Functional Properties of Neurons in the Primate Tongue Primary Motor Cortex During Swallowing

1997 ◽  
Vol 78 (3) ◽  
pp. 1516-1530 ◽  
Author(s):  
Ruth E. Martin ◽  
Gregory M. Murray ◽  
Pentti Kemppainen ◽  
Yuji Masuda ◽  
Barry J. Sessle
Keyword(s):  
Motor Cortex ◽  
Functional Properties ◽  
Primary Motor Cortex ◽  
Motor Behavior ◽  
Critical Role ◽  
Electromyographic Activity ◽  
Related Activity ◽  
Tongue Protrusion ◽  
Significant Difference ◽  
Tongue Movements

Martin, Ruth E., Gregory M. Murray, Pentti Kemppainen, Yuji Masuda, and Barry J. Sessle. Functional properties of neurons in the primate tongue primary motor cortex during swallowing. J. Neurophysiol. 78: 1516–1530, 1997. Recent studies conducted in our laboratory have suggested that the tongue primary motor cortex (i.e., tongue-MI) plays a critical role in the control of voluntary tongue movements in the primate. However, the possible involvement of tongue-MI in semiautomatic tongue movements, such as those in swallowing, remains unkown. Therefore the present study was undertakein in attempts to address whether tongue-MI plays a role in the semiautomatic tongue movements produced during swallowing. Extracellular single neuron recordings were obtained from tongue-MI, defined by intracortical microstimulation (ICMS), in two awake monkeys as they performed three types of swallowing (swallowing of a juice reward after successful tongue task performance, nontask-related swallowing of a liquid bolus, and nontask-related swallowing of a solid bolus) as well as a trained tongue-protrusion task. Electromyographic activity was recorded simultaneously from various orofacial and laryngeal muscles. In addition, the afferent input to each tongue-MI neuron and ICMS-evoked motor output characteristics at each neuronal recording site were determined. Neurons were considered to show swallow and/or tongue-protrusion task-related activity if a statistically significant difference in firing rate was seen in association with these behaviors compared with that observed during a control pretrial period. Of a total of 80 neurons recorded along 40 microelectrode penetrations in the ICMS-defined tongue-MI, 69% showed significant alterations of activity in relation to the swallowing of a juice reward, whereas 66% exhibited significant modulations of firing in association with performance of the trained tongue-protrusion task. Moreover, 48% showed significant alterations of firing in relation to both swallowing and the tongue-protrusion task. These findings suggest that the region of cortex involved in swallowing includes MI and that tongue-MI may play a role in the regulation of semiautomatic tongue movement, in addition to trained motor behavior. Swallow-related tongue-MI neurons exhibited a variety of swallow-related activity patterns and were distributed throughout the ICMS-defined tongue-MI at sites where ICMS evoked a variety of types of tongue movements. These findings are consistent with the view that multiple efferent zones for the production of tongue movements are activated in swallowing. Many swallow-related tongue-MI neurons had an orofacial mechanoreceptive field, particularly on the tongue dorsum, supporting the view that afferent inputs may be involved in the regulation of the swallowing synergy.

Functional properties of single neurons in the face primary motor cortex of the primate. II. Relations with trained orofacial motor behavior

1992 ◽  
Vol 67 (3) ◽  
pp. 759-774 ◽  
Author(s):  
G. M. Murray ◽  
B. J. Sessle
Keyword(s):  
Motor Cortex ◽  
Cortical Neurons ◽  
Primary Motor Cortex ◽  
Motor Behavior ◽  
Afferent Input ◽  
Single Neurons ◽  
Tongue Protrusion ◽  
Input And Output ◽  
The Face ◽  
Tongue Movements

