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Biomedical subjects

G Westling

Publications and source records attributed to G Westling.

At least 19 recordsLinked to original sources

Development of human precision grip. IV. Tactile adaptation of isometric finger forces to the frictional condition.

The adaptation of the grip forces to the frictional condition between the digits and an object relies on feedforward sensorimotor mechanisms that use tactile afferent input to intermittently update a sensorimotor memory that controls the force coordination, i.e., the ratio between grip force (normal to the grip surface) and load force (tangential to the grip surface). The present study addressed the development of these mechanisms. Eighty-nine children and 15 adults lifted an instrumented object with exchangeable grip surfaces measuring the grip and load forces. Particularly in trials with high friction (sandpaper), the youngest children used a high grip force to load force ratio. Although this large safety margin against slips indicated an immature capacity to adapt to the frictional condition, higher grip forces were produced for more slippery material (silk versus sandpaper). The safety margin decreased during the first 5 years of age, in parallel with a lower variability in the grip force and a better adaptation to the current frictional condition. The youngest children (18 months) could adapt the grip force to load force ratio to the frictional condition in a series of lifts when the same surface structure was presented in blocks of trials, but failed when the surface structure was unpredictably changed between subsequent lifts. The need for repetitive presentation suggests a poor capacity to form a sensorimotor memory representation of the friction or an immature capacity to control the employed ratio from this representation. The memory effects, reflected by the influences of the frictional condition in the previous trial, gradually increased with age. Older children required a few lifts and adults only one lift to update their force coordination to a new friction. Hence, the present finding suggests that young children use excessive grip force, a strategy to avoid frictional slips, to compensate for an immature tactile control of the precision grip.

Adolescent

Corticospinal control during reach, grasp, and precision lift in man.

Transcranial magnetic brain stimulation (TMS) was used to assess the influence of the corticospinal system on motor output in seven human subjects during a task in which they had to reach out, grasp, and lift an object. Stimuli, directed at the hand area of the motor cortex, were delivered at eight defined points during the task: during reach, at grip closure, during object manipulation, during the parallel isometric increase in grip and load forces, during the lifting movement, and while the object was held in air. The amplitudes of short-latency EMG responses evoked by TMS in six arm and hand muscles showed a striking modulation across the different phases of the task. This modulation may well reflect phasic changes in corticospinal excitability because: (1) it did not simply reflect phasic changes in muscular activity associated with task performance, (2) it could vary inversely with the amplitude of the background EMG, and (3) it was only obtained with weak TMS intensities, below threshold for evoking responses in hand muscles of the relaxed subject. Our results suggest that the cortical representations of extrinsic hand muscles, which act to orientate the hand and finger tips, were subjected to a strong excitatory drive throughout the reach. This drive was also observed for brachioradialis and anterior deltoid, which contribute to transport of the hand. In contrast, the intrinsic hand muscles appear to receive their strongest cortical input as the digits closed around the object, and just after the subject first touched the object at the onset of manipulation. The isometric parallel increase in load and grip forces necessary to lift the object, which is normally triggered by tactile contact, was delayed by TMS delivered late during the reach. TMS at this time may disrupt processing necessary to control this critical phase of the task.

Adult

Time-varying enhancement of human cortical excitability mediated by cutaneous inputs during precision grip.

1. We have investigated the afferent neurogram, muscular activity and mechanical responses while subjects restrained, with a precision grip, an object subjected to pulling loads directed away from the hand. At unpredictable times 'ramp-and-hold' loads of 1 N were delivered at a rate of ca 80 N s-1. The load ramp produced a sharp increase in multiunit activity recorded from cutaneous afferents of the median nerve. The first response in the EMG of distal hand muscles commenced at 51 +/- 2.4 ms (mean +/- S.D.); a further steep increase in activity began about 20 ms later, and this was associated with a marked augmentation of the grip force increase. 2. In four subjects, transcranial magnetic stimulation (TMS) was delivered to the contralateral motor cortex in 1000 out of a total of 1500 loading trials. The time of the stimulus was randomly selected to occur either at one of nine defined points (separated by 20 ms) before and after the computer command triggering the load force increase, or during steady periods of grip. 3. In most hand and arm muscles, there was a powerful facilitation of the short-latency EMG responses evoked by TMS delivered 40-140 ms after the load force command. The amplitudes of the largest responses (TMS delivered at 80-100 ms) were 850% higher on average than those observed when subjects gripped the unloaded object or when they restrained the statically loaded object. This large modulation was only obtained with stimulus intensities that were subthreshold in the relaxed subject. 4. The modulation was not simply a reflection of the time-varying level of motoneuronal activity during the loading trial. In most muscles, changes in the amplitude of the TMS-evoked responses were disproportionately larger than the corresponding modulation of the background EMG activity. At its maximum, the modulation in the TMS-evoked response was nearly 300% larger. Furthermore, the strength of the TMS-evoked responses did not strictly co-vary with amplitude of background EMG, i.e. inverse relationships were seen. 5. Since motor responses to the loading of the object depend on cutaneous afferent input from the gripping digits, the results demonstrate an interaction between the effects of these inputs and those of TMS. A possible site of this interaction is the primary motor cortex; the strong modulation of the responses to TMS could reflect variation in the excitability of cortical neurons mediated by the cutaneous afferent input. However, such excitability changes lagged the predicted onset of cortical excitation in a manner suggesting that the earliest 20 ms of the subjects' EMG responses to the load increase are subcortical in origin. In contrast, the results are consistent with a cortical mediation of the subsequent powerful boosting of the EMG responses associated with the robust grip force response.

