Importance of cutaneous feedback in maintaining a secure grip during manipulation of hand-held objects

Previous research has shown that grip and load forces are modulated simultaneously during manipulation of a hand-held object. This close temporal coupling suggested that both forces are controlled by an internal model within the CNS that predicts the changes in tangential force on the fingers. The objective of the present study was to examine how the internal model would compensate for the loss of cutaneous sensation through local anesthesia of the index and thumb. Ten healthy adult subjects (5 men and 5 women aged 20-57 yr) were asked to grasp, lift, and hold stationary, a 250 g object for 20 s. Next, the subjects were asked to perform vertical oscillatory movements over a distance of 20 cm at a rate of 1.0 Hz for 30 s. Eleven trials were performed with intact sensation, and 11 trials after a local ring-block anesthesia of the index and thumb with bupivacain (5 mg/ml). During static holding, loss of cutaneous sensation produced a significant increase in the safety margin. However, the grip force declined significantly over the 20-s static hold period. During oscillatory arm movements, grip and load forces were continuously modulated together in a predictive manner as suggested by Flanagan and Wing. Again, the grip force declined over the 30-s movement, and 7/10 subjects dropped the object at least once. With intact sensation, the object was never dropped; but with the fingers anesthetized, it was dropped on 36% of the trials, and a significant slip occurred on a further 12%. The mean correlation between the grip and load forces for all subjects deteriorated from 0.71 with intact sensation to 0.48 after digital anesthesia. However, a cross-correlation calculated between the grip and load forces indicated that the phase lag was approximately zero both with and without digital anesthesia. Taken together, the data from the present study suggest that cutaneous afferents are required for setting and maintaining the background level of the grip force in addition to their phasic slip-detection function and their role in adapting the grip force/load force ratio to the friction on initial contact with an object. Finally, at a more theoretical level, they correct and maintain an internal model of the physical properties of hand-held objects.     

Digit cooling influences grasp efficiency during manipulative tasks

A commonly experienced effect of cold is a sensation of numbness and loss of sensibility in the fingers. Intact tactile sensibility of the grasping digits is essential for the efficient scaling of grip force level during the manipulation of hand-held objects. We investigated whether or not cooling of the grasping digits affects scaling of the grip force magnitude in relation to the loads resulting from continuous vertical arm movements performed with a grasped instrumented object. Maxima and minima of load force occurred at the lower and upper turning point of the movement cycle, respectively, and were accompanied by maximum and minimum peaks in grip force occurring close in time prior to and following digit cooling, respectively. Thus, digit cooling did not influence the ability to adjust the grip force profile in anticipation of movement-induced fluctuations in load force. However, subjects established significantly higher grip forces against the hand-held object following digit cooling and generated a 10–70% higher ratio between grip and load forces at the upper and lower turning points of the movement cycle. It is thought that the impaired economical scaling of grip force level is the result of reduced sensory feedback from the grasping fingers during digit cooling. The results provide further evidence to support the suggestion that cutaneous afferent input plays a subordinate role in the predictive temporal regulation of the grip force profile, but is used to adapt economically the force level to the actual loading requirements during dynamic object manipulation. Electronic Publication     

