neurones associated with the motor areas studied, our findings also suggest that motor deficit severity is likely to increase as a lesion occupies progressively more posterior regions of the IC.

Remodelling of sensorimotor maps in paraplegia: a functional magnetic resonance imaging study after a surgical nerve transfer

The adult mammalian brain has the capacity of reorganising its neural connections in response to lesions/modifications of the peripheral and central nervous system. We show in vivo, using functional magnetic resonance imaging (fMRI), that in paraplegics the lower-limb sensorimotor cortex is invaded by the arm representation. This functional reshaping appears to be reversible. Indeed, surgical transfer of the ulnar nerve to the ipsilateral quadriceps and hip muscles allowed their contraction in a paraplegic patient. During fMRI, these voluntary movements activated the hip and thigh representation in sensorimotor cortex. We suggest that the functional recovery of the lower-limb functional maps might have been driven by the restored somatosensory inputs from the reactivated periphery. The voluntary movements of the lower-limbs are regained through the ‘re-awakening’ of the corresponding sensorimotor cortex.

FMRI Evaluation of Somatotopic Representation in Human Primary Motor Cortex

We used fMRI to map foot, elbow, fist, thumb, index finger, and lip movements in 30 healthy subjects. For each movement type confidence intervals of representational sites in the primary motor cortex (M1) were evaluated. In order to improve the precision of their anatomical localization and to optimize the mapping of cortical activation sites, we used both the assessment of locations in the conventional 3D system and a 2D projection method. In addition to the computation of activation maxima of activation clusters within the precentral gyrus, centers of gravity were determined. Both methods showed a high overlap of their representational confidence intervals. The 2D-projection method revealed statistically significant distinct intralimb locations, e.g., elbow versus index finger movements and index finger versus thumb movements. Increased degree of complexity of finger movements resulted in a spread of the somatotopic location toward the arm representation. The 2D-projection method-based fMRI evaluation of limb movements showed high precision and was able to reveal differences in intralimb movement comparisons. fMRI activation revealed a clear somatotopic order of movement representation in M1 and also reflected different degrees of complexity of movement.

Growth of new brainstem connections in adult monkeys with massive sensory loss.

Somatotopic maps in the cortex and the thalamus of adult monkeys and humans reorganize in response to altered inputs. After loss of the sensory afferents from the forelimb in monkeys because of transection of the dorsal columns of the spinal cord, therapeutic amputation of an arm or transection of the dorsal roots of the peripheral nerves, the deprived portions of the hand and arm representations in primary somatosensory cortex (area 3b), become responsive to inputs from the face and any remaining afferents from the arm. Cortical and subcortical mechanisms that underlie this reorganization are uncertain and appear to be manifold. Here we show that the face afferents from the trigeminal nucleus of the brainstem sprout and grow into the cuneate nucleus in adult monkeys after lesions of the dorsal columns of the spinal cord or therapeutic amputation of an arm. This growth may underlie the large-scale expansion of the face representation into the hand region of somatosensory cortex that follows such deafferentations.

Tremor-frequency (3–6 Hz) activity in the sensorimotor arm representation of the internal segment of the globus pallidus in patients with Parkinson's disease

Neurons in the internal segment of the globus pallidus (GPi) oscillate at approximately the frequency of parkinsonian tremor. However, the correlation of that activity with tremor has not previously been studied. We now describe the relationship between single neuron activity in the arm sensorimotor portion of GPi and upper extremity tremor in patients with Parkinson's disease. There was a significant concentration of power in the tremor-frequency range (3–6 Hz) for 11/44 GPi neurons. However, pallidal tremor-frequency activity correlated significantly with electromyogram (EMG) activity during tremor for only a single GPi neuron. These data are most consistent with the hypothesis that the output of neurons in GPi is transformed in thalamus by a non-linear mechanism, before transmission via the cortex to the spinal motorneurons that drive movement.

Subcortical contributions to massive cortical reorganizations.

Subcortical contributions to massive cortical reorganizations.

Cortical networks for visual reaching: physiological and anatomical organization of frontal and parietal lobe arm regions.

The functional and structural properties of the dorsolateral frontal lobe and posterior parietal proximal arm representations were studied in macaque monkeys. Physiological mapping of primary motor (MI), dorsal premotor (PMd), and posterior parietal (area 5) cortices was performed in behaving monkeys trained in an instructed-delay reaching task. The parietofrontal corticocortical connectivities of these same areas were subsequently examined anatomically by means of retrograde tracing techniques. Signal-, set-, movement-, and position-related directional neuronal activities were distributed nonuniformly within the task-related areas in both frontal and parietal cortices. Within the frontal lobe, moving caudally from PMd to the MI, the activity that signals for the visuo-spatial events leading to target localization decreased, while the activity more directly linked to movement generation increased. Physiological recordings in the superior parietal lobule revealed a gradient-like distribution of functional properties similar to that observed in the frontal lobe. Signal- and set-related activities were encountered more frequently in the intermediate and ventral part of the medial bank of the intraparietal sulcus (IPS), in area MIP. Movement-and position-related activities were distributed more uniformly within the superior parietal lobule (SPL), in both dorsal area 5 and in MIP. Frontal and parietal regions sharing similar functional properties were preferentially connected through their association pathways. As a result of this study, area MIP, and possibly areas MDP and 7m as well, emerge as the parietal nodes by which visual information may be relayed to the frontal lobe arm region. These parietal and frontal areas, along with their association connections, represent a potential cortical network for visual reaching. The architecture of this network is ideal for coding reaching as the result of a combination between visual and somatic information.

Topographic organization of corticospinal projections from the frontal lobe: motor areas on the medial surface of the hemisphere

We examined the topographic organization of corticospinal neurons in the four premotor areas on the medial wall of the hemisphere of macaques. These motor areas include the supplementary motor area (SMA) and three areas buried within the cingulate sulcus: the caudal cingulate motor area on the dorsal bank (CMAd), the caudal cingulate motor area on the ventral bank (CMAv), and the rostral cingulate motor area (CMAr). In one set of animals, we injected one fluorescent tracer into lower cervical segments of the spinal cord and another fluorescent tracer into lower lumbosacral segments to define the topographic organization of arm and leg representation within each premotor area. Similarly, in another set of animals, we injected different tracers into upper cervical and lower cervical segments to provide an indication of the topographic organization of proximal and distal arm representation within the arm representation of each premotor area. We found that all four of the premotor areas on the medial wall project to cervical and lumbosacral segments of the spinal cord. Three of these areas (SMA, CMAd, and CMAv) are like the primary motor cortex in having distinct arm and leg representations. The arm representation in each of the four motor areas on the medial wall contains separate regions that project densely to upper or to lower cervical segments. This observation suggests that each motor area contains distinct proximal and distal representations of the arm. Surprisingly, the size of the distal representation is comparable to or larger than the size of the proximal representation in each motor area. Thus, contrary to some previous hypotheses, the anatomical substrate exists for the premotor areas on the medial wall to be involved in the control of distal, as well as proximal arm movements. Our results provide a new map for guiding the exploration of the motor functions of the medial wall of the hemisphere. Furthermore, the observations of the present study support our suggestion that each of the premotor areas may be an important source of descending commands for the generation and control of movement.