Adolescent

Development of human precision grip. II. Anticipatory control of isometric forces targeted for object's weight.

The development of anticipatory control during lifts with the precision grip was examined in 100 children aged 1 to 15 years and in 15 adults. The children were instructed to lift an instrumented test object by using the precision grip between the thumb and index finger. The employed grip force, load force (vertical lifting force), vertical position and their corresponding time derivatives (i.e., grip and load force rates and acceleration) were recorded. The weight of the object was varied between trials to access the influence of the object's weight in the previous trial on the isometric force output. Already by the second year, children began to use information pertaining to the object's weight in the previous lift, i.e., they began to use an anticipatory control strategy. This occurred concomitant to the development of mainly bell shaped force rate profiles (Forssberg et al. 1991). The succeeding development of a more mature anticipatory control was gradual and adult-like capacity was not reached until 8-11 years of age.

Adolescent

Development of human precision grip. III. Integration of visual size cues during the programming of isometric forces.

Recent evidence has shown that visual and haptical size information can be used by adults to estimate the weight of the object, forming the basis of the force programming during precision grip (Gordon et al. 1991a, b,). The present study examined the development of the capacity to use visual size information. In the first experiment, 30 children (age 1-7 years) and 10 adults performed a series of lifts with two boxes presented in an unpredictable order. The boxes were equal in weight but unequal in size and were attached to an instrumented grip handle which measured the employed grip force, load force, position and their corresponding time derivatives. The isometric force development was not influenced by the box size before the age of 3. However, the children aged 3 years and older demonstrated greater visual influences on the force programming than adults. To determine more precisely when children began to use visual size information, a second experiment in which the size and weight covaried was performed on 15 children. Children still did not use the size information during the force programming until the later half of the third year. It is concluded that this ability, probably involving associative transformations between the size and weight of objects, emerges around one year after anticipatory control based on somatosensory information pertaining to the weight of the object.

Adolescent

Independent control of human finger-tip forces at individual digits during precision lifting.

1. Subjects lifted an object with two parallel vertical grip surfaces and a low centre of gravity using the precision grip between the tips of the thumb and index finger. The friction between the object and the digits was varied independently at each digit by changing the contact surfaces between lifts. 2. With equal frictional conditions at the two grip surfaces, the finger-tip forces were about equal at the two digits, i.e. similar vertical lifting forces and grip forces were used. With different frictions, the digit touching the most slippery surface exerted less vertical lifting force than the digit in contact with the rougher surface. Thus, the safety margins against slips were similar at the two digits whether they made contact with surfaces of similar or different friction. 3. During digital nerve block, large and variable safety margins were employed, i.e. the finger-tip forces did not reflect the surface conditions. Slips occurred more frequently than under normal conditions (14% of all trials with nerve block, <5% during normal conditions), and they only occasionally elicited compensatory adjustments of the finger-tip forces and then at prolonged latencies. 4. The partitioning of the vertical lifting force between the digits was thus dependent on digital afferent inputs and resulted from active automatic regulation and not just from the mechanics of the task. 5. The safety margin employed at a particular digit was mainly determined by the frictional conditions encountered by the digit, and to a lesser degree by the surface condition at the same digit in the previous lift (anticipatory control), but was barely influenced by the surface condition at the other digit. 6. It was concluded that the finger-tip forces were independently controlled for each digit according to a 'non-slip strategy'. The findings suggest that the force distribution among the digits represents a digit-specific lower-level neural control establishing a stable grasp. This control relies on digit-specific afferent inputs and somatosensory memory information. It is apparently subordinated to a higher-level control that is related to the total vertical lifting and normal forces required by the lifting task and the relevant physical properties of the manipulated object.

