Functional properties of grasping-related neurons in the ventral premotor area F5 of the macaque monkey

We investigated the motor and visual properties of F5 grasping neurons, using a controlled paradigm that allows the study of the neuronal discharge during both observation and grasping of many different three-dimensional objects with and without visual guidance. All neurons displayed a preference for grasping of an object or a set of objects. The same preference was maintained when grasping was performed in the dark without visual feedback. In addition to the motor-related discharge, about half of the neurons also responded to the presentation of an object or a set of objects, even when a grasping movement was not required. Often the object evoking the strongest activity during grasping also evoked optimal activity during its visual presentation. Hierarchical cluster analysis indicated that the selectivity of both the motor and the visual discharge of the F5 neurons is determined not by the object shape but by the grip posture used to grasp the object. Because the same paradigm has been used to study the properties of hand-grasping neurons in the dorsal premotor area F2, and in the anterior intraparietal area (AIP), a comparison of the functional properties of grasping-related neurons in the three cortical areas (F5, F2, AIP) is addressed for the first time.     

Responses of mirror neurons in area F5 to hand and tool grasping observation

Mirror neurons are a distinct class of neurons that discharge both during the execution of a motor act and during observation of the same or similar motor act performed by another individual. However, the extent to which mirror neurons coding a motor act with a specific goal (e.g., grasping) might also respond to the observation of a motor act having the same goal, but achieved with artificial effectors, is not yet established. In the present study, we addressed this issue by recording mirror neurons from the ventral premotor cortex (area F5) of two monkeys trained to grasp objects with pliers. Neuron activity was recorded during the observation and execution of grasping performed with the hand, with pliers and during observation of an experimenter spearing food with a stick. The results showed that virtually all neurons responding to the observation of hand grasping also responded to the observation of grasping with pliers and, many of them to the observation of spearing with a stick. However, the intensity and pattern of the response differed among conditions. Hand grasping observation determined the earliest and the strongest discharge, while pliers grasping and spearing observation triggered weaker responses at longer latencies. We conclude that F5 grasping mirror neurons respond to the observation of a family of stimuli leading to the same goal. However, the response pattern depends upon the similarity between the observed motor act and the one executed by the hand, the natural motor template.     

Facilitation of neuronal activity in somatosensory and posterior parietal cortex during prehension

In order to study prehension in a reproducible manner, we trained monkeys to perform a task in which rectangular, spherical, and cylindrical objects were grasped, lifted, held, and lowered in response to visual cues. The animal’s hand movements were monitored using digital video, together with simultaneously recorded spike trains of neurons in primary somatosensory cortex (S-I) and posterior parietal cortex (PPC). Statistically significant task-related modulation of activity occurred in 78% of neurons tested in the hand area; twice as many cells were facilitated during object acquisition as were depressed. Cortical neurons receiving inputs from tactile receptors in glabrous skin of the fingers and palm, hairy skin of the hand dorsum, or deep receptors in muscles and joints of the hand modulated their firing rates during prehension in consistent and reproducible patterns. Spike trains of individual neurons differed in duration and amplitude of firing, the particular hand behavior(s) monitored, and their sensitivity to the shape of the grasped object. Neurons were classified by statistical analysis into groups whose spike trains were tuned to single task stages, spanned two successive stages, or were multiaction. The classes were not uniformly distributed in specific cytoarchitectonic fields, nor among particular somatosensory modalities. Sequential deformation of parts of the hand as the task progressed was reflected in successive responses of different members of this population. The earliest activity occurred in PPC, where 28% of neurons increased firing prior to hand contact with objects; such neurons may participate in anticipatory motor control programs. Activity shifted rostrally to S-I as the hand contacted the object and manipulated it. The shape of the grasped object had the strongest influence on PPC cells. The results suggest that parietal neurons monitor hand actions during prehension, as well as the physical properties of the grasped object, by shifting activity between populations responsive to hand shaping, grasping, and manipulatory behaviors. Received: 1 October 1998 / Accepted: 4 May 1999     

