Kelly J. Cole

Sensory-motor coordination during grasping and manipulative actions.

Goal-directed grasping and manipulation of objects are human skills that depend on automatic sensory control in which predictive feed-forward mechanisms integrate somatosensory and visual signals with sensory-motor memory systems. Memory representations of physical and task-relevant properties of the object play a pivotal role. Anticipatory strategies are crucial when purposeful actions arise from learned relationships between afferent patterns and efferent commands. The development of even elementary precision grip skills is a protracted process not concluded until early adolescence. Not surprisingly, the neural control of manual actions engages most central nervous system areas known to be involved in motor control.

Control of multimovement coordination: sensorimotor mechanisms in speech motor programming.

The present paper provides some hypotheses concerning the role of sensorimotor mechanisms in the coordination and programming of multimovement behaviors. The primary database is from experiments on the control of speech, a motor behavior that inherently requires multimovement coordination. From these data, it appears that coordination may be implemented by calibrated, sensorimotor actions which couple multiple movements for the accomplishment of common functional goals. The data from speech and select observations in other motor systems also reveal that these sensorimotor linkages are task-dependent and may underlie the intermovement motor equivalence that characterizes many natural motor behaviors. In this context, it is hypothesized also that motor learning may involve the calibration of these intermovement sensorimotor actions. These observations in turn provide some alternative perspectives on the concept of a motor program, primarily suggesting that individual movements and muscle contractions are not wholly prespecified, but shaped by sensorimotor adjustments.

Friction at the digit-object interface scales the sensorimotor transformation for grip responses to pulling loads

When restraining a mechanically “active” object (one that exerts unpredictable changes in loading forces) with a precision grip of the digits, we maintain a stable grasp by modulating our grip force using somatosensory information related to the loading forces. The response to ramp load increases consists of an initial fast rise in grip force (“catch-up”) followed by a secondary response that steadily increases the grip force in parallel with the load force (“tracking”). The sizes of these response components scale in proportion to the loading rate. However, maintaining a stable grasp without employing an exceedingly large grip force may require further scaling of this load to grip sensorimotor transformation based on two additional factors: (1) the friction at the digit-object interface and (2) the grip force present at the start of the load increase. The present experiments sought to determine whether such scaling occurs and to characterize its control. Subjects restrained a manipulandum held between the tips of the thumb and index finger. At unpredictable times a pulling force appeared, directed away from the subjects hand. Each pull had a trapezoidal load profile beginning and ending at 0 N with 4-N/s ramps; each ramp was 1 s in duration. The texture of the gripped surfaces varied among sandpaper, suede, and rayon, which represented increasingly slippery surfaces. The grip force at the start of the load ramp (intertrial grip force), and the amplitudes of the catch up and secondary grip responses scaled in proportion to the inverse friction. We interpret these results to indicate a uniform scaling of the transformations controlling the intertrial grip force, the catch up response, and the secondary response. Initial state information from tactile cues available upon object contact appeared to update the frictional scaling value. This conclusion is based on observations of immediate changes in the intertriai grip force upon contact with a new surface, and because differences in force-rate profiles appeared virtually by the onset of the catch-up response. Similarly, the intertriai grip force also constituted initial state information. The size of the catch-up and secondary grip force responses varied inversely with the size of the intertrial grip force. These scalings of the load to grip force sensorimotor transformation for friction and intertrial grip force level appear to be functionally adaptive, because they contribute to a stable grasp (prevent object slips) while avoiding exceedingly large safety margins.

COORDINATION OF INDEX FINGER MOVEMENTS

The purpose of this investigation was to describe the patterns of coordination among the joint motions of the index finger, and among the EMGs of index finger muscles. Index finger movements involving all three joints were varied in speed and direction. Joint motions were recorded along with fine-wire EMG from all the muscles that insert into the index finger. We observed nearly linear relationships for angular position between the two interphalangeal (IP) joints, and between the metacarpophalangeal (MP) and proximal IP (PIP) joints regardless of movement, speed and direction. The activities of the extrinsic flexors were of similar magnitude and were highly correlated when they acted as agonists but were poorly correlated when they acted as antagonists to the movement. Extrinsic extensor muscles behaved in this way also. The activation patterns of the intrinsic musculature correlated weakly except for extension movements voluntarily limited to the IP joints. We conclude that the highly coordinated action of the extrinsic flexors during flexion contribute importantly to the linked motions of the IP joints in part because these muscles span two or all the three index finger joints. Hence, interjoint movement patterns appear not to arise solely from restraints imposed by passive tissues, especially for fast flexion movements. The weakly correlated intrinsic muscle activity does not uncouple the flexion motions at the PIP and DIP joints because these muscles exert extensor torques at both IP joints. However, the actions of the intrinsic muscles are necessary for stabilizing the MP joint in flexion postures during IP motion and in producing motions voluntarily limited to the MP joint.

