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Movement and electromyographic disorders associated with cerebellar dysmetria.

The objective of these experiments was to determine whether dysmetric elbow flexions, which occurred during cerebellar dysfunction, had the same kinematic and electromyographic characteristics as movements of the same amplitude and velocity performed under normal conditions. Reversible cerebellar lesions were produced by cooling through two probes implanted on either side of the dentate nucleus in five Cebus albifrons monkeys. Normal, fast, and accurate elbow flexions had single-peaked velocities and a bi- or triphasic EMG pattern in agonist and antagonist muscles. During cerebellar dysfunction movements became ataxic. Ataxic movements were classified into two categories: those with oscillations (tremor) during the movement and those without oscillations. A terminal tremor occurred after both types of movements. Oscillations during movements were more likely to occur when a constant force loaded the antagonist. Addition of mass to the handle attenuated or abolished the oscillations. Movements with oscillations reached the target with increased variability of end position, whereas movements without oscillations were often hypermetric. The movement parameters and EMG patterns associated with flexions without oscillations during the movement were studied in detail. A characteristic of these movements was that the acceleration and deceleration phases were asymmetric. Compared with control movements of the same peak velocity, they had smaller magnitudes of acceleration and larger magnitudes of deceleration. The large deceleration was abnormal because it initiated the terminal tremor. The disorder in acceleration was associated with agonist EMG activity that was less abrupt in onset, smaller in magnitude, and more prolonged in duration. The disorder in deceleration was associated with delayed onset of phasic antagonist EMG activity. The results show that hypermetric arm movements without oscillations have different properties than those of normal movements of similar velocity and amplitude. Thus it is unlikely that dysmetria results from inappropriate selection or triggering of an otherwise normal motor program. We conclude that normal function of the cerebellum is necessary for the generation of agonist and antagonist muscle activity that is both of the appropriate magnitude and timing to control the dynamic phase of arm movements.

Animals↗

Cerebellar neuronal activity related to whole-arm reaching movements in the monkey.

1. Three monkeys were trained to make whole-arm reaching movements from a common central starting position toward eight radially arranged targets disposed at 45 degrees intervals. A sample of 312 cerebellar neurons with proximal-arm receptive fields or discharge related to shoulder or elbow movements was studied in the task. The sample included 69 Purkinje cells, 115 unidentified cortical cells, 65 interpositus neurons, and 63 dentate units. 2. The reaching task was divided into three movement-related epochs: a reaction time, a movement time, and holding over the target. All neurons demonstrated significant changes in discharge during one or more of these three epochs. Almost all of the cells (95%) showed a significant change in activity during the movement, whereas 68-69% of the cells showed significant changes from premovement activity during the reaction time and holding periods. 3. During the combined reaction time-movement period, 231/312 cells were strongly active in the task. Of these, 151 cells (65.4%) demonstrated unimodal directional responses. Sixty-three had a reciprocal relation to movement direction, whereas 88 showed only graded increases or decreases in activity. A further 37 cells (16.0%) were nondirectional, with statistically uniform changes in discharge in all eight directions. The remaining 43 cells (18.6%) showed significant differences in activity for different directions of movement, but their response patterns were not readily classifiable. 4. The proportion of directional versus nondirectional cells was consistent across the four cell populations. However, graded response patterns were more common and reciprocal responses less common among Purkinje and dentate neurons than among unidentified cortical cells and interpositus neurons. 5. The distribution of preferred directions of the population of cerebellar neurons covered all possible movement directions away from the common central starting position in the horizontal plane. When the preferred direction of each cell in the sample population was aligned, the mean direction-related activity of the cerebellar population formed a bell-shaped tuning curve for the activity recorded during both the reaction time and the movement, as well as during the time the arm maintained a fixed posture over the targets. A vector representation also showed that the overall activity of the cerebellar population during normal reaching arm movements generated a signal that varied with movement direction. 6. These results demonstrate that the cerebellum generates a signal that varies with the direction of movement of the proximal arm during normal aimed reaching movements and is consistent with a role in the control of the activity of muscles or muscle groups generating these movements.