Grasping with the left and right hand: a kinematic study

The goal of the present study was to compare prehension movements of the dominant and the non-dominant hand. Twenty right-handed volunteers (age 20–30 years) reached forward to grasp a cylindrical object, which was lifted and then placed into a target position in a retraction–insertion movement. The movements were performed at three different velocities (normal, deliberately fast, or slowly) both, under visual control, and in a no-vision condition. Analysis of the kinematic data revealed that the speed of hand transport influenced pre-shaping of both hands in a similar way. In the visual condition, the grip aperture increased about linearly with peak transport velocity, while it increased non-linearly with shorter movement duration. Comparison of the regression parameters showed that these relationships were nearly identical for both hands. The dominant hand was faster in inserting the object into the target position. Otherwise, no significant inter-manual differences were found. During prehension without visual control, the fingers opened more and movement duration was prolonged. Except for a larger grip aperture of the dominant hand at the end of the acceleration phase, the kinematic data of both hands were again comparable. This invariance was in contrast to performance in fine motor skills such as a pegboard test and drawing movements, where there was a clear advantage of the dominant hand. The similar pre-shaping of both hands during prehension is discussed with regard to a common motor representation of grasping.     

The effect of illusory size on force production when grasping objects

Milner and Goodale (1995) have proposed that visuomotor and perceptual processes are mediated by discrete visual systems that reflect the functional independence of action and perception. The visuomotor system is proposed to be insensitive to pictorial illusions of object size, whereas the perceptual system is reliably "tricked" by such figures. Brenner and Smeets (1996) and Jackson and Shaw (2000) demonstrated that grasp preshaping, but not grasping force, is immune to the Ponzo visual illusion, suggesting that not all visuomotor processes operate independently of the perceptual system. The present study investigated the effect of illusory object size on prehension kinematics and grasping dynamics (i.e., grip force and load force) as well as perceptual judgements of object size. Unlike previous investigations, object mass was held constant independent of changes in size. The Ponzo figure reliably affected perceptual estimates of object size, but this effect was restricted to one form of the illusion. Some aspects of the prehension movement were sensitive to veridical but not illusory object size (peak grip aperture, peak grip force, peak vertical wrist acceleration), whereas other movement parameters demonstrated illusory size effects (movement time, peak wrist velocity). Still other movement parameters were not sensitive to veridical or illusory object size (peak load force). Together the data suggest that certain prehension components are immune to pictorial illusions of object size, whereas others are not. Complex interactions between the perceptual and visuomotor systems appear to underlie the anticipatory scaling of grasping forces in prehension.     

Neurons related to goal-directed motor acts in inferior area 6 of the macaque monkey

Summary A new class of neurons was identified in the rostral part of inferior area 6 in the macaque monkey (Macaca nemestrina). These neurons fire in relation to motor acts which have a particular aim such as reaching, grasping or holding. The same neuron discharges when the animal uses the right hand, the left hand and, frequently, also the mouth. Furthermore most of these neurons specify how the aim can be achieved (e.g. precision grip vs whole hand prehension). Different types of goal-related neurons form a vocabulary of simple motor acts localized in inferior area 6.     

Cerebellar nuclear cell activity during antagonist cocontraction and reciprocal inhibition of forearm muscles

Monkeys were trained to exert a maintained isometric pinch with the thumb and forefinger. This task reliably elicited a simultaneous cocontraction of the forearm muscles. The same monkeys were also taught to insert the open hand into a manipulandum, flex and extend the wrist 35 and 15 degrees, respectively, and maintain an isometric wrist position against a mechanical stop for 1 s. This second task comprised two conditions: a dynamic or movement phase and a static or isometric phase. Movement always involved a wrist displacement of 50 degrees. Although some forearm muscles demonstrated bidirectional activity during the wrist displacement phase, all the wrist and finger muscles were alternatively active in isometric flexion or extension. Of the neurons in the dentate and interposed nuclei that consistently changed discharge during repeated isometric prehension, over 90% (61/67) of the neurons increased activity during this cocontraction of forearm muscles. About 70% (47/67) of these same nuclear cells discharged with a reciprocal pattern of firing during alternating wrist flexion-extension movements. Forty-six neurons had sustained and reciprocal discharge during the maintained isometric wrist postures. No differences were seen between the activity patterns of dentate and interposed cells with respect to either the prehension task or the reciprocal wrist-movement task. The discharge frequency of some dentate and interpositus neurons could be correlated with prehensile force as well as velocity of wrist movement and torque developed by wrist muscles. Correlation coefficients were calculated between nuclear cell discharge and the amplitude of the surface EMGs of the flexors and extensors of the wrist and fingers during the wrist-movement task. Sixteen nuclear cells showed low-order, but reliably positive, correlations with one of the two forearm muscle groups (mean r = 0.33). In contrast, a sample of seven Purkinje cells recorded during the same task demonstrated low-order correlations that were negative in sign (mean r = -0.30) between discharge frequency and one of the two forearm EMGs.     

