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

Summary 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 move ment of the object were measured. Most subjects report 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. Furthermore, they did not perceive any difference between the objects' weight. Together, these results suggest that haptic exploration may be used to convey size information and further support the hypoth esis that size-related information may be combined with other sensory information in the programming of the precision grip.     

Old age impairs the use of arbitrary visual cues for predictive control of fingertip forces during grasp

Old age impairs the ability to form new associations for declarative memory, but the ability to acquire and retain procedural memories remains relatively intact. Thus, it is unclear whether old age affects the ability to learn the visuomotor associations needed to set efficient fingertip forces for handling familiar objects. We studied the ability for human subjects to use visual cues (color) about the mechanical properties (texture or weight) of a grasped object to control fingertip forces during prehension. Old and young adults (mean age 77 years and 22 years, respectively) grasped and lifted an object that varied in texture at the gripped surfaces (experiment 1: sandpaper versus acetate surface materials) or weight (experiment 2: 200 g versus 400 g). The object was color-coded according to the mechanical property in the "visual cue" condition, and the mechanical property varied unpredictably across lifts in the "no visual cue" condition. In experiment 1 (texture), the young adults' grip force (force normal to the gripped surface) when the object lifted from the support surface was 24% smaller when the surfaces were color-coded. The old adults' grip force did not vary between the visual conditions despite their accurate reports of the grip surface colors prior to each lift. Comparable findings were obtained in experiment 2, when object weight was varied and peak grip force rate prior to object lift-off was measured. Furthermore old and young subjects alike used about 2 N of grip force when lifting the 200 g object in experiment 2. Therefore, the old adults' failure to adjust grip force when the color cue was present cannot be attributed to a general inability or unwillingness to use low grip force when handling objects. We conclude that old age affects the associative learning that links visual identification of an object with the fingertip forces for efficiently handling the object. In contrast, old and young subjects' grip force was influenced by the preceding lift, regardless of visual cues, which supports existing theories that multiple internal representations govern predictive control of fingertip forces during prehension.     

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

Summary 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.     

The size of the visual size cue used for programming manipulative forces during precision grip

We used a perturbation technique to quantify the contribution of visual size cues to the programming of target force when lifting an object. Our results indicate that the nervous system attaches a reasonable weight to visual size cues when programming the target grip force for a novel object. In a subsequent lift of the same object, however, the confidence attached to the visual size cue fell dramatically. It is not clear whether the decrease in the use of size information was accelerated by the presence of a cue conflict or whether the fall represents the normal shift towards the use of a memory-based representation for programming grip force. In a second experiment, we used the "size-weight illusion" to explore the relationship between the verbal report of an object's weight and the programming of the grip and load force. We found that erroneous motor programming (as indexed by a number of measures) was neither necessary nor sufficient for the size-weight illusion to occur. These findings call for a re-evaluation of a previous explanation for the size-weight illusion. We suggest that the illusion arises because the cognitive system attempts to rationalise the fact that objects of apparently equal density but different size feel as if they have the same weight.     

Roles of glabrous skin receptors and sensorimotor memory in automatic control of precision grip when lifting rougher or more slippery objects

Summary To be successful, precision manipulation of small objects requires a refined coordination of forces excerted on the object by the tips of the fingers and thumb. The present paper deals quantitatively with the regulation of the coordination between the grip force and the vertical lifting force, denoted as the load force, while small objects were lifted, positioned in space and replaced by human subjects using the pinch grip. It was shown that the grip force changed in parallel with the load force generated by the subject to overcome various forces counteracting the intended manipulation. The balance between the two forces was adapted to the friction between the skin and the object providing a relatively small safety margin to prevent slips, i.e. the more slippery the object the higher the grip force at any given load force. Experiments with local anaesthesia indicated that this adaptation was dependent on cutaneous afferent input. Afferent information related to the frictional condition could influence the force coordination already about 0.1 s after the object was initially gripped, i.e. approximately at the time the grip and load forces began to increase in parallel. Further, secondary, adjustments of the force balance could occur later in response to small short-lasting slips, revealed as vibrations in the object. The new force balance following slips was maintained, indicating that the relationship between the two forces was set on the basis of a memory trace. Its updating was most likely accounted for by tactile afferent information entering intermittently at inappropriate force coordination, e.g. as during slips. The latencies between the onset of such slips and the appearance of the adjustments (0.06–0.08 s) clearly indicated that the underlying neural mechanisms operated highly automatically.     

