Visuospatial properties of ventral premotor cortex

In macaque ventral premotor cortex, we recorded the activity of neurons that responded to both visual and tactile stimuli. For these bimodal cells, the visual receptive field extended from the tactile receptive field into the adjacent space. Their tactile receptive fields were organized topographically, with the arms represented medially, the face represented in the middle, and the inside of the mouth represented laterally. For many neurons, both the visual and tactile responses were directionally selective, although many neurons also responded to stationary stimuli. In the awake monkeys, for 70% of bimodal neurons with a tactile response on the arm, the visual receptive field moved when the arm was moved. In contrast, for 0% the visual receptive field moved when the eye or head moved. Thus the visual receptive fields of most "arm + visual" cells were anchored to the arm, not to the eye or head. In the anesthetized monkey, the effect of arm position was similar. For 95% of bimodal neurons with a tactile response on the face, the visual receptive field moved as the head was rotated. In contrast, for 15% the visual receptive field moved with the eye and for 0% it moved with the arm. Thus the visual receptive fields of most "face + visual" cells were anchored to the head, not to the eye or arm. To construct a visual receptive field anchored to the arm, it is necessary to integrate the position of the arm, head, and eye. For arm + visual cells, the spontaneous activity, the magnitude of the visual response, and sometimes both were modulated by the position of the arm (37%), the head (75%), and the eye (58%). In contrast, to construct a visual receptive field that is anchored to the head, it is necessary to use the position of the eye, but not of the head or the arm. For face + visual cells, the spontaneous activity and/or response magnitude was modulated by the position of the eyes (88%), but not of the head or the arm (0%). Visual receptive fields anchored to the arm can encode stimulus location in "arm-centered" coordinates, and would be useful for guiding arm movements. Visual receptive fields anchored to the head can likewise encode stimuli in "head-centered" coordinates, useful for guiding head movements. Sixty-three percent of face + visual neurons responded during voluntary movements of the head. We suggest that "body-part-centered" coordinates provide a general solution to a problem of sensory-motor integration: sensory stimuli are located in a coordinate system anchored to a particular body part.     

Three-dimensional organization of otolith-ocular reflexes in rhesus monkeys. I. Linear acceleration responses during off-vertical axis rotation

1. The dynamic properties of otolith-ocular reflexes elicited by sinusoidal linear acceleration along the three cardinal head axes were studied during off-vertical axis rotations in rhesus monkeys. As the head rotates in space at constant velocity about an off-vertical axis, otolith-ocular reflexes are elicited in response to the sinusoidally varying linear acceleration (gravity) components along the interaural, nasooccipital, or vertical head axis. Because the frequency of these sinusoidal stimuli is proportional to the velocity of rotation, rotation at low and moderately fast speeds allows the study of the mid-and low-frequency dynamics of these otolith-ocular reflexes. 2. Animals were rotated in complete darkness in the yaw, pitch, and roll planes at velocities ranging between 7.4 and 184 degrees/s. Accordingly, otolith-ocular reflexes (manifested as sinusoidal modulations in eye position and/or slow-phase eye velocity) were quantitatively studied for stimulus frequencies ranging between 0.02 and 0.51 Hz. During yaw and roll rotation, torsional, vertical, and horizontal slow-phase eye velocity was sinusoidally modulated as a function of head position. The amplitudes of these responses were symmetric for rotations in opposite directions. In contrast, mainly vertical slow-phase eye velocity was modulated during pitch rotation. This modulation was asymmetric for rotations in opposite direction. 3. Each of these response components in a given rotation plane could be associated with an otolith-ocular response vector whose sensitivity, temporal phase, and spatial orientation were estimated on the basis of the amplitude and phase of sinusoidal modulations during both directions of rotation. Based on this analysis, which was performed either for slow-phase eye velocity alone or for total eye excursion (including both slow and fast eye movements), two distinct response patterns were observed: 1) response vectors with pronounced dynamics and spatial/temporal properties that could be characterized as the low-frequency range of "translational" otolith-ocular reflexes; and 2) response vectors associated with an eye position modulation in phase with head position ("tilt" otolith-ocular reflexes). 4. The responses associated with two otolith-ocular vectors with pronounced dynamics consisted of horizontal eye movements evoked as a function of gravity along the interaural axis and vertical eye movements elicited as a function of gravity along the vertical head axis. Both responses were characterized by a slow-phase eye velocity sensitivity that increased three- to five-fold and large phase changes of approximately 100-180 degrees between 0.02 and 0.51 Hz. These dynamic properties could suggest nontraditional temporal processing in utriculoocular and sacculoocular pathways, possibly involving spatiotemporal otolith-ocular interactions. 5. The two otolith-ocular vectors associated with eye position responses in phase with head position (tilt otolith-ocular reflexes) consisted of torsional eye movements in response to gravity along the interaural axis, and vertical eye movements in response to gravity along the nasooccipital head axis. These otolith-ocular responses did not result from an otolithic effect on slow eye movements alone. Particularly at high frequencies (i.e., high speed rotations), saccades were responsible for most of the modulation of torsional and vertical eye position, which was relatively large (on average +/- 8-10 degrees/g) and remained independent of frequency. Such reflex dynamics can be simulated by a direct coupling of primary otolith afferent inputs to the oculomotor plant. (ABSTRACT TRUNCATED)     

