The role of vision on hand preshaping during reach to grasp

During reaching to grasp objects with different shapes hand posture is molded gradually to the object's contours. The present study examined the extent to which the temporal evolution of hand posture depends on continuous visual feedback. We asked subjects to reach and grasp objects with different shapes under five vision conditions (VCs). Subjects wore liquid crystal spectacles that occluded vision at four different latencies from onset of the reach. As a control, full-vision trials (VC5) were interspersed among the blocked vision trials. Object shapes and all VCs were presented to the subjects in random order. Hand posture was measured by 15 sensors embedded in a glove. Linear regression analysis, discriminant analysis, and information theory were used to assess the effect of removing vision on the temporal evolution of hand shape. We found that reach duration increased when vision was occluded early in the reach. This was caused primarily by a slower approach of the hand toward the object near the end of the reach. However, vision condition did not have a significant effect on the covariation patterns of joint rotations, indicating that the gradual evolution of hand posture occurs in a similar fashion regardless of vision. Discriminant analysis further supported this interpretation, as the extent to which hand posture resembled object shape and the rate at which hand posture discrimination occurred throughout the movement were similar across vision conditions. These results extend previous observations on memory-guided reaches by showing that continuous visual feedback of the hand and/or object is not necessary to allow the hand to gradually conform to object contours.     

Distorting the visual size of the hand affects hand pre-shaping during grasping

Vision of the body is known to affect somatosensory perception (e.g. proprioception or tactile discrimination). However, it is unknown whether visual information about one’s own body size can influence bodily action. We tested this by measuring the maximum grip aperture (MGA) parameter of grasping while eight subjects viewed a real size, enlarged or shrunken image of their hand reaching to grasp a cylinder. In the enlarged view condition, the MGA decreased relative to real size view, as if the grasping movement was actually executed with a physically larger hand, thus requiring a smaller grip aperture to grasp the cylinder. Interestingly, MGA remained smaller even after visual feedback was removed. In contrast, no effect was found for the reduced view condition. This asymmetry may reflect the fact that enlargement of body parts is experienced more frequently than shrinkage, notably during normal growth. In conclusion, vision of the body can significantly and persistently affect the internal model of the body used for motor programming.     

Contribution of primate magnocellular red nucleus to timing of hand preshaping during reaching to grasp

Magnocellular red nucleus (RNm) is involved in controlling goal-directed limb movements such as reaching to grasp. We tested two hypotheses related to RNm's role in controlling reach-to-grasp movements. One hypothesis is that forelimb RNm neurons are grasp specific, and the other is that they specify the timing of metacarpi-phalangeal (MCP) extension to preshape the hand during the appropriate phase of the reach. We recorded single-unit discharge while monkeys performed two behavioral tasks that elicited similar reaches but differed in grasp. One task consisted of a reach with a precision grasp that elicited independent use of thumb and forefinger; the other included a whole-hand grasp that elicited concerted use of the four fingers. Most RNm neurons tested were engaged strongly during both the whole-hand and precision tasks, and the magnitude of discharge modulation did not differ between tasks. Thus most RNm neurons are not grasp specific but, instead, may contribute to behavioral features common to the two tasks. Two methods were used to investigate relations between single-unit discharge and kinematic data from the same individual trials of the whole-hand and precision tasks for a subset of forelimb RNm neurons. One method focused on correlations between parameters of RNm discharge and the duration, amplitude, and velocity of rotation of forelimb joints for each of the tasks. The second method compared between-task differences in times of peak neuronal discharge to between-task differences in times of rotations of forelimb joints. Parameters of reach-related RNm discharge were more frequently correlated with parameters of MCP extension than with parameters of rotation of wrist, elbow, and shoulder joints. Analyses of temporal relations between discharge and kinematic data during both the whole-hand and precision tasks indicate that discharge was time locked most frequently to MCP extension and, to a lesser extent, elbow extension during both tasks. We conclude that RNm may command muscle synergies that provide a basic preshape of the hand at the appropriate phase of limb transport. In addition, the timing of RNm's contribution to hand preshaping varies with the behavioral requirements of the task.     