1. The previous paper has described in detail the input and output features of single neurons located at sites within primate face motor cortex from which intracortical microstimulation (ICMS, less than or equal to 20 microA) evoked tongue movements at the lowest threshold ("tongue-MI" sites); for comparative purposes, we also reported on the input and output features of a smaller number of neurons recorded at sites from which ICMS could evoke jaw movements ("jaw-MI" sites), facial movements ("face-MI" sites), or, at a few sites, tongue movements and, at the same threshold intensity, either a jaw movement or a facial movement. 2. Our findings of an extensive and diverse representation of sites within face motor cortex of monkeys for the generation of elemental components of tongue movement, and the relatively few sites from which jaw-closing movements could be evoked, were consistent with our recent observations that reversible, cooling-induced inactivation of the face motor cortex severely impaired the performance by monkeys of a tongue-protrusion task but had only relatively minor effects on the performance of a biting task. In an attempt to establish a neuronal correlate for these different behavioral relations, the present study has documented the task-related activities of those single neurons that were characterized in the previous paper in terms of afferent input and ICMS-defined output features. 3. Each task required the development and maintenance by each monkey of a fixed force level for a minimum period of time to obtain a fruit-juice reward. During one or both of these tasks, we characterized the activities of 231 single face motor cortical neurons that were located at the above-mentioned ICMS-defined sites. Neurons were said to be related to a particular task if they showed statistically significant differences in firing rates during the task in comparison with a control pretrial period (PTP). 4. In tongue-MI, there was a significantly higher proportion of neurons (63% of 156 neurons tested) that were related to the tongue-protrusion task than to the biting task (15% of 65). However, in jaw-MI the proportion of neurons that were biting task-related (63% of 19) was significantly higher than the proportion related to the tongue-protrusion task (11% of 9); the proportion of biting task-related neurons at ICMS-defined jaw-closing sites was also higher than that at jaw-opening sites.(ABSTRACT TRUNCATED AT 400 WORDS)

scholarly journals Unilateral nasal obstruction affects motor representation development within the face primary motor cortex in growing rats

2017 ◽  
Vol 122 (6) ◽  
pp. 1494-1503 ◽  
Author(s):  
Yasunori Abe ◽  
Chiho Kato ◽  
Karin Harumi Uchima Koecklin ◽  
Hidemasa Okihara ◽  
Takayoshi Ishida ◽  
...  
Keyword(s):  
Motor Cortex ◽  
Nasal Obstruction ◽  
Primary Motor Cortex ◽  
Respiratory Pattern ◽  
Arterial Oxygen ◽  
Electromyographic Activity ◽  
Motor Representation ◽  
Significant Difference ◽  
The Face ◽  
Experimental Group

Postnatal growth is influenced by genetic and environmental factors. Nasal obstruction during growth alters the electromyographic activity of orofacial muscles. The facial primary motor area represents muscles of the tongue and jaw, which are essential in regulating orofacial motor functions, including chewing and jaw opening. This study aimed to evaluate the effect of chronic unilateral nasal obstruction during growth on the motor representations within the face primary motor cortex (M1). Seventy-two 6-day-old male Wistar rats were randomly divided into control ( n = 36) and experimental ( n = 36) groups. Rats in the experimental group underwent unilateral nasal obstruction after cauterization of the external nostril at 8 days of age. Intracortical microstimulation (ICMS) mapping was performed when the rats were 5, 7, 9, and 11 wk old in control and experimental groups ( n = 9 per group per time point). Repeated-measures multivariate ANOVA was used for intergroup and intragroup statistical comparisons. In the control and experimental groups, the total number of positive ICMS sites for the genioglossus and anterior digastric muscles was significantly higher at 5, 7, and 9 wk, but there was no significant difference between 9 and 11 wk of age. Moreover, the total number of positive ICMS sites was significantly smaller in the experimental group than in the control at each age. It is possible that nasal obstruction induced the initial changes in orofacial motor behavior in response to the altered respiratory pattern, which eventually contributed to face-M1 neuroplasticity. NEW & NOTEWORTHY Unilateral nasal obstruction in rats during growth periods induced changes in arterial oxygen saturation (SpO2) and altered development of the motor representation within the face primary cortex. Unilateral nasal obstruction occurring during growth periods may greatly affect not only respiratory function but also craniofacial function in rats. Nasal obstruction should be treated as soon as possible to avoid adverse effects on normal growth, development, and physiological functions.

Functional properties of single neurons in the face primary motor cortex of the primate. I. Input and output features of tongue motor cortex

1992 ◽  
Vol 67 (3) ◽  
pp. 747-758 ◽  
Author(s):  
G. M. Murray ◽  
B. J. Sessle
Keyword(s):  
Motor Cortex ◽  
Primary Motor Cortex ◽  
Afferent Input ◽  
Single Neurons ◽  
Tongue Movement ◽  
Tongue Protrusion ◽  
The Face ◽  
Tongue Movements ◽  
Low Threshold ◽  
Organizational Features

