Age-related changes in multi-finger interactions in adults during maximum voluntary finger force production tasks

This study aimed to continue our characterization of finger strength and multi-finger interactions across the lifespan to include those in their 60s and older. Building on our previous study of children, we examined young and elderly adults during isometric finger flexion and extension tasks. Sixteen young and 16 elderly, gender-matched participants produced maximum force using either a single finger or all four fingers in flexion and extension. The maximum voluntary finger force (MVF), the percentage contributions of individual finger forces to the sum of individual finger forces during four-finger MVF task (force sharing), and the non-task finger forces during a task finger MVF task (force enslaving), were computed as dependent variables. Force enslaving during finger extension was greater than during flexion in both young and elderly groups. The flexion-extension difference was greater in the elderly than the young adult group. The greater independency in flexion may result from more frequent use of finger flexion in everyday manipulation tasks. The non-task fingers closer to a task finger produced greater enslaving force than non-task fingers farther from the task finger. The force sharing pattern was not different between age groups. Our findings suggest that finger strength decreases over the aging process, finger independency for flexion increases throughout development, and force sharing pattern remains constant across the lifespan.     

Characteristics of finger force production during one- and two-hand tasks

Relations among finger forces were studied during one-hand and two-hand isometric maximal force production tasks in right- and left-handers. We particularly focused on the phenomena of force deficit during one-hand multi-finger tasks and of bilateral force deficit during two-hand tasks. Ten healthy subjects (five of them left-handed) performed maximal voluntary force production tasks with different finger combinations involving fingers of one of the hands or of both hands together. In one-hand tasks, finger enslaving (forces produced by fingers that were not instructed to produce force) was larger in the dominant hand, while force deficit (drop in individual finger peak force during multi-finger tasks) showed no differences between the hands. An additional drop in finger forces was seen in two-hand tests (bilateral deficit). The magnitude of the bilateral deficit for a hand was larger for tasks involving fewer fingers within the hand and more fingers in the other hand, with a ceiling effect. Smaller bilateral deficit was seen in tasks involving symmetrical finger combinations. In two-hand tasks that could potentially lead to the generation of large total moments in the frontal plane, the hand that was expected to generate larger moments showed larger bilateral deficit, so that the magnitude of the total moment was reduced. These observations suggest that force deficit within a hand and bilateral deficit have different origins but their effects are combined at a certain level of the multi-finger control hierarchy. Bilateral deficit may display task dependence reflecting, in particular, the principle of minimization of secondary moments. A double-representation, mirror-image hypothesis is suggested to provide a neurophysiological basis for the observed patterns of bilateral deficit.     

Stabilization of the total force in multi-finger pressing tasks studied with the 'inverse piano' technique

When one finger changes its force, other fingers of the hand can show unintended force changes in the same direction (enslaving) and in the opposite direction (error compensation). We tested a hypothesis that externally imposed changes in finger force predominantly lead to error compensation effects in other fingers thus stabilizing the total force. A novel device, the “inverse piano”, was used to impose controlled displacements to one of the fingers over different magnitudes and at different rates. Subjects (n = 10) pressed with four fingers at a constant force level and then one of the fingers was unexpectedly raised. The subjects were instructed not to interfere with possible changes in the finger forces. Raising a finger caused an increase in its force and a drop in the force of the other three fingers. Overall, total force showed a small increase. Larger force drops were seen in neighbors of the raised finger (proximity effect). The results showed that multi-finger force stabilizing synergies dominate during involuntary reactions to externally imposed finger force changes. Within the referent configuration hypothesis, the data suggest that the instruction “not to interfere” leads to adjustments of the referent coordinates of all the individual fingers.     