Role of primate magnocellular red nucleus neurons in controlling hand preshaping during reaching to grasp

Reaching to grasp is of fundamental importance to primate motor behavior and requires coordinating hand preshaping with limb transport and grasping. We aimed to clarify the role of cerebellar output via the magnocellular red nucleus (RNm) to the control of reaching to grasp. Rubrospinal fibers originating from RNm constitute one pathway by which cerebellar output influences spinal circuitry directly. We recorded discharge from individual forelimb RNm neurons while monkeys performed a reach-to-grasp task and two tasks that were similar to the reach-to-grasp task in trajectory, amplitude, and direction but did not include a grasp. One of these, the device task, elicited reaches while holding a handle, and the other, the free-reach task, elicited reaches that did not require any specific hand use for task performance. The results demonstrate that coordinated whole-limb reaching movements are associated with large discharge modulations of RNm neurons predominantly when hand use is included. Therefore RNm neurons can at best only make a minor contribution to the control of reaching movements that lack hand use. We evaluated relations between the discharge of individual RNm neurons and electromyographic (EMG) activity of forelimb muscles during the reach-to-grasp task by comparing times of peak RNm discharge to times of peak EMG activity. The results are consistent with the view that RNm discharge may contribute to EMG activity of both distal and proximal muscles during reaching to grasp especially digit extensor and limb elevation muscles. Relations between the discharge of individual RNm neurons and movements of the metacarpi-phalangeal (MCP), wrist, elbow, and shoulder joints during individual trials of task performance were quantified by parametric correlation analyses on a subset of neurons studied during the reach-to-grasp and free-reach tasks. The results indicate that MCP extensions were consistently preceded by bursts of RNm discharge, and strong correlations were observed between parameters of discharge and the duration, velocity, and amplitude of corresponding MCP extensions. In contrast, relations between discharge and movements of proximal joints were poorly represented, and RNm discharge was not related to the speed of limb transport. Based on our data and those of others, we hypothesize that cerebellar output via RNm is specialized for controlling hand use and conclude that RNm may contribute to the control of hand preshaping during reaching to grasp by activating muscle synergies that produce the appropriate MCP extension at the appropriate phase of limb transport.     

Periodic modulation of motor-unit activity in extrinsic hand muscles during multidigit grasping

We recently examined the extent to which motor units of digit flexor muscles receive common input during multidigit grasping. This task elicited moderate to strong motor-unit synchrony (common input strength, CIS) across muscles (flexor digitorum profundus, FDP, and flexor pollicis longus, FPL) and across FDP muscle compartments, although the strength of this common input was not uniform across digit pairs. To further characterize the neural mechanisms underlying the control of multidigit grasping, we analyzed the relationship between firing of single motor units from these hand muscles in the frequency domain by computing coherence. We report three primary findings. First, in contrast to what has been reported in intrinsic hand muscles, motor units belonging to different muscles and muscle compartments of extrinsic digit flexors exhibited significant coherence in the 0- to 5- and 5- to 10-Hz frequency ranges and much weaker coherence in the higher 10-20 Hz range (maximum 0.0025 and 0.0008, respectively, pooled across all FDP compartment pairs). Second, the strength and incidence of coherence differed considerably across digit pairs. Third, contrary to what has been reported in the literature, across-muscle coherence can be stronger and more prevalent than within-muscle coherence, as FPL-FDP2 (thumb-index digit pair) exhibited the strongest and most prevalent coherence in our data (0.010 and 43% at 3 Hz, respectively). The heterogeneous organization of common input to these muscles and muscle compartments is discussed in relation to the functional role of individual digit pairs in the coordination of multiple digit forces in grasping.     

Effects of object shape and visual feedback on hand configuration during grasping

Normal subjects gradually preshape their hands during a grasping movement in order to conform the hand to the shape of a target object. The evolution of hand preshaping may depend on visual feedback about arm and hand position as well as on target shape and location at specific times during the movement. The present study manipulated object shape in order to produce differentiable patterns of finger placement along two orthogonal "dimensions" (flexion/extension and abduction/adduction), and manipulated the amount of available visual information during a grasp. Normal subjects were asked to reach to and grasp a set of objects presented in a randomized fashion at a fixed spatial location in three visual feedback conditions: Full Vision (both hand and target visible), Object Vision (only the object was visible but not the hand) and No Vision (vision of neither the hand nor the object during the movement). Flexion/extension angles of the metacarpophalangeal and proximal interphalangeal joints of the index, ring, middle and pinkie fingers as well as the abduction/adduction angles between the index-middle and middle-ring fingers were recorded. Kinematic analysis revealed that as visual feedback was reduced, movement duration increased and time to peak aperture of the hand decreased, in accord with previously reported studies. Analysis of the patterns of joint flexion/extension and abduction/adduction per object shape revealed that preshaping based on the abduction/adduction dimension occurred early during the reach for all visual feedback conditions (~45% of normalized movement time). This early preshaping across visual feedback conditions suggests the existence of mechanisms involved in the selection of basic hand configurations. Furthermore, while configuration changes in the flexion/extension dimension resulting in well-defined hand configurations occurred earlier during the movement in the Object Vision and No Vision conditions (45%), those in the Full Vision condition were observed only after 75% of the movement, as the moving hand entered the central region of the visual field. The data indicate that there are at least two control mechanisms at work during hand preshaping, an early predictive phase during which grip selection is attained regardless of availability of visual feedback and a late responsive phase during which subjects may use visual feedback to optimize their grasp.     