Tactile impairments cannot explain the effect of age on a grasp and lift task

Abstract This experiment addressed the often-posed theory that age-related declines in manual dexterity result from diminished tactile function. We measured the time ’young’ subjects (n=33; mean=45 years) and ’old’ subjects (n=33; mean=74 years) needed to grip (thumb and index finger), lift, and transport a small metal sphere when vision was permitted and when blindfolded. Subjects began each trial by reaching for the sphere and were instructed to complete the entire task quickly. In the absence of visual information, placement of the finger and thumb for a secure grip and lift cannot be performed efficiently without tactile information. If age-related tactile changes are functionally significant for this task, then without visual information the ’old’ group should show a disproportionate increase in the duration of the grip and lift phase of the task compared to the ’young’ group. Perceptual thresholds for tactile pressure stimuli (Semmes-Weinstein filaments) confirmed well-known age-related changes. Age and vision effects were manifest mainly during the grip-lift phase (time from object contact to lift-off from its support surface), with the expected finding that the ’old’ group required more time than ’young’ group, regardless of visual condition. The main finding was that the ’grip-lift’ duration in the ’no-vision’ condition was about twice the duration observed in the ’vision’ condition for both age groups (ratios of 2.1 and 2.3 for ’young’ and ’old’, respectively). This similar relative slowing for the two groups fails to support the hypothesis that old adults’ ability to grip and lift the object was limited by changes in the availability or use of tactile information.

Grip-force responses to unanticipated object loading: load direction reveals body- and gravity-referenced intrinsic task variables

Humans preserve grasp stability by automatically regulating the grip forces when loads are applied tangentially to the grip surfaces of a manipulandum held in a precision grip. The effects of the direction of the load force in relation to the palm, trunk, and gravity were investigated in blindfolded subjects. Controlled, tangential load-forces were delivered in an unpredictable manner to the grip surface in contact with the index finger either in the distal and proximal directions (away from and toward the palm) or in the ulnar and radial directions (transverse to the palm). The hand was oriented in: (1) a standard position, with the forearm extended horizontally and anteriorly in intermediate pronosupination; (2) an inverted position, reversing the direction of radial and ulnar loads in relation to gravity; and (3) a horizontally rotated position, in which distal loads were directed toward the trunk. The amplitude of the grip-force responses (perpendicular to the grip surface) varied with the direction of load in a manner reflecting frictional anisotropies at the digit-object interface; that is, the subjects automatically scaled the grip responses to provide similar safety margins against frictional slips. For all hand positions, the time from onset of load increase to start of the gripforce increase was shorter for distal loads, which tended to pull the object out of the hand, than for proximal loads. Furthermore, this latency was shorter for loads in the direction of gravity, regardless of hand position. Thus, shorter latencies were observed when frictional forces alone opposed the load, while longer latencies occurred when gravity also opposed the load or when the more proximal parts of the digits and palm were positioned in the path of the load. These latency effects were due to different processing delays in the central nervous system and may reflect the preparation of a default response in certain critical directions. The response to loads in other directions would incur delays required to implement a new frictional scaling and a different muscle activation pattern to counteract the load forces. We conclude that load direction, referenced to gravity and to the hands geometry, represents intrinsic task variables in the automatic processes that maintain a stable grasp on objects subjected to unpredictable load forces. In contrast, the grip-force safety margin against frictional slips did not vary systematically with respect to these task variables. Instead, the magnitude of the grip-force responses varied across load direction and hand orientation according to frictional differences providing similar safety margins supporting grasp stability.

