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Much of our normal behavior depends on the sequential execution of multiphased movements, or the execution of multiple movements arranged in a correct temporal order. This article deals with the issue of motor selection to arrange multiple movements in an appropriate temporal order, rather than the issue of constructing spatio-temporal structures in a single action. Planning, generating, and controlling the sequential motor behavior involves multiple cortical and subcortical neural structures. Studies on human subjects and nonhuman primates, however, have revealed that the medial motor areas in the frontal cortex and the basal ganglia play particularly important roles in the temporal sequencing of multiple movements. Cellular activity observed in the supplementary and presupplementary motor areas while performing specifically designed motor tasks suggests the way in which these areas take part in constructing the time structure for the sequential execution of multiple movements.

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Recent studies have presented evidence that the prefrontal cortex plays a crucial role in every aspect of the cognitive processes necessary for behavioral planning: processing and integration of perceived or memorized information, associative learning, reward-based behavioral control, behavioral selection/decision-making and behavioral guidance. We propose that the creation of novel information is the means by which the prefrontal cortex operates to achieve executive control over behavioral planning. The prefrontal cortex is the site of operation of nodal points, where neural circuits integrate currently available or memorized information to generate the information that is necessary to perform an action. The prefrontal cortex also regulates the flow of information through multiple nodes to meet behavioral demands.

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To construct an animal model of strategy formation, we designed a maze path-finding task. First, we asked monkeys to capture a goal in the maze by moving a cursor on the screen. Cursor movement was linked to movements of each wrist. When the animals learned the association between cursor movement and wrist movement, we established a start and a goal in the maze, and asked them to find a path between them. We found that the animals took the shortest pathway, rather than approaching the goal randomly. We further found that the animals adopted a strategy of selecting a fixed intermediate point in the visually presented maze to select one of the shortest pathways, suggesting a visually based path planning. To examine their capacity to use that strategy flexibly, we transformed the task by blocking pathways in the maze, providing a problem to solve. The animals then developed a strategy of solving the problem by planning a novel shortest path from the start to the goal and rerouting the path to bypass the obstacle.

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We examined the location and spatial distribution of cingulate cortical cells projecting to the forelimb areas of the primary motor cortex (MI), supplementary motor area (SMA), and pre-supplementary motor area (pre-SMA) using a multiple retrograde labeling technique in the monkeys (Macaca fuscata). The forelimb areas of the MI, SMA and pre-SMA were physiologically identified, based on the findings of intracortical microstimulation (ICMS) and single cell recording. Three different tracers, diamidino yellow (DY), fast blue (FB), and wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP), were injected into each of the three motor areas in the same monkey. Retrogradely labeled cells in the cingulate cortex were plotted with an automated plotting system. Cells projecting to the forelimb area of the MI were distributed in the two separate regions situated rostrocaudally in the dorsal and ventral banks of the cingulate sulcus, namely the rostral cingulate motor area (CMAr) and caudal cingulate motor area (CMAc). These two regions corresponded to the forelimb areas identified by the ICMS in the same animal. The distribution of projection cells to the SMA overlapped extensively with that of projection cells to the MI. Although the MI received relatively sparse inputs from the CMAr than from the CMAc, the SMA received inputs from the CMAr and its adjacent areas as much as from the CMAc. The projection cells to the pre-SMA were distributed in the anterior portion of the cingulate cortex, including the anterior part of the CMAr and in a small part of the cingulate gyrus. These findings indicate that the MI and SMA share a considerable common information from the cingulate cortex, including the CMAr and CMAc, whereas the pre-SMA receives a different set of information from the anterior part of the cingulate cortex.