Control of fingertip forces in multidigit manipulation

Previous studies of control of fingertip forces in skilled manipulation have focused on tasks involving two digits, typically the thumb and index finger. Here we examine control of fingertip actions in a multidigit task in which subjects lifted an object using unimanual and bimanual grasps engaging the tips of the thumb and two fingers. The grasps resembled those used when lifting a cylindrical object from above; the two fingers were some 4.25 cm apart and the thumb was approximately 5.54 cm from either finger. The three-dimensional forces and torques applied by each digit and the digit contact positions were measured along with the position and orientation of the object. The vertical forces applied tangential to the grasp surfaces to lift the object were synchronized across the digits, and the contribution by each digit to the total vertical force reflected intrinsic object properties (geometric relationship between the object's center of mass and the grasped surfaces). Subjects often applied small torques tangential to the grasped surfaces even though the object could have been lifted without such torques. The normal forces generated by each digit increased in parallel with the local tangential load (force and torque), providing an adequate safety margin against slips at each digit. In the present task, the orientations of the force vectors applied by the separate digits were not fully constrained and therefore the motor controller had to choose from a number of possible solutions. Our findings suggest that subjects attempt to minimize (or at least reduce) fingertip forces while at the same time ensure that grasp stability is preserved. Subjects also avoid horizontal tangential forces, even at a small cost in total force. Moreover, there were subtle actions exerted by the digits that included changes in the distribution of vertical forces across digits and slight object tilt. It is not clear to what extent the brain explicitly controlled these actions, but they could serve, for instance, to keep tangential torques small and to compensate for variations in digit contact positions. In conclusion, we have shown that when lifting an object with a three-digit grip, the coordination of fingertip forces, in many respects, matches what has been documented previously for two-digit grasping. At the same time, our study reveals novel aspects of force control that emerge only in multidigit manipulative tasks.     

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.     

The effects of muscimol inactivation of small regions of motor and somatosensory cortex on independent finger movements and force control in the precision grip

This study investigated the effects of inactivating small regions of the primary somatosensory (SI) and motor (MI) cortex on the control of finger forces in a precision grip. A monkey was trained to grasp and lift a computer-controlled object between the thumb and index finger and to hold it stationary within a narrow position window for 2 s. The grip force applied perpendicular to the object surface, the lifting or load force applied tangentially in the vertical direction, and the vertical displacement were sampled at 100 Hz. Also, the ability of the monkey to extract small pieces of food from narrow wells of a Klüver board was analyzed from video-tape. Preliminary single-unit recordings and microstimulation studies were used to map the extent of the thumb and index-finger representation within SI and MI. Two local injections of 1 μl each (5 μg/μl) of the GABA_A-agonist muscimol were used to inactivate the thumb and index region of either the pre- or post-central gyrus. The precision grip was differently affected by muscimol injection into either SI or MI. MI injections produced a deficit in the monkey’s ability to perform independent finger movements and a general weakness in the finger muscles. Whole-hand grasping movements were inappropriately performed in an attempt to grasp either the instrumented object or morsels of food. Although the effect seemed strongest on intrinsic hand muscles, a clear deficit in digit extension was also noted. As a result, the monkey was unable to lift and maintain the object within the position window for the required 2 s, and, over time, the grip force decreased progressively until the animal stopped working. Following SI injections, the most obvious effect was a loss of finger coordination. In grasping, the placement of the fingers on the object was often abnormal and the monkey seemed unable to control the application of prehensile and lifting forces. However, the detailed analysis of forces revealed that a substantial increase in the grip force occurred well before any deficit in the coordination of finger movements was noted. This observation suggests that cutaneous feedback to SI is essential for the fine control of grip forces. Received: 05 October 1998 / Accepted: 30 March 1999     

Prehension synergies in three dimensions

The goal of this study was to investigate the conjoint changes of digit forces/moments in 3 dimensions during static prehension under external torques acting on the object in one plane. The experimental paradigm was similar to holding a book vertically in the air where the center of mass of the book is located farther from the hand than the points of digit contacts. Three force and 3 moment components from each digit were recorded during static prehension of a customized handle. Subjects produced forces and moments in all 3 directions, although the external torques were exerted on the handheld object about only the Z-axis. The 3-dimensional response to a 2-dimensional task was explained by the cause-effect chain effects prompted by the noncollinearity of the normal forces of the thumb and the 4 fingers (represented by the "virtual finger"). Because the forces are not collinear (not along the same line), they generate moments of force about X- and Y-axes that are negated by the finger forces along the Y- and X-directions. The magnitudes of forces produced by lateral fingers (index and little) with longer moment arms were larger compared with the central fingers (middle and ring). At the virtual finger (an imaginary digit whose mechanical action is equivalent to the summed action of the 4 fingers) level, the relative contribution of different fractions of the resistive moment produced by subjects did not depend on the torque magnitude. We conclude that the CNS 1) solves a planar prehension task by producing forces and moments in all 3 directions, 2) uses mechanical advantage of fingers, and 3) shares the total torque among finger forces and moments in a particular way disregarding the torque magnitude.     