Adolescent

Development of human precision grip. I: Basic coordination of force.

The coordination of manipulative forces was examined while children and adults repeatedly lifted a small object between the thumb and index finger. Grip force, load force (vertical lifting force), grip force rate and the vertical position of the test object were continuously measured. In adults, the force generation was highly automatized and was nearly invariant between trials. After a preload phase in which the grip was established, the grip and load forces increased in parallel under isometric conditions until the load force overcame the force of gravity and the object started to move. During this loading phase, the force rate profiles were essentially bell shaped and single peaked, suggesting that the force increases were programmed as one coordinated event. Children below the age of two exhibited a prolonged preload phase and a loading phase during which the grip and load forces did not increase in parallel. A major increase in grip force preceded the increase in load force, and at the start of the loading phase, the grip force was usually several Newtons (N). The force rate profiles were multi peaked with stepwise force increases most likely allowing peripheral feedback to play an important role in the control of the forces. After the age of two, the grip force increased less during the preload phase. The loading phase was more regularly characterized by a parallel increase of the grip force and load force and the duration of the various phases decreased. The older children programmed the forces in one force rate pulse indicating the emergence of an anticipatory strategy. Yet, the mature coordination of forces was not fully developed until several years later.(ABSTRACT TRUNCATED AT 250 WORDS)

Adolescent

Visual size cues in the programming of manipulative forces during precision grip.

A size-weight illusion was used to examine the role of visual cues in the programming of manipulative forces during the lifting of test objects utilizing the precision grip. Three different boxes of equal weight and unequal size were lifted. These were equipped with an instrumented grip handle to measure the employed grip force, load force (vertical lifting force), force rates and vertical movement. All fifteen subjects participating in the study reported that the smallest box was the heaviest, which is consistent with size-weight illusion predictions. However, the rate of increase of the isometric grip and load forces initially during the lift, the peaks of the grip and load force and the vertical acceleration were all found to increase with the box size. Thus, despite the conscious perception indicating a heavier weight for the small object, the motor program was scaled for a lighter weight. Yet, no differences were found in grip force during the static phase of the lift, where weight related information was apparently available via sensory feedback. Previous studies have reported that the programming of the precision grip is based on somatosensory information gained during previous lifts (Johansson and Westling 1984, 1988a, b). The present study suggests that visual cues are integrated in the programming of manipulative forces during precision grip.

Adult

The integration of haptically acquired size information in the programming of precision grip.

Recent evidence for the use of visual cues in the programming of the precision grip has been given by Gordon et al. (1991). Visually invoked size-related information influenced the physical forces used to produce a lift, even when it was not consistent with other sensory information. In the present study, blind-folded subjects were required to feel the size of an object by haptic exploration prior to lifting it. Two boxes of equal weight and unequal size were used for the lift objects and were attached to an instrumented (grip) handle. Grip force and load force, their rates, and the vertical movement of the object were measured. Most subjects reported that the small box was heavier, which is consistent with size-weight illusion predictions. However, peak grip force, grip force rate, peak load force, and load force rate were greater for the large box when the boxes were randomly presented, but not when the same boxes were lifted consecutively. If subjects did not feel the box prior to a lift, these parameters were scaled in between those normally employed for the large and small box. Most subjects apparently programmed the parallel increase of the grip and load force during the loading phase as one force rate pulse. This represented a "target strategy" in which an internal neural representation of the objects weight determined the actual target parameter (i.e. just enough force required to overcome gravity). The other subjects exhibited a slower stepwise increase in grip and load force rate. The subjects choosing this "probing strategy" did not scale the force parameters differently for the two boxes.(ABSTRACT TRUNCATED AT 250 WORDS)

Adult

Integration of sensory information during the programming of precision grip: comments on the contributions of size cues.