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To plan an action, we must first select an object to act on and the body part (or parts) to use to accomplish our intention. To plan the motor task of reaching, we specify both the target to reach for and the arm to use. In the process of planning and preparing a motor task, information about the motor target and the arm to use must be integrated before a motor program can be formulated to generate the appropriate limb movement. One of the structures in the brain that is probably involved in integrating these two sets of information is the premotor area in the cerebral cortex of primates. The lateral sector of the dorsal premotor cortex is known to receive both visual and somatosensory input, and we show here that neurons in this area gather information about both the target and the body part, while subsequent activity specifies the planned action.

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To study how neurons in the medial motor areas participate in performing sequential multiple movements that are individually separated in time, we analyzed neuronal activity in the supplementary (SMA) and presupplementary (pre-SMA) motor areas. Monkeys were trained to perform three different movements separated by waiting times, in four or six different orders. Initially each series of movements was learned during five trials guided by visual signals that indicated the correct movements. The monkeys subsequently executed the three movements in the memorized order without the visual signals. Three types of neuronal activity were of particular interest; these appeared to be crucially involved in sequencing the multiple motor tasks in different orders. First, we found activity changes that were selective for a particular sequence of the three movements that the monkeys were prepared to perform. The sequence-selective activity ceased when the monkeys initiated the first movement. Second, we found interval-selective activity that appeared in the interval between one particular movement and the next. Third, we found neuronal activity representing the rank order of three movements arranged chronologically; that is, the activity differed selectively in the process of preparing the first, second, or third movements in individual trials. The interval-selective activity was more prevalent in the SMA, whereas the rank-order selective activity was more frequently recorded in the pre-SMA. These results suggest how neurons in the SMA and pre-SMA are involved in sequencing multiple movements over time.

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This study examined neuronal activity in the prefrontal cortex (PF) involved in the process of motor selection in accordance with two behavioral rules. We trained two monkeys to select a target based on the integration of memorized and current sensory information. Initially, a sample cue (triangle or circle) appeared at one of three locations (top, left, or right) for 1 s. After a 3-s delay, one of two types of choice cue appeared. The first type asked the monkeys to reach for a target by matching the location (location-matching task). The second type asked the monkeys to reach for a target by matching the shape (shape-matching task). The choice cue for location matching consisted of either three circles or three triangles, and the choice cue for shape matching consisted of a circle and a triangle. When the color of the choice cue changed from red to green 1.5 s later (GO signal), the monkeys touched the correct object to obtain a reward. We found cue-, delay-, choice-, and movement-related neuronal activity in the lateral prefrontal cortex. During the sample cue presentation and delay periods, we found selective neuronal activity for the location or shape of the sample cue. Shape-selective neurons were located more anteriorly in the ventral bank of the principal sulcus and inferior convexity area, whereas location-selective neurons were more posteriorly. After the choice cue appeared, we found three main types of neuronal activity in the critical period when the subject selected the future target: 1) activity reflecting past sensory information (the location or shape of the sample cue presented 3 s earlier), 2) activity selective for the configuration of the current choice cue, and 3) activity reflecting the properties (location or shape) of the future target. During the motor-response period, we found neuronal activity selective for the location or shape of the reaching target. When muscimol was microinjected into the ventral bank of principal sulcus and inferior convexity area, the performance of both tasks was impaired. Furthermore, we found that the wealth of neuronal activity in the PF that seemed to play a role in motor selection was rarely seen in the primary motor cortex.

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To investigate functional differences between the rostral and caudal parts of the dorsal premotor cortex (PMd), we first examined the effects of intracortical microstimulation (ICMS) while monkeys were performing oculomotor and limb motor tasks or while they were at rest. We found that saccades were evoked from the rostral part (PMdr) whereas ICMS in the caudal part (PMdc) predominantly produced forelimb or body movements. Subsequently, we examined neuronal activity in relation to the performance of visually cued and memorized saccades while monkeys reached an arm toward a visual target. We found that roughly equal numbers of PMdr neurons were active during performance of the oculomotor and limb motor tasks. In contrast, the majority of PMdc neurons were related preferentially to arm movements and not to saccades. In the subsequent analysis, we found that the oculomotor effects evoked in the PMdr differ from the effects evoked in either the frontal eye field (FEF) or supplementary eye field (SEF). These findings suggest that the PMdr is involved in oculomotor as well as limb motor behavior. However, the oculomotor involvement of the PMdr seems to have a functional aspect different from that operating in the FEF and SEF.