Two virtual fingers in the control of the tripod grasp

To investigate the organization of multi-fingered grasping, we asked subjects to grasp an object using three digits: the thumb, the index finger, and the middle or ring finger. The object had three coarse flat contact surfaces, whose locations and orientations were varied systematically. Subjects were asked to grasp and lift the object and then to hold it statically. We analyzed the grasp forces in the horizontal plane that were recorded during the static hold period. Static equilibrium requires that the forces exerted by the three digits intersect at a common point, the force focus. The directions of the forces exerted by the two fingers opposing the thumb depended on the orientation of the contact surfaces of both fingers but not on the orientation of the contact surface of the thumb. The direction of the thumb's force did not depend on the orientation of the contact surfaces of the two fingers and depended only weakly on the orientation of the thumb's contact surface. In general, the thumb's force was directed to a point midway between the two fingers. The results are consistent with a hierarchical model of the control of a tripod grasp. At the first level, an opposition space is created between the thumb and a virtual finger located approximately midway between the two actual fingers. The directions of the forces exerted by the two fingers are constrained to be mirror symmetric about the opposition axis. The actual directions of finger force are elaborated at the next level on the basis of stability considerations.     

Finger control in the tripod grasp

The present study aimed to determine whether grasping is based on either (1) synchronous finger movements producing stereotyped types of grasp, or (2) independently controlled finger movements producing variable final finger postures. Participants reached for and grasped sphere-shaped objects of three sizes. They were allowed to select three different grasp configurations: a "pinch" grip (thumb and index finger), a "middle" grip (thumb and middle finger) and a "tripod" grip (thumb and index plus middle finger). Object distance from the subject was varied in order to verify whether finger control and final finger postures varied according to the degree of accuracy required by target object distance. All the participants always selected the tripod grip when reaching for the large and medium size objects. The pinch grip was used by half of the participants when reaching for the small object, but only in 17% of the trials. Target object distance did not appear to influence the type of selected grip. The tripod grip was found to consist of two different components: an aperture component (opening and closing the gap between the thumb and opposition finger) and a finger separation component (increasing and decreasing the gap between the index and middle fingers). The timing of the aperture component was the same for the index and middle fingers. In contrast, the timing of the finger separation was weakly coupled with the aperture components. Moreover, the relative spatial position among the three fingers during and at the end of grasp varied according to object size. When grasping the large object, both the index finger and the middle finger were in opposition to the thumb. In contrast, when grasping the small object, the index finger was less in opposition to the thumb with respect to the middle finger. The final spatial position of the thumb relative to the starting position was independent of object size, whereas those of the index and middle fingers varied with object size. The results support the notion that grasp is accomplished by using two virtual fingers formed by the thumb and one or more other fingers that synchronously open and close on the object along the opposition space [Arbib 1990; in: Jeannerod M (ed) Attention and performance XIII: motor representation and control. Lawrence Erlbaum, Hillsdale, pp 111–138]. This suggests a degree of coupling between the control of the virtual fingers. Electronic Publication     

Distributing vertical forces between the digits during gripping and lifting: the effects of rotating the hand versus rotating the object