Finger interdependence and unintentional force drifts: Lessons from manipulations of visual feedback

We explored the phenomenon of unintentional finger force drift by using visual feedback on the force produced either by explicitly instructed (master) finger pairs or by non-instructed (enslaved) finger pairs. In particular, we drew contrasting predictions from two hypotheses: that force drifts represented consequences of drifts in effector referent coordinates at the level of individual fingers vs. at the level of finger modes (hypothetical variables accounting for the finger force interdependence). Subjects performed accurate force production with two fingers of a hand, index-ring or middle-little. They received visual feedback on the force produced either by the master fingers or by the other two, enslaved, fingers. The feedback scale was adjusted to ensure that the subjects did not know the difference between these two, randomly presented, conditions. Under feedback on the master finger force, enslaved force showed a consistent drift upward. Under feedback on the enslaved finger force, master force showed a consistent drift downward. The subjects were unaware of the force drifts, which could reach over 35% of the initial force magnitude. The data support the hypothesis on drifts in the referent coordinate at the level of individual digits, not finger modes, as the origin of unintentional force drifts. The consistent increase in the relative amount of force produced by the enslaved fingers suggests that the commonly used methods to quantify enslaving should include relatively brief force production tasks.     

Dependence of finger flexion force on the posture of the nonperforming fingers during key pressing tasks

The influence of different positions of the nonperforming (idle) fingers on the maximal force contraction of flexion (master) fingers during key pressing tasks was investigated. Ten participants performed maximal voluntary flexion contractions with various combinations of the index, middle, ring, and little fingers while the idle fingers rested on or were lifted away from the supporting surface. The effect of idle finger posture on total finger force production of master fingers was dependent on finger combination. In general, force production by master fingers was higher when the idle fingers were lifted away from the supporting surface than when they rested on it. The average increase in total force production by master fingers caused by the lifting of idle fingers was +12.4% (from -8.3% to +30.2%). Force-production capability of individual master fingers can be facilitated (as high as 34.1%), unchanged, or depressed (as high as -29.0%) by lifting the idle fingers. The effect of idle finger posture on finger force production of master fingers led to changes in force deficit. Neural, anatomical, and mechanical factors might account for the dependence of finger flexion force of master fingers on the posture of the idle fingers.     

Experimental finger dyspraxia

Subjects were required to perform discrete finger movements in accordance with a pre-arranged sequence of instructions. In all cases, any movement made by any finger was recorded by means of a constant-speed kymograph equipped with levers designed to record separately the movements of the individual fingers. This experiment was carried out under three conditions: (a) with no vision of the hand; (b) with direct vision of the hand; and (c) with the hand presented in mirror-image. It was found that, whereas deprivation of visual control was without effect on the efficiency of finger movements, presentation of the hand in mirror-image gave rise to significant increases in reaction time of three fingers and to an increase in the percentages of prior and substitute movements of other digits. Phenomena akin to depersonalisation were reported in some cases. It was also found that the rank order of mean reaction times of the five digits was approximately constant under all conditions and that the two fingers with the shortest reaction times were preceded by the fewest movements of other digits. These two digits were also the most frequently moved in advance when movements of other digits were requested and it is suggested that they have a certain “signpost function” in guiding identification of the remaining fingers. Some implications of these results for an understanding of “finger agnosia” are briefly indicated in an Appendix.     

Is the principle of minimization of secondary moments validated during various fingertip force production conditions?

During the application of fingertip forces with simultaneous flexion of the four fingers, namely index, middle, ring, and little fingers, a stable force sharing among fingers is adopted. Several studies have hypothesized that this stable force sharing is established to minimize unnecessary rotational moments (different from the main flexion moments). This principle labeled "minimization of secondary moments" is presented in the literature as a principle used by the central nervous system to solve musculoskeletal redundancy. However, this principle has only been tested with one solicited degree of freedom and in one finger posture. Our study tests this principle with various degrees of freedom solicited as secondary moments and in two different finger postures. Participants (n=6) were asked to apply a downward vertical force using their four fingers with the forearm placed in two different configurations: a "horizontal" condition (involving flexion/extension and pronation/supination at the wrist joint) and a "vertical" condition (involving flexion/extension and radial/ulnar deviation at the wrist joint). Additionally, two finger postures were tested in each forearm configuration: in the first, the distal inter-phalangeal joints (DIP) were extended and the proximal inter-phalangeal joints (PIP) highly flexed. In the second finger posture, both DIP and PIP joints were flexed. The resultant four-finger force and the relative involvement of each finger in the resultant four-finger force (force sharing) were analyzed. Results showed that the finger postures did not influence the finger force sharing, showing that the minimization of the secondary moment principle was stable among the finger joint angle configurations. Nonetheless, the relative involvement of each finger was dependent on the secondary degree of freedom solicited (pronation/supination vs. radial/ulnar). The modifications of the finger force sharing between the "horizontal" and "vertical" conditions were in accordance with the principle of minimization of the secondary moments.     