Evidence has recently been given by Gordon et al. (1991a, b) for the use of visually and haptically acquired information in the programming of lifts with the precision grip. The size-related information influences the development of manipulative forces prior to the lift-off, and the force output for larger objects is adjusted for a heavier weight even if the weight of the objects is kept the same. However, the size influences on the force output were small compared to the relative effects of the expected weight in previous trials (Johansson and Westling 1988). In the present study, both the size and weight of objects were changed between consecutive lifts to more fully determine the strength of visual size cues. During most trials, the size and weight covaried (i.e. the weight was proportional to the volume). However, in some trials, only the size was switched while the weight was kept the same to create a mismatch between the size and weight. The forces were still appropriately scaled towards an expected weight proportional to the volume of the object. It was concluded that visual size cues are highly purposeful. The effects were much larger than previously reported and were similar in magnitude to the effects based upon the expected weight. Thus, the small effects reported in the previous experiments may have been a result of conflicting "size-weight" information.

Adolescent

Measurement of contractile and electrical properties of single human thenar motor units in response to intraneural motor-axon stimulation.

1. A method is described for measuring contractile properties of single human motor units. Conventional human microneurographic techniques were adapted to stimulate individual motor axons in the median nerve, with the use of negative current pulses and a tungsten microelectrode, while recording motor-unit electromyographic activity (EMG) and isometric force responses from the thenar muscles. 2. EMG signals were recorded from both proximal and distal thenar muscle surfaces. Force was recorded in two directions (thumb flexion and abduction). This allowed calculation of the direction and magnitude of resultant force exerted by each unit. 3. Data accepted as originating from a single unit satisfied all the traditional "all-or-none" criteria. Additional criteria also required the following: 1) a wide safety margin between the threshold for unit activation and the current intensity needed to elicit responses from other units; 2) that the characteristic direction in which each unit generated force did not change during the recording period; and 3) whenever F-responses were encountered, the second EMG waveform was identical to the first--a highly improbable event if more than one unit had been excited. 4. Respiration and blood pressure waves introduced baseline fluctuations that distorted the force measurements. These fluctuations were minimized by synchronizing stimuli to the pulse pressure cycle and resetting the baseline electronically just before stimulus onset. 5. Combining motor-axon stimulation at a site remote from the muscle with electronic resetting of the force baseline and delivery of stimuli at fixed intervals after the pulse pressure waves allowed the full time course of human motor-unit twitch and tetanic force and EMG signals to be recorded accurately without signal averaging.

Axons

Twitch properties of human thenar motor units measured in response to intraneural motor-axon stimulation.

1. The twitch properties of human thenar motor units were examined in response to intraneural motor-axon stimulation. Force components of thumb abduction and flexion were measured before and after tetanic stimulation. The magnitude, direction, and time derivatives of resultant forces, together with axon conduction velocities, were calculated for each unit. 2. Various indexes of contraction and relaxation rate were measured including contraction time (time from force onset to peak), one-half relaxation time (time from peak force to one-half that value), normalized maximum contraction and normalized maximum relaxation rates (peak positive and negative time derivatives of the force signal normalized to twitch force), and the times at which these maximum rates occurred. 3. For different units, the directions of resultant forces were approximately evenly distributed between thumb abduction and flexion. At the onset of the experiment, initial twitch forces ranged from 3 to 34 mN, contraction times from 35 to 80 ms, and one-half relaxation times from 25 to 108 ms. 4. Resultant twitch forces were positively correlated to normalized maximum relaxation rates, but not to other rate indexes or to conduction velocity. The various contraction rate measures were correlated to each other, but generally not to relaxation rates. 5. After the first test involving tetanic stimulation, the twitches of most units were potentiated and slowed, especially their relaxation phase. However, the extent of these changes varied considerably between units. In general, units with weak initial forces potentiated most, some up to three-fold. These changes in twitch properties were denoted posttetanic twitch potentiation.(ABSTRACT TRUNCATED AT 250 WORDS)

Axons

A comparison of human thenar motor-unit properties studied by intraneural motor-axon stimulation and spike-triggered averaging.