Animals↗

Proprioceptive consequences of tendon vibration during movement.

1. Previous studies have used tendon vibration to investigate kinesthetic illusions in the isometric limb and end point control in the moving limb. These previous studies have shown that vibration distorts the perceptions of static joint angle and movement and causes systematic errors in the end point of movement. In this paper we describe the effects of tendon vibration during movement while human subjects performed a proprioceptively coordinated motor task. In an earlier study we showed that the CNS coordinates this motor task-a movement sequence-with proprioceptive information related to the dynamic position and velocity of the limb. 2. When performing this movement sequence, each subject sat at a table and opened the right hand as the right elbow was passively rotated in the extension direction through a prescribed target angle. Vision of the arm was prevented, and the movement velocity was changed randomly from trial to trial, leaving proprioception as the only useful source of kinematic information with which to perform the task. 3. In randomly occurring trials, vibration was applied to the tendon of the biceps brachii, a muscle that lengthens during elbow extension. In some experiments the timing of tendon vibration was varied with respect to the onset of elbow rotation, and in other experiments the frequency of vibration was varied. In each experiment we compared the accuracy of the subject's response (i.e., the elbow angle at which the subject opened the hand) in trials with tendon vibration with the accuracy in trials without tendon vibration. 4. The effect of tendon vibration depended on the frequency of vibration. When the biceps tendon was vibrated at 20 Hz, subjects opened the hand after the elbow passed through the target angle ("overshooting"). Overshooting is consistent with an underestimate of the actual displacement or velocity of the elbow. Vibration at 30 Hz had little or no effect on the elbow angle at hand opening. Vibration at 40 Hz caused subjects to open the hand before the elbow reached the target angle ("undershooting"). Undershooting is consistent with an overestimate of the actual displacement or velocity of the elbow. The size of the error depended on the velocity of the passively imposed elbow rotation. 5. The effect of tendon vibration also depended on the timing of vibration. If 40-Hz vibration began at the onset of movement, the subject undershot the target. If 40-Hz vibration started 5 s before movement onset and continued throughout the movement, the undershoot error increased in magnitude. However, if 40-Hz vibration started 5 s before movement onset and then stopped at movement onset, the subject overshot the target. When vibration was shut off during movement, a transition occurred from an over-shooting error to an undershooting error at a time that depended on the velocity of elbow rotation. 6. In a separate experiment, subjects were instructed to match either the perceived dynamic position or the perceived velocity of rotation imposed on the right elbow by actively rotating the left elbow. In both matching tasks, tendon vibration produced oppositely directed errors depending on the frequency of vibration. Vibration at 20 Hz produced a perception of decreased elbow velocity and a bias in dynamic position in the flexion direction, and vibration at 40 Hz produced the opposite perceptions. 7. We conclude that muscle spindle afferents, which are activated by tendon vibration, are an important source of the dynamic position and velocity information that the CNS uses to coordinate this movement sequence task. The observed effects of vibration timing and frequency suggest that perceptual changes evoked by vibration cannot be explained by the simple summation of sensory input evoked by movement and by vibration. Rather, the bias in perception produced by vibration appears to be related to the difference between vibration- and movement-evoked activity in muscle spindle afferents.

Adult↗

Temporary inactivation in the primate motor thalamus during visually triggered and internally generated limb movements.

