Marco Santello

THE CONTROL OF TIMING AND AMPLITUDE OF EMG ACTIVITY IN LANDING MOVEMENTS IN HUMANS

The control of self‐initiated falls from different heights was studied. The objective of the study was to investigate in a quantitative manner the modulation of EMG timing (i.e. onset from take‐off and duration from onset to touch‐down) and amplitude (before and after foot contact) as a function of fall height. The muscles studied were m. soleus and m. tibialis anterior. Kinematic (ankle joint angle) and kinetic (ground reaction force) variables were also measured. Six subjects took part in the experiments that consisted of ten landings from each of five heights (0.2, 0.4, 0.6, 0.8 and 1 m) onto a force platform. We found a consistent pattern of co‐contraction before and after touch‐down across the fall heights studied. In both muscles, the onset of pre‐landing EMG activity occurred at a longer latency following take‐off when landing from greater heights. The absolute EMG duration was affected to a lesser extent by increasing fall height. These findings suggest that the onset of muscle activity of the muscles studied prior to foot contact is timed relative to the expected time of foot contact. Pre‐ and post‐landing EMG amplitude tended to increase with height. Despite a doubling in the magnitude of ground reaction force, the amplitude of ankle joint rotation caused by the impact remained constant across heights. These findings suggest that the observed pattern of co‐contraction is responsible for increasing ankle joint stiffness as fall height is increased. The attainment of an appropriate level of EMG amplitude seems to be controlled by (a) timing muscle activation at a latency timed from the expected instant of foot contact and (b) varying the rate at which EMG builds up.

Modelling natural and artificial hands with synergies

We report on recent work in modelling the process of grasping and active touch by natural and artificial hands. Starting from observations made in human hands about the correlation of degrees of freedom in patterns of more frequent use (postural synergies), we consider the implications of a geometrical model accounting for such data, which is applicable to the pre-grasping phase occurring when shaping the hand before actual contact with the grasped object. To extend applicability of the synergy model to study force distribution in the actual grasp, we introduce a modified model including the mechanical compliance of the hands musculotendinous system. Numerical results obtained by this model indicate that the same principal synergies observed from pre-grasp postural data are also fundamental in achieving proper grasp force distribution. To illustrate the concept of synergies in the dual domain of haptic sensing, we provide a review of models of how the complexity and heterogeneity of sensory information from touch can be harnessed in simplified, tractable abstractions. These abstractions are amenable to fast processing to enable quick reflexes as well as elaboration of high-level percepts. Applications of the synergy model to the design and control of artificial hands and tactile sensors are illustrated.

Choice of Contact Points during Multidigit Grasping: Effect of Predictability of Object Center of Mass Location

It has been shown that when subjects can predict object properties [e.g., weight or center of mass (CM)], fingertip forces are appropriately scaled before the object is lifted, i.e., before somatosensory feedback can be processed. However, it is not known whether subjects, in addition to these anticipatory force mechanisms, exploit the ability to choose where digits can be placed to facilitate object manipulation. We addressed this question by asking subjects to reach and grasp an object whose CM was changed to the left, center, or right of the object in either a predictable or unpredictable manner. The only task requirement was to minimize object roll during lift. We hypothesized that subjects would modulate contact points but only when object CM location could be predicted. As expected, object roll was significantly smaller in the predictable condition. This experimental condition was also associated with statistically distinct spatial distributions of contact points as a function of object CM location but primarily when large torques had to be counteracted, i.e., for right and left CM locations. In contrast, when subjects could not anticipate CM location, a “default” distribution of contact points was used, this being statistically indistinguishable from that adopted for the center CM location in the predictable condition. We conclude that choice of contact points is integrated with anticipatory force control mechanisms to facilitate object manipulation. These results demonstrate that planning of digit placement is an important component of grasp control.