Integration of hand and finger location in external spatial coordinates for tactile localization

Tactile stimulus location is automatically transformed from somatotopic into external spatial coordinates, rendering information about the location of touch in three-dimensional space. This process is referred to as tactile remapping. Whereas remapping seems to occur automatically for the hands and feet, the fingers may constitute an exception in that some studies have implied purely somatotopic coding of touch to the fingers. When participants judge the order of two tactile stimuli, they often err when the stimulated body parts (usually the two hands) are crossed, presumably because somatotopic and external coordinates are in conflict in crossed postures. Using this task, we investigated, first, whether the fingers are unlike other limbs with regard to spatial coding, by testing whether crossing effects, indicative of external coding, were observable when stimulating two fingers, either on the same or on different hands. Second, we investigated the interaction of hand and finger posture in tactile localization of finger stimuli. Crossing effects emerged when fingers and hands were crossed, suggesting external coding for all body parts. Crossing effects were larger when both hand and finger were located in the hemifield opposite to their body side, and smaller when only hand or finger lay in the opposite hemifield. We suggest that tactile location is estimated by integrating the external location of all relevant body parts, here of a finger and its belonging hand, and that such integrative coding may represent a general principle for body part processing as well as for tool use.     

The effect of hand position and pattern motion on temporal order judgments

Subjects made temporal order judgments (TOJs) of tactile stimuli presented to the fingerpads. The subjects judged which one of two locations had been stimulated first. The tactile stimuli were patterns that simulated movement across the fingerpads. Although irrelevant to the task, the direction of movement of the patterns biased the TOJs. If the pattern at one location moved in the direction of the second location, the subjects tended to judge the first location as leading the second location. If the pattern moved in the opposite direction, that location was judged as trailing. In a series of experiments, the effect of the spatial position of the hands and fingers on TOJs and the perception of the direction of pattern movement were examined. Changing the position of the hands so that the patterns no longer moved directly toward each other reduced or eliminated the effect of motion on TOJs. In a variation of Aristotle's illusion, the moving patterns were presented to crossed and uncrossed fingers. The results indicated that, contrary to Aristotle's illusion, the subjects processed the moving patterns relative to an environmental framework, rather than to the local direction of motion on the fingerpads. Presenting the patterns to crossed hands produced results similar to those obtained with crossed fingers: The subjects processed the patterns according to an environmental framework.     

The influence of response grouping on free-choice decision making in a response selection task

Previous research has demonstrated an advantage for the preparation of fingers on one hand over the preparation of fingers on two hands, and for the preparation of homologous fingers over that of non-homologous fingers. In the present study, we extended the precuing effects observed with finger responses to response selection under free-choice conditions. Participants were required to choose from a range of possible responses following the presentation of a precue that indicated which response to prepare (go-to precue) or prevent (no-go-to precue). In Experiment 1 the choice was between homologous and non-homologous finger responses on the hand opposite to the precue while in Experiment 2 the choice was between finger responses on the same or different hand to the precue. In the go-to precue condition, the frequency of homologous finger choices was more frequent than non-homologous finger responses. Similarly, participants chose finger responses on the same hand as the precue regardless of whether they were instructed to prepare or prevent the precued response. The hand effect bias was stronger than the finger effect bias. These findings are consistent with the Grouping Model (Adam, Hommel, & Umilta, 2003).     

Finger flexor motor control patterns during active flexion: an in vivo tendon force study

An in vivo tendon force measurement system was used to evaluate index finger flexor motor control patterns during active finger flexion. During open carpal tunnel release surgery (N=12) the flexor digitorum profundus (FDP) and flexor digitorum superficilias (FDS) tendons were instrumented with buckle force transducers and participants performed finger flexion at two different wrist angles (0 degrees or 30 degrees ). During finger flexion, there was concurrent change of metacarpophalangeal (MCP) and proximal interphalangeal (PIP) joint angles, but the FDP and FDS tendon force changes were not concurrent. For the FDS tendon, no consistent changes in force were observed across participants at either wrist angle. For the FDP tendon, there were two force patterns. With the wrist in a neutral posture, the movement was initiated without force from the finger flexors, and further flexion (after the first 0.5s) was carried out with force from the FDP. With the wrist in a flexed posture, the motion was generally both initiated and continued using FDP force. At some wrist postures, finger flexion was initiated by passive forces which were replaced by FDP force to complete the motion.     