To better understand the contribution of cerebellar- and basal ganglia-receiving areas of the thalamus [ventral posterolateral nucleus, pars oralis (VPLo), area X, ventral lateral nucleus, pars oralis (VLo), or ventral anterior nucleus, pars parvicellularis (VApc)] to movements based on external versus internal cues, we temporarily inactivated these individual nuclei in two monkeys trained to make visually triggered (VT) and internally generated (IG) limb movements. Infusions of lignocaine centered within VPLo caused hemiplegia during which movements of the contralateral arm rarely were performed in either task for a short period of time ( approximately 5-30 min). When VT responses were produced, they had prolonged reaction times and movement times and a higher incidence of trajectory abnormalities compared with responses produced during the preinfusion baseline period. In contrast, those IG responses that were produced remained relatively normal. Infusions centered within area X never caused hemiplegia. The only deficits observed were an increase in reaction time and movement amplitude variability and a higher incidence of trajectory abnormalities during VT trials. Every other aspect of both the VT and IG movements remained unchanged. Infusions centered within VLo reduced the number of movements attempted during each block of trials. This did not appear to be due to hemiplegia, however, as voluntary movements easily could be elicited outside of the trained tasks. The other main deficit resulting from inactivation of VLo was an increased reaction time in the VT task. Finally, infusions centered within VApc caused IG movements to become slower and smaller in amplitude, whereas VT movements remained unchanged. Control infusions with saline did not cause any consistent deficits. This pattern of results implies that VPLo and VLo play a role in the production of movements in general regardless of the context under which they are performed. They also suggest that VPLo contributes more specifically to the execution of movements that are visually triggered and guided, whereas area X contributes specifically to the initiation of such movements. In contrast, VApc appears to play a role in the execution of movements based on internal cues. These results are consistent with the hypothesis that specific subcircuits within the cerebello- and basal ganglio-thalamo-cortical systems preferentially contribute to movements based on external versus internal cues.

Analysis of Variance↗

Head waving in Aplysia californica. I. Behavioural characterization of searching movements.

Searching is an important component of several types of goal-directed behaviour. In soft-bodied animals, searching behaviour can appear quite complex because the range of body movement is not constrained by joints, limbs or muscles with discrete areas of origin and insertion. In addition, animals exhibiting this type of behaviour utilize their maximum freedom of movement. In this paper, we describe the head-waving searching behaviour of the sea hare Aplysia californica by characterizing patterns of movements and the changes in body shape that underlie these movements. A bout of head waving consists of a series of discrete movements separated by pauses. Each discrete movement lasts 4-10s and is directed either horizontally or vertically with respect to stationary part of the animal. Large movements, such as bending from the extreme right to the extreme left, consist of a series of these shorter movements separated by pauses lasting 1-2s. In all head-waving movements, the transverse axis of the head is kept relatively parallel with the substratum. Thus, vertical movements require only bending of the body, whereas horizontal movements require twisting of the body, particularly when the posture is more erect. During head waving, the anterior two-thirds of the body is free to move, but most of the bending occurs immediately posterior to the head region. There is no periodicity of movements within a bout of head waving, although isolated instances of repeated movements are sometimes observed. Therefore, although the individual movements during head waving are relatively simple, the absence of a patterned sequence accounts for the complexity of the overall behaviour. These observations both constrain models of the neural organization of head waving and provide criteria for categorizing head-waving movements in further behavioural and physiological studies.

Animals↗

Making arm movements within different parts of space: the premotor and motor cortical representation of a coordinate system for reaching to visual targets.

The activity of 156 individual arm-related neurons was studied in the premotor cortex (area 6) while monkeys made arm movements of similar directions within different parts of 3-dimensional space. This study was aimed at describing the relationship between premotor cortical cell activity and direction of arm movement and assessing the coordinate system underlying this relationship. We found that the activity of 152 (97.4%) cells varied in an orderly fashion with the direction of movement, in at least some region of the work space. Premotor cortical cells fired most for a given preferred direction and less for other directions of movement. These preferred directions covered the directional continuum in a uniform fashion across the work space. It was found that, as movements of similar directions were made within different parts of the work space, the cells' preferred directions changed their orientation. Although these changes had different magnitudes for different cells, at the population level, they followed closely the changes in orientation of the arm necessary to move the hand from one to another part of the work space. This shift of cells' preferred direction with the orientation of the arm in space has been observed with similar characteristics in the motor cortex (see Caminiti et al., 1990). In both premotor and motor cortices, neuronal movement population vectors provide a good description of movement direction. Unlike the individual cell preferred directions upon which they are based, movement population vectors did not change their spatial orientation across the work space, suggesting that they remain good predictors of movement direction regardless of the region of space in which movements are made. The firing frequency of both premotor and motor cortical neurons varied significantly with the position occupied by the hand in space. These static positional effects were observed in 88.5% of premotor and 91.8% of motor cortical cells. In a second task, monkeys made movements from differing origins to a common end point. This task was performed within 3 different parts of space and was aimed at dissociating movement direction from movement end point. It was found that in both premotor and motor cortices virtually all cells were related to the direction and not to the end point of movement. These data suggest that premotor and motor cortices use common mechanisms for coding arm movement direction. They also provide a basis for understanding the coordinate transformation required to move the hand toward visual targets in space.