Influence of fatigue on hand muscle coordination and EMG-EMG coherence during three-digit grasping

Fingertip force control requires fine coordination of multiple hand muscles within and across the digits. While the modulation of neural drive to hand muscles as a function of force has been extensively studied, much less is known about the effects of fatigue on the coordination of simultaneously active hand muscles. We asked eight subjects to perform a fatiguing contraction by gripping a manipulandum with thumb, index, and middle fingers while matching an isometric target force (40% maximal voluntary force) for as long as possible. The coordination of 12 hand muscles was quantified as electromyographic (EMG) muscle activation pattern (MAP) vector and EMG-EMG coherence. We hypothesized that muscle fatigue would cause uniform changes in EMG amplitude across all muscles and an increase in EMG-EMG coherence in the higher frequency bands but with an invariant heterogeneous distribution across muscles. Muscle fatigue caused a 12.5% drop in the maximum voluntary contraction force (P < 0.05) at task failure and an increase in the SD of force (P < 0.01). Although EMG amplitude of all muscles increased during the fatiguing contraction (P < 0.001), the MAP vector orientation did not change, indicating that a similar muscle coordination pattern was used throughout the fatiguing contraction. Last, EMG-EMG coherence (0-35 Hz) was significantly greater at the end than at the beginning of the fatiguing contraction (P < 0.01) but was heterogeneously distributed across hand muscles. These findings suggest that similar mechanisms are involved for modulating and sustaining digit forces in nonfatiguing and fatiguing contractions, respectively.     

Analysis of hand forces in health and disease during maximum isometric grasping of cylinders

An analysis of force distribution in the hand during maximum isometric grasping actions is reported in a detailed and accurate manner. A microcomputer-controlled instrument which measures all 12 phalangeal forces of fingers simultaneously, in a single attempt at squeezing a cylindrical object, is described. The study involved 20 normal subjects of different weights and age groups grasping tubes of 50 mm, 75 mm, 90 mm and 110 mm diameters. Normal grasp forces decreased significantly with the increase in tube diameter, with the force being concentrated more on the distal segments of the fingers than on the proximal and middle segments. The mean percentage contributions of finger forces to total grip strength, from index to little fingers, were 31, 33, 22 and 24 per cent, respectively. The study was extended to cover leprotic and paralytic hands to assess their functional capabilities. In the case of leprosy subjects, the grip strength decreased with the severity of the disease and was only about 50 per cent of that of normal subjects. In hemiplegics, the grip strength was only about one-eighth of the normal values. The above assessment procedure provides baseline data which could sere as guidelines to a clinician in assessing the severity of the disease and observing the patient's recovery following the treatment. It would also be useful in the design of hand-operated controls and prosthetic arms.     

Planning and control of hand orientation in grasping movements

Humans grasp objects in a way that facilitates the intended use of the object. We examined how humans grasp a circular control knob in order to turn it in different directions and by different extents. To examine the processes involved in anticipatory planning of grasps, we manipulated advance information about the location of the control knob and the target of the knob-turn. The forearm orientation at the time of grasping depended strongly on the knob-turn, with the direction of the knob-turn having a stronger effect than the extent of the knob-turn. However, the variability of the forearm orientations after the knob-turn remained considerable. Anticipatory forearm orientations began early during the grasping movement. Advance information had no influence on the trajectory of the grasp but affected reaction times and the duration of the grasp. From the results, we conclude that (1) grasps are selected in anticipation of the upcoming knob rotation, (2) the desired hand location and forearm orientation at the time of grasping are specified before the onset of the grasp, and (3) an online programming strategy is used to schedule the preparation of the knob-turn during the execution of the grasp.     