Generalisability of sensory gating during passive movement of the legs

Movement-related gating of somatosensory evoked potentials in the upper limb is restricted mainly to nerve stimulation supplying the moved limb segment. In the lower limb, this principle may not be followed. Tibial nerve (stimulation at the knee) somatosensory evoked potentials (SEPs) and soleus H reflexes exhibit quite similar patterns of modulation during movement. We hypothesised that movement-related gating of initial SEPs in the leg would be generalised from ipsilateral to contralateral leg movement and that such sensory gating would not be generalised to modalities with no functional relevance to the movement. Somatosensory, visual, and auditory evoked potentials (SEPs, VEPs, and AEPs) were recorded from scalp electrodes during unilateral passive movement. Short-latency tibial nerve SEPs, representing the first cortical components, and soleus H reflexes in both the moved leg and the stationary leg were attenuated compared to non-movement controls (p<0.05). Neither VEPs nor middle latency AEPs were modulated (p>0.05). We conclude that sensory gating occurs during contralateral movement. This gating is absent in other sensory modalities with no apparent functional relationship to the imposed movement.     

Eye position effects on the neuronal activity of dorsal premotor cortex in the macaque monkey

Visual inputs to the brain are mapped in a retinocentric reference frame, but the motor system plans movements in a body-centered frame. This basic observation implies that the brain must transform target coordinates from one reference frame to another. Physiological studies revealed that the posterior parietal cortex may contribute a large part of such a transformation, but the question remains as to whether the premotor areas receive visual information, from the parietal cortex, readily coded in body-centered coordinates. To answer this question, we studied dorsal premotor cortex (PMd) neurons in two monkeys while they performed a conditional visuomotor task and maintained fixation at different gaze angles. Visual stimuli were presented on a video monitor, and the monkeys made limb movements on a panel of three touch pads located at the bottom of the monitor. A trial begins when the monkey puts its hand on the central pad. Then, later in the trial, a colored cue instructed a limb movement to the left touch pad if red or to the right one if green. The cues lasted for a variable delay, the instructed delay period, and their offset served as the go signal. The fixation spot was presented at the center of the screen or at one of four peripheral locations. Because the monkey's head was restrained, peripheral fixations caused a deviation of the eyes within the orbit, but for each fixation angle, the instructional cue was presented at nine locations with constant retinocentric coordinates. After the presentation of the instructional cue, 133 PMd cells displayed a phasic discharge (signal-related activity), 157 were tonically active during the instructed delay period (set-related or preparatory activity), and 104 were active after the go signal in relation to movement (movement-related activity). A large proportion of cells showed variations of the discharge rate in relation to limb movement direction, but only modest proportions were sensitive to the cue's location (signal, 43%; set, 34%; movement, 29%). More importantly, the activity of most neurons (signal, 74%; set, 79%; movement, 79%) varied significantly (analysis of variance, P < 0.05) with orbital eye position. A regression analysis showed that the neuronal activity varied linearly with eye position along the horizontal and vertical axes and can be approximated by a two-dimensional regression plane. These data provide evidence that eye position signals modulate the neuronal activity beyond sensory areas, including those involved in visually guided reaching limb movements. Further, they show that neuronal activity related to movement preparation and execution combines at least two directional parameters: arm movement direction and gaze direction in space. It is suggested that a substantial population of PMd cells codes limb movement direction in a head-centered reference frame.     