Trajectory of human movement during sit to stand: a new modeling approach based on movement decomposition and multi-phase cost function

The ability to predict accurately the weights of objects is essential for skilled and dexterous manipulation. A potentially important source of information about object weight is through the observation of other people lifting objects. Here, we tested the hypothesis that when watching an actor lift an object, people naturally learn the object’s weight and use this information to scale forces when they subsequently lift the object themselves. Participants repeatedly lifted an object in turn with an actor. Object weight unpredictably changed between 2 and 7 N every 5th to 9th of the actor’s lifts, and the weight lifted by the participant always matched that previously lifted by the actor. Even though the participants were uninformed about the structure of the experiment, they appropriately adapted their lifting force in the first trial after a weight change. Thus, participants updated their internal representation about the object’s weight, for use in action, when watching a single lift performed by the actor. This ability presumably involves the comparison of predicted and actual sensory information related to actor’s actions, a comparison process that is also fundamental in action.     

Material evidence: interaction of well-learned priors and sensorimotor memory when lifting objects

Skilled object lifting requires the prediction of object weight. When lifting new objects, such prediction is based on well-learned size-weight and material-density correlations, or priors. However, if the prediction is erroneous, people quickly learn the weight of the particular object and can use this knowledge, referred to as sensorimotor memory, when lifting the object again. In the present study, we explored how sensorimotor memory, gained when lifting a given object, interacts with well-learned material-density priors when predicting the weight of a larger but otherwise similar-looking object. Different groups of participants 1st lifted 1 of 4 small objects 10 times. These included a pair of wood-filled objects and a pair of brass-filled objects where 1 of each pair was covered in a wood veneer and the other was covered in a brass veneer. All groups then lifted a larger, brass-filled object with the same covering as the small object they had lifted. For each lift, we determined the initial peak rate of change of vertical load-force rate and the load-phase duration, which provide estimates of predicted object weight. Analysis of the 10th lift of the small cube revealed no effects of surface material, indicating participants learned the appropriate forces required to lift the small cube regardless of object appearance. However, both surface material and core material of the small cube affected the 1st lift of the large block. We conclude that sensorimotor memory related to object density can contribute to weight prediction when lifting novel objects but also that long-term priors related to material properties can influence the prediction.     

The intermanual transfer of anticipatory force control in precision grip lifting is not influenced by the perception of weight

Passing objects from one hand to the other occurs frequently in our daily life. What kind of information about the weight of the object is transferred between the holding and lifting hand? To examine this, we asked people to hold (and heft) an object in one hand and then pick it up with the other. The objects were presented in the context of a size–weight illusion: that is, two objects of different sizes but the same weight were used. One group of participants held one of the objects in their left hand and then picked it up with their right. Another group of participants simply picked up the objects from a table. Thus, the former group had on-line information about the weight of the object, whereas the latter did not. Both groups showed a strong and equivalent size–weight illusion throughout the experiment. At the same time, the group that lifted the objects from the hefting hand applied equal grip force to the small and large object right from the start; in contrast, the group lifting the objects from the table, initially applied more grip force to the large than to the small object before eventually applying the same force to both. In two additional groups, a delay period was imposed between the lifting of the first and the second hands. The force parameters employed by these last two groups were virtually identical to those used by the group that lifted the object directly from the other hand. These results suggest that the initial calibration of grip force uses veridical information about the weight of the object provided by the other hand. This veridical information about weight is available on-line and is retained in memory for later access. The perceived weight of the object is basically ignored in forming grasping forces.     