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This article describes the benefits of exercise, including why exercise improves blood pressure control, how it may reduce the risk for cancer, and increase mental health, and how it may improve bone mineral density. The article also explores the gender-independent and gender-specific benefits of exercise for women.

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We compared the effects of intracortical microstimulation (ICMS) of the lateral wall of the intraparietal sulcus (LIP) with those of ICMS of the frontal eye field (FEF) on monkeys performing oculomotor tasks. When ICMS was applied during a task that involved fixation, contraversive saccades evoked in the LIP and FEF appeared similar. When ICMS was applied to the FEF at the onset of voluntary saccades, the evoked saccades collided with the ongoing voluntary saccade so that the trajectory of voluntary saccade was compensated by the stimulus. Thus the resultant saccade was redirected and came close to the endpoint of saccades evoked from the fixation point before the start of voluntary saccade. In contrast, when ICMS was applied to the LIP at the onset of voluntary saccades, the resultant saccade followed a trajectory that was different from that evoked from the FEF. In that case, the colliding saccades were redirected toward an endpoint that was close to the endpoint of saccades evoked when animals were already fixating at the target of the voluntary saccade. This finding suggests that the colliding saccade was directed toward an endpoint calculated with reference to the target of the voluntary saccade. We hypothesize that, shortly before initiation of voluntary saccades, a dynamic process occurs in the LIP so that the reference point for calculating the saccade target shifts from the fixation point to the target of a voluntary saccade. Such predictive updating of reference points seems useful for immediate reprogramming of upcoming saccades that can occur in rapid succession.

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We investigated the interrelationship between the supplementary motor area (SMA) thalamocortical projection neurons and the pallidothalamic and cerebellothalamic territories in the monkey (Macaca fuscata) using a combination of three tracers in a triple labeling paradigm. Thalamic labeling was analyzed following injections of the anterograde tracers, biotinylated dextran amine (BDA) into the internal segment of the globus pallidus (GPi) and wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP) into the contralateral cerebellar interpositus and dentate nuclei. In addition, the retrograde tracer cholera toxin subunit B (CTB) was injected into the physiologically identified hand/arm representation of SMA. The tissue was processed sequentially using different chromogens in order to visualize all three tracers in a single section. We found that the SMA thalamocortical neurons occupied a wide band extending from the ventral anterior nucleus pars principalis (VApc) through the ventral lateral nucleus pars oralis (VLo) and the ventral lateral nucleus pars medialis (VLm) and into to the ventral lateral nucleus pars caudalis (VLc) including a portion of ventral posterior lateral nucleus pars oralis (VPLo) and nucleus X. The heaviest CTB labeling was found in VLo with dense plexuses of BDA labeled pallidothalamic fibers and swellings often observed superimposed upon retrogradely labeled CTB cells. In addition, dense foci of cerebellothalamic WGA-HRP anterograde label were observed coinciding with the occasional retrogradely CTB labeled neurons in VLc and transitional zones between VApc, VLo and VPLo. Our light microscopic results suggest that the SMA receives thalamic inputs with afferents largely derived from GPi and minor inputs originating from the cerebellum.

