Understanding finger coordination through analysis of the structure of force variability

 Most common motor acts involve highly redundant effector systems. Understanding how such systems are controlled by the nervous system is a long-standing scientific challenge. Most proposals for solving this problem are based on the assumption that a particular solution, which optimizes additional constraints, is selected by the nervous system out of the many possible solutions. This study attempts to address this question in the context of coordinating individual finger forces to produce a controlled total force oscillation between 5% and 35% of each subject's maximum force of voluntary contraction, under two different combinations of four fingers. The structure of variability of individual finger forces was evaluated with respect to hypotheses that, at each instance in time, subjects attempt to: (1) stabilize the value of total force and (2) stabilize the total moment created by the fingers about the long axis passing through the forearm and midline of the hand. The results provide evidence that a range of goal-equivalent finger force combinations is generated to stabilize the values of total force and the total moment. The control of total force was specified explicitly by the task. However, it was stabilized only near the time of peak force. In contrast, the total moment was stabilized throughout most of the force cycle. The results lead to the suggestion that successful task performance is achieved, not by selecting a single optimal solution, but by discovering an appropriate control law that selectively stabilizes certain combinations of degrees of freedom relevant to the task while releasing from control other combinations. Received: 2 February 2001 / Accepted in revised form: 21 June 2001     

Hand function: peripheral and central constraints on performance.

The hand is one of the most fascinating and sophisticated biological motor systems. The complex biomechanical and neural architecture of the hand poses challenging questions for understanding the control strategies that underlie the coordination of finger movements and forces required for a wide variety of behavioral tasks, ranging from multidigit grasping to the individuated movements of single digits. Hence, a number of experimental approaches, from studies of finger movement kinematics to the recording of electromyographic and cortical activities, have been used to extend our knowledge of neural control of the hand. Experimental evidence indicates that the simultaneous motion and force of the fingers are characterized by coordination patterns that reduce the number of independent degrees of freedom to be controlled. Peripheral and central constraints in the neuromuscular apparatus have been identified that may in part underlie these coordination patterns, simplifying the control of multi-digit grasping while placing certain limitations on individuation of finger movements. We review this evidence, with a particular emphasis on how these constraints extend through the neuromuscular system from the behavioral aspects of finger movements and forces to the control of the hand from the motor cortex.     

Indices of nonlinearity in finger force interaction

Experiments with force production by subsets of fingers within the human hand have shown that finger interaction may be significantly nonlinear. In particular, this nonlinearity is reflected in the phenomenon of force deficit, a drop of the peak force of a finger when several fingers act simultaneously. We describe nonlinear effects in force relations within finger pairs, triplets, etc. Finger forces are represented as the sums of components resulting from force interactions within all subsets of the explicitly involved (master) fingers. The values of these components computed at extreme values of control signals, zero and unity, are taken as indices of such elementary force interactions. Indices of the first order reflect purposeful force production by a single master finger and its effects on forces produced by other fingers (enslaving). Indices of the second order reflect additional influences from pairs of simultaneously recruited master fingers, etc. Force interaction indices were computed based on finger forces measured in earlier experiments. Signs of indices alternated with their order, being positive for the indices of the first order (enslaving), negative for the indices of the second order (force deficit), positive for the indices of the third order, and mostly negative for the indices of the fourth order. Indices of the third and fourth orders reflect phenomena of force interaction not reported earlier. The study emphasizes the importance of nonlinear interactions among finger forces and introduces a set of independent indices that can be used to quantify such interactions in different subpopulations and their possible changes with practice and/or rehabilitation of the hand function.     