During pinch grip we partition the vertical tangential forces at the digits according to the friction at the grip surfaces, and the mass distribution of the object. However, we cannot predictively partition the vertical forces to adjust to new frictional conditions after viewing a 180-deg rotation of an object with different textures at each grip surface. Hence, the processes that lead to predictive force partitioning may not access object representations, thereby suggesting that these processes are digit-specific. If this is true, then we should fail to predictively partition our fingertip forces when we rotate our hand. We tested this prediction by comparing the effects of object rotation with hand rotation for repeated lifts of an object that had one slippery grip surface and one rough grip surface. Subjects did not predictively redistribute the vertical tangential forces upon grasping the rotated object. Following object rotation, the vertical tangential force trajectories during the first 100 ms after contact indicated that 12/15 subjects failed to anticipate the reversed digit-friction relationships. All subjects appropriately partitioned the vertical tangential forces between the digits by the second lift after object rotation, confirming previous reports that sensory signals update the memory associated with lifting the object. In contrast, after hand rotation, 13/15 subjects anticipated the new digit-friction relationships and upon grasping the object immediately generated a steep rise in the vertical force trajectory at the rough surface. They also delayed the initial rise in vertical tangential force at the digit encountering the low-friction surface by approximately 65 ms. Thus, anticipatory partitioning of vertical fingertip forces is not strictly digit-specific. Internally driven motor plans can access the relevant memories or internal models for predictively partitioning the vertical tangential forces. It is not clear if this process involves rotating internal representations of fingertip force directly, or if the forces are derived after internally rotating a representation of the object. In contrast to the robust effects of vision on reach kinematics, or on wrist and finger configuration, visual signals about object rotation and orientation apparently do not influence vertical tangential fingertip forces.     

Independent movements of the digits in grasping

Reaching out for an object is often considered to consist of the control of two components: transporting the hand to the object's position, and scaling the grip to the object's size. We recently proposed an alternative view. According to this view, grasping consists of controlling the digits, not the hypothetical transport and grip. This alternative view assumes that the opening of the hand emerges from the trajectories of the digits. We therefore studied the movements of the digits in grasping. We asked subjects to grasp disks (diameters ranging from 5 to 8 cm) at marked positions with two digits. The positions were at opposite sides of the disk, at the same distance from the starting position, so that the orientation of the surface was the same for both digits. The subjects grasped the disks either with the index finger and thumb of the dominant hand, with the same digits of the non-dominant hand, or bimanually with both index fingers. Our predictions are: that the well-known relation between object size and grip aperture holds for each digit; that the same relation holds if the object is grasped with two hands instead of with the thumb and finger of one hand; that maximum deviation, variability and duration of the digit movements are related; and that variations in the timing of the maximum deviation of one digit are independent of those in the other digit. In accordance with our predictions, we found that the maximum deviation of both digits increased with 0.75 times the object radius, independent of the hand(s) used. The movements of the thumb were more variable than those of the index finger, which was reflected by a larger deviation earlier in the movement. The timing of the maximum deviation of the two digits was independent. These results on the digits' movements are consistent with our view that grasping can be understood as the largely independent movements of the digits. The results are not in conflict with the hypothesis that the grip is controlled during grasping, but can only be explained by extending that hypothesis post hoc. Electronic Publication     

Effects of grasping force magnitude on the coordination of digit forces in multi-finger prehension

The study addresses three main questions: (1) Does the magnitude of the grasping force affect the prehension synergies, i.e., conjoint changes of finger forces and moments? (2) Do individual finger forces scale with the total grasping forces (‘scale-invariance hypothesis’)? (3) How specification of the grasping force magnitude affects the inverse optimization of digit forces. Subjects (n = 7) grasped with minimal force an instrumented handle and maintained it at rest in the air. Then, the subjects doubled the initial grasping force. The forces and moments exerted by individual digits were recorded with six-component sensors. External torques that the subjects should resist (9 in total) varied among the trials from 0 to 0.46 Nm both in clockwise and counterclockwise directions. After the force doubling, the moments of the normal forces (M ⁿ) increased in the pronation effort tasks (PR-tasks) and decreased in the supination effort tasks (SU-tasks). The changes in the moments of the tangential forces (M ^t) were opposite to the M ⁿ changes; the moments increased in the SU-tasks and decreased in the PR-tasks. The opposite effects of force doubling on the M ^ts in the SU-tasks and PR-tasks were a consequence of the unidirectional changes of the thumb tangential forces: in all the tasks the contribution of the thumb tangential force to the total tangential force increased after the grasping force doubling (and the total contribution of the four fingers decreased). The decrease of the virtual finger (VF) tangential force was mainly due to the decrease of the index finger force (VF is an imagined finger that exerts the same force and moment as all the fingers together). In the non-zero torque tasks the individual finger forces did not scale proportionally with the grasping force, the sharing percentage of the individual finger forces in the VF normal force changed with the grasping force increase. The root mean square differences between the actual finger sharing percentages in the VF force and the sharing percentages predicted from optimization procedures in which different cost functions were used were in all cases smaller after the doubling than before the doubling. Hence the answers to the three questions formulated above are: (1) the alteration of the grasping force magnitude induces complex coordinated changes of all digit forces and moments; (2) the scale invariance hypothesis is confirmed only for the zero-torque tasks and rejected for the non-zero tasks, and (3) the specification of the grasping force magnitude at the level of twice the initial grasping force—which essentially restricts the control task to the object tilt prevention—improves the accuracy of the employed optimization procedures.     