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Both supplementary and presupplementary motor areas are crucial for the temporal organization of multiple movements. J. Neurophysiol. 80: 3247-3260, 1998. To study the involvement of the supplementary (SMA) and presupplementary (pre-SMA) motor areas in performing sequential multiple movements that are individually separated in time, we injected muscimol, a gamma-aminobutyric acid agonist, bilaterally into the part of each area that represents the forelimb. Two monkeys were trained to perform three different movements, separated by a waiting time, in four or six different orders. First, each series of movements was learned during five trials guided by visual signals that indicated the correct movements. The monkeys subsequently executed the three movements in the memorized order, without the visual signals. After the injection of muscimol (3 microliter, 5 micrograms/microliters in 10 min) into either the SMA or pre-SMA bilaterally, the animals started making errors in performing the sequence of movements correctly from memory. However, when guided with a visual signal, they could select and perform the three movements correctly. The impaired memory-based sequencing of movements worsened progressively with time until the animals could not perform the task. Yet they still could associate the visual signals with the different movements at that stage. In control experiments on two separate monkeys, we found that injections of the same amount of muscimol into either the SMA or pre-SMA did not cause problems with nonsequential reaching movement regardless of whether it was visually triggered or self-initiated. These results support the view that both the SMA and pre-SMA are crucially involved in sequencing multiple movements over time.

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Task-dependent selectivity of movement-related neuronal activity in the primate prefrontal cortex. J. Neurophysiol. 80: 3392-3397, 1998. We studied movement-related neuronal activity in the dorsolateral prefrontal cortex from the perspective of a general role for the prefrontal cortex in controlling motor behavior to achieve a specific goal according to the requirements of a given task. Monkeys were trained to perform two delayed motor tasks. The first task involved reaching for a target that matched the shape of a cue. The second task involved reaching for a target that matched the location of the cue. A majority (54%) of 175 movement-related prefrontal neurons exhibited preference for either the target shape or the type of task requirements. Sixty-four neurons (36%) were selectively active while reaching for a circle or a triangle. On the other hand, the activity of 59 neurons (34%) depended on whether the task required matching the shape or the location. These properties, characterizing the movement-related neuronal activity in the prefrontal cortex, rarely were found in the arm area of the primary motor cortex. Only 1 of 130 movement-related neurons (0.8%) showed task selectivity, and none showed target-shape selectivity.

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Most natural actions are chosen voluntarily from many possible choices. An action is often chosen based on the reward that it is expected to produce. What kind of cellular activity in which area of the cerebral cortex is involved in selecting an action according to the expected reward value? Results of an analysis in monkeys of cellular activity during the performance of reward-based motor selection and the effects of chemical inactivation are presented. We suggest that cells in the rostral cingulate motor area, one of the higher order motor areas in the cortex, play a part in processing the reward information for motor selection.

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The involvement of N-methyl-D-aspartate (NMDA) and non-NMDA glutamate receptors in mediating the excitatory responses of neurons in the primary motor cortex (MI) to electrical stimulation of the supplementary motor area (SMA) and the somatosensory cortex (SI) was examined in monkeys performing a trained motor task. During the task, a total of 109 MI neurons were identified and classified as movement related (91), motor set related (7), or mixed (11). Subsequently, the influence of receptor antagonists on the stimulus-evoked and task-related activities of these neurons was examined. The selective NMDA antagonist D-2-amino-5-phosphonovaleric acid (APV) and the selective non-NMDA antagonist 6-cyano-7-nitroquinoxaline-2,3-dione (CNQX) were applied iontophoretically through multibarreled micropipettes. One barrel was used for extracellular unit recording. The excitatory response evoked by SI stimulation was suppressed by CNQX in the vast majority (83%) of the motor task related neurons, and only 10% were suppressed by APV. On the other hand, the response evoked by SMA stimulation was suppressed by APV in 56% of the neurons and by CNQX in 54%. APV and CNQX had parallel effects on the stimulus-evoked responses and the task-related neuronal activity. These results indicate that NMDA and non-NMDA receptors are both involved in mediating the excitatory responses of MI neurons to input from the SMA and SI. On the other hand, the data suggest a greater contribution of non-NMDA receptors in response to SI input and greater involvement of NMDA receptors in mediating the response to SMA input, especially among set-related MI neurons.