Balancing a force on the fingertip of a two-dimensional finger model without intrinsic muscles

A slightly flexed human middle finger can balance an external force on the fingertip. Internal stabilization is also possible, which means that the externally unloaded finger can be kept stiff. We want to analyse whether in these situations the intrinsic hand muscles are needed. Distances from tendons to flexion axes are taken from the literature and are substituted in the moment equilibrium equations of a two-dimensional finger model. Diagrams illustrate the statically indeterminate problem of solving tendon forces. The possibilities for equilibrium without intrinsics appear to depend mainly on four tendon-to-joint distances. These distances determine to which of two groups a finger belongs: (1) one in which intrinsics are not necessary for internal stabilization nor for balancing a force on the fingertip in any direction in the sagittal plane; (2) one in which, without intrinsics, internal stabilization is impossible and only dorso-distally directed forces on the fingertip can be balanced.     

Age-related changes in finger coordination in static prehension tasks.

We studied age-related changes in the performance of maximal and accurate submaximal force and moment production tasks. Elderly and young subjects pressed on six dimensional force sensors affixed to a handle with a T-shaped attachment. The weight of the whole system was counterbalanced with another load. During tasks that required the production of maximal force or maximal moment by all of the digits, young subjects were stronger than elderly. A greater age-related deficit was seen in the maximal moment production tests. During maximal force production tests, elderly subjects showed larger relative involvement of the index and middle fingers; they moved the point of thumb force application upward (toward the index and middle fingers), whereas the young subjects rolled the thumb downward. During accurate force/moment production trials, elderly persons were less accurate in the production of both total moment and total force. They produced higher antagonistic moments, i.e., moment by fingers that acted against the required direction of the total moment. Both young and elderly subjects showed negative covariation of finger forces across repetitions of a ramp force production task. In accurate moment production tasks, both groups showed negative covariation of two components of the total moment: those produced by the normal forces and those produced by the tangential forces. However, elderly persons showed lower values of the indexes of both finger force covariation and moment covariation. We conclude that age is associated with an impaired ability to produce both high moments and accurate time profiles of moments. This impairment goes beyond the well-documented deficits in finger and hand force production by elderly persons. It involves worse coordination of individual digit forces and of components of the total moment. Some atypical characteristics of finger forces may be viewed as adaptive to the increased variability in the force production with age.     

Finger Flexor Force Influences Performance in Senior Male Air Pistol Olympic Shooting

The ability to stabilize the gun is crucial for performance in Olympic pistol shooting and is thought to be related to the shooters muscular strength. The present study examines the relation between performance and finger flexor force as well as shoulder abduction isometric force in senior male air pistol shooting. 46 Spanish national level shooters served as test subjects of the study. Two maximal force tests were carried out recording handgrip and deltoid force data under competition conditions, during the official training time at national Spanish championships. Performance was measured as the total score of 60 shots at competition. Linear regressions were calculated to examine the relations between performance and peak and average finger flexor forces, peak and average finger flexor forces relative to the BMI, peak and average shoulder abduction isometric forces, peak shoulder abduction isometric force relative to the BMI. The connection between performance and other variables such as age, weight, height, BMI, experience in years and training hours per week was also analyzed. Significant correlations were found between performance at competition and average and peak finger flexor forces. For the rest of the force variables no significant correlations were found. Significant correlations were also found between performance at competition and experience as well as training hours. No significant correlations were found between performance and age, weight, height or BMI. The study concludes that hand grip strength training programs are necessary for performance in air pistol shooting.     

In vivo forces generated by finger flexor muscles do not depend on the rate of fingertip loading during an isometric task

Risk factors for activity-related tendon disorders of the hand include applied force, duration, and rate of loading. Understanding the relationship between external loading conditions and internal tendon forces can elucidate their role in injury and rehabilitation. The goal of this investigation is to determine whether the rate of force applied at the fingertip affects in vivo forces in the flexor digitorum profundus (FDP) tendon and the flexor digitorum superficialis (FDS) tendon during an isometric task. Tendon forces, recorded with buckle force transducers, and fingertip forces were simultaneously measured during open carpal tunnel surgery as subjects (N=15) increased their fingertip force from 0 to 15N in 1, 3, and 10s. The rates of 1.5, 5, and 15N/s did not significantly affect FDP or FDS tendon to fingertip force ratios. For the same applied fingertip force, the FDP tendon generated more force than the FDS. The mean FDP to fingertip ratio was 2.4+/-0.7 while the FDS to tip ratio averaged 1.5+/-1.0 (p<0.01). The fine motor control needed to generate isometric force ramps at these specific loading rates probably required similar high activation levels of multiple finger muscles in order to stabilize the finger and control joint torques at the force rates studied. Therefore, for this task, no additional increase in muscle force was observed at higher rates. These findings suggest that for high precision, isometric pinch maneuvers under static finger conditions, tendon forces are independent of loading rate.     