An artificial grasping evaluation system for the paralysed hand

Neuromuscular electrical stimulation (NMES) has been used in upper limb rehabilitation towards restoring motor hand function. Quantitative evaluation of the artificially generated movement is necessary to achieve proper muscle activation. Custom-made gloves instrumented with force and position transducers were used to evaluate artificial quadriplegic grasping for a drinking activity. In spite of different sensor position, stimulation parameter dependence and lack of repeatability, grasp patterns achieved with the application of NMES follow the same patterns previously obtained with normal subjects, regarding force distribution among fingers and the shape of force curves. Larger forces were exerted by the thumb (average ranged from 2.8 to 4.5 N) following by index or long finger (average ranged from 1.8 to 3 N). The forces exerted ranged within the same interval as those previously measured and were sufficient to grasp an object of 10 N. Finger position achieved by interphalangeal joint status indicated the opening size of the hand throughout the range of movement. The instrumented gloves offer an alternative force and position feedback system for use in cylindrical grasp evaluation. The gloves can be used in a closed-loop control system, allowing on-line adjustment or in a clinical application to evaluate the results of a rehabilitation programme.     

Coordination of intrinsic and extrinsic hand muscle activity as a function of wrist joint angle during two-digit grasping

Fingertip forces result from the activation of muscles that cross the wrist and muscles whose origins and insertions reside within the hand (extrinsic and intrinsic hand muscles, respectively). Thus, tasks that involve changes in wrist angle affect the moment arm and length, hence the force-producing capabilities, of extrinsic muscles only. If a grasping task requires the exertion of constant fingertip forces, the Central Nervous System (CNS) may respond to changes in wrist angle by modulating the neural drive to extrinsic or intrinsic muscles only or by co-activating both sets of muscles. To distinguish between these scenarios, we recorded electromyographic (EMG) activity of intrinsic and extrinsic muscles of the thumb and index finger as a function of wrist angle during a two-digit object hold task. We hypothesized that changes in wrist angle would elicit EMG amplitude modulation of the extrinsic and intrinsic hand muscles. In one experimental condition we asked subjects to exert the same digit forces at each wrist angle, whereas in a second condition subjects could choose digit forces for holding the object. EMG activity was significantly modulated in both extrinsic and intrinsic muscles as a function of wrist angle (both p < 0.05) but only for the constant force condition. Furthermore, EMG modulation resulted from uniform scaling of EMG amplitude across all muscles. We conclude that the CNS controlled both extrinsic and intrinsic muscles as a muscle synergy. These findings are discussed within the theoretical frameworks of synergies and common neural input across motor nuclei of hand muscles.     

Prediction of object contact during grasping

The maximum grip aperture (MGA) during prehension is linearly related to the size of objects to be grasped and is adapted to the haptically sensed object size when there is a discrepancy between visual and haptic information. We have investigated what information is used to drive this adaptation process and how the onset of fingertip forces on the object is triggered. Subjects performed a reach-to-grasp task, where the object seen and the object grasped physically never were the same. We measured the movements of the index finger and the thumb and the contact forces between each fingertip and the object. The subjects’ adaptation of the MGA was unrelated both to different fingertip velocities at the moment of object contact, or the fingertip forces. Instead, the ‘timing’ of contact between the fingers and the object was most consistently influenced by introducing a size discrepancy. Specifically, if the object was larger than expected, the moment of contact occurred earlier, and if the object was decreased in size, then the contact occurred later. During adaptation, these timing differences were markedly reduced. Also, the motor command for applying forces on the object seemed to be released in anticipation of the predicted moment of contact. We therefore conclude that the CNS dynamically predicts when contact between the fingertips and objects occur and that aperture adaptation is primarily driven by timing prediction errors.     

Modulation of grasping forces during object transport

Subjects held an instrumented object in a tripod grasp and moved it in the horizontal plane in various directions. The contact forces at the digits were measured and the grip force was decomposed into 2 components: a manipulating force responsible for accelerating the object and a grasping force responsible for holding the object steady. The grasping forces increased during the movement, reaching a peak near the time of peak velocity. The grasping forces also exhibited directional tuning, but this tuning was idiosyncratic for each subject. Although the overall grip forces should be modulated with acceleration, the load force did not vary during the task. Therefore the increase in the grasping force is not required to prevent slip. Rather, it is suggested that grasping force increases during translational motion to stabilize the orientation of grasped objects.     

Trajectory of the index finger during grasping

The trajectory of the index finger during grasping movements was compared to the trajectories predicted by three optimization-based models. The three models consisted of minimizing the integral of the weighted squared joint derivatives along the path (inertia-like cost), minimizing torque change, and minimizing angular jerk. Of the three models, it was observed that the path of the fingertip and the joint trajectories, were best described by the minimum angular jerk model. This model, which does not take into account the dynamics of the finger, performed equally well when the inertia of the finger was altered by adding a 20 g weight to the medial phalange. Thus, for the finger, it appears that trajectories are planned based primarily on kinematic considerations at a joint level.     