The effects of delay on the kinematics of grasping

 We examined the effect on manual prehension of introducing a 5-s delay between viewing a target object and initiating a grasping movement. Subjects were tested in four conditions: three involved grasping the object and a fourth involved estimating its size. In the main experimental condition (Open Loop Delay), subjects viewed a target object for 300 ms, but did not initiate a grasping movement until an auditory signal was presented 5 s later. In this condition, subjects had to rely on stored visual information for guiding their grasp after the delay. In another condition (Open Loop), subjects initiated their grasping movement as soon as the target appeared. In both of these open-loop conditions, subjects reached out and grasped the object without seeing their hand. In the third grasping condition (Closed Loop), the target object and the hand were visible throughout the reach. In the three grasping conditions, subjects were instructed to pick up the object across its width using their index finger and thumb. In a final condition (Perceptual Estimation), subjects gave a manual estimate of the object’s width with their index finger and thumb after viewing the object for 300 ms. In all four conditions, subjects were presented with a target object in which the height, length and width were independently varied from trial to trial. The results of the experiment indicated that reaching and grasping movements made in the Open-Loop and Closed-Loop conditions did not differ in any kinematic measures. In contrast, when subjects performed in the Open-Loop Delay condition, their reaches took significantly longer and achieved peak velocity proportionately earlier. As well, their maximum grip aperture was significantly larger. In addition, reaching movements in all three grasping conditions were affected by both the object’s width (the ’relevant’ dimension) and height. The manual estimates in the Perceptual-Estimation condition, however, reflected only the object’s width. These results, together with evidence from other studies, suggest that motor actions performed after a delay use different transformations than those used for ’real-time’ grasping. We argue that the stored visual information used to drive delayed actions arises from a perceptual rather than a visuomotor analysis of the target object. Received: 10 July 1998 / Accepted: 4 December 1998     

Depression of neuronal firing rates in somatosensory and posterior parietal cortex during object acquisition in a prehension task

Prehension is an object-oriented behavior consisting of four components: reach, grasp, manipulation, and release. To determine how such actions are represented in primary somatosensory (S-I) and posterior parietal cortex (PPC), we used digital video to synchronize spike trains of neurons recorded in Brodmann's areas 3b, 1, 2, 5, and 7 with the hand kinematics as monkeys performed a prehension task. Statistical analyses indicated that one-third of task-modulated neurons showed significantly depressed firing rates during object acquisition and/or manipulation. This population was dominated by neurons innervated by deep receptors that sensed extension movements of the fingers, or by tactile receptors in hairy skin sensing stretch. Grasp-inhibited responses were the most common type. Tonic firing rates of these cells dropped significantly during approach as the hand was preshaped for grasping, or at contact when grasp was initiated, and persisted until hand motion ceased or as the grip relaxed. Maximum suppression of firing occurred at grasp completion. Their lack of specificity for particular hand behaviors formed the inhibitory counterpart of broadly tuned cells that fired prolonged bursts during grasp and manipulatory stages of prehension. The remainder of the task-inhibited population showed biphasic responses. Firing rates were significantly depressed during grasping and manipulation when the hand interacted directly with the object, but were enhanced prior to contact, when the hand was preshaped (approach-tuned), or upon relaxation of grasp and release of the object from the hand (loweror relax-tuned). Grasp-inhibited responses occurred primarily in S-I, whereas biphasic inhibitory activity was recorded mainly in PPC. Suppression of activity within these populations may thereby increase the saliency of excitatory responses to acquisition and manipulation of objects. Reduction of firing during prehension might also signal the flexed postures used to retain objects in the hand, rather than a generalized gating of sensory information. The similarity of responses to active and passive extension movements suggests that the inhibitory responses may provide important postural and motor information about the hand kinematics when performing skilled tasks.     