The integration of size and weight cues for perception and action: evidence for a weight–size illusion

Humans routinely estimate the size and weight of objects. Yet, when lifting two objects of equal weight but different size, they often perceive the smaller object as being heavier. This size–weight illusion (SWI) is known to have a lesser effect on motor control of object lifting. How the nervous system combines “weight” and “size” cues with prior experience and whether these cues are differentially integrated for perception and sensorimotor action is still not fully understood. Therefore, we assessed not only whether the experience of size biases weight perception, but also if experience of weight biases the size perception of objects. Further, to investigate differences between perceptual and motor systems for cue-experience integration, participants haptically explored the weight of an object with one hand and then shaped the aperture of their other hand to indicate its perceived size. Results—First, next to a SWI, healthy adults (N = 21) perceived lighter objects as being smaller and heavier objects as being larger, demonstrating a weight–size illusion (WSI). Second, participants were more susceptible to either the SWI or WSI. Third, aperture of the non-exploring hand was scaled to perceived weight and not to physical size. Hand openings were consistently smaller than physical size, with SWI-sensitive participants being significantly more affected than WSI-sensitive subjects. We conclude: first, both size and weight perceptions are biased by prior experience. Weight perception is biased by expectations of size, while size perception is influenced by the expectancy of weight. Second, humans have the tendency to use one cue predominantly for both types of perception. Third, combining perceived weight with expected size influenced hand motor control, while online haptic feedback was largely ignored. Finally, we present a processing model underlying the size–weight cue integration for the perceptual and motor system.     

Influences of object weight and instruction on grip force adjustments

Summary This study investigated the influence of object weight and instructions on grip force responses in humans. Using a precision grip, subjects lifted a small instrumented test object to a predetermined height. Prior to each set of 40 trials, subjects were verbally instructed to either hold or let go of the object in response to any change in weight. Unpredictably on some trials (< 20%), a sudden sustained increase (load) or decrease (unload) in vertical load was applied to the object. Grip responses to these induced weight changes were evaluated by measuring grip force, object position, and associated electromyographic (EMG) activity. Grip force changes for a load were over three times greater than those for an unload. Such asymmetry may reflect everyday grasp and manipulation in a gravity-influenced world. Grip force adjustments to loads following hold instructions were on the average somewhat larger than those following let go instructions, but there was no influence of instructions on responses to unloads. These findings contrast with more robust influences of verbal instruction on automatic postural and proximal upper limb responses and also may suggest that grip force adjustments are influenced to a greater extent by intrinsic task variables than by extrinsic volitional intent. Such organization appears tailored to functional task requirements in natural environmental contexts.     

Anticipatory scaling of grip forces when lifting objects of everyday life

The ability to predict and anticipate the mechanical demands of the environment promotes smooth and skillful motor actions. Thus, the finger forces produced to grasp and lift an object are scaled to the physical properties such as weight. While grip force scaling is well established for neutral objects, only few studies analyzed objects known from daily routine and none studied grip forces. In the present study, eleven healthy subjects each lifted twelve objects of everyday life that encompassed a wide range of weights. The finger pads were covered with force sensors that enabled the measurement of grip force. A scale registered load forces. In a control experiment, the objects were wrapped into paper to prevent recognition by the subjects. Data from the first lift of each object confirmed that object weight was anticipated by adequately scaled forces. The maximum grip force rate during the force increase phase emerged as the most reliable measure to verify that weight was actually predicted and to characterize the precision of this prediction, while other force measures were scaled to object weight also when object identity was not known. Variability and linearity of the grip force–weight relationship improved for time points reached after liftoff, suggesting that sensory information refined the force adjustment. The same mechanism seemed to be involved with unrecognizable objects, though a lower precision was reached. Repeated lifting of the same object within a second and third presentation block did not improve the precision of the grip force scaling. Either practice was too variable or the motor system does not prioritize the optimization of the internal representation when objects are highly familiar.     

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

Summary 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.     