1. Measurements of twitch contractile properties of human motor units recorded by spike-triggered averaging may be distorted by partial fusion between twitches, because motor units seldom fire at rates below 8-10 Hz. The effects of this fusion were examined by comparing the responses of 27 human thenar motor units when their motor axons were stimulated at 1, 8, and 10 Hz. 2. Resultant forces were calculated from the abduction and flexion force components, together with various contraction and relaxation rate indexes as reported previously. Values for single twitches were compared to measurements made from the unfused force fluctuations ("apparent twitches") of the same units recorded during 8 and 10 Hz stimulation. 3. For all units, stimulation at 8 and 10 Hz caused partial twitch fusion. At 10 Hz, mean values for "apparent twitch" forces, contraction times (CT), and one-half relaxation times (1/2RT) were reduced to 44, 76, and 52% of the corresponding values measured from separate twitches evoked by 1 Hz stimulation. Similar but smaller reductions were seen at 8 Hz. 4. Slow units, with initial twitch CT greater than 60 ms, showed significantly more distortion of all "apparent twitch" parameters when stimulated at both 8 and 10 Hz, compared to fast units (less than 50 ms). 5. The potentiated abduction force component data were compared with abduction forces obtained previously by spike-triggered averaging from the same muscle group. Mean force obtained by spike-triggered averaging ("STA twitch" force: 21 mN) was significantly larger than that measured in abduction in response to either 1 or 10 Hz motor-axon stimulation (14 mN, 6 mN, respectively).(ABSTRACT TRUNCATED AT 250 WORDS)

Axons

Coordinated isometric muscle commands adequately and erroneously programmed for the weight during lifting task with precision grip.

Small objects were lifted from a table, held in the air, and replaced using the precision grip between the index finger and thumb. The adaptation of motor commands to variations in the object's weight and sensori-motor mechanisms responsible for optimum performance of the transition between the various phases of the task were examined. The lifting movement involved mainly a flexion of the elbow joint. The grip force, the load force (vertical lifting force) and the vertical position were measured. Electromyographic activity (e.m.g.) was recorded from four antagonist pairs of hand/arm muscles primarily influencing the grip force or the load force. In the lifting series with constant weight, the force development was adequately programmed for the current weight during the loading phase (i.e. the phase of parallel increase in the load and grip forces during isometric conditions before the lift-off). The grip and load force rate trajectories were mainly single-peaked, bell-shaped and roughly proportional to the final force. In the lifting series with unexpected weight changes between lifts, it was established that these force rate profiles were programmed on the basis of the previous weight. Consequently, with lifts programmed for a lighter weight the object did not move at the end of the continuous force increase. Then the forces increased in a discontinous fashion until the force of gravity was overcome. With lifts programmed for a heavier weight, the high load and grip force rates at the moment the load force overcame the force of gravity caused a pronounced positional overshoot and a high grip force peak, respectively. In these conditions the erroneous programmed commands were automatically terminated by somatosensory signals elicited by the start of the movement. A similar triggering by somatosensory information applied to the release of programmed motor commands accounting for the unloading phase (i.e. the parallel decrease in the grip and load forces after the object contacted the table following its replacement). These commands were always adequately programmed for the weight.

Adolescent

Programmed and triggered actions to rapid load changes during precision grip.

A test object (grip apparatus) was held at its upper part using a precision grip. Small balls were dropped into a target cup at the bottom of the apparatus. The grip force, the load force (vertical lifting force) and the vertical movement were measured. Electromyographic activity (e.m.g.) was recorded from four antagonist pairs of hand/arm muscles primarily influencing the grip force or the load force. The balls were dropped either by the subject during a bimanual task, or unexpectedly by the experimenter. When the subject dropped the ball, preparatory actions occurred before the rapid increase in the vertical load caused by the impact. These actions appeared ca. 150 ms prior to the impact and involved a grip force increase and a lifting movement of the grip apparatus. The e.m.g. activity increased in all eight of the hand and arm muscles, indicating a general stiffening of the hand/arm system prior to the impact. Furthermore, the preparatory actions were programmed adequately for the size of the load force step at the impact, i.e. an adequate safety margin to prevent slips was preserved during the critical period of the impact. Thus, variations in this step caused by changes in (i) the weight of ball, (ii) the weight of the grip apparatus and (iii) the length of the drop were adequately taken into account during the programming of these actions. In addition, the frictional condition between the skin and the grip surface was also taken into account. The relevant sensory information apparently was obtained during the handling of the ball and the grip apparatus prior to the drop. There were also task-related automatic muscle responses triggered by the impact. These responses, which also served to stiffen the hand/arm system, were most pronounced during unexpected load changes, but they appeared too late to prevent slips. However, if no overall slip occurred, the triggered responses were functional in the sense that they helped to quickly restore the safety margin and the vertical position of the object.

Adult

Responses in glabrous skin mechanoreceptors during precision grip in humans.