Orofacial and thumb-index finger ramp-and-hold isometric force dynamics in young neurotypical adults

The relation among several parameters of the ramp-and-hold isometric force contraction (peak force and dF/dt_max during the initial phase of force recruitment, and the proportion of hold-phase at target) was quantified for the right and left thumb-index finger pinch, and lower lip midline compression in 40 neurotypical right-handed young adults (20 female/20 males) using wireless force sensors and data acquisition technology developed in our laboratory. In this visuomotor control task, participants produced ramp-and-hold isometric forces as ‘rapidly and accurately’ as possible to end-point target levels at 0.25, 0.5, 1 and 2 Newtons presented to a computer monitor in a randomized block design. Significant relations were found between the parameters of the ramp-and-hold lip force task and target force level, including the peak rate of force change (dF/dt_max), peak force, and the criterion percentage of force within ±5% of target during the contraction hold phase. A significant performance advantage was found among these force variables for the thumb-index finger over the lower lip. The maximum voluntary compression force (MVCF) task revealed highly significant differences in force output between the thumb-index fingers and lower lip (∼4.47–4.70 times greater for the digits versus lower lip), a significant advantage of the right thumb-index finger over the non-dominant left thumb-index finger (12% and 25% right hand advantage for males and females, respectively), and a significant sex difference (∼1.65–1.73 times greater among males).     

Directional Coordination of Thumb and Finger Forces during Precision Pinch

The human opposable thumb enables the hand to perform dexterous manipulation of objects, which requires well-coordinated digit force vectors. This study investigated the directional coordination of force vectors generated by the thumb and index finger during precision pinch. Fourteen right-handed, healthy subjects were instructed to exert pinch force on an externally stabilized apparatus with the pulps of the thumb and index finger. Subjects applied forces to follow a force-ramp profile that linearly increased from 0 to 12 N and then decreased to 0 N, at a rate of ±3 N/s. Directional relationships between the thumb and index finger force vectors were quantified using the coordination angle (CA) between the force vectors. Individual force vectors were further analyzed according to their projection angles (PAs) with respect to the pinch surface planes and the shear angles (SAs) within those planes. Results demonstrated that fingertip force directions were dependent on pinch force magnitude, especially at forces below 2 N. Hysteresis was observed in the force-CA relationship for increasing and decreasing forces and fitted with exponential models. The fitted asymptotic values were 156.0±6.6° and 150.8±9.3° for increasing and decreasing force ramps, respectively. The PA of the thumb force vector deviated further from the direction perpendicular to the pinching surface planes than that of the index finger. The SA showed that the index finger force vector deviated in the ulnar-proximal direction, whereas the thumb switched its force between the ulnar-proximal and radial-proximal directions. The findings shed light on the effects of anatomical composition, biomechanical function, and neuromuscular control in coordinating digit forces during precision pinch, and provided insight into the magnitude-dependent force directional control which potentially affects a range of dexterous manipulations.     

Tangential finger forces use mechanical advantage during static grasping

When grasping and manipulating objects, the central controller utilizes the mechanical advantage of the normal forces of the fingers for torque production. Whether the same is valid for tangential forces is unknown. The main purpose of this study was to determine the patterns of finger tangential forces and the use of mechanical advantage as a control mechanism when dealing with objects of nonuniform finger positioning. A complementary goal was to explore the interaction of mechanical advantage (moment arm) and the role a finger has as a torque agonist/antagonist with respect to external torques (±0.4 N m). Five 6-df force/torque transducers measured finger forces while subjects held a prism handle (6 cm width × 9 cm height) with and without a single finger displaced 2 cm (handle width). The effect of increasing the tangential moment arm was significant (p < .01) for increasing tangential forces (in >70% of trials) and hence creating greater moments. Thus, the data provides evidence that the grasping system as a rule utilizes mechanical advantage for generating tangential forces. The increase in tangential force was independent of whether the finger was acting as a torque agonist or antagonist, revealing their effects to be additive.     