Sprawling locomotion in the lizard Sceloporus clarkii: speed modulation of motor patterns in a walking trot

Previous kinematic analyses in Sceloporus clarkii have shown that increased speed during trotting is attained by retracting the femur relatively faster (decreasing retraction time relative to stride duration) while all other aspects of axial and limb movements occur simply faster (scaling with stride duration). Thus, most of the limb muscles must be modulated to move the joints absolutely faster, while muscles effecting femoral retraction must be modulated differently to retract the femur relatively faster to increase speed. This prediction was examined by analyzing motor patterns in several key leg muscles in the spiny lizard running over a threefold increase in speed during a trot. The prediction is borne out in the limb muscles where the limb adductor (flexor tibialis), knee extender (femorotibialis), and plantar flexor of the ankle (gastrocnemius) have similar patterns of motor modulation that are different from that of the femoral retractor (caudofemoralis). To modulate a muscle to move simply faster (scaled with speed) the offset of the motor pattern is moved relatively earlier to decrease burst duration, while the intensity of electromyographical activation is ramped up. Increasing the relative speed of action is done by activating the muscle earlier, increasing the duration of the burst, and increasing the relative level of activation. Comparisons to other studies illustrate that the confounding effects that stance and swing duration have on stride duration with speed have important consequences for functional interpretations and that scaling locomotory data to stance duration is a more appropriate and useful convention because it relates information directly to the duty cycle when the propulsive effects of motor modulation are transmitted to the substrate. The iliocostalis in Sceloporus clarkii has a pattern of activity indicating that it functions to rotate the pelvis to aid the contralateral duty cycle. This is strikingly different from the function of the iliocostalis in the monitor lizard. Differences in axial function and differences among lizards in postures of the foot and crus during locomotion indicate that there are different ways that lizards run and that the functional and anatomical diversity of modes of locomotion in lizards is greater than is recognized at present.     

Step-tracking movements of the wrist. IV. Muscle activity associated with movements in different directions

We examined the patterns of muscle activity associated with multiple directions of step-tracking movements of the wrist in humans and monkeys. Human subjects made wrist movements to 12 different targets that required varying amounts of flexion-extension and radial-ulnar deviation. Wrist muscles displayed two patterns of electromyographic (EMG) modulation as movement direction changed: amplitude graded and temporally shifted. The amplitude-graded pattern was characterized by modulation of the quantity of muscle activity that occurred during two distinct time periods, an agonist burst interval that began before movement onset and an antagonist burst interval that began just after movement onset. The timing of muscle activity over the two intervals showed little variation with changes in movement direction. For some directions of movement, EMG activity was present over both time intervals, resulting in "double bursts." Modulation of activity during the agonist burst interval was particularly systematic and was well fit by a cosine function. In contrast, the temporally shifted pattern was characterized by a gradual change in the timing of a single burst of muscle activity. The burst occurred at a time intermediate between the agonist and antagonist burst intervals. The temporally shifted pattern was seen less frequently than the amplitude-graded pattern and was present only in selected wrist muscles for specific directions of movement. Monkeys made wrist movements to 8-16 different targets that required varying amounts of flexion-extension and radial-ulnar deviation. These movements were performed more slowly than those of human subjects. The wrist muscles of the monkeys we examined displayed the amplitude-graded pattern of activity but not the temporally shifted pattern. Stimulation of individual wrist muscles in monkeys resulted in wrist movements that were markedly curved, particularly for the wrist extensors. These results indicate that step-tracking movements of the wrist are generated mainly by using the amplitude-graded pattern to modulate muscle activity. We propose that this pattern reflects a central process that decomposes an intended movement into an agonist, "propulsive" component and an antagonist, "braking" component. Separate bursts of muscle activity then are generated to control each component. On the other hand, we argue that the temporally shifted pattern may function to reduce the amount of movement curvature associated with the activation of wrist muscles.     