Changes in pinch force with bidirectional load forces

The forces used to grasp an object were measured while positive (push) and negative (pull) load forces were applied to the hand under varying frictional conditions. Subjects held between the tips of their thumb and index finger a manipulandum composed of two symmetrically mounted disks. The manipulandum was connected to the stage of an electromagnetic linear motor that generated load forces under computer control. In the first experiment, subjects held the position of the manipulandum constant while the motor generated forces in first the positive and then the negative direction. The motor force at which the manipulandum slipped from the fingers was measured in the second experiment. In both experiments, friction was varied by changing the surface (sandpaper, suede, or plastic) of the manipulandum disks. The pinch forces produced by subjects were linearly related to changes in motor force in both the positive and negative directions, with the slope of this relation varying as a function of the surface properties of the manipulandum. The modulation of pinch force with motor force was influenced, however, by the direction of the load force; higher forces were produced in response to negative load forces. Slip forces varied as a function of pinch force and surface texture; higher forces were associated with materials with lower coefficients of friction. These findings suggest that the friction between the skin and an object being grasped changes as a function of the direction of force that the object applies to the skin, possibly due to the anisotropic nature of glabrous skin, and that this mechanical property contributes to variations in pinch force.     

Sequential and Biomechanical Factors Constrain Timing and Motion in Tapping

The authors examined how timing accuracy in tapping sequences is influenced by sequential effects of preceding finger movements and biomechanical interdependencies among fingers. Skilled pianists tapped sequences at 3 rates; in each sequence, a finger whose motion was more or less independent of other fingers' motion was preceded by a finger to which it was more or less coupled. Less independent fingers and those preceded by a more coupled finger showed large timing errors and change in motion because of the preceding finger's motion. Motion change correlated with shorter intertap intervals and increased with rate. Thus, timing of sequence elements is not independent of the motion trajectories that individuals use to produce them. Neither motion nor its relation to timing is invariant across rates.     

Oculomotor interference during manual response preparation: evidence from the response-cueing paradigm

Preparation provided by visual location cues is known to speed up behavior. However, the role of concurrent saccades in response to visual cues remains unclear. In this study, participants performed a spatial precueing task by pressing one of four response keys with one of four fingers (two of each hand) while eye movements were monitored. Prior to the stimulus, we presented a neutral cue (baseline), a hand cue (corresponding to left vs. right positions), or a finger cue (corresponding to inner vs. outer positions). Participants either remained fixated on a central fixation point or moved their eyes freely. The results demonstrated that saccades during the cueing interval altered the pattern of cueing effects. Finger cueing trials in which saccades were spatially incompatible (vs. compatible) with the subsequently required manual response exhibited slower manual RTs. We propose that interference between saccades and manual responses affects manual motor preparation.     

Determinants of two-choice reaction-time patterns for same-hand and different-hand finger pairings

Several studies of two-choice reaction times have compared situations in which the alternative responses are fingers from one hand (the same-hand pairing) or one finger from each hand (the different-hand pairing). Two patterns of results have been obtained: (a) equivalent reaction times for the same-hand and different-hand pairings and (b) faster reaction times for the different-hand pairing. Previously, these outcomes have been attributed to the adoption of different response-preparation strategies when response pairs are constant (low response-pair uncertainty) versus when they are varied from trial to trial (high response-pair uncertainty). However, response-pair uncertainty has been confounded with whether only the two relevant fingers were placed on response keys or whether more than two fingers were. Experiment 1 of the present study demonstrated that finger placement, rather than response-pair uncertainty, determines which reaction-time pattern is obtained. Experiments 2 and 3 investigated the nature of the finger-placement effect by placing the fingers that were irrelevant for the task on response keys or on immovable blocks. The experiments indicated that the crucial factor is the number of fingers on active response keys, with the type of preparation being different when only two fingers are on keys rather than when more than two fingers are.     