Simultaneous action execution and observation optimise grasping actions

Action observation and execution share overlapping neural resonating mechanisms. In the present study, we sought to examine the effect of the activation of this system during concurrent movement observation and execution in a prehension task, when no a priori information about the requirements of grasping action was available. Although it is known that simultaneous activation by observation and execution influences motor performance, the importance of the delays of these two events and the specific effect of movement observation itself (and not the prediction of the to-be-observed movement) on action performance are poorly known. Fine-grained kinematic analysis of both the transport and grasp components of the movement should provide knowledge about the influence of movement observation on the precision and the performance of the executed movement. The experiment involved two real participants who were asked to grasp a different side of a single object that was composed of a large and a small part. In the first experiment, we measured how the transport component and the grasp component were affected by movement observation. We tested whether this influence was greater if the observed movement occurred just before the onset of movement (200 ms) or well before the onset of movement (1 s). In a second experiment, to reproduce the previous experiment and to verify the specificity of the grasping movements, we also included a condition consisting of pointing towards the object. Both experiments showed two main results. A general facilitation of the transport component was found when observing a simultaneous action, independent of its congruency. Moreover, a specific facilitation of the grasp component was present during the observation of a congruent action when movement execution and observation were nearly synchronised. While the general facilitation may arise from a competition between the two participants as they reached for the object, the specific facilitation of the grasp component seems to be directly related to mirror neuron system activity induced by action observation itself. Moreover, the time course of the events appears to be an essential factor for this modulation, implying the transitory activation of the mirror neuron system.     

Multijoint grasping movements

Studies of human prehension have revealed characteristic patterns of grasping kinematics. We sought to gain insight into the determinants of those patterns by means of a computer simulation and accompanying behavioral experiment concerning multijoint, planar grasping behavior. The simulation was based on a recent theory of posture-based motion planning which hypothesizes that movement preparation entails time-limited, multiple task-constraint satisfaction. Prehension was modeled with a stick-figure animation involving 12 series of 81 grasping movements. Circular objects to be grasped were located at three angles (45 degrees, 90 degrees, and 135 degrees) and at three distances (20 cm, 30 cm, and 40 cm) relative to the initial location of the hand in the workplane. Additionally, three object sizes (2 cm, 4 cm, and 6 cm in diameter) and three initial aperture sizes (0.3 cm, 3.3 cm, and 7.0 cm) were used. Analyses of the simulated grasping movements focused on the time course of the hand opening, the tangential velocity of the wrist, and the rotations of the joints in the arm, hand, and fingers. The results showed that the model accurately mimicked detailed kinematics of prehension observed in earlier studies. With respect to the frequently reported relationship between object size and hand opening, the simulations further revealed an effect of initial aperture. This predicted effect was confirmed in an experiment in which four participants performed analogous planar grasping tasks. An analysis of the time course of the opening of the hand showed that maximum aperture covaried with initial aperture. A conclusion of this work is that a major determinant of grasping kinematics is avoidance of collisions with objects that are to be grasped.     

Selectivity for the shape, size, and orientation of objects for grasping in neurons of monkey parietal area AIP

In this study, we mainly investigated the visual selectivity of hand-manipulation-related neurons in the anterior intraparietal area (area AIP) while the animal was grasping or fixating on three-dimensional (3D) objects of different geometric shapes, sizes, and orientations. We studied the activity of 132 task-related neurons during the hand-manipulation tasks in the light and in the dark, as well as during object fixation. Seventy-seven percent (101/132) of the hand-manipulation-related neurons were visually responsive, showing either lesser activity during manipulation in the dark than during that in the light (visual-motor neurons) or no activation in the dark (visual-dominant neurons). Of these visually responsive neurons, more than half (n = 66) responded during the object-fixation task (object-type). Among these, 55 were tested for their shape selectivity during the object-fixation task, and many (n = 25) were highly selective, preferring one particular shape of the six different shapes presented (ring, cube, cylinder, cone, sphere, and square plate). For 28 moderately selective object-type neurons, we performed multidimensional scaling (MDS) to examine how the neurons encode the similarity of objects. The results suggest that some moderately selective neurons responded preferentially to common geometric features shared by similar objects (flat, round, elongated, etc.). Moderately selective nonobject-type visually responsive neurons, which did not respond during object fixation, were found by MDS to be more closely related to the handgrip than to the object shape. We found a similar selectivity for handgrip in motor-dominant neurons that did not show any visual response. With regard to the size of the objects, 16 of 26 object-type neurons tested were selective for both size and shape, whereas 9 object-type neurons were selective for shape but not for size. Seven of 12 nonobject-type and all (8/8) of the motor-dominant neurons examined were selective for size, and almost all of them were also selective for objects. Many hand-manipulation-related neurons that preferred the plate and/or ring were selective for the orientation of the objects (17/20). These results suggest that the visual responses of object-type neurons represent the shape, size, and/or orientation of 3D objects, whereas those of the nonobject-type neurons probably represent the shape of the handgrip, grip size, or hand-orientation. The activity of motor-dominant neurons was also, in part, likely to represent these parameters of hand movement. This suggests that the dorsal visual pathway is concerned with the aspect of form, orientation, and/or size perception that is relevant for the visual control of movements.     