Modulation of cerebral somatosensory evoked potentials arising from tibial and sural nerve stimulation during rhythmic active and passive movements of the human lower limb

The magnitudes of cerebral somatosensory evoked potentials (SEPs), following stimulation of cutaneous or muscle afferents in the upper limb, are reduced during active and passive movements of the fingers. The generalizability of such a movement effect was tested for lower limb events. We measured SEP magnitudes following activation of cutaneous (sural) and mixed (tibial) nerves during the flexion phase of active and passive rhythmic movements of the human lower limb. In eight volunteers, 150 SEPs per condition were recorded from Cz' referenced to Fpz'. Compared to stationary controls, both active and passive movements significantly depressed the early SEP components (P1-N1) [mean values, to 12.8%, 9.9% respectively for tibial nerve and to 29.6%, 25.6% for sural nerve stimulation, p < 0.05]. The attenuation was still observed when only one leg was moved and with stimulation at an earlier point in the flexion phase of movement. Visual fixation did not significantly affect P1-N1 amplitudes, compared to eyes closed. As previously shown, soleus H reflexes with stable M waves were significantly depressed during the movements (p < 0.05). The general construct may be that centripetal flow initiated from somatosensory receptors during limb movement leads to modulation of both spinal and cortical responses following large diameter cutaneous or muscle afferent activation.     

Movement-related potential measures of different modes of movement selection in Parkinson's disease

Movement-related potentials were recorded preceding self-paced voluntary movements in patients with Parkinson's disease and in healthy subjects of the same age group. We compared the Readiness Potential preceding joystick movements in a fixed direction and preceding joystick movements in freely selected directions. In normal subjects the Readiness Potential amplitude was higher preceding freely selected movements than preceding movements in a fixed direction. The Readiness Potential in Parkinson patients failed to be modified by the different modes of movement selection. The modulation of the Readiness Potential by different ways of preparing for movement might be due to the supplementary motor area (SMA) being more strongly engaged by tasks requiring internal control of movements than by tasks that are externally structured. The results suggest that this task-dependent variation of SMA activity is reduced in Parkinson's disease. A failing capacity to adapt SMA activity to different task demands has previously been suggested by evidence from positron emission tomography studies using similar tasks.     

Morphometric ultrastructural evaluation of the axonal endings in the neuromuscular junctions of pigeons after long lasting limitation of movement

To analyze the discharge patterns of the reticulospinal (R-S) neurons associated with four-limb movement, we recorded the unit spikes of 108 R-S neurons in 18 thalamic cats. (1) Unit spikes of R-S neurons exhibited alternating firings during leg movements, not only stepping on the treadmill but also upon passive flexion and extension movement by the experimenter's hand. (2) R-S neurons manifested firing patterns associated with diagonal, reciprocal and quadrupedal leg movements. About half of the neurons showed reciprocal patterns upon bilateral forelimb movements; spikes were increased when the ipsilateral forelimb was in a backward position; they were decreased when that leg was in a forward position. In contrast, the spikes were increased when the contralateral forelimb was placed forward and decreased when it was backward. About 15% of the R-S neurons showed discharge patterns correlated with quadrupedal leg movements. Firing increased when the left forelimb and right hindlimb were placed backward and the left hindlimb and right forelimb were forward. In contrast, when the position of all 4 limbs was reversed, firing rates decreased. (3) When brief touch stimulation was applied to the skin around the leg, bursting spikes were obtained; these were suppressed upon touching the skin of the contralateral limb. Even after transection of the muscle nerves, alternating firings were observed. (4) Local anesthesia to the shoulder joint resulted in a marked reduction of spontaneous discharges and alternating firings. (5) Our results indicate that afferents of joints and of cutaneous origins in individual limbs ascend to the brainstem reticular formation, that integrative action is organized as pattern generation in that region, and that this patterned information is sent to the spinal cord via the reticulospinal tracts.     