Analysis of Variance↗

[Biomechanical study on orthodontic tooth movement by means of numerical simulation. Effects of principal stresses in periodontal membrane].

The effect of biomechanical factors on tooth movement has not been clarified in a quantitative manner. This study was designed to investigate the stresses affecting tooth movement, using a numerical simulation. The influence of decrease in Young's modulus of the periodontal membrane on canine retraction was also examined through the simulation. A two-dimensional finite element model was constructed based on the average anatomic morphology of Japanese canine. A numerical simulation program based on the finite element method was developed for the orthodontic tooth movement. The stresses in the periodontal membrane were evaluated. The principal stress of which absolute is larger was selected as a reference stress. Each nodal point at the alveolar bone-periodontal membrane interface was repositioned in the direction of reference stresses, in response to discrepancy between assumed thresholds and the reference stresses. Moment to force (M/F) ratios at the bracket position of this model were examined for evaluating force conditions. Simulation of tooth movement were executed under three force conditions with different M/F ratios for distal movements and two force conditions for vertical movements. Three types of canine retraction, tipping movement, bodily movement and root movement, were displayed with the numerical simulation. Extrusion and intrusion were also displayed. Analytic movements of the canine were close to the actual tooth movements that have been reported, utilizing the principal stresses with the thresholds of the maximum and minimum principal stresses being about +0.4 gf/mm2 and -0.4 gf/mm2. The decrease in the Young's modulus of the periodontal membrane changed bodily movement to tipping movement under the same force conditions. These results indicate that the value and the direction of the principal stress in the periodontal membrane are key determinants of tooth movement and this numerical simulation is useful for investigating the influence of the biomechanical factors on tooth movement.

Dental Stress Analysis↗

[Biomechanic study of sternal movements in adult man (author's transl)].

The study of the sternal movements by the use of biometry mensurations of 62 young adult men allows to precise the respiratory variations of the Louis's angle and sternal displacements in the sagittal plane. The sternal movements are successivly considered in standing and horizontal positions, at three respiratory moments corresponding to the respiratory rest, the forced inspiration and the forced expiration. The sternal displacements in the sagittal plane are the resultant of backward and forward movements and elevation and lowering movements. In standing position the amplitude of the forward movements can be compared to these of the backward movements while the amplitude of the elevation movements is four times more important than that of the lowering movements. In horizontal position the amplitude of the forward movements is twice more important than that of the backward movements and the amplitude of the elevation movements is twice more important than that of the lowering movements. The correlation research between the sternal movements and the vital capacity allows to precise that the vital capacity appears essentially in correlation with the maximum of the amplitude of the Louis's angle movements in horizontal position and with the forward and backward sternal movements in standing and horizontal positions.

Adult↗

On the relations between the direction of two-dimensional arm movements and cell discharge in primate motor cortex.