Factors affecting higher-order movement planning: a kinematic analysis of human prehension

Summary Past studies of the kinematics of human prehension have shown that varying object size affects the maximum opening of the hand, while varying object distance affects the kinematic profile of the reaching limb. These data contributed to the formulation of a theory that the reaching and grasping components of human prehension reflect the output of two independent, though temporally coupled, motor programs (Jeannerod 1984). In the first experiment of the present study, subjects were required to reach out and grasp objects, with or without on-line, visual feedback. Object size and distance were covaried in a within-subjects design, and it was found that both grip formation and reach kinematics were affected by the manipulation of either variable. These data suggest that the control mechanisms underlying transport of the limb and grip formation are affected by similar task constraints. It was also observed that when visual feedback was unavailable after movement onset subjects showed an exaggerated opening of their hands, although grip size continued to be scaled for object size. The question remained as to whether the larger opening of the hand during no-feedback trials reflected the lack of opportunity to fine-tune the opening of the hand on-line, or the adoption of a strategy designed to increase tolerance for initial programming errors. To address this question, a second experiment was carried out in which we manipulated the predictability of visual feedback by presenting feedback and no-feedback trials in a random order. In contrast to the situation in which feedback and no-feedback trials were presented in separate blocks of trials (Exp. 1), in the randomly ordered series of trials presented in Exp. 2, subjects always behaved as if they were reaching without vision, even on trials where visual feedback was continuously available. These findings suggest that subjects adopt different strategies on the basis of the predictability of visual feedback, although there is nothing to suggest that this takes place at a conscious, or voluntary, level. The results of both experiments are consistent with the notion of a hierarchically-organized motor control center, responsible for optimizing performance under a variety of conditions through the coordination of different effector systems and the anticipation of operating constraints.     

The effect of a pictorial illusion on closed-loop and open-loop prehension

It has been proposed that movements to visible and remembered targets are sensitive to qualitatively different types of visual information. When the target is continuously visible, prehensile movements are thought to reflect veridical object size, whereas memory-dependent prehension is sensitive to the perceived size of the object. This hypothesis was explored by assessing the influence of illusory target width on prehension kinematics in three visual conditions: closed-loop (CL; full vision during the response), open-loop brief-delay (OL; visual occlusion coincident with the movement initiation cue) and open-loop 3-s delay (OL3; visual occlusion 3 s prior to movement initiation). To modulate illusory target width, objects were placed on backgrounds consisting of three forms of the Müller-Lyer (ML) figure. Peak grip aperture was sensitive to the ML figure in the OL and OL3, but not CL conditions, suggesting that perceptual information is used to modulate this grasping parameter when the movement is programmed and executed on the basis of visual memory. Peak-aperture velocity was affected by the ML illusion in all three visual conditions, suggesting that perceived object size might be important for modulating this aspect of prehension, independent of memory requirements. The different sensitivity of grip aperture and aperture velocity to illusory target width in the CL condition suggests that grasp preshaping might reflect multiple visuomotor processes. The results of this study are consistent with the tenets of the two-stream model of visual processing.     