Wipe and flexion reflexes of the frog. I. Kinematics and EMG patterns

1. We evaluated the hypothesis that the neural control of complex motor behaviors is simplified by building movement sequences from a series of simple neural "building blocks." In particular, we compared two reflex behaviors of the frog, flexion withdrawal and the hindlimb-hindlimb wipe reflex, to determine whether a single neural circuit that coordinates flexion withdrawal is incorporated as the first element in a sequence of neural circuits comprising the wipe. The neural organization of these two reflexes was compared using a quantitative analysis of movement kinematics and muscle activity patterns [electromyograms (EMGs)]. 2. The three-dimensional coordinates of the position of the foot over time and the angular excursion of hip, knee, and ankle joints were recorded using a WATSMART infrared emitter-detector system. These data were quantified using principal-components analysis to provide a measure of the shape (eigenvalues) and orientation (eigen-vector coefficients) of the movement trajectories. The latencies and magnitudes of EMGs of seven muscles acting at the hip, knee, and ankle were analyzed over the interval from EMG onset to movement onset, and EMG magnitudes during the initial flexion of the limb. These variables were compared during flexion withdrawal and the initial flexion movement of the limb during the hindlimb-hindlimb wipe reflex (before the onset of the frequently rhythmic portion when the stimulus is removed) when the two reflexes were elicited from comparable stimulus locations. 3. In both the flexion reflex and the initial movement segment of the wipe reflex, the foot moves along a relatively straight line. However, the foot is directed to a more rostral and lateral position during flexion than during wipe. All three joints flex during flexion withdrawal, whereas during the wipe, the knee and ankle joints flex but the angular excursion of the hip joint may vary. The different orientations of the movement trajectories are associated with EMG patterns that differ in both timing and magnitude between the two reflexes. 4. The differences in the kinematics and EMG patterns of the two reflexes during unrestrained movements make it unlikely that the neural circuit that coordinates flexion withdrawal is incorporated as the first element in the sequence of neural circuits underlying the wipe reflex. 5. Unlike the wipe reflex, during flexion withdrawal there is no apparent constraint on the accuracy of placement at the end of the movement, yet the animals nevertheless achieved consistent final positions of both the foot and of each joint. The implications of these findings with respect to the controlled variables are discussed.     

Characterization of a directional selective inhibitory input from the medial terminal nucleus to the pretectal nuclear complex in the rat

The receptive field properties of neurons in the medial terminal nucleus of the accessory optic system (MTN) that project to the ipsilateral nucleus of the optic tract (NOT) and dorsal terminal nucleus (DTN), as identified by antidromic electrical activation, were analysed in the anaesthetized rat. The great majority (88%) of MTN neurons that were antidromically activated from NOT and DTN preferred downward directed movement of large visual stimuli while the remaining cells preferred upward directed stimulus movement. Distinct retrograde tracer injections into the NOT/DTN and the ipsilateral inferior olive (IO) revealed that no MTN neurons project to both targets. MTN neurons projecting to the ipsilateral NOT/DTN were predominantly found in the ventral part of the MTN, whereas those projecting to the IO were found in the dorsal part of the MTN. In situ hybridization for glutamic acid decarboxylase (GAD) mRNA was used as a marker for GABAergic neurons. Up to 98% of MTN neurons retrogradely labelled from the ipsilateral NOT/DTN also expressed GAD mRNA. Earlier studies have shown that MTN neurons that prefer upward directed stimulus movements are segregated from MTN neurons that prefer downward directed stimulus movements. It also has been demonstrated that directionally selective neurons in the NOT/DTN prefer horizontal stimulus movements and receive an inhibitory input from ipsilateral MTN. Our results indicate that this input is mediated by GABAergic cells in the ventral part of MTN, which to a large extent prefer downward directed stimulus movements, and that the great majority of MTN neurons that prefer upward directed stimulus movements project to other targets one of which possibly is the IO.     

Dorsal spinocerebellar tract neuronal activity in the intact chronic cat

The ability to electrophysiologically identify the axonal projections of lumbar neurons recorded in chronic unanesthetized intact awake animals is a formidable but essential requirement toward understanding ascending sensory transmission under naturally occurring conditions. Chronic immobilization procedures previously introduced by Morales et al. (1981) for intracellular studies of motoneurons are modified and then integrated with procedures for antidromic cellular identification and extracellular recording of upper (or lower) dorsal lumbar spinocerebellar tract (DSCT) neuronal activity, in conjunction with behavioral state recording and drug microiontophoresis. These implant procedures provide up to 6 months of stable recording conditions and, when combined with other techniques, allow individual DSCT neurons to be monitored over multiple cycles of sleep and wakefulness, following the induction into and recovery from barbiturate anesthesia and/or during the juxtacellular microiontophoretic ejection of inhibitory or excitatory amino acid neurotransmitters. The combination of such techniques allows a comprehensive examination of synaptic transmission through the DSCT and other lumbar sensory pathways in the intact normally respiring cat and its modulation during the general anesthetic state. These techniques permit investigations of the supraspinal controls impinging on lumbar sensory tract neurons during wakefulness and other behavioral states such as active sleep.     