Effect of human grip strategy on force control in precision tasks

Alternate grip strategies are often used for object manipulation in individuals with sensorimotor deficits. To determine the effect of grip type on force control, ten healthy adult subjects were asked to grip and lift a small manipulandum using a traditional precision grip (lateral pinch), a pinch grip with the fingers oriented downwards (downward pinch) and a key grip between the thumb and the side of the index finger. The sequence of grip type and hand used was varied randomly after every ten lifts. Each of the three grips resulted in different levels of force, with the key grip strategy resulting in the greatest grip force and the downward pinch grip using the least amount of grip force to lift the device. Cross-correlation analysis revealed that the ability to scale accurately the rate of grip force and load force changes was lowest in the downward pinch grip. This was also associated with a more variable time-shift between the two forces, indicating that the precise anticipatory control when lifting an object is diminished in this grip strategy. There was a difference between hands across all grips, with the left non-dominant hand using greater grip force during the lift but not the hold phase. Further, in contrast with the right hand, the left hand did not reduce grip force during the lift or the hold phase over the ten lifts, suggesting that the non-dominant hand did not quickly learn to optimise grip force. These findings suggest that the alternate grip strategies used by patients with limited fine motor control, such as following stroke, may partly explain the disruption of force control during object manipulation.     

Failure to disrupt the ‘sensorimotor’ memory for lifting objects with a precision grip

When repetitively lifting an object with mechanical properties that vary from lift-to-lift, the fingertip forces for gripping and lifting are influenced strongly by the previous lift, revealing a ‘sensorimotor’ memory. Two recent reports indicate that the sensorimotor memory for grip force is easily disrupted by an unrelated task like a strong pinch or vibration, even when the lift was performed with the hand contralateral to the vibration or preceding pinch. These findings indicate that this memory may reflect sensory input or muscle contraction levels, rather than object properties or the specific task of gripping and lifting. Here we report that the predictive scaling of lift force was not disrupted by conditioning tasks that featured exerting a vertical isometric force with the upper extremity. When subjects lifted a 2 N object repetitively the peak lift force rate was 26.4 N/s. The lift force rate increased to 36.1 N/s when the 2 N object was lifted (regardless of hand) after lifting the 8 N object with the right hand, which reveals the expected ‘sensorimotor’ memory. The lift force rate did not increase (24.8 vs. 26.4 N/s for the control condition) when a bout of isometric exertion (9.8 N) in the vertical direction with the distal right forearm preceded lifts of the 2 N object. This finding was confirmed with another isometric task designed to more closely mimic lifting an object with a precision grip. This difference in the sensitivity of grip versus lift force to a preceding isometric contraction indicates that separate sensorimotor memories contribute to the predictive scaling of the commands for gripping and lifting an object.     

Lifting a familiar object: visual size analysis, not memory for object weight, scales lift force

The brain can accurately predict the forces needed to efficiently manipulate familiar objects in relation to mechanical properties such as weight. These predictions involve memory or some type of central representation, but visual analysis of size also yields accurate predictions of the needed fingertip forces. This raises the issue of which process (weight memory or visual size analysis) is used during everyday life when handling familiar objects. Our aim was to determine if subjects use a sensorimotor memory of weight, or a visual size analysis, to predictively set their vertical lift force when lifting a recently handled object. Two groups of subjects lifted an opaque brown bottle filled with water (470 g) during the first experimental session, and then rested for 15 min in a different room. Both groups were told that they would lift the same bottle in their next session. However, the experimental group returned to lift a slightly smaller bottle filled with water (360 g) that otherwise was identical in appearance to the first bottle. The control group returned to lift the same bottle from the first session, which was only partially filled with water so that it also weighed 360 g. At the end of the second session subjects were asked if they observed any changes between sessions, but no subject indicated awareness of a specific change. An acceleration ratio was computed by dividing the peak vertical acceleration during the first lift of the second session by the average peak acceleration of the last five lifts during the first session. This ratio was >1 for the control subjects 1.30 (SEM 0.08), indicating that they scaled their lift force for the first lift of the second session based on a memory of the (heavier) bottle from the first session. In contrast, the acceleration ratio was 0.94 (0.10) for the experimental group (P < 0.011). We conclude that the experimental group processed visual cues concerning the size of the bottle. These findings raise the possibility that even with familiar objects we predict fingertip forces using an on-line visual analysis of size (along with memory of density), rather than accessing memory related to object weight.     