1. We have recently demonstrated that reversible, cooling-induced inactivation of the face motor cortex results in a severe impairment in the ability of monkeys (Macaca fascicularis) to perform a tongue-protrusion task but produces only relatively minor effects on the performance of a biting task by the same monkeys. To establish a neuronal correlate for these different behavioral relations, the present study has detailed the afferent input and intracortical microstimulation (ICMS)-defined output features of a population of face motor cortical neurons, and in a subsequent study we have documented the activities of the same population of neurons during the performance of the tongue-protrusion and biting tasks. 2. Of the 231 single neurons recorded within the face motor cortex, 163 were located at sites from which ICMS (less than or equal to 20 microA) could evoke tongue movements (i.e., "tongue-MI" sites) at the lowest threshold for eliciting orofacial movements. The remainder were located at sites from which ICMS evoked jaw movements ("jaw-MI" sites), face movements ("face-MI" sites), or at a few sites, tongue movements and, at the same threshold intensity, either a jaw movement or a facial movement. 3. We confirmed the general organizational features of the face motor cortex that have been defined in previous studies, but we documented in detail the organizational features for tongue-MI. Thus we found that tongue movements were well represented, whereas jaw-closing movements were poorly represented; the representations for face, jaw, and tongue movements were overlapped; the same ICMS-evoked tongue movement could be multiply represented within tongue-MI; tongue-MI was characterized by a prominent input from superficial mechanosensory afferents, whereas there was little evidence for deep input; a close spatial match was found between ICMS-defined motor output and somatosensory afferent input for tongue-MI. 4. A variety of tongue movements could be evoked by ICMS at tongue-MI sites and were categorized into protrusion, retrusion, laterally directed, and other types of tongue movement. Low-threshold (i.e., less than or equal to 5 microA) ICMS-defined tongue-MI sites, which were considered to represent "efferent zones" projecting relatively directly to motoneurons, were reconstructed three dimensionally to provide insights into the spatial organization of tongue-MI. Examples of each of the four low-threshold efferent-zone categories were usually found throughout the ICMS-defined tongue-MI without any apparent preferential distribution. Furthermore, different low-threshold efferent-zone categories had close spatial relationships to each other in cortex.(ABSTRACT TRUNCATED AT 400 WORDS)

Reorganization of Remote Cortical Regions After Ischemic Brain Injury: A Potential Substrate for Stroke Recovery

2003 ◽  
Vol 89 (6) ◽  
pp. 3205-3214 ◽  
Author(s):  
S. B. Frost ◽  
S. Barbay ◽  
K. M. Friel ◽  
E. J. Plautz ◽  
R. J. Nudo
Keyword(s):  
Motor Cortex ◽  
Primary Motor Cortex ◽  
Motor Behavior ◽  
Premotor Cortex ◽  
Critical Role ◽  
Retrieval Task ◽  
Gradual Improvement ◽  
Intact Tissue ◽  
Ventral Premotor Cortex ◽  
Hand Area

Although recent neurological research has shed light on the brain's mechanisms of self-repair after stroke, the role that intact tissue plays in recovery is still obscure. To explore these mechanisms further, we used microelectrode stimulation techniques to examine functional remodeling in cerebral cortex after an ischemic infarct in the hand representation of primary motor cortex in five adult squirrel monkeys. Hand preference and the motor skill of both hands were assessed periodically on a pellet retrieval task for 3 mo postinfarct. Initial postinfarct motor impairment of the contralateral hand was evident in each animal, followed by a gradual improvement in performance over 1–3 mo. Intracortical microstimulation mapping at 12 wk after infarct revealed substantial enlargements of the hand representation in a remote cortical area, the ventral premotor cortex. Increases ranged from 7.2 to 53.8% relative to the preinfarct ventral premotor hand area, with a mean increase of 36.0 ± 20.8%. This enlargement was proportional to the amount of hand representation destroyed in primary motor cortex. That is, greater sparing of the M1 hand area resulted in less expansion of the ventral premotor cortex hand area. These results suggest that neurophysiologic reorganization of remote cortical areas occurs in response to cortical injury and that the greater the damage to reciprocal intracortical pathways, the greater the plasticity in intact areas. Reorganization in intact tissue may provide a neural substrate for adaptive motor behavior and play a critical role in postinjury recovery of function.

scholarly journals Protective role of mirtazapine in adult female Mecp2+/− mice and patients with Rett syndrome