Impulses in single tactile units innervating the human glabrous skin were recorded percutaneously from the median nerve using tungsten electrodes. The units were classified as belonging to one of the four categories: fast adapting with small receptive fields (FA I), fast adapting with large receptive fields (FA II), slowly adapting with small fields (SA I), and slowly adapting with large fields (SA II). A small test object was lifted, positioned in space and replaced using the precision grip between fingers and thumb. The grip force, the load force (vertical lifting force), the vertical movements of the object and vibrations (accelerations) in the object were recorded. After being virtually silent between lifts, the FA I units whose fields contacted the object became highly active during the initial period of grip force increase (initial response). This was also true for most SA I units. Accordingly, most of the skin deformation changes took place at low grip forces (below ca. 1 N). Later, while the load and grip forces increased in parallel during isometric conditions, the FA I and SA I units continued firing but generally at declining impulse rates. As long as the object was held in the air, the SA I units generally maintained firing with a tendency to adaptation. A minority of the FA I unit also discharged, especially during periods of pronounced physiological muscle tremor. The SA I units usually became silent when the grip and load forces in parallel declined to zero during isometric conditions after the object had contacted the table. However, during the very release of the grip the FA I units and some SA I units showed brief burst discharges (release response). The FA II units responded distinctly to the mechanical transients associated with the start of the vertical movement and especially with the sudden cessation of movement at the terminal table contact. FA II units whose end organs were remotely located in relation to the skin areas in contact with the object also responded. Most FA II units also discharged at the initial touch and at the release of the object, albeit less reliably than the type I units. In addition to weak dynamic responses during the phase of isometric force increase, the SA II units showed comparatively strong tonic responses while the object was held during static conditions. High firing rates also were maintained during long-lasting lifts. Moreover, it was established that the signals in SA II afferents were related to the three dimensional force profile in the grip.(ABSTRACT TRUNCATED AT 400 WORDS)

Adolescent

Signals in tactile afferents from the fingers eliciting adaptive motor responses during precision grip.

While human subjects lift small objects using the precision grip between the tips of the fingers and thumb the ratio between the grip force and the load force (i.e. the vertical lifting force) is adapted to the friction between the object and the skin. The present report provides direct evidence that signals in tactile afferent units are utilized in this adaptation. Tactile afferent units were readily excited by small but distinct slips between the object and the skin revealed as vibrations in the object. Following such afferent slip responses the force ratio was upgraded to a higher, stable value which provided a safety margin to prevent further slips. The latency between the onset of the a slip and the appearance of the ratio change (74 +/- 9 ms) was about half the minimum latency for intended grip force changes triggered by cutaneous stimulation of the fingers. This indicated that the motor responses were automatically initiated. If the subjects were asked to very slowly separate their thumb and the opposing finger while the object was held in air, grip force reflexes originating from afferent slip responses appeared to counteract the voluntary command, but the maintained upgrading of the force ratio was suppressed. In experiments with weak electrical cutaneous stimulation delivered through the surfaces of the object it was established that tactile input alone could trigger the upgrading of the force ratio. Although, varying in responsiveness, each of the three types of tactile units which exhibit a pronounced dynamic sensitivity (FA I, FA II and SA I units) could reliably signal these slips. Similar but generally weaker afferent responses, sometimes followed by small force ratio changes, also occurred in the FA I and the SA I units in the absence of detectable vibrations events. In contrast to the responses associated with clear vibratory events, the weaker afferent responses were probably caused by localized frictional slips, i.e. slips limited to small fractions of the skin area in contact with the object. Indications were found that the early adjustment to a new frictional condition, which may appear soon (ca. 0.1-0.2 s) after the object is initially gripped, might depend on the vigorous responses in the FA I units during the initial phase of the lifts (see Westling and Johansson 1987). The role of the tactile input in the adaptation of the force coordination to the frictional condition is discussed.

Adaptation, Physiological

Significance of cutaneous input for precise hand movements.

During precision manipulation of small objects a refined coordination of forces exerted on the object by the tips of the fingers and thumb is required. An essential feature of this coordination is that the grip forces change in parallel with the load forces produced to overcome various forces counteracting the intended manipulation. The balance between the grip forces and load forces tending to cause slips is adapted to the friction between the skin and the object, providing a relatively small safety margin to prevent slips. This adaptation, which is dependent on tactile afferent input from the fingers, works highly automatically. Tactile afferent signals account for an initial adjustment of the force balance to the current frictional condition, taking place soon after the object is initially touched. Moreover, brief bursts of action potentials elicited in dynamically sensitive units by small slips can trigger a change in this balance, resulting in an increased safety margin for the prevention of further slips. This change includes a rapid reflex response (latency 60-80 msec) together with updating of a sensorimotor memory maintaining the new force balance.

Action Potentials