Threshold position control of arm movement with anticipatory increase in grip force

The grip force holding an object between fingers usually increases before or simultaneously with arm movement thus preventing the object from sliding. We experimentally analyzed and simulated this anticipatory behavior based on the following notions. (1) To move the arm to a new position, the nervous system shifts the threshold position at which arm muscles begin to be recruited. Deviated from their activation thresholds, arm muscles generate activity and forces that tend to minimize this deviation by bringing the arm to a new position. (2) To produce a grip force, with or without arm motion, the nervous system changes the threshold configuration of the hand. This process defines a threshold (referent) aperture (R_a) of appropriate fingers. The actual aperture (Q_a) is constrained by the size of the object held between the fingers whereas, in referent position R_a, the fingers virtually penetrate the object. Deviated by the object from their thresholds of activation, hand muscles generate activity and grip forces in proportion to the gap between the Q_a and R_a. Thus, grip force emerges since the object prevents the fingers from reaching the referent position. (3) From previous experiences, the system knows that objects tend to slide off the fingers when arm movements are made and, to prevent sliding, it starts narrowing the referent aperture simultaneously with or somewhat before the onset of changes in the referent arm position. (4) The interaction between the fingers and the object is accomplished via the elastic pads on the tips of fingers. The pads are compressed not only due to the grip force but also due to the tangential inertial force (“load”) acting from the object on the pads along the arm trajectory. Compressed by the load force, the pads move back and forth in the gap between the finger bones and object, thus inevitably changing the normal component of the grip force, in synchrony with and in proportion to the load force. Based on these notions, we simulated experimental elbow movements and grip forces when subjects rapidly changed the elbow angle while holding an object between the index finger and the thumb. It is concluded that the anticipatory increase in the grip force with or without correlation with the tangential load during arm motion can be explained in neurophysiological and biomechanical terms without relying on programming of grip force based on an internal model.     

Effects of visual uncertainty on grasping movements

To successfully lift an object, a person’s fingers must be moved to locations where forces can be applied that are sufficient for maintaining contact and that allow for easy object manipulation. Obtaining such finger positions becomes more difficult when there is perceptual uncertainty about the location of the hand and object. However, knowledge about the amount of uncertainty could be incorporated into grasp plans to mitigate its effect. For example, during peripheral viewing the fingers could open wider to avoid colliding with or missing the object. The goal of this study is to determine the degree to which people incorporate their understanding of visual uncertainty when making a precision grasp. To investigate, subjects reached to a spatially fixed object whose retinal location was varied by fixating points 0–80° to the left of the object. This manipulation controlled the visual uncertainty of the hand and target without affecting the kinematic demands of the task. We found that people systematically changed their grasping behavior as a function of the amount of visual uncertainty in the task. Specifically, subjects’ maximum grip aperture increased linearly with target eccentricity. Moreover, the effect of visual uncertainty on finger trajectories could be captured by a single dimension of change along an axis. Together, these findings suggest that the sensorimotor system estimates visual uncertainty and behaviorally adjusts for it during grasping movements.     