2020 ◽  
Vol 12 (1) ◽  
Author(s):  
Javier Flores Gutiérrez ◽  
Claudio De Felice ◽  
Giulia Natali ◽  
Silvia Leoncini ◽  
Cinzia Signorini ◽  
...  
Keyword(s):  
Motor Cortex ◽  
Motor Learning ◽  
Disease Progression ◽  
Rett Syndrome ◽  
Adult Female ◽  
Primary Motor Cortex ◽  
Barrel Cortex ◽  
Motor Behavior ◽  
Clinical Severity ◽  
Long Term Treatment

Abstract Background Rett syndrome (RTT), an X-linked neurodevelopmental rare disease mainly caused by MECP2-gene mutations, is a prototypic intellectual disability disorder. Reversibility of RTT-like phenotypes in an adult mouse model lacking the Mecp2-gene has given hope of treating the disease at any age. However, adult RTT patients still urge for new treatments. Given the relationship between RTT and monoamine deficiency, we investigated mirtazapine (MTZ), a noradrenergic and specific-serotonergic antidepressant, as a potential treatment. Methods Adult heterozygous-Mecp2 (HET) female mice (6-months old) were treated for 30 days with 10 mg/kg MTZ and assessed for general health, motor skills, motor learning, and anxiety. Motor cortex, somatosensory cortex, and amygdala were analyzed for parvalbumin expression. Eighty RTT adult female patients harboring a pathogenic MECP2 mutation were randomly assigned to treatment to MTZ for insomnia and mood disorders (mean age = 23.1 ± 7.5 years, range = 16–47 years; mean MTZ-treatment duration = 1.64 ± 1.0 years, range = 0.08–5.0 years). Rett clinical severity scale (RCSS) and motor behavior assessment scale (MBAS) were retrospectively analyzed. Results In HET mice, MTZ preserved motor learning from deterioration and normalized parvalbumin levels in the primary motor cortex. Moreover, MTZ rescued the aberrant open-arm preference behavior observed in HET mice in the elevated plus-maze (EPM) and normalized parvalbumin expression in the barrel cortex. Since whisker clipping also abolished the EPM-related phenotype, we propose it is due to sensory hypersensitivity. In patients, MTZ slowed disease progression or induced significant improvements for 10/16 MBAS-items of the M1 social behavior area: 4/7 items of the M2 oro-facial/respiratory area and 8/14 items of the M3 motor/physical signs area. Conclusions This study provides the first evidence that long-term treatment of adult female heterozygous Mecp2tm1.1Bird mice and adult Rett patients with the antidepressant mirtazapine is well tolerated and that it protects from disease progression and improves motor, sensory, and behavioral symptoms.

Laterality of Movement-Related Activity Reflects Transformation of Coordinates in Ventral Premotor Cortex and Primary Motor Cortex of Monkeys

2007 ◽  
Vol 98 (4) ◽  
pp. 2008-2021 ◽  
Author(s):  
Kiyoshi Kurata
Keyword(s):  
Motor Cortex ◽  
Neuronal Activity ◽  
Primary Motor Cortex ◽  
Premotor Cortex ◽  
Visuomotor Transformation ◽  
Related Activity ◽  
Reaching Task ◽  
Ventral Premotor Cortex ◽  
Cortical Motor ◽  
Target Locations

The ventral premotor cortex (PMv) and the primary motor cortex (MI) of monkeys participate in various sensorimotor integrations, such as the transformation of coordinates from visual to motor space, because the areas contain movement-related neuronal activity reflecting either visual or motor space. In addition to relationship to visual and motor space, laterality of the activity could indicate stages in the visuomotor transformation. Thus we examined laterality and relationship to visual and motor space of movement-related neuronal activity in the PMv and MI of monkeys performing a fast-reaching task with the left or right arm, toward targets with visual and motor coordinates that had been dissociated by shift prisms. We determined laterality of each activity quantitatively and classified it into four types: activity that consistently depended on target locations in either head-centered visual coordinates (V-type) or motor coordinates (M-type) and those that had either differential or nondifferential activity for both coordinates (B- and N-types). A majority of M-type neurons in the areas had preferences for reaching movements with the arm contralateral to the hemisphere where neuronal activity was recorded. In contrast, most of the V-type neurons were recorded in the PMv and exhibited less laterality than the M-type. The B- and N-types were recorded in the PMv and MI and exhibited intermediate properties between the V- and M-types when laterality and correlations to visual and motor space of them were jointly examined. These results suggest that the cortical motor areas contribute to the transformation of coordinates to generate final motor commands.