The Stability of Precision Grip Force in Older Adults

The exceedingly large grip forces that many older adults employ when lifting objects with a precision pinch grip (Cole, 1991) may compensate for a reduced capability to produce a stable isometric force. That is, their grip force may fluctuate enough from moment to moment to yield grip forces that approach the force at which the object would slip from grasp. We examined the within-trial variability of isometric force in old (68-85 years, n = 13) and young (n = 11) human subjects (a) when they were asked to produce a constant pinch force at three target levels (0.49, 2.25, and 10.5 N) with external support of the arm, hand, and force transducer and (b) when they were asked to grasp, lift, and hold a small test object with a precision grip. Pinch force produced in the first task was equally stable across the two subject groups during analysis intervals that lasted 4 s. The elderly subjects produced grip forces when lifting objects that averaged twice as much as those produced by the young subjects. The force variability during the static (hold) phase of the lift for the old subjects was comparable with that used by the young subjects, after adjusting for the difference in grip force. The failure to observe less stable grip force in older adults contradicts a similar recent study. Differences in task (isometric grip force versus isometric abduction torque of a single digit) may account for this conflict, however. Thumb and finger forces for grip are produced through coactivation of many muscles and thus promote smooth force output through temporal summation of twitches. We conclude that peripheral reorganization of muscle in older adults does not yield increased instability of precision grip force and therefore does not contribute directly to increased grip forces in this population. However, force instability may affect other grip configurations (e.g., lateral pinch) or manipulation involving digit abduction or adduction forces.

Physiologic loading of the anterior cruciate ligament does not activate quadriceps or hamstrings in the anesthetized cat

We attempted to elicit quadriceps and hamstring elec tromyographic responses in seven chloralose-anesthe tized cats by loading the ACL with controlled anterior displacement of the tibia on the femur using rigid fixa tion and an MTS testing machine. We did not detect reflex activity in the quadriceps or hamstring muscles of any of the cats in response to anterior tibial displace ments of up to 4 mm, with rise times ranging from 1.0 to 0.1 seconds. In four of the cats we loaded the ACL using a wire loop. Loads of up to 125 N (4 to 5 times body weight) produced no reflex activity in any of the four animals, although we consistently observed mono synaptic reflex responses to tendon taps. Whole nerve recordings from the posterior articular nerve revealed substantial activity from afferents in response to tug ging on the ACL, although we could not differentiate receptors in the ACL from those in other periarticular tissues. Thus, while traction on the intact ACL causes signals in the afferent nerves, those signals are not translated into direct monosynaptic reflexes.

Handling objects in old age: forces and moments acting on the object

We measured the external moments and digit-tip force directions acting on a freely moveable object while it was grasped and manipulated by old (OA) and young (YA) adults. Participants performed a grasp and lift task and a precision orientation (key-slot) task with a precision (thumb-finger) grip. During the grasp-lift task the OA group misaligned their thumb and finger contacts and produced greater grip force, greater external moments on the object around its roll axis, and oriented force vectors differently compared with the YA group. During the key-slot task, the OA group was more variable in digit-tip force directions and performed the key-slot task more slowly. With practice the OA group aligned their digits, reduced their grip force, and minimized external moments on the object, clearly demonstrating that the nervous system monitored and actively manipulated one or more variables related to object tilt. This was true even for the grip-lift task, a task for which no instructions regarding object orientation were given and which could tolerate modest amounts of object tilt without interfering with task goals. Although the OA group performed the key-slot task faster with experience, they remained slower than the YA group. We conclude that with old age comes a reduced ability to control the forces and moments applied to objects during precision grasp and manipulation. This may contribute to the ubiquitous slowing and deteriorating manual dexterity in healthy aging.

Effects of transcranial direct current stimulation in combination with motor practice on dexterous grasping and manipulation in healthy older adults

Transcranial anodal stimulation (tDCS) over primary motor cortex (M1) improves dexterous manipulation in healthy older adults. However, the beneficial effects of anodal tDCS in combination with motor practice on natural and clinically relevant functional manual tasks, and the associated changes in the digit contact forces are not known. To this end, we studied the effects of 20 min of tDCS applied over M1 for the dominant hand combined with motor practice (MP) in a sham‐controlled crossover study. We monitored the forces applied to an object that healthy elderly individuals grasped and manipulated, and their performances on the Grooved Pegboard Test and the Key‐slot task. Practice improved performance on the Pegboard test, and anodal tDCS + MP improved retention of this performance gain when tested 35 min later, whereas similar performance gains degraded in the sham group after 35 min. Interestingly, grip force variability on an isometric precision grip task performed with visual feedback of precision force increased following anodal tDCS + MP, but not sham tDCS + MP. This finding suggests that anodal tDCS over M1 might alter the descending drive to spinal motor neurons involved in the performance of isometric precision grip task under visual feedback leading to increased fluctuations in the grip force exerted on the object. Our results demonstrate that anodal stimulation in combination with motor practice helps older adults to retain their improved performance on a functionally relevant manual task in healthy older adults.