Neural bases of hand synergies

The human hand has so many degrees of freedom that it may seem impossible to control. A potential solution to this problem is “synergy control” which combines dimensionality reduction with great flexibility. With applicability to a wide range of tasks, this has become a very popular concept. In this review, we describe the evolution of the modern concept using studies of kinematic and force synergies in human hand control, neurophysiology of cortical and spinal neurons, and electromyographic (EMG) activity of hand muscles. We go beyond the often purely descriptive usage of synergy by reviewing the organization of the underlying neuronal circuitry in order to propose mechanistic explanations for various observed synergy phenomena. Finally, we propose a theoretical framework to reconcile important and still debated concepts such as the definitions of “fixed” vs. “flexible” synergies and mechanisms underlying the combination of synergies for hand control.

Anticipatory planning and control of grasp positions and forces for dexterous two-digit manipulation

Dexterous object manipulation requires anticipatory control of digit positions and forces. Despite extensive studies on sensorimotor learning of digit forces, how humans learn to coordinate digit positions and forces has never been addressed. Furthermore, the functional role of anticipatory modulation of digit placement to object properties remains to be investigated. We addressed these questions by asking human subjects (12 females, 12 males) to grasp and lift an inverted T-shaped object using precision grip at constrained or self-chosen locations. The task requirement was to minimize object roll during lift. When digit position was not constrained, subjects could have implemented many equally valid digit position-force coordination patterns. However, choice of digit placement might also have resulted in large trial-to-trial variability of digit position, hence challenging the extent to which the CNS could have relied on sensorimotor memories for anticipatory control of digit forces. We hypothesized that subjects would modulate digit placement for optimal force distribution and digit forces as a function of variable digit positions. All subjects learned to minimize object roll within the first three trials, and the unconstrained device was associated with significantly smaller grip forces but larger variability of digit positions. Importantly, however, digit load force modulation compensated for position variability, thus ensuring consistent object roll minimization on each trial. This indicates that subjects learned object manipulation by integrating sensorimotor memories with sensory feedback about digit positions. These results are discussed in the context of motor equivalence and sensorimotor integration of grasp kinematics and kinetics.

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 objects 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.

Visual and non-visual control of landing movements in humans

1 The role of vision in controlling leg muscle activation in landing from a drop was investigated. Subjects (n= 8) performed 10 drops from four heights (0.2, 0.4, 0.6 and 0.8 m) with and without vision. Drop height was maintained constant throughout each block of trials to allow adaptation. The aim of the study was to assess the extent to which proprioceptive and vestibular information could substitute for the lack of vision in adapting landing movements to different heights. 2 At the final stages of the movement, subjects experienced similar peak centre of body mass (CM) displacements and joint rotations, regardless of the availability of vision. This implies that subjects were able to adapt the control of landing to different heights. The amplitude and timing of electromyographic signals from the leg muscles scaled to drop height in a similar fashion with and without vision. 3 However, variables measured throughout the execution of the movement indicated important differences. Without vision, landings were characterised by 10 % larger ground reaction forces, 10 % smaller knee joint rotations, different time lags between peak joint rotations, and more variable ground reaction forces and times to peak CM displacement. 4 We conclude that non‐visual sensory information (a) could not fully compensate for the lack of continuous visual feedback and (b) this non‐visual information was used to reorganise the motor output. These results suggest that vision is important for the very accurate timing of muscle activity onset and the kinematics of landing.

Anticipatory Control of Grasping: Independence of Sensorimotor Memories for Kinematics and Kinetics