The activity of single cells in the motor cortex was recorded while monkeys made arm movements in eight directions (at 45 degrees intervals) in a two-dimensional apparatus. These movements started from the same point and were of the same amplitude. The activity of 606 cells related to proximal arm movements was examined in the task; 323 of the 606 cells were active in that task and were studied in detail. The frequency of discharge of 241 of the 323 cells (74.6%) varied in an orderly fashion with the direction of movement. Discharge was most intense with movements in a preferred direction and was reduced gradually when movements were made in directions farther and farther away from the preferred one. This resulted in a bell-shaped directional tuning curve. These relations were observed for cell discharge during the reaction time, the movement time, and the period that preceded the earliest changes in the electromyographic activity (approximately 80 msec before movement onset). In about 75% of the 241 directionally tuned cells, the frequency of discharge, D, was a sinusoidal function of the direction of movement, theta: D = b0 + b1 sin theta + b2cos theta, or, in terms of the preferred direction, theta 0: D = b0 + c1cos (theta - theta0), where b0, b1, b2, and c1 are regression coefficients. Preferred directions differed for different cells so that the tuning curves partially overlapped. The orderly variation of cell discharge with the direction of movement and the fact that cells related to only one of the eight directions of movement tested were rarely observed indicate that movements in a particular direction are not subserved by motor cortical cells uniquely related to that movement. It is suggested, instead, that a movement trajectory in a desired direction might be generated by the cooperation of cells with overlapping tuning curves. The nature of this hypothetical population code for movement direction remains to be elucidated.

Animals↗

[Study on human eye movements induced by blinks--report 1. Three kinds of blinking].

The eye movement associated with blinking was studied with a magnetic search coil method. Lid movements and eye movements associated with spontaneous blinks, voluntary blinks, and reflex blinks induced by puffs of air or flashes of light were analysed. Eye movements associated with spontaneous blinks and voluntary blinks had initial downward and adducted movements followed by upward and abducted movements of the eyeball. Reflex blinks initially showed the same downward and adducted movements, but upward and abducted movements were not constantly obtained in given subjects. The velocity-amplitude ratio was equal among the three kinds of blinks in both horizontal and vertical movements. The velocity-amplitude ratio of the eye movements associated with blinks and visually-guided saccadic eye movements was compared. The horizontal and vertical eye movements associated with blinks were slower than those of saccadic eye movements in the same subject. Therefore, the eye movements associated with the blinks may have a different neural mechanism in the brain from that of the saccadic eye movement system.

Adult↗

Threshold control of arm posture and movement adaptation to load.

We addressed the fundamental questions of which variables underlie the control of arm movement and how they are stored in motor memory, reproduced and modified in the process of adaptation to changing load conditions. Such variables are defined differently in two major theories of motor control (internal models and threshold control). To resolve the controversy, these theories were tested (experiment 1) based on their ability to explain why active movement away from a stable posture is not opposed by stabilizing mechanisms (the posture-movement problem). The internal model theory suggests that the system counteracts the opposing forces by increasing the muscle activity in proportion to the distance from the initial posture (position-dependent EMG control). In contrast, threshold control fully excludes these opposing forces by shifting muscle activation thresholds and thus resetting the stabilizing mechanisms to a new posture. Subjects were sitting, holding the vertical handle of a double-joint manipulandum with their right hand and were facing a computer screen on which the handle and target to be reached were displayed. In response to an auditory signal, subjects quickly moved the handle from an initial position to one of two (frontal and sagittal) targets. No load was applied during the movement but in separate trials, a brief perturbation was applied to the handle by torque motors controlling the manipulandum. Perturbations were applied prior to or 3 s after movement offset, in the latter case in one of eight directions. The EMG activity of the majority of the seven recorded muscles was at zero level before movement onset and returned to zero level after movement offset. Those muscles that remained active before or after the movement could be made silent whereas previously silent muscles could be activated after a small passive displacement (several millimeters) elicited by perturbations in appropriate directions. Results showed that the activation thresholds of motoneurons of arm muscles were reset from the initial to a final position and that EMG activity was not position-dependent. These results were inconsistent with the internal model theory but confirmed the threshold control theory. Then the ability of threshold control theory to explain rapid movement adaptation to a position-dependent load was investigated (experiment 2 and 3). Subjects produced fast movement to the frontal target with and without a position-dependent load applied to the handle. Trials were organized in blocks alternating between the load and no-load condition (20 blocks in total, with randomly chosen number of five to ten trials in each). Subjects were instructed "do not correct" in experiment 2 and "correct" movement errors during the trial in experiment 3. Five threshold arm configurations underlying the movement production and adaptation were identified. When instructed "do not correct", movement precision was fully restored on average after two trials. No significant improvement was observed as the experiment progressed despite the fact that the same load condition was repeated after one block of trials. Thus, in each block, the adaptation was made anew, implying that subjects relied on short-term memory and could not recall the threshold arm configurations they specified to accurately reach the same target in the same load condition in previous blocks. When instructed to "correct" within each trial, precision was restored faster, on average after one trial. Major aspects of the production and adaptation of arm movement (including the kinematics, movement errors, instruction-dependent behavior, and absence of position-related EMG activity) are explained in terms of threshold control.