Four-channel telemetry system for in vivo measurement of hip joint forces

The long-term loosening of artificial hip joints remains a serious clinical problem. Optimization of implant design and material will improve the fixation, but it requires a detailed knowledge of the forces which act on the implant. A four-channel telemetric transmitter was developed and arranged completely inside a hermetically closed artificial hip joint. This permits long term in vivo measurements of the three-dimensional forces without endangering the patient. The external telemetry system consists of an inductive power supply, an RF receiver, a microcomputer with hardware extension and a VHS video system. The personal computer offers real-time data processing of three orthogonal force components as well as slow motion analysis of recorded measurements. After several years of animal tests, two instrumented prostheses were implanted in the first patient (male) in May and August of 1988. In March 1990 a third prosthesis was implanted in a second patient (female). Joint force measurements have regularly been performed from the first post-operative day until now for several kinds of activity.     

How Variability and Effort Determine Coordination at Large Forces

Motor control is a challenging task for the central nervous system, since it involves redundant degrees of freedom, nonlinear dynamics of actuators and limbs, as well as noise. When an action is carried out, which factors does your nervous system consider to determine the appropriate set of muscle forces between redundant degrees-of-freedom? Important factors determining motor output likely encompass effort and the resulting motor noise. However, the tasks used in many previous motor control studies could not identify these two factors uniquely, as signal-dependent noise monotonically increases as a function of the effort. To address this, a recent paper introduced a force control paradigm involving one finger in each hand that can disambiguate these two factors. It showed that the central nervous system considers both force noise and amplitude, with a larger weight on the absolute force and lower weights on both noise and normalized force. While these results are valid for the relatively low force range considered in that paper, the magnitude of the force shared between the fingers for large forces is not known. This paper investigates this question experimentally, and develops an appropriate Markov chain Monte Carlo method in order to estimate the weightings given to these factors. Our results demonstrate that the force sharing strongly depends on the force level required, so that for higher force levels the normalized force is considered as much as the absolute force, whereas the role of noise minimization becomes negligible.     

Biodynamic response of human fingers in a power grip subjected to a random vibration

BACKGROUND: Knowledge of the biodynamic response (BR) of the human hand-arm system is an important part of the foundation for the measurement and assessment of hand-transmitted vibration exposure. This study investigated the BR of human fingers in a power grip subjected to a random vibration. METHOD: Ten male subjects were used in the experiment. Each subject applied three coupling actions to a simulated tool handle at three different finger grip force levels. RESULTS AND CONCLUSIONS: The BR is practically independent of the hand coupling actions for frequencies at or above 100 Hz. Above 50 Hz, the BR is correlated to finger and hand sizes. Increasing the finger coupling force significantly increases the BR. Therefore, hand forces should be measured and used when assessing hand-transmitted vibration exposure. The results also show that under a constant-velocity vibration, the finger vibration power absorption at frequencies above 200 Hz is approximately twice that at frequencies below 100 Hz. This suggests that the frequency weighting specified in the current ISO 5349-1 (2001) may underestimate the high frequency effect on vibration-induced finger disorders.     