The effect of motion on tactile and visual temporal order judgments

Four experiments were conducted, three with tactile stimuli and one with visual stimuli, in which subjects made temporal order judgments (TOJs). The tactile stimuli were patterns that moved laterally across the fingerpads. The subject's task was to judge which finger received the pattern first. Even though the movement was irrelevant to the task, the subjects' TOJs were greatly affected by the direction of movement of the patterns. Accuracy in judging temporal order was enhanced when the patterns moved in a direction that was consistent with the temporal order of presentation--for example, when the movement on each fingerpad was from right to left and the temporally leading site of stimulation was to the right of the temporally trailing site of stimulation. When movement was inconsistent with the temporal order of presentation, accuracy was considerably reduced, often well below chance.The bias in TOJs was unaffected by training or by presenting the stimuli to fingers on opposite hands. In a fourth experiment, subjects judged the temporal order of visual stimuli that, like the tactile stimuli, moved in a direction that was either consistent or inconsistent with the TOJ. The results were similar to those obtained with tactile stimuli. It is suggested that the bias may be affected by attentional mechanisms and by apparent motion generated between the two sites on the skin.     

Determinants of Two-Choice Reaction-Time Patterns for Same-Hand and Different-Hand Finger Pairings

Several studies of two-choice reaction times have compared situations in which the alternative responses are fingers from one hand (the same-hand pairing) or one finger from each hand (the different-hand pairing). Two patterns of results have been obtained: (a) equivalent reaction times for the same-hand and different-hand pairings and (b) faster reaction times for the different-hand pairing. Previously, these outcomes have been attributed to the adoption of different response-preparation strategies when response pairs are constant (low response-pair uncertainty) versus when they are varied from trial to trial (high response-pair uncertainty). However, response-pair uncertainty has been confounded with whether only the two relevant fingers were placed on response keys or whether more than two fingers were. Experiment 1 of the present study demonstrated that finger placement, rather than response-pair uncertainty, determines which reaction-time pattern is obtained. Experiments 2 and 3 investigated the nature of the finger-placement effect by placing the fingers that were irrelevant for the task on response keys or on immovable blocks. The experiments indicated that the crucial factor is the number of fingers on active response keys, with the type of preparation being different when only two fingers are on keys rather than when more than two fingers are.     

Difference of motor overflow depending on the impaired or unimpaired hand in stroke patients

The aim of this study was to investigate the patterns of contralateral motor overflow (i.e. mirror movement) between the homologous body parts on the right and left side, in stroke patients during single-finger and multi-finger maximum force production tasks. Forty subjects, including stroke (n = 20) and normal subjects (n = 20), participated in this study. The stroke subjects maximally pressed force sensors with their fingers in a flexed position using a single (index, middle, ring, or little) or all fingers (all 4 fingers) using the impaired (IH) or unimpaired (UIH) hand, while the non-patient subjects used their right hands for the same tasks. The maximal voluntary forces in the ipsilateral and unintended pressing forces of each contralateral finger were recorded during the tasks. The magnitude of motor overflow to the contralateral side was calculated using the index of contralateral independence (CI). During the single finger tasks, the finger CI was significantly decreased in the UIH (91%) compared with that in the IH (99%) or normal hands (99%). Likewise, the multiple finger tasks showed that the CI was significantly lower in the UIH (84%) compared with that in the IH (96%) or normal hands (99%). However, the maximal forces were significantly lower in the IH relative to those in the UIH and normal hands. These data demonstrate that stroke patients have greater motor overflow from the UIH to the IH.     

Proprioceptive space perception after anaesthetization of the mid-interphalangeal joint of the finger

The effect of a digital block on the voluntary positioning of the index finger from a point of extreme extension through 45° flexion was examined in 12 subjects. A xylocaine ring block of the second proximal interphalangeal joint was used to eliminate the joint receptors while weights were attached to the finger. An inverse relationship was found between weight and distance moved when the subject was required to move the index finger of the nonpreferred hand to the required position. The results indicate that the muscular system is ancillary to the joint system in proprioceptive space perception.