Adaptation, Physiological↗

Cortical areas involved in virtual movement of phantom limbs: comparison with normal subjects.

OBJECTIVE: To demonstrate that amputees performing "virtual" movements of their amputated limb activate cortical areas previously devoted to their missing limb, we studied amputees with functional magnetic resonance imaging (fMRI) and positron emission tomographic (PET) scans and compared the results with those of normal volunteers performing imaginary movements during fMRI acquisitions. METHODS: Ten amputees (age range, 33-92 yr; average age, 49 yr; six men and four women; eight upper-limb and two lower-limb amputations) able to move their phantom limb at will were studied by fMRI (all patients) and PET scan (seven patients). The time between amputation and fMRI and PET studies ranged from 1 to 27 years (average, 13 yr). Patients were asked to perform virtual movements of the amputated limb and normal movements of the contralateral normal limb according to the functional images acquisition procedure. Movements of the stump were also used to differentiate stump cortical areas from virtual movement-activated areas. Ten right-handed volunteers, age- and sex-matched to the amputees, were also studied by fMRI. All volunteers were asked to perform four tasks during their fMRI study: imaginary movements of their right arm (1 task) and foot (1 task) and real movements of their left arm (1 task) and foot (1 task). RESULTS: In amputees, virtual movements of the missing limbs produced contralateral primary sensorimotor cortex activation on both fMRI and PET scans. These activation areas, different from the stump activation areas, were similar in location to contralateral normal limb-activated areas. Quantitatively, in two amputees who claimed to be able to perform both slow and fast virtual movements, regional cerebral blood flow measured by PET scan in the precentral gyrus increased significantly during fast movements in comparison with slow virtual movements. In normal subjects, significant differences between real versus imaginary fMRI activations were found (for both foot and hand movements); imaginary right hand and foot tasks activated primarily the contralateral supplementary motor areas, with no significant activation detected in the contralateral precentral or postcentral gyri. CONCLUSION: Primary sensorimotor cortical areas can be activated by phantom-limb movements and thus can be considered functional for several years or decades after amputation. In this study, we found that the location of the activation of these areas is comparable to that of activations produced by normal movements in control subjects or in amputees.

Adult↗

Intermanual interactions during initiation and production of rhythmic and discrete movements in individuals lacking a corpus callosum.

Three individuals lacking a corpus callosum, two due to callosotomy and one agenesis, and three age-matched healthy controls were tested on a bimanual task in which a discrete or rhythmic arm movement was initiated following a visual signal while the other arm produced continuous, rhythmic movements. The control participants initiated the secondary, rhythmic movement in phase with the ongoing rhythmic base movement and the two limbs were coupled in an inphase mode across the duration of the trial. In contrast, the acallosal individuals failed to show phase entrainment at the initiation of the secondary, rhythmic movements. Moreover, the callosotomy patients exhibited weak coupling between the rhythmically moving limbs while the individual with callosal agenesis consistently synchronized in an antiphase mode. The control participants exhibited increased perturbation of the ongoing base movement when initiating a discrete movement; for the acallosal participants, the base movement was similarly perturbed in both secondary movement conditions. These results are consistent with the hypothesis that intermanual interactions observed during bimanual movements arise from various levels of control, and that these are distinct for discrete and rhythmic movements. Temporal coupling during rhythmic movements arises in large part from transcallosal interactions between the two hemispheres. The imposition of a secondary movement may transiently disrupt an ongoing rhythmic movement even in the absence of the corpus callosum. This may reflect subcortical interactions associated with response initiation, or, due to dual task demands, a transient shift in attentional resources.