Quantitative analysis of the movements of cytoplasmic granules in polarized fibroblasts

Movements of cytoplasmic organelles were analyzed in Vero fibroblasts. In the cells polarized at the edge of an experimental wound, cytoplasmic granules moved randomly (Brownian motions) and by separate jumps (saltatory movements). The displacement of granules by the Brownian motions exceeded by more than an order of magnitude that of the mitochondria similar by weight. Lipid droplets moved predominantly by saltations, whereas mitochondria and lysosomes moved much less often. In a front part of the polarized cells, the main directions of saltatory movements were from the nucleus to the leading edge of a cell and back, whereas the tangential movements (across the long axis of a cell) were less than 1%. 90% of saltatory movements occurred in the area starting 10-12 microm from the nucleus and ending 10-12 microm from the leading edge of a cell. The average rate of saltatory movements of the granules (2.38 microm/s) was identical in both directions. The average length of the track was 7.49 microm; the maximum track length reached 30 microm. An increase in the granule diameter from 0.3 to 1.4 microm resulted in a minor (statistically insignificant) decrease in the average rate of the movements. The average rate of saltatory movements of mitochondria was 1.00 microm/s, and the average track length was 6.04 microm. Therefore, mitochondria, in contrast to lipid droplets, are rigidly fixed in the cytoplasm, and the force holding mitochondria is equal to the force produced by the microtubule-associated motors. Taking into account the characteristic of the centrifugal saltations, we suggest that they are mediated by an unusual dynein.     

La manipulation d'indices kinesthésiques et tactiles lors d'une tache de reproduction de mouvement.

Reexamined the effect of several variables on the performance of the kinesthetic system. A classical movement duplication task was used in which each of the 48 18-38 yr old Ss was required to duplicate a passive criterion movement of one arm by positioning the same arm. Six factors were manipulated in a randomized design with fixed effects: mode of duplication (passive, active), tactile sensation (limb uncovered, limb covered, limb covered with forced air currents), kinesthetic cues for reproduction (either final arm position or distance moved), the sector of presentation of the criterion movement (43-76°, 87-217°, 128-264° in a horizontal arm adduction movement), information load (1, 2, or 4 criterion demonstrations of the movement), and the starting position of duplication. Constant and variable errors were used as dependent variables. Results confirm that active duplication was superior to passive, concerning variable, but not constant error. Duplication of final arm position was better than duplication of distance moved, also concerning only variable error. Two significant interactions (mode of duplication by kinesthetic cues, and kinesthetic cues by starting position of duplication) were found. Results indicate that final location cue and starting position of duplication are important markers in accurate reproduction. It is suggested that joint afference is transformed into a motor plan compatible with active duplication. (English summary) (25 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

The structure of somatosensory information for human postural control

The goal of the present study was to determine the properties of the somatosensory stimulus that alter its temporal coupling to body sway. Six standing subjects were tested while touching a metal plate positioned either directly in front of or lateral to the subject. In each condition, the plate moved 4 mm at 0.2 Hz in either the medial-lateral (ML) or anterior-posterior direction (AP). The results showed that coupling between body sway and touch plate movement was strongest when the touch plate moved in a direction along the longitudinal axis of the arm. Coupling strength was weaker when the touch plate moved perpendicular to the longitudinal axis of the arm. The results consistently show that a radial expansion stimulus was more effective than a lamellar-type stimulus at the fingertip. Moreover, somatosensory information from a surface is interpreted in terms of the orientation of the contact limb and the potential degrees of freedom available through its movement.     