We have recently provided evidence for anticipatory grasp control mechanisms in the kinematic domain by showing that subjects modulate digit placement on an object based on its center of mass (CM) when it can be anticipated (Lukos et al., 2007). This behavior relied on sensorimotor memories about digit contact points and forces required for optimal manipulation. We found that accurate sensorimotor memories depended on the acquisition of implicit knowledge about object properties associated with repeated manipulations of the same object. Whereas implicit knowledge of object properties is essential for anticipatory grasp control, the extent to which subjects can use explicit knowledge to accurately scale digit forces in an anticipatory manner is controversial. Additionally, it is not known whether subjects are able to use explicit knowledge of object properties for anticipatory control of contact points. We addressed this question by asking subjects to grasp and lift an object while providing explicit knowledge of object CM location as visual or verbal cues. Contact point modulation and object roll, a measure of anticipatory force control, were assessed using blocked and random CM presentations. We found that explicit knowledge of object CM enabled subjects to modulate contact points. In contrast, subjects could not minimize object roll in the random condition to the same extent as in the blocked when provided with a verbal or visual cue. These findings point to a dissociation in the effect of explicit knowledge of object properties on grasp kinematics versus kinetics, thus suggesting independent anticipatory processes for grasping.

Compensatory Motor Control After Stroke: An Alternative Joint Strategy for Object-Dependent Shaping of Hand Posture

Efficient grasping requires planned and accurate coordination of finger movements to approximate the shape of an object before contact. In healthy subjects, hand shaping is known to occur early in reach under predominantly feedforward control. In patients with hemiparesis after stroke, execution of coordinated digit motion during grasping is impaired as a result of damage to the corticospinal tract. The question addressed here is whether patients with hemiparesis are able to compensate for their execution deficit with a qualitatively different grasp strategy that still allows them to differentiate hand posture to object shape. Subjects grasped a rectangular, concave, and convex object while wearing an instrumented glove. Reach-to-grasp was divided into three phases based on wrist kinematics: reach acceleration (reach onset to peak horizontal wrist velocity), reach deceleration (peak horizontal wrist velocity to reach offset), and grasp (reach offset to lift-off). Patients showed reduced finger abduction, proximal interphalangeal joint (PIP) flexion, and metacarpophalangeal joint (MCP) extension at object grasp across all three shapes compared with controls; however, they were able to partially differentiate hand posture for the convex and concave shapes using a compensatory strategy that involved increased MCP flexion rather than the PIP flexion seen in controls. Interestingly, shape-specific hand postures did not unfold initially during reach acceleration as seen in controls, but instead evolved later during reach deceleration, which suggests increased reliance on sensory feedback. These results indicate that kinematic analysis can identify and quantify within-limb compensatory motor control strategies after stroke. From a clinical perspective, quantitative study of compensation is important to better understand the process of recovery from brain injury. From a motor control perspective, compensation can be considered a model for how joint redundancy is exploited to accomplish the task goal through redistribution of work across effectors.

Force-Independent Distribution of Correlated Neural Inputs to Hand Muscles During Three-Digit Grasping

The ability to modulate digit forces during grasping relies on the coordination of multiple hand muscles. Because many muscles innervate each digit, the CNS can potentially choose from a large number of muscle coordination patterns to generate a given digit force. Studies of single-digit force production tasks have revealed that the electromyographic (EMG) activity scales uniformly across all muscles as a function of digit force. However, the extent to which this finding applies to the coordination of forces across multiple digits is unknown. We addressed this question by asking subjects (n = 8) to exert isometric forces using a three-digit grip (thumb, index, and middle fingers) that allowed for the quantification of hand muscle coordination within and across digits as a function of grasp force (5, 20, 40, 60, and 80% maximal voluntary force). We recorded EMG from 12 muscles (6 extrinsic and 6 intrinsic) of the three digits. Hand muscle coordination patterns were quantified in the amplitude and frequency domains (EMG-EMG coherence). EMG amplitude scaled uniformly across all hand muscles as a function of grasp force (muscle x force interaction: P = 0.997; cosines of angle between muscle activation pattern vector pairs: 0.897-0.997). Similarly, EMG-EMG coherence was not significantly affected by force (P = 0.324). However, coherence was stronger across extrinsic than that across intrinsic muscle pairs (P = 0.0039). These findings indicate that the distribution of neural drive to multiple hand muscles is force independent and may reflect the anatomical properties or functional roles of hand muscle groups.