Agenesis of Corpus Callosum↗

Discrimination between active and passive head movements by macaque ventral and medial intraparietal cortex neurons.

An important prerequisite for effective motor action is the discrimination between active and passive body movements. Passive movements often require immediate reflexes, whereas active movements may demand suppression of the latter. The vestibular system maintains correct body and head posture in space through reflexes. Since vestibular inputs have been reported to be largely suppressed in the vestibular nuclei during active head movements, we investigated whether head movement-related signals in the primate parietal cortex, a brain region involved in self-motion perception, could support both reflex functions and self-movement behaviour. We employed a paradigm that made available direct comparison of neuronal discharge under active and passive movement conditions. In this study, we demonstrate that a population of intraparietal (VIP (ventral) and MIP (medial)) cortex neurons change their preferred directions during horizontal head rotations depending on whether animals have performed active movements, or if they were moved passively. In other neurons no such change occurred. A combination of these signals would provide differential information about the active or passive nature of an ongoing movement. Moreover, some neurons' responses clearly anticipated the upcoming active head movement, providing a possible basis for vestibular-related reflex suppression. Intraparietal vestibular neurons thus distinguish between active and passive head movements, and their responses differ substantially from those reported in brainstem vestibular neurons, regarding strength, timing, and direction selectivity. We suggest that the contextual firing characteristics of these neurons have far-reaching implications for the suppression of reflex movements during active movement, and for the representation of space during self-movement.

Action Potentials↗

Role of the cerebellar flocculus region in the coordination of eye and head movements during gaze pursuit.

The contribution of the flocculus region of the cerebellum to horizontal gaze pursuit was studied in squirrel monkeys. When the head was free to move, the monkeys pursued targets with a combination of smooth eye and head movements; with the majority of the gaze velocity produced by smooth tracking head movements. In the accompanying study we reported that the flocculus region was necessary for cancellation of the vestibuloocular reflex (VOR) evoked by passive whole body rotation. The question addressed in this study was whether the flocculus region of the cerebellum also plays a role in canceling the VOR produced by active head movements during gaze pursuit. The firing behavior of 121 Purkinje (Pk) cells that were sensitive to horizontal smooth pursuit eye movements was studied. The sample included 66 eye velocity Pk cells and 55 gaze velocity Pk cells. All of the cells remained sensitive to smooth pursuit eye movements during combined eye and head tracking. Eye velocity Pk cells were insensitive to smooth pursuit head movements. Gaze velocity Pk cells were nearly as sensitive to active smooth pursuit head movements as they were passive whole body rotation; but they were less than half as sensitive ( approximately 43%) to smooth pursuit head movements as they were to smooth pursuit eye movements. Considered as a whole, the Pk cells in the flocculus region of the cerebellar cortex were <20% as sensitive to smooth pursuit head movements as they were to smooth pursuit eye movements, which suggests that this region does not produce signals sufficient to cancel the VOR during smooth head tracking. The comparative effect of injections of muscimol into the flocculus region on smooth pursuit eye and head movements was studied in two monkeys. Muscimol inactivation of the flocculus region profoundly affected smooth pursuit eye movements but had little effect on smooth pursuit head movements or on smooth tracking of visual targets when the head was free to move. We conclude that the signals produced by flocculus region Pk cells are neither necessary nor sufficient to cancel the VOR during gaze pursuit.

Animals↗

Co-ordinated mandibular and head-neck movements during rhythmic jaw activities in man.