Neurophysiology of prehension. II. Response diversity in primary somatosensory (S-I) and motor (M-I) cortices

Prehension responses of 76 neurons in primary somatosensory (S-I) and motor (M-I) cortices were analyzed in three macaques during performance of a grasp and lift task. Digital video recordings of hand kinematics synchronized to neuronal spike trains were compared with responses in posterior parietal areas 5 and AIP/7b (PPC) of the same monkeys during seven task stages: 1) approach, 2) contact, 3) grasp, 4) lift, 5) hold, 6) lower, and 7) relax. S-I and M-I firing patterns signaled particular hand actions, rather than overall task goals. S-I responses were more diverse than those in PPC, occurred later in time, and focused primarily on grasping. Sixty-three percent of S-I neurons fired at peak rates during contact and/or grasping. Lift, hold, and lowering excited fewer S-I cells. Only 8% of S-I cells fired at peak rates before contact, compared with 27% in PPC. M-I responses were also diverse, forming functional groups for hand preshaping, object acquisition, and grip force application. M-I activity began < or =500 ms before contact, coinciding with the earliest activity in PPC. Activation of specific muscle groups in the hand was paralleled by matching patterns of somatosensory feedback from S-I needed for efficient performance. These findings support hypotheses that predictive and planning components of prehension are represented in PPC and premotor cortex, whereas performance and feedback circuits dominate activity in M-I and S-I. Somatosensory feedback from the hand to S-I enables real-time adjustments of grasping by connections to M-I and updates future prehension plans through projections to PPC.     

A detailed analysis of the planning and execution of prehension movements by three adolescents with spastic hemiparesis due to cerebral palsy

Assuming that primary symptoms of motor disorders can best be distinguished from signs of adaptation through behavioral analyses on an individual basis, the present study provides a detailed analysis of the prehension movements of three adolescents with mild spastic hemiparesis of different etiology. We investigated the extent to which the hemiparetic participants took their movement limitations into account when planning and performing sequences of prehension movements. We examined three indices of flexibility in grip planning in conjunction with an analysis of arm-joint coordination patterns as the movements unfolded. Participants were asked to repeatedly grasp a square object of which the position was gradually changed leftwards or rightwards. In half the trials the goal of the task was to lift the object, in the other half it had to be rotated back-and-forth. Trunk, arm, and hand movements were recorded with two synchronized 3-D motion-tracking systems. The movements of the hemiparetic participants were compared with the average performance of 11 control participants of which the collective data were taken to represent a typical control participant. Whereas one hemiparetic participant (GV) maintained a single grasping pattern throughout the experiment, the other two (CV and LC) only partially persevered in previously adopted grasping patterns. The shoulder contributed more and the wrist contributed less to this perseverance. No effects of task goal were found on grip selection. However, two hemiparetic participants (CV and LC) did tune their hand displacement to the task that followed the grasps. Taken collectively, the results show that the hemiparetic participants took their limitations into account when performing movements, but not when planning movements.     

The role of proprioception in the control of prehension movements: a kinematic study in a peripherally deafferented patient and in normal subjects

In this study we investigated the role of proprioception in the control of prehension movements, with particular reference to the grasp component. Grasp and transport kinematics were studied in a peripherally deafferented patient and in five healthy subjects. Two experiments were carried out: the prehension experiment and the grasp perturbation experiment. In the prehension experiment both the patient and the control subjects were required to reach and grasp three objects of different size, located at three different distances, both with and without visual feedback. In the grasp perturbation experiment a mechanical perturbation was applied to the fingers during prehension movements, again executed with and without visual feedback. In the prehension experiment temporal parameters of the patient's movements were generally slowed, with greater variability on some measures. However, over the first phase of the movement the pattern of the patient's hand opening and transport acceleration, scaled to object size and distance, was the same as that of controls, both with and without visual feedback. On the contrary, during the final phase of the movement (the finger closure phase and deceleration) the patient's performance differed significantly from the controls. These phases were abnormally lengthened and frequent movement adjustments were observed. In the grasp perturbation experiment the patient was not able to compensate for the perturbations applied to the fingers, even with visual feedback. The data allowed us to investigate also the respective contribution of proprioception and of vision of the hand in the control of prehension. We compared prehension kinematics in two conditions: (a) with visual but no proprioceptive feedback (in the patient) and (b) with proprioceptive but no visual feedback (in the controls). In both experiments proprioceptive control was more efficient than visual control. The results of this study are interpreted in favour of the strict dependence of prehension control on proprioception. The first phase of the movement, however, can be appropriately planned and executed without the necessity of either proprioceptive or visual information about the hand.     

Primary motor cortical activity related to the weight and texture of grasped objects in the monkey.