Changing directions of forthcoming arm movements: neuronal activity in the presupplementary and supplementary motor area of monkey cerebral cortex

1. To understand roles played by two cortical motor areas, the presupplementary motor area (pre-SMA) and supplementary motor area (SMA), in changing planned movements voluntarily, cellular activity was examined in two monkeys (Macaca fuscata) trained to perform an arm-reaching task in which they were asked to press one of two target buttons (right or left) in three different task modes. 2. In the first mode (visual), monkeys were visually instructed to result and press either a right or left key in response to a forth coming trigger signal. In the second mode (stay), monkeys were required to wait for the trigger signal and press the same target key as pressed in preceding trials. In the third mode (shift), a 50 Hz auditory cue instructed the monkey to shift the target of the future reach from the previous target to the previous nontarget. 3. While the monkeys were performing this task, we recorded 399 task-related cellular activities from the SMA and the pre-SMA. Among them, we found a group of neurons that exhibited activity changes related specifically to shift trials (shift-related cells). The following properties characterized these 112 neurons. First, they exhibited activity changes after the onset of the 50-Hz auditory cue and before the movement execution when the monkeys were required to change the direction of forthcoming movement. Second, they were not active when the monkeys pressed the same key without changing the direction of the movements. Third, they were not active when the monkeys received the 50-Hz auditory cue but failed to change the direction of the movements by mistake. These observations indicate that the activity of shift-related cells is related to the redirection of the forthcoming movements, but not to the auditory instruction itself or to the location of the target key or the direction of the forthcoming movements. 4. Although infrequently, monkeys made errors in the stay trials and changed directions of the reach voluntarily. In that case, a considerably high proportion of shift-related neurons (12 of 19) exhibited significant activity changes long before initiation of the reach movement. These long-lasting activities were not observed during the preparatory period in correct stay trials, but resembled the shift-related activity observed when the target shift was made toward the same direction. Thus these activity changes were considered to be also related to the process of changing the intended movements voluntarily. 5. We found another population of neurons that showed activity modulation when the target shift was induced by the visual instruction in visual trials (visually guided shift-related neurons). These neurons were active when the light-emitting diode (LED) guided the forthcoming reach to the previous nontarget but not to the previous target. Therefore their activity was not a simple visual response to the LED per se. A majority of them also showed shift-related activity in shift trials (19 of 22 in monkey 2). 6. Neurons exhibiting the shift-related activity were distributed differentially among the two areas. In the pre-SMA, 31% of the neurons recorded showed the shift-related activity, whereas in the SMA, only 7% showed such an activity. These results suggest that pre-SMA and SMA play differential roles in updating the motor plans in accordance with current requirements.     

The role of inertial sensitivity in motor planning

To achieve a given motor task a single trajectory must be chosen from the infinite set of possibilities consistent with the task. To investigate such motor planning in a natural environment, we examined the kinematics of reaching movements made around a visual obstacle in three-dimensional space. Within each session, the start and end points of the movement were uniformly varied around the obstacle. However, the distribution of the near points, where the paths came closest to the obstacle, showed a strong anisotropy, clustering at the poles of a preferred axis through the center of the obstacle. The preferred axes for movements made with the left and right arms were mirror symmetric about the midsagittal plane, suggesting that the anisotropy stems from intrinsic properties of the arm rather than extrinsic visual factors. One account of these results is a sensitivity model of motor planning, in which the movement path is skewed so that when the hand passes closest to the obstacle, the arm is in a configuration that is least sensitive to perturbations that might cause collision. To test this idea, we measured the mobility ellipse of the arm. The mobility minor axis represents the direction in which the hand is most inertially stable to a force perturbation. In agreement with the sensitivity model, the mobility minor axis was not significantly different from the preferred near point axis. The results suggest that the sensitivity of the arm to perturbations, as determined by its inertial stability, is taken into account in the planning process.     