A mode hypothesis for finger interaction during multi-finger force-production tasks

 Finger forces are known to change involuntarily during multi-finger force-production tasks, even when a finger's involvement in a task is not consciously changed (the enslaving effect). Furthermore, during maximal force-production (MVC) tests, the force produced by a given finger in a multi-finger task is smaller than the force generated by this finger in its single-finger MVC test (the force-deficit effect). A set of hypothetical control variables – modes – is introduced. Modes can be estimated based on individual finger forces during single-finger MVC tests. We show that a simple formal model based on modes with only one free parameter accounts for finger forces during a variety of multi-finger MVC tests. The free parameter accounts for the force-deficit effect, and its value depends only on the number of explicitly involved fingers. This approach offers a simple framework for the analysis of finger interaction during multi-finger actions. Received: 7 December 2001 / Accepted in revised form: 17 April 2002 Correspondence to: F. Danion (e-mail: danion@laps.univ-mrs.fr, Tel.: +33-491-172265, Fax: +33-491-172252)     

Biomechanical Analysis of the Human Finger Extensor Mechanism during Isometric Pressing

This study investigated the effects of the finger extensor mechanism on the bone-to-bone contact forces at the interphalangeal and metacarpal joints and also on the forces in the intrinsic and extrinsic muscles during finger pressing. This was done with finger postures ranging from very flexed to fully extended. The role of the finger extensor mechanism was investigated by using two alternative finger models, one which omitted the extensor mechanism and another which included it. A six-camera three-dimensional motion analysis system was used to capture the finger posture during maximum voluntary isometric pressing. The fingertip loads were recorded simultaneously using a force plate system. Two three-dimensional biomechanical finger models, a minimal model without extensor mechanism and a full model with extensor mechanism (tendon network), were used to calculate the joint bone-to-bone contact forces and the extrinsic and intrinsic muscle forces. If the full model is assumed to be realistic, then the results suggest some useful biomechanical advantages provided by the tendon network of the extensor mechanism. It was found that the forces in the intrinsic muscles (interosseus group and lumbrical) are significantly reduced by 22% to 61% due to the action of the extensor mechanism, with the greatest reductions in more flexed postures. The bone-to-bone contact force at the MCP joint is reduced by 10% to 41%. This suggests that the extensor mechanism may help to reduce the risk of injury at the finger joints and also to moderate the forces in intrinsic muscles. These apparent biomechanical advantages may be a result of the extensor mechanism''s distinctive interconnected fibrous structure, through which the contraction of the intrinsic muscles as flexors of the MCP joint can generate extensions at the DIP and PIP joints.     

An integrative approach to the biomechanical function and neuromuscular control of the fingers

The exquisite mechanical functionality and versatility of the human hand emerges from complex neuro-musculo-skeletal interactions that are not completely understood. I have found it useful to work within a theoretical/experimental paradigm that outlines the fundamental neuro-musculo-skeletal components and their interactions. In this integrative paradigm, the laws of mechanics, the specifications of the manipulation task, and the sensorimotor signals define the interactions among hand anatomy, the nervous system, and manipulation function. Thus, our collaborative research activities emphasize a firm grounding in the mechanics of finger function, insistence on anatomical detail, and meticulous characterization of muscle activity. This overview of our work on precision pinch (i.e., the ability to produce and control fingertip forces) presents some of our findings around three Research Themes: Mechanics-based quantification of manipulation ability; Anatomically realistic musculoskeletal finger models; and Neural control of finger muscles. I conclude that (i) driving the fingers to some limit of sensorimotor performance is instrumental to elucidating motor control strategies; (ii) that the cross-over of tendons from flexors to extensors in the extensor mechanism is needed to produce force in every direction, and (iii) the anatomical routing of multiarticular muscles makes co-contraction unavoidable for many tasks. Moreover, creating realistic and clinically useful finger models still requires developing new computational means to simulate the viscoelastic tendinous networks of the extensor mechanism, and the muscle-bone-ligament interactions in complex articulations. Building upon this neuromuscular biomechanics paradigm is of immense clinical relevance: it will be instrumental to the development of clinical treatments to preserve and restore manual ability in people suffering from neurological and orthopedic conditions. This understanding will also advance the design and control of robotic hands whose performance lags far behind that of their biological counterparts.     

Microcomputer programs for DNA sequence analysis.