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The purpose of this study was to identify the brain regions activated in relation to oculomotor sequence learning. Nine healthy subjects participated in the study, which consisted of three positron emission tomography scans. In the initial learning task, subjects were instructed to track a sequence of seven successive positions of visual targets and to memorize the order of the targets as well as their spatial locations. In the saccade task, subjects were instructed to track visual targets presented at random locations. In the control task, subjects were instructed to gaze at a fixation point. Fields showing significant regional cerebral blood flow change were determined from task-minus-control subtraction images. We determined that fields in the pre-supplementary motor area (pre-SMA), the intraparietal cortex, and the prefrontal cortex were activated not only in the learning-minus-control images but also in the learning-minus-saccade images. Although prefrontal and parietal activations were bilateral, pre-SMA activation was confined to the left hemisphere. The results indicate that these fields function as a part of the neural network involved in the learning of sequential saccadic eye movements.

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We explored the ventral part of the premotor cortex (PMV) with intracortical microstimulation (ICMS) while monkeys performed a visual fixation task, to see whether the PMV is involved in oculomotor control. ICMS evoked saccades from a small-restricted region in the PMV, without evoking movements in the limbs, neck, or body. We found the saccade-evoking site in the PMV in a total of three hemispheres in two monkeys. Quantitative analysis of the effects of eye position on saccades evoked by microstimulation of the PMV characterized the evoked saccades as goal directed. The nature of the saccades evoked in the PMV contrasted with the fixed vector nature of saccades evoked by ICMS of the frontal eye field. We also found that neurons in this restricted area of the PMV were active while the animals were performing a saccade task that required them to make saccades toward targets without arm movements. These data provide evidence for the presence of an oculomotor-specific subregion within the PMV. This subregion and the surrounding skeletomotor-representing regions of the PMV seem to coordinate oculomotor and skeletomotor control in performing goal-directed motor tasks.

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We trained two monkeys to perform a fixation task. Intracortical microstimulation (ICMS) was applied to the monkey frontal eye field (FEF) while monkeys were fixating on one of five fixation LEDs. The ICMS was applied in two different manners. Under the single stimulation condition, ICMS was delivered to either right or left FEF. Under the paired stimulation condition, bilateral FEF were successively stimulated with an interval of 30-250 ms. The single stimulation elicited contraversive saccades. As reported previously, these saccades were not much affected by initial eye positions, maintaining the same vector. In contrast, the paired stimulation elicited double-step saccades. The first of the paired stimulation elicited constant vector saccades, but the second of the paired stimulation evoked saccades whose vector varied greatly depending on the eye position at the start of individual saccades. The second saccades, starting from various initial positions, were directed to the endpoint of saccades that were elicited from the same FEF site under the single stimulation condition. Endpoints of second saccades varied little despite variations of intervals of the stimulation pairs, ranging from 60 to 150 ms. On the basis of these observations, we propose a novel view that the FEF is involved in directing saccades to an internally referenced visual target.

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We recorded 200 neurons from the ventral part of the premotor cortex (PMv) and 110 neurons from the primary motor cortex (MI) of a monkey performing a visually cued arm-reaching task with a delay. We compared neuronal activity in the premovement period while the monkey reached the target with the eyes fixating on either a left or right fixation target. Our data demonstrate that about half of the movement-related activity in the PMv was modulated by the direction of gaze. In contrast, a vast majority of the activity of MI neurons and about half of PMv neurons were not influenced by the direction of gaze. We further analyzed the movement-related activity during the reaching movement to targets at the top, bottom, left, and right of each fixation point. The magnitude of activity of neurons showing the gaze-direction selectivity was primarily determined by the position of the reaching target relative to the eye-fixation target, and not by the position of the target relative to the animal's body. These data suggest that a part of the coordinate transformation of the motor command signals concerning the direction of reaching from the retinotopic to body-centered frame of reference may occur at the level of premotor cortex but not in MI.

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