Motor synergies for dampening hand vibration during human walking

This study investigated the motion required to carry a cup filled with water without spilling it, which is a common human dexterous task. This task requires the individual to dampen hand vibration while walking. We hypothesize that a reduction in hand jerk and a constant cup angle are required to achieve this task. We measured movements while human subjects carried a cup with water (WW task) and with stones (WS task) using a three-dimensional position measurement system and then analyzed joint coordination. We empirically confirmed that the value of hand jerk and the variance in cup angle in the WW task were smaller than those in the WS task. We used uncontrolled manifold (UCM) analysis to quantify joint coordination corresponding to the motor synergy required to reduce the hand jerk and variance of the cup angle. UCM components, which did not affect the hand jerk and cup angle, were larger than orthogonal components, which directly affected the hand jerk and cup angle in the WW task. These results suggest that there is a coordinated control mechanism that reduces hand jerk and maintains a constant cup angle when carrying a cup filled with water without spilling it. In addition, we suggest that humans adopt a flexible and coordinated control strategy of allowing variance independent of the variables that should be controlled to achieve this dexterous task.     

Gaze strategies during visually-guided versus memory-guided grasping

Vision plays a crucial role in guiding motor actions. But sometimes we cannot use vision and must rely on our memory to guide action—e.g. remembering where we placed our eyeglasses on the bedside table when reaching for them with the lights off. Recent studies show subjects look towards the index finger grasp position during visually-guided precision grasping. But, where do people look during memory-guided grasping? Here, we explored the gaze behaviour of subjects as they grasped a centrally placed symmetrical block under open- and closed-loop conditions. In Experiment 1, subjects performed grasps in either a visually-guided task or memory-guided task. The results show that during visually-guided grasping, gaze was first directed towards the index finger’s grasp point on the block, suggesting gaze targets future grasp points during the planning of the grasp. Gaze during memory-guided grasping was aimed closer to the blocks’ centre of mass from block presentation to the completion of the grasp. In Experiment 2, subjects performed an ‘immediate grasping’ task in which vision of the block was removed immediately at the onset of the reach. Similar to the visually-guided results from Experiment 1, gaze was primarily directed towards the index finger location. These results support the 2-stream theory of vision in that motor planning with visual feedback at the onset of the movement is driven primarily by real-time visuomotor computations of the dorsal stream, whereas grasping remembered objects without visual feedback is driven primarily by the perceptual memory representations mediated by the ventral stream.     

Contact points during multidigit grasping of geometric objects

We investigated the choice of contact points during multidigit grasping of different objects. In Experiment 1, cylinders were grasped and lifted. Participants were either instructed as to the number of fingers they should use, ranging from a two-finger grasp to a five-finger grasp, or could grasp with their preferred number of fingers. We found a strong relationship between the position of the fingertips on the object and the number of fingers used. In general, variability in the choice of contact points was low within- as well as between participants. The virtual finger, defined as the geometric mean position of fingers opposing the thumb, was in almost perfect opposition to the thumb, suggesting the formation of a functional unit using all contributing fingers in the grasp. In Experiment 2, four more complex shapes (rectangle, hexagon, pentagon, curved object) were grasped. Although we found some moderate between-participant variability in the choice of contact points, the within-participant variability was again remarkably low. In both experiments, participants showed a strong preference to use four or five fingers during grasping when left with free choice. Taken together, our findings suggest a preplanning of the grasping movement and that grasping results from a coordinated interplay between the fingers contributing to the grasp that cannot be understood as independent digit movements.     

The organization of digit contact timing during grasping

While the process of hand preshaping during grasping has been studied for over a decade, there is relatively little information regarding the organization of digit contact timing (DCT). This dearth of information may be due to the assumption that DCT while grasping exhibits few regularities or to the difficulty in obtaining information through traditional movement recording techniques. In this study, we employed a novel technique to determine the time of digit contacts with the target object at a high precision rate in normal healthy participants. Our results indicate that, under our task conditions, subjects tend to employ a radial to ulnar pattern of DCT which may be modulated by the shape of the target object. Moreover, a number of parameters, such as the total contact time, the frequency of first contacts by the thumb and index fingers and the number of simultaneous contacts, are affected by the relative complexity of the target object. Our data support the notion that a great deal of information about the object’s physical features is obtained during the early moments of the grasp.