Recent observations in man of concomitant mandibular and head movements during single maximal jaw-opening/-closing tasks suggest a close functional relationship between the mandibular and the head-neck motor systems. This study was aimed at further testing of the hypothesis of a functional integration between the human jaw and neck regions. Spatiotemporal characteristics of mandibular and associated head movements were evaluated for 3 different modes of rhythmic jaw activities: self-paced continuous maximal jaw-opening/-closing movements, paced continuous maximal jaw-opening/-closing movements at 50 cycles/minute, and unilateral chewing. Mandibular and head-neck movements were simultaneously recorded in 12 healthy young adults, by means of a wireless opto-electronic system for 3-D movement recordings, with retro-reflective markers attached to the lower (mandible) and upper (head) incisors. The results showed that rhythmic mandibular movements were paralleled by head movements. An initial change in head position (head extension) was seen at the start of the first jaw-movement cycle, and this adjusted head position was retained during the following cycles. In addition to this prevailing head extension, the maximal jaw-opening/-closing cycles were paralleled by head extension-flexion movements, and in general the start of these head movements preceded the start of the mandibular movements. The results support the idea of a functional relationship between the temporomandibular and the cranio-cervical neuromuscular systems. We therefore suggest a new concept for human jaw function, in which "functional jaw movements" are the result of activation of jaw as well as neck muscles, leading to simultaneous movements in the temporomandibular, atlanto-occipital, and cervical spine joints.

Adult↗

Movement representation in the dorsal and ventral premotor areas of owl monkeys: a microstimulation study.

We used intracortical microstimulation to investigate the lateral premotor cortex and neighboring areas in 14 hemispheres of owl monkeys, focusing on the somatotopic distribution of evoked movements, thresholds for forelimb movements, and the relative representation of proximal and distal forelimb movements. We elicited movements from the dorsal and ventral premotor areas (PMD, PMV), the caudal and rostral divisions of primary motor cortex (M1c, M1r), the frontal eye field (FEF), the dorsal oculomotor area (OMD; area 8b), the supplementary motor area (SMA), and somatosensory cortex (areas 3a and 3b). Area PMD was composed of architectonically distinguishable caudal and rostral subdivisions (PMDc, PMDr). Stimulation of PMD elicited movements of the hindlimb, forelimb, neck and upper trunk, face, and eyes. Hindlimb and forelimb movements were represented in the caudalmost part of PMDc. Face, neck, and eye movements were represented in the lateral and rostral parts of PMDc and in PMDr. Stimulation of PMV elicited forelimb and orofacial movements, but not hindlimb movements. Both proximal and distal forelimb movements were elicited from PMDc and PMV, although PMD stimulation elicited mainly shoulder and elbow movements, while PMV stimulation evoked primarily wrist and digit movements. Distal movements were evoked more frequently from PMV than from M1r or M1c. Across cases, the median forelimb thresholds for PMDc and PMV were 60 and 36 microA, respectively, values that differ significantly from each other and from the value of 11 microA obtained for M1r. Our observations indicate that premotor cortex is much more responsive to electrical stimulation than commonly thought, and contains a large territory from which eye movements can be elicited. These results suggest that in humans, much of the electrically excitable cortex located on the precentral gyrus, including cortex sometimes considered part of the frontal eye field, is probably homologous to the premotor cortex of nonhuman primates.

Animals↗

Motor cortex and the distributed anatomy of finger movements.

Voluntary movements are thought to be controlled via a well-ordered, spatially discrete, somatotopic map in the primary motor cortex (M1). We examined this hypothesis in monkeys trained to perform visually-cued, individuated flexion and extension movements of each digit and of the wrist. Single neurone recordings in M1 during such finger movements revealed two unexpected features. First, single M1 neurones often discharge during instructed movements of multiple digits. Second, neurones active during any particular instructed movement are distributed widely throughout the same M1 territory as neurones active during any other movement. Reversible, partial inactivation of the M1 hand representation produced by injection of 5-10 microg muscimol at one site impaired the monkeys' ability to perform finger movements, but no relationship was evident between the particular finger movements that were affected and the mediolateral location of the injection site along the central sulcus. Thus each finger movement is represented by activity distributed widely in the M1 upper extremity representation. If not controlled from spatially segregated M1 regions, movements of different fingers might be controlled by groups of spatially scattered but physiologically similar neurones. Cluster analysis of M1 neurones demonstrated a large group that discharged during most finger movements, and a small group that paused during most movements. Distinct functional groups of M1 neurones that might control particular finger movements were identified inconsistently. We therefore hypothesize that M1 neurones are a very diverse network controlling finger movements.

Animals↗