Internal forces during object manipulation

Internal force is a set of contact forces that does not disturb object equilibrium. The elements of the internal force vector cancel each other and, hence, do not contribute to the resultant (manipulation) force acting on the object. The mathematical independence of the internal and manipulation forces allows for their independent (decoupled) control realized in robotic manipulators. To examine whether in humans internal force is coupled with the manipulation force and what grasping strategy the performers utilize, the subjects (n=6) were instructed to make cyclic arm movements with a customized handle. Six combinations of handle orientation and movement direction were tested. These involved: parallel manipulations (1) VV task (vertical orientation and vertical movement) and (2) HH task (horizontal orientation and horizontal movement); orthogonal manipulations (3) VH task (vertical orientation and horizontal movement) and (4) HV task (horizontal orientation and vertical movement); and diagonal manipulations (5) DV task (diagonal orientation and vertical movement) and (6) DH task (diagonal orientation and horizontal movement). Handle weight (from 3.8 to 13.8 N), and movement frequency (from 1 to 3 Hz) were systematically changed. The analysis was performed at the thumb-virtual finger level (VF, an imaginary finger that produces a wrench equal to the sum of wrenches produced by all the fingers). At this level, the forces of interest could be reduced to the internal force and internal moment. During the parallel manipulations, the internal (grip) force was coupled with the manipulation force (producing object acceleration) and the thumb-VF forces increased or decreased in phase: the thumb and VF worked in synchrony to grasp the object more strongly or more weakly. During the orthogonal manipulations, the thumb-VF forces changed out of phase: the plots of the internal force vs. object acceleration resembled an inverted letter V. The HV task was the only task where the relative phase (coupling) between the normal forces of the thumb and VF depended on oscillation frequency. During the diagonal manipulations, the coupling was different in the DV and DH tasks. A novel observation of substantial internal moments is described: the moments produced by the normal finger forces were counterbalanced by the moments produced by the tangential forces such that the resultant moments were close to zero. Implications of the findings for the notion of grasping synergies are discussed.     

Task-dependent modulation of multi-digit force coordination patterns

When grasping and holding an object with five digits under a variety of task constraints, subjects use well-defined force coordination patterns, i.e., consistent force covariations and in-phase synchronization among all digit pairs. The question arises as to whether these force coordination patterns are default mechanisms for controlling multi-digit force production or whether they are specific to lifting and holding an object. To address this question, we asked subjects to grasp a manipulandum and exert forces with five digits simultaneously so as to match a force template measured from an actual object grasp, lift, and hold task (GLH). Unlike GLH, the force production task (FP) lacked the constraint of having to maintain object stability against gravity. The amplitude of individual finger forces and force covariations were similar for both tasks (with the exception of the little finger, which tended to produce less force in FP). Nonetheless, when multiple grip forces were not required to hold the manipulandum against gravity (FP), there was a significantly lower tendency for forces to be synchronized with higher intertrial variability of phase differences between forces exerted by all digit-pairs. Furthermore, the tendency for force phase differences to cluster at 0 degrees was lower for FP than GLH. These results suggest that some aspects of the control of multi-digit grasping, i.e., force synchronization, are specific to object lift and hold rather than to the production of multi-digit forces. Modeling work suggests that motor unit synchronization might play an important role in the modulation of force synchronization patterns.     

Thumb and finger forces produced by motor units in the long flexor of the human thumb

The uncommonly good proprioceptive performance of the long flexor of the thumb, flexor pollicis longus (FPL), may add significantly to human manual dexterity. We investigated the forces produced by FPL single motor units during a weak static grip involving all digits by spike-triggered averaging from single motor units, and by averaging from twitches produced by intramuscular stimulation. Nine adult subjects were studied. The forces produced at each digit were used to assess how forces produced in FPL are distributed to the fingers. Most FPL motor units produced very low forces on the thumb and were positively correlated with the muscle force at recruitment. Activity in FPL motor units commonly loaded the index finger (42/55 units), but less commonly the other fingers ( P < 0.001). On average, these motor units produced small but significant loading forces on the index finger (∼5.3% of their force on the thumb) with the same time-to-peak force as the thumb (∼50 ms), but had no significant effect on other fingers. However, intramuscular stimulation within FPL did not produce significant forces in any finger. Coherence at 2–10 Hz between the thumb and index finger force was twice that for the other finger forces and the coherence to the non-index fingers was not altered when the index finger did not participate in the grasp. These results indicate that, within the long-term coordinated forces of all digits during grasping, FPL motor units generate forces highly focused on the thumb with minimal peripheral transfer to the fingers and that there is a small but inflexible neural coupling to the flexors of the index finger.     