scholarly journals Corticocortical Connections of the Rostral Forelimb Area in Rats: A Quantitative Tract-Tracing Study

2020 ◽  
Author(s):  
Edward T Urban ◽  
Mariko Nishibe ◽  
Scott Barbay ◽  
David J Guggenmos ◽  
Randolph J Nudo
Keyword(s):  
Motor Cortex ◽  
Primary Motor Cortex ◽  
Motor Behavior ◽  
Premotor Cortex ◽  
Relative Strength ◽  
Cortical Region ◽  
Corticocortical Connections ◽  
Cortical Connections ◽  
Efferent Projections ◽  
Biotinylated Dextran

AbstractThe rostral forelimb area (RFA) in the rat is considered to be a premotor cortical region based primarily on its efferent projections to the primary motor cortex. The purpose of the present study was to identify corticocortical connections of RFA, and to describe the relative strength of connections with other cortical areas. This will allow us to better understand the broader cortical network in which RFA participates, and thus, determine its function in motor behavior. In the present study, the RFA of adult male Long-Evans rats (n=6) was identified using intracortical microstimulation techniques and injected with the tract tracer, biotinylated dextran amine (BDA). In post-mortem tissue, location of BDA-labeled terminal boutons and neuronal somata were plotted and superimposed on cortical field boundaries. The results demonstrated that the RFA has dense to moderate reciprocal connections with primary motor cortex, the frontal cortex medial and lateral to RFA, primary somatosensory cortex (S1), and lateral somatosensory areas. Importantly, S1 connections were dense to moderate in dysgranular zones, but sparse to negligible in granular zones. Cortical connections of RFA in rat are strikingly similar to cortical connections of the ventral premotor cortex in non-human primates, suggesting that these areas share similar functions.

Neuronal Activity Patterns in Primate Primary Motor Cortex Related to Trained or Semiautomatic Jaw and Tongue Movements

2002 ◽  
Vol 87 (5) ◽  
pp. 2531-2541 ◽  
Author(s):  
Dongyuan Yao ◽  
Kensuke Yamamura ◽  
Noriyuki Narita ◽  
Ruth E. Martin ◽  
Gregory M. Murray ◽  
...  
Keyword(s):  
Motor Cortex ◽  
Primary Motor Cortex ◽  
Activity Patterns ◽  
Rhythmic Activity ◽  
Firing Frequency ◽  
Orofacial Movements ◽  
Jaw Opening ◽  
The Face ◽  
Tongue Movements ◽  
Opening Phase

The present study was undertaken to determine the firing patterns and the mechanoreceptive field (RF) properties of neurons within the face primary motor cortex (face-MI) in relation to chewing and other orofacial movements in the awake monkey. Of a total of 107 face-MI neurons recorded, 73 of 74 tested had activity related to chewing and 47 of 66 neurons tested showed activity related to a trained tongue task. Of the 73 chewing-related neurons, 52 (71.2%) showed clear rhythmic activity during rhythmic chewing. A total of 32 (43.8%) also showed significant alterations in activity in relation to the swallowing of a solid food (apple) bolus. Many of the chewing-related neurons (81.8% of 55 tested) had an orofacial RF, which for most was on the tongue dorsum. Tongue protrusion was evoked by intracortical microstimulation (ICMS) at most (63.6%) of the recording sites where neurons fired during the rhythmic jaw-opening phase, whereas tongue retraction was evoked by ICMS at most (66.7%) sites at which the neurons firing during the rhythmic jaw-closing phase were recorded. Of the 47 task-related neurons, 21 of 22 (95.5%) examined also showed chewing-related activity and 29 (61.7%) demonstrated significant alteration in activity in relation to the swallowing of a juice reward. There were no significant differences in the peak firing frequency among neuronal activities related to chewing, swallowing, or the task. These findings provide further evidence that face-MI may play an important role not only in trained orofacial movements but also in chewing as well as swallowing, including the control of tongue and jaw movements that occur during the masticatory sequence.