1. Two monkeys were trained to grasp an object between the thumb and index finger and lift it to a vertical distance of 12-25 mm. Up to 12 different conditions defined by different combinations of object weights (15, 65, and 115 g) and four surface textures (oiled metal, smooth metal, fine and coarse sandpaper) were used. The apparatus was equipped to measure grip (prehensile) force, vertical (load) force, and object displacement. 2. The monkeys appropriately scaled the grip force for the weight and the coefficient of friction of the object. However, during the dynamic phase of the task (grasping and lifting), the monkeys increased the prehensile force in multiple steps, suggesting that they relied on sensory feedback from the fingers to attain an adequate grip force to lift the object rather than programming the lift in advance. 3. Single-unit activity of 248 neurons was recorded in the hand area of the primary motor cortex while the monkeys performed the task. Of 208 neurons tested for cutaneous and proprioceptive receptive fields (RFs), 96 were sensitive to cutaneous stimulation of the glabrous skin of the hand, whereas 82 received proprioceptive input from wrist and finger muscles. The concentration of neurons with cutaneous input was significantly greater in the rostral bank of the central sulcus compared with cells with proprioceptive RFs, which were more concentrated in the convexity of the precentral gyrus. 4. From the global sample, 199 cells were tested with the three object weights, and 128 of these with at least two surface textures were used in combination with the object weights. The discharge of 58/199 (29%) cells was modulated with the object weight. Cells with cutaneous (20/84, 24%) and proprioceptive (23/71, 32%) RFs were about equally responsive to the object weight. 5. A greater number of motor cortical neurons were influenced by surface texture than by object weight. Of 128 cells tested with at least two surface textures, 67 (52%) showed a modulation of their activity as a function of texture. A significantly greater proportion of neurons with cutaneous RFs (40/63, 63%) showed differential activity as a function of object texture than cells receiving proprioceptive input (21/47, 45%). 6. Weight- and texture-related neurons were not distributed equally in the rostrocaudal dimension of the motor cortex. Only 8% of texture-related cells were located in the convexity of the precentral gyrus, whereas 30% of weight-related neurons were recorded from this rostral zone.(ABSTRACT TRUNCATED AT 400 WORDS)     

Functional magnetic resonance adaptation reveals the involvement of the dorsomedial stream in hand orientation for grasping

Reach-to-grasp actions require coordination of different segments of the upper limbs. Previous studies have examined the neural substrates of arm transport and hand grip components of such actions; however, a third component has been largely neglected: the orientation of the wrist and hand appropriately for the object. Here we used functional magnetic resonance imaging adaptation (fMRA) to investigate human brain areas involved in processing hand orientation during grasping movements. Participants used the dominant right hand to grasp a rod with the four fingers opposing the thumb or to reach and touch the rod with the knuckles without visual feedback. In a control condition, participants passively viewed the rod. Trials in a slow event-related design consisted of two sequential stimuli in which the rod orientation changed (requiring a change in wrist posture while grasping but not reaching or looking) or remained the same. We found reduced activation, that is, adaptation, in superior parieto-occipital cortex (SPOC) when the object was repeatedly grasped with the same orientation. In contrast, there was no adaptation when reaching or looking at an object in the same orientation, suggesting that hand orientation, rather than object orientation, was the critical factor. These results agree with recent neurophysiological research showing that a parieto-occipital area of macaque (V6A) is modulated by hand orientation during reach-to-grasp movements. We suggest that the human dorsomedial stream, like that in the macaque, plays a key role in processing hand orientation in reach-to-grasp movements.     

Interference of grasping observation during prehension, a behavioural study

During the last 10 years a considerable number of neurophysiological and functional imaging studies have provided evidence that observation and execution of movements activate common representations. Furthermore, several behavioural studies suggest that action observation can influence the performance of movements. Recently it was shown that viewing incongruent movements interferes with the execution of non-object oriented sinusoidal arm movements (Kilner et al. in Curr Biol 13:522–525, 2003). In the current study, we investigated whether interference of action observation also occurs during goal-directed prehension movements. Participants were required to grasp cubes of different sizes while simultaneously observing an actor performing grasping or pointing movements. The actors’ movement could be directed at objects that were identical, or different in size to the cube grasped by the participant. The results showed that maximum grip aperture was affected by observation of grasping towards larger objects. No effect of object size was found during observation of pointing movements. These results suggest that observation of grasping movements can interfere with the on-line control of prehension movements and provides further evidence for overlapping networks for grasping observation and execution.