Selective recruitment of single motor units in human flexor digitorum superficialis muscle during flexion of individual fingers

Flexor digitorum superficialis (FDS) is an extrinsic multi-tendoned muscle which flexes the proximal interphalangeal joints of the four fingers. It comprises four digital components, each with a tendon that inserts onto its corresponding finger. To determine the degree to which these digital components can be selectively recruited by volition, we recorded the activity of a single motor unit in one component via an intramuscular electrode while the subject isometrically flexed each of the remaining fingers, one at a time. The finger on which the unit principally acted was defined as the 'test finger' and that which flexed isometrically was the 'active' finger. Activity in 79 units was recorded. Isometric finger flexion forces of 50% maximum voluntary contraction (MVC) activated less than 50% of single units in components of FDS acting on fingers that were not voluntarily flexed. With two exceptions, the median recruitment threshold for all active–test finger combinations involving the index, middle, ring and little finger test units was between 49 and 60% MVC (60% MVC being the value assigned to those not recruited). The exceptions were flexion of the little finger while recording from ring finger units (median: 40% MVC), and vice versa (median: 2% MVC). For all active–test finger combinations, only 35/181 units were activated when the active finger flexed at less than 20% MVC, and the fingers were adjacent for 28 of these. Functionally, to recruit FDS units during grasping and lifting, relatively heavy objects were required, although systematic variation occurred with the width of the object. In conclusion, FDS components can be selectively activated by volition and this may be especially important for grasping at high forces with one or more fingers.     

Digit force adjustments during finger addition/removal in multi-digit prehension

We explored adjustments in multi-digit coordinated action on a hand-held object with finger addition and removal. The subjects (n = 7) kept a vertically oriented handle at rest using a prismatic grasp as if holding a glass of liquid and then either added one finger to the grasp, the index (I) or little (L) finger, or removed one finger. Three external torques were applied on the apparatus: clockwise, counterclockwise, and no torque. The individual digit forces and moments were recorded with six-component sensors. The change in grasping force, normal force of the thumb and virtual finger (VF, an imagined finger that generates the same mechanical effect as all fingers together), depended on the function of the manipulated finger, i.e. on whether the finger resisted external torque (torque agonist) or assisted it (torque antagonist). There was a significant increase of the grasping force when an antagonist was added or when an agonist was removed. These force increases were not necessary for slipping prevention: the normal forces prior to the manipulation were large enough to prevent slipping. All other finger manipulations exhibited no significant change in the grip force, except for the antagonist removal during the supination efforts (after removing the I finger the grasping force decreased). In contrast, the changes in the tangential force of the thumb depended on the manipulated finger, not on the finger function with respect to external torque. There was a significant thumb tangential force increase when the I finger was added or when the L finger was removed; opposite changes were seen when the L finger was added or the I finger was removed. The changes of the virtual finger (VF) tangential force were equal and opposite to the thumb tangential force alterations; these opposite changes caused changes in the moments, these forces generated. The changes in the moments of the tangential forces were counterbalanced by the opposite changes in the moments of normal forces such that the total moment remained constant and the handle orientation was maintained. At the level of individual finger (IF) forces two strategies of error compensation were found: (a) local error compensation—the opposite action of the neighboring finger, i.e. force decrease in response to a force increase (finger addition), and vice versa and (b) distant error compensation—similar action by a finger that is a torque antagonist to the manipulated finger. During the transient periods, the changes in the thumb and VF forces were simultaneous and equal in magnitude. The normal forces increased or decreased concurrently while the changes in the tangential forces were opposite in direction. The data support the existence of chain effects in the digit force adjustments to finger addition or removal. We conclude that the digit force adjustments during the object manipulation are controlled mainly in a feed-forward manner. The obtained data agree with the principle of superposition reported previously. The findings agree with earlier reports on the limited ability of CNS to organize synergies at two levels of a control hierarchy simultaneously.     