Age-associated modulation of apoptosis and activation in murine B lymphocytes

We examined neuronal activity in three motor cortical areas while a rhesus monkey adapted to novel visuomotor transforms. The monkey moved a joystick that controlled a cursor on a video screen. Each trial began with the joystick centered. Next, the cursor appeared in one of eight positions, arranged in a circle around a target stimulus at the center of the screen. To receive reinforcement, the monkey moved the joystick so that the cursor contacted the target continuously for Is. The video monitor provided continuous visual feedback of both cursor and target position. With those elements of the task constant, we modified the transform between joystick movement and that of the cursor at the beginning of a block of trials. Neuronal activity was studied as the monkey adapted to these novel joystick-cursor transforms. Some novel tasks included spatial transforms such as single-axis inversions, asymmetric double-axis inversions and angular deviations (also known as rotations). Other tasks involved changes in the spatiotemporal pattern and magnitude of joystick movement. As the monkey adapted to various visuomotor tasks, 209 task-related neurons (selected for stable background activity) showed significant changes in their task-related activity: 88 neurons in the primary motor cortex (M1), 32 in the supplementary motor cortex (M2), and 89 in the caudal part of the dorsal premotor cortex (PMdc). Slightly more than half of the sample in each area showed significant changes in the magnitude of activity modulation during adaptation, with the number of increases approximately equaling the number of decreases. These data support the prediction that changes in task-related neuronal activity can be observed in M1 during motor adaptation, but fail to support the hypothesis that M1 and PMdc differ in this regard. When viewed in population averages, motor cortex continued to change its activity for at least dozens of trials after performance reached a plateau. This slow, apparently continuing change in the pattern and magnitude of task-related activity may reflect the initial phases of consolidating the motor memory for preparing and executing visuomotor skills.     

A morphological study of cat dorsal spinocerebellar tract neurons after intracellular injection of horseradish peroxidase

This work represents an attempt to elucidate structural features of electrophysiologically characterized, individual cat dorsal spinocerebellar tract (DSCT) neurons by using intracellular application of horseradish peroxidase (HRP). Intracellular recordings and HRP injections were made in DSCT neurons of the Clarke's column in cat lumbar (L3) spinal cord. The units were identified by antidromic invasion following electrical stimulation of the ipsilateral dorsolateral funiculus at C1. In addition, sensory inputs to the DSCT neurons were determined by natural (adequate) stimuli applied to the hind limb with intact innervation. The morphological analysis is based on data obtained from 19 well-stained electrophysiologically identified neurons located in Clarke's column. Thirteen of these units received excitatory sensory inputs from muscle receptors, two were activated by cutaneous afferents only, and four had a convergent (muscle + cutaneous) input. The DSCT--muscle cells were equivalent to the large Clarke cells (class C of Leowy, '70). Their dendrites were oriented primarily in the rostro--caudal direction (up to 2500 micron) and appeared generally smooth except for some branchlets. In four of these cells, the axon was traced into the lateral funiculus. In light microscopic analysis there was no evidence that axon collaterals arose from these axons during the initial trajectory through the spinal grey matter. The four DSCT--convergent neurons were similar in shape to the DSCT--muscle units although they appeared to have somewhat smaller cell bodies. Of the two DSCT--cutaneous neurons one was found to be of the B type, with the dendritic tree having fewer branches and oriented mainly in the medio--lateral direction. The other cell, however, turned out to be similar in appearance to the C type Clarke neurons.     

Directional organization of eye movement and visual signals in the floccular lobe of the monkey cerebellum

The floccular lobe of the monkey is critical for the generation of visually-guided smooth eye movements. The present experiments reveal physiological correlates of the directional organization in the primate floccular lobe by examining the selectivity for direction of eye motion and visual stimulation in the firing of individual Purkinje cells (PCs) and mossy fibers. During tracking of sinusoidal target motion along different axes in the frontoparallel plane, PCs fell into two classes based on the axis that caused the largest modulation of simple-spike firing rate. For "horizontal" PCs, the response was maximal during horizontal eye movements, with increases in firing rate during pursuit toward the side of recording (ipsiversive). For "vertical" PCs, the response was maximal during eye movement along an axis just off pure vertical, with increases in firing rate during pursuit directed downward and slightly contraversive. During pursuit of target motion at constant velocity, PCs again fell into horizontal and vertical classes that matched the results from sinusoidal tracking. In addition, the directional tuning of the sustained "eye velocity" and transient "visual" components of the neural responses obtained during constant velocity tracking were very similar. PCs displayed very broad tuning approximating a cosine tuning curve; the mean half-maximum bandwidth of their tuning curves was 170-180 degrees. Other cerebellar elements, related purely to eye movement and presumed to be mossy fibers, exhibited tuning approximately 40 degrees narrower than PCs and had best directions that clustered around the four cardinal directions. Our data indicate that the motion signals encoded by PCs in the monkey floccular lobe are segregated into channels that are consistent with a coordinate system defined by the vestibular apparatus and eye muscles. The differences between the tuning properties exhibited by PCs compared with mossy fibers indicate that a spatial transformation occurs within the floccular lobe.