Computer programs are described which allow (a) analysis of DNA sequences to be performed on a laboratory microcomputer or (b) transfer of DNA sequences between a laboratory microcomputer and another computer system, such as a DNA library. The sequence analysis programs are interactive, do not require prior experience with computers and in many other respects resemble programs which have been written for larger computer systems (1-7). The user enters sequence data into a text file, accesses this file with the programs, and is then able to (a) search for restriction enzyme sites or other specified sequences, (b) translate in one or more reading frames in one or both directions in order to find open reading frames, or (c) determine codon usage in the sequence in one or more given reading frames. The results are given in table format and a restriction map is generated. The modem program permits collection of large amounts of data from a sequence library into a permanent file on the microcomputer disc system, or transfer of laboratory data in the reverse direction to a remote computer system.     

Analysis of Finger Muscular Forces using a Wearable Hand Exoskeleton System

In this paper,the finger muscular forces were estimated and analyzed through the application of inverse dynamics-based static optimization,and a hand exoskeleton system was designed to pull the fingers and measure the dynamics of the hand.To solve the static optimization,a muscular model of the hand flexors was derived.The experimental protocol was devised to analyze finger flexors in order to evaluate spasticity of the clenched fingers;muscular forces were estimated while the flexed fingers were extended by the exoskeleton with external loads applied.To measure the finger joint angles,the hand exoskeleton system was designed using four-bar linkage structure and potentiometers.In addition,the external loads to the fingertips were generated by cable driven actuators and simultaneously measured by loadcells which were located at each phalanx.The experiments were performed with a normal person and the muscular forces estimation results were discussed with reference to the physical phenomena.     

Age effects on force produced by intrinsic and extrinsic hand muscles and finger interaction during MVC tasks.

Finger-pressing forces are produced by activation of the intrinsic hand muscles, which are finger specific, and the extrinsic muscles that connect to multiple fingers. We tested a hypothesis of greater weakening of intrinsic hand muscles with age and quantified associated indexes of finger interaction such as enslaving (force production by unintended fingers) and force deficit (loss of finger force in multifinger tasks compared with single-finger tasks). Twelve young (23-35 yr old) and 12 elderly (70-95 yr old) men and women performed single-finger and four-finger maximal pressing tasks, in which force was applied at the proximal phalanges (PP, the intrinsic muscles are major focal force generators) and at the distal phalanges (DP, the extrinsic muscles are focal force generators). The decline in the peak force with age was greater at PP (30%) than at DP (19%). Larger indexes of finger interaction were observed at PP (enslaving = 17.2 +/- 9.4%, force deficit = 36.1 +/- 11.1%) than at DP (enslaving = 14.9 +/- 8.8%, force deficit = 27.7 +/- 10.8%) across ages and genders. We conclude that intrinsic hand muscles show disproportionate weakening with age. The greater indexes of finger interaction in PP tests with greater involvement of intrinsic hand muscles suggest that the finger interactions are predominantly of a central origin across ages and genders.     

Tactile feedback plays a critical role in maximum finger force production

This study investigates the role of cutaneous feedback on maximum voluntary force (MVF), finger force deficit (FD) and finger independence (FI). FD was calculated as the difference between the sum of maximal individual finger forces during single-finger pressing tasks and the maximal force produced by those fingers during an all-finger pressing task. FI was calculated as the average non-task finger forces normalized by the task-finger forces and subtracted from 100 percent. Twenty young healthy right-handed males participated in the study. Cutaneous feedback was removed by administering ring block digital anesthesia on the 2nd, 3rd, 4th and 5th digits of the right hands. Subjects were asked to press force sensors with maximal effort using individual digits as well as all four digits together, with and without cutaneous feedback. Results from the study showed a 25% decrease in MVF for the individual fingers as well as all the four fingers pressing together after the removal of cutaneous feedback. Additionally, more than 100% increase in FD after the removal of cutaneous feedback was observed in the middle and ring fingers. No changes in FI values were observed between the two conditions. Results of this study suggest that the central nervous system utilizes cutaneous feedback and the feedback mechanism plays a critical role in maximal voluntary force production by the hand digits.