Selective deficits of grip force control during object manipulation in patients with reduced sensibility of the grasping digits

Persons with impaired manual sensibility frequently report problems to use the hand in manipulative tasks, such as using tools or buttoning a shirt. At least two control processes determine grip forces during voluntary object manipulation. Anticipatory force control specifies the motor commands on the basis of predictions about physical object properties and the consequences of our own actions. Feedback sensory information from the grasping digits, representing mechanical events at the skin-object interface, automatically modifies grip force according to the actual loading requirements and updates sensorimotor memories to support anticipatory grip force control. We investigated grip force control in nine patients with moderately impaired tactile sensibility of the grasping digits and in nine sex- and age-matched healthy controls lifting and holding an instrumented object. In healthy controls grip force was adequately scaled to the weight of the object to be lifted. The grip force was programmed to smoothly change in parallel with load force over the entire lifting movement. In particular, the grip force level was regulated in an economical way to be always slightly higher than the minimum required to prevent the object slipping. The temporal coupling between the grip and load force profiles achieved a high precision with the maximum grip and load forces coinciding closely in time. For the temporal regulation of the grip force profile patients with impaired tactile sensibility maintained the close co-ordination between proximal arm muscles, responsible for the lifting movement and the fingers stabilising the grasp. Maximum grip force coincided with maximum acceleration of the lifting movement. However, patients employed greater maximum grip forces and greater grip forces to hold the object unsupported when compared with controls. Our results give further evidence to the suggestion that during manipulation of objects with known physical properties the anticipatory temporal regulation of the grip force profile is centrally processed and less under sensory feedback control. In contrast, sensory afferent information from the grasping fingers plays a dominant role for the efficient scaling of the grip force level according to actual loading requirements.     

Nondigital afferent input in reactive control of fingertip forces during precision grip

Sensory inputs from the digits are important in initiating and scaling automatic reactive grip responses that help prevent frictional slips when grasped objects are subjected to destabilizing load forces. In the present study we analyzed the contribution to grip-force control from mechanoreceptors located proximal to the digits when subjects held a small manipulandum between the tips of the thumb and index finger. Loads of various controlled amplitudes and rates were delivered tangential to the grip surfaces at unpredictable times. Grip forces (normal to the grip surfaces) and the position of the manipulandum were recorded. In addition, movements of hand and arm segments were assessed by recording the position of markers placed at critical points. Subjects performed test series during normal digital sensibility and during local anesthesia of the index finger and thumb. To grade the size of movements of tissues proximal to the digits caused by the loadings, three different conditions of arm and hand support were used; (1) in the hand-support condition the subjects used the three ulnar fingers to grasp a vertical dowel support and the forearm was supported in a vacuum cast; (2) in the forearm-support condition only the forearm was supported; finally, (3) in the no-support condition the arm was free. With normal digital sensibility the size of the movements proximal to the digits had small effects on the grip-force control. In contrast, the grip control was markedly influenced by the extent of such movements during digital anesthesia. The poorest control was observed in the hand-support condition, allowing essentially only digital movements. The grip responses were either absent or attenuated, with greatly prolonged onset latencies. In the forearm and no-support conditions, when marked wrist movements took place, both the frequency and the strength of grip-force responses were higher, and the grip response latencies were shorter. However, the performance never approached normal. It is concluded that sensory inputs from the digits are dominant in reactive grip control. However, nondigital sensory input may be used for some grip control during impaired digital sensibility. Furthermore, the quality of the control during impaired sensibility depends on the extent of movements evoked by the load in the distal, unanesthetized parts of the arm. The origin of these useful sensory signals is discussed.