Abstract 2806: Does Motor Cortex Activity Predict Motor Recovery? An fMRI Study of Subacute Lacunar Stroke

Stroke ◽  
2012 ◽  
Vol 43 (suppl_1) ◽  
Author(s):  
Assia Jaillard ◽  
Chantal Delon martin ◽  
Leeanne Carey ◽  
Laurent Lalamalle ◽  
Marc J Hommel ◽  
...  
Keyword(s):  
Motor Cortex ◽  
Primary Motor Cortex ◽  
Motor Recovery ◽  
Small Sample ◽  
Stroke Onset ◽  
Stroke Recovery ◽  
Finger Tapping ◽  
Lacunar Stroke ◽  
Related Activity ◽  
Recovery Score

Background: While primary motor cortex (M1) has been demonstrated to be crucial for motor recovery in a recent meta-analysis including fMRI and TMS studies, other functional neuroimaging studies have found that activity in a broader sensorimotor cortical network correlate with motor recovery. The heterogeneity of stroke lesions and the small sample size characterizing many studies could account for these discrepancies. Hypothesis : The strength of task-related activity in primary motor cortex predicts motor recovery in a clinically homogenous population of acute lacunar stroke patients. Methods: We used fMRI to investigate the neural mechanisms of stroke recovery. We studied 18 stroke patient (4 females, 14 males) after their first single lacunar stroke (7 right , 11 left hemisphere). The lesions caused pure hemiparesia one week after stroke onset (mean 7.2 days; range 2 -15). Lesions were limited to the deep territory of the anterior choroid artery, involving the corticospinal tract at the level of the internal capsule or the corona radiata ( Figure 1 ). Patients were matched to 18 healthy controls for age and sex. Motor impairment was assessed using the NIH Stroke Scale (NIHSS), the Fugl-Meyer Scale (FMS), Finger Tapping Score (FTS), Purdue Pegboard and simple reaction times 7 days and 6 months after stroke. At 6 months, a global motor recovery score was computed using the FMS and the FTS to assess motor recovery. Functional MRI scans were obtained using a self-paced finger tapping (FT) task implemented as a block design alternating right FT, left FT and rest. Data were processed using SPM8. In the first level analysis “FT minus fixation” contrasts were computed for the impaired hand. At the second level, multiple regression was used to assess the effect of the motor recovery score on the FT-related motor activity (threshold p<0.05 FWE; extent threshold k=5). Age and FT rate recorded during the experiment were included as covariates in the second level model. Results: As a group, the patients showed good recovery at 6 months. Both patients and controls exhibited a typical pattern of FT task-related activity. Activity in primary motor cortex predicted motor recovery at 6 months, after adjustment for age and FT rate. MNI coordinates = [-34,-14,48] See Figure 1 . Conclusions: Primary motor cortex activity, measured soon after stroke onset, predicts motor recovery assessed at 6 months post-stroke. fMRI measurements made in the early phase of stroke recovery could be useful to derive prognostic biomarkers in both clinical practice and clinical trials investigating novel treatments, such as stem cell administration.

scholarly journals Layer specificity of inputs from supplementary motor area and dorsal premotor cortex to primary motor cortex in macaque monkeys

2019 ◽  
Vol 9 (1) ◽  
Author(s):  
Taihei Ninomiya ◽  
Ken-ichi Inoue ◽  
Eiji Hoshi ◽  
Masahiko Takada
Keyword(s):  
Motor Cortex ◽  
Primary Motor Cortex ◽  
Supplementary Motor Area ◽  
Motor Behavior ◽  
Premotor Cortex ◽  
Distribution Patterns ◽  
Motor Area ◽  
Tracer Injection ◽  
Macaque Monkeys ◽  
Motor Information

AbstractThe primate frontal lobe processes diverse motor information in parallel through multiple motor-related areas. For example, the supplementary motor area (SMA) is mainly involved in internally-triggered movements, whereas the premotor cortex (PM) is highly responsible for externally-guided movements. The primary motor cortex (M1) deals with both aspects of movements to execute a single motor behavior. To elucidate how the cortical motor system is structured to process a variety of information, the laminar distribution patterns of signals were examined between SMA and M1, or PM and M1 in macaque monkeys by using dual anterograde tract-tracing. Dense terminal labeling was observed in layers 1 and upper 2/3 of M1 after one tracer injection into SMA, another tracer injection into the dorsal division of PM resulted in prominent labeling in the deeper portion of layer 2/3. Weaker labeling was also visible in layer 5 in both cases. On the other hand, inputs from M1 terminated in both the superficial and the deep layers of SMA and PM. The present data indicate that distinct types of motor information are arranged in M1 in a layer-specific fashion to be orchestrated through a microcircuit within M1.

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