Opposite perceptual and sensorimotor responses to a size-weight illusion

The perceptual size-weight illusion (SWI) occurs when two different-sized objects with equal mass are lifted in sequence: the smaller object is consistently reported to feel heavier than the larger object even after repeated lifting attempts. Here we explored the relationship between sensorimotor and perceptual responses to a SWI in which the smaller of the two target objects in fact weighed slightly less (2.7 N) than the larger object (3.2 N). For 20 consecutive lifts, participants consistently reported that the small-light object felt heavier than the large-heavy object; however, concurrently measured lifting dynamics showed exactly the opposite pattern: peak grip force, peak grip force rate, peak load force, and peak load force rate were all significantly greater for the large-heavy object versus the small-light object. The difference in peak load rate between the two objects was greatest for the initial lift but decreased significantly beyond that point, suggesting that the sensorimotor system used sensory feedback to correct for initial over- and underestimations of object mass. Despite these adjustments to lifting dynamics over the early trials, the difference between the judged heaviness of the two objects did not change. The findings clearly demonstrate that the sensorimotor and perceptual systems utilize distinctly different mechanisms for determining object mass.     

Object size effects on initial lifting forces under microgravity conditions

Individuals usually report for two objects of equal mass but different volume that the larger object feels lighter. This so-called size-weight illusion has been investigated for more than a century. The illusion is accompanied by increased forces, used to lift the larger object, resulting in a higher initial lifting speed and acceleration. The illusion holds when subjects know that the mass of the two objects is equal and it is likely that this also counts for the enlarged initial effort in lifting a larger box. Why should this happen? Under microgravity, subjects might be able to eliminate largely the weight-related component of the lifting force. Then, if persistent upward scaling of the weight-related force component had been the main cause of the elevated initial lifting force under normal gravity, this elevated force might disappear under microgravity. On the other hand, the elevated initial lifting effort in the large box would be preserved if it had been caused mainly by a persistent upward scaling of the force component, necessary to accelerate the object. To test whether the elevated initial lifting effort either persists or disappears under microgravity, a lifting experiment was carried out during brief periods of microgravity in parabolic flights. Subjects performed whole-body lifting movements with their feet strapped to the floor of the aircraft, using two 8-kg boxes of different volume. The subjects were aware of the equality of the box masses. The peak lifting forces declined almost instantaneously with approx. a factor 9 in the first lifting movements under microgravity compared with normal gravity, suggesting a rapid adaptation to the loss of weight. Though the overall speed of the lifting movement decreased under microgravity, the mean initial acceleration of the box over the first 200 ms of the lifting movement remained higher (P=0.030) in the large box (1.87±0.127 m/s²) compared with the small box (1.47±0.122 m/s²). Under normal gravity these accelerations were 3.30±0.159 m/s² and 2.67±0.159 m/s², respectively (P=0.008). A comparable trend was found in the initial lifting forces, being significant in the pooled gravity conditions (P=0.036) but not in separate tests on the normal gravity (P=0.109) and microgravity (P=0.169) condition. It is concluded that the elevated initial lifting effort with larger objects holds during short-term exposure to microgravity. This suggests that upward scaling of the force component, required to accelerate the larger box, is an important factor in the elevated initial lifting effort (and the associated size-weight illusion) under normal gravity.     

Prehension in the pigeon

Summary Eating in the pigeon involves a series of jaw movements some of which serve a prehensile function; i.e., they are utilized in the grasping and manipulation of objects. These prehensile behaviors are extremely brief (30–80 ms), produce an adjustment of jaw opening amplitude to the size of the food object, are mediated by an effector system involving a relatively small number of muscles and are amenable to both reflexive and voluntary control. This combination of structural simplicity and functional complexity suggests that the pigeon's jaw movements may provide a useful model system for the study of motor control mechanisms in targeted movements. The present report provides a classification of jaw opening movements occurring during eating and a preliminary determination of the extent to which each movement class is scaled to the size of the food object. Jaw movements were monitored during responses to spherical food pellets of six different sizes (3.2–11.1 mm in diameter) using a transducing system which produces a continuous record of gape (i.e., interbeak distance). Assignment to movement classes was then carried out using a computer-assisted scoring program. Functions relating jaw opening amplitude to target size were determined for each movement class. Four jaw movement classes were identified: Prepecks (just prior to pecking), Grasps (opening movements made during pecking but prior to contact with the target), Mandibulations (movements serving to position and transport the object within the buccal cavity) and Swallows. For two of these movement classes (Grasps, Mandibulations) jaw opening amplitude is scaled to pellet size but the scaling functions differ in ways that reflect the functional requirements of the two behaviors. However, for both movements, the data suggest that information about initial gape is used to control opening amplitude. It is concluded that during Grasping, the adjustment of opening amplitude to stimulus size involves visual inputs and open-loop control mechanisms, while for Mandibulation, that adjustment involves tactile input and closed-loop mechanisms.     

Posture-based or trajectory-based movement planning: a comparison of direct and indirect pointing movements

Various models have been proposed in the literature to explain the control of human arm movements. To make a quantitative comparison between the predictions of various models, we tested subjects for movements to targets on a vertical screen in various conditions. Subjects were asked to move directly from one target to another, or to move by a via-point, at various movement velocities and in a condition with a weight of 0.6 kg attached to the forearm. This set of experimental data was used for comparison with the predictions by various posture-based and trajectory-based models on 3-D movement planning and control. Small but significant effects of starting position and path towards the target were found on the torsion of the arm at the end of the movement. No effects of movement velocity and weight attached to the forearm were found. The experimental results differed significantly from the predictions by any of the models considered. Of the models considered, Donders law best predicts the experimental data. Our data indicate that future tests of models for motor control (1) should compare the predictions of not just one, but several models to a data set, and (2) should include not only planar, but rather 3-D movements in such a comparison.     

Unconscious updating of grasp motor program

Grasp modification during prehension movements was studied in response to slight variations of somesthetic information about object size. Three experiments were carried out. In experiment 1 eight subjects were required to reach and grasp an object whose size could either increase or decrease, whereas its visual image remained unmodified. The object size was changed during the experiment with uninformed subjects after a block of trials during which visual and somesthetic information were congruent. At the end of the experiment subjects were required to reproduce the size of the object with their fingers (matching test). Results showed that maximal grip aperture during prehension as well as finger aperture in the matching test were modified according to variation in object size, although no subject realized that the object had changed during the experiment. Grasp time was also altered by object size change. Greater and earlier adaptation in maximal grip aperture, as well as perturbation of grasp time, were observed for decrease than for increase in object size. However, complete compensation was never reached for both parameters. Constant confidence in vision could have prevented both complete compensation and conscious detection of object change. This was investigated in two additional experiments. In experiment 2 visual information was made unreliable by informing subjects about variation in grasped object size. This led to greater and earlier modification in maximal grip aperture than in experiment 1. Grasp time was kept almost constant regardless of size variation. In experiment 3 vision of the stimulus was prevented and no information on change in object size was given to subjects. The results of experiment 3 were similar to those of experiment 1, although modification in maximal grip aperture was larger for increase in object size. Correspondingly, grasp time was more affected by increase than by decrease in object size. The results of the three experiments suggest that kinematic parameters usually considered as dependent on object properties, such as maximal grip aperture, were modified in order to compensate perturbation of temporal parameters. This modification induced a pragmatic knowledge of object size (as showed by the results of the matching test), although awareness was not reached.