The representation of predictive force control and internal forward models: evidence from lesion studies and brain imaging

Force control on the basis of prediction avoids time delays from sensory feedback during motor performance. Thus, self-produced loads arising from gravitational and inertial forces during object manipulation can be compensated for by simultaneous anticipatory changes in grip force. It has been suggested that internal forward models predict the consequences of our movements, so that grip force can be programmed in anticipation of movement-induced loads. The cerebellum has been proposed as the anatomical correlate of such internal models. Here, we present behavioural data from patients with cerebellar damage and data from brain imaging in healthy subjects further elucidating the role of the cerebellum in predictive force control. Patients with cerebellar damage exhibited clear deficits in the coupling between grip force and load. A positron-emission-tomography (PET) paradigm that separated the process of the grip force/load coupling from the isolated production of similar grip forces and loads was developed. Interaction and conjunction analyses revealed a strong activation peak in the ipsilateral posterior cerebellum particularly devoted to the predictive coupling between grip force and load. Both approaches clearly demonstrate that the cerebellum plays a major role in force prediction that cannot be compensated for by other sensorimotor structures in case of cerebellar disease. However, evidence suggests that also extra-cerebellar structures may significantly contribute to predictive force control: (1) grip force/load coupling may also be impaired after cerebral and peripheral sensorimotor lesions, (2) a coupling-related activation outside the cerebellum was observed in our PET study, and (3) the scaling of the grip force level and the dynamic grip force coupling are dissociable aspects of grip force control.     

Deficits of anticipatory grip force control after damage to peripheral and central sensorimotor systems

Healthy subjects adjust their grip force economically to the weight of a hand-held object. In addition, inertial loads, which arise from arm movements with the grasped object, are anticipated by parallel grip force modulations. Internal forward models have been proposed to predict the consequences of voluntary movements. Anesthesia of the fingers impairs grip force economy but the feedforward character of the grip force/load coupling is preserved. To further analyze the role of sensory input for internal forward models and to characterize the consequences of central nervous system damage for anticipatory grip force control, we measured grip force behavior in neurological patients. We tested a group of stroke patients with varying degrees of impaired fine motor control and sensory loss, a single patient with complete and permanent differentation from all tactile and proprioceptive input, and a group of patients with amyotrophic lateral sclerosis (ALS) that exclusively impairs the motor system without affecting sensory modalities. Increased grip forces were a common finding in all patients. Sensory deficits were a strong but not the only predictor of impaired grip force economy. The feedforward mode of grip force control was typically preserved in the stroke patients despite their central sensory deficits, but was severely disturbed in the patient with peripheral sensory deafferentation and in a minority of stroke patients. Moderate deficits of feedforward control were also obvious in ALS patients. Thus, the function of the internal forward model and the precision of grip force production may depend on a complex anatomical and functional network of sensory and motor structures and their interaction in time and space.     

Motor learning of cue-dependent pull-force changes during an isometric precision grip task

The “raspberry task” represents a precision grip task that requires continuous adjustment of grip and pull forces. During this task subjects grip a specialized grip rod and have to increase the pull force linearly while the rod is locked. The aim of this study was to determine whether an associated, initially neutral cue is able to evoke pull-force changes in the raspberry task. A standard delay paradigm was used to study cued pull-force changes during an ongoing movement resulting in unloading. Pull force and EMG activity of hand and arm muscles were recorded from 13 healthy, young subjects. The cue was associated with a complex change in motor behavior.In this task, cued force changes take place more rapidly than in protective reflex systems (in median after the second presentation of the cueing stimulus). A cued force change was detectable in two-thirds of paired trials. Although the force change is produced by a decrease of the EMG activity in several grip- and pull-force-producing muscles, the most significant effect in the majority of the subjects was an increase of the activity of the flexor carpi ulnaris muscle which antagonises corresponding pull-force-producing muscles. Cued force changes require adequately and precisely controlled activation of the muscle groups involved in the movement.     

Generalization as a behavioral window to the neural mechanisms of learning internal models

In generating motor commands, the brain seems to rely on internal models that predict physical dynamics of the limb and the external world. How does the brain compute an internal model? Which neural structures are involved? We consider a task where a force field is applied to the hand, altering the physical dynamics of reaching. Behavioral measures suggest that as the brain adapts to the field, it maps desired sensory states of the arm into estimates of force. If this neural computation is performed via a population code, i.e., via a set of bases, then activity fields of the bases dictate a generalization function that uses errors experienced in a given state to influence performance in any other state. The patterns of generalization suggest that the bases have activity fields that are directionally tuned, but directional tuning may be bimodal. Limb positions as well as contextual cues multiplicatively modulate the gain of tuning. These properties are consistent with the activity fields of cells in the motor cortex and the cerebellum. We suggest that activity fields of cells in these motor regions dictate the way we represent internal models of limb dynamics.     

CUE: The continuous unified electronic diary method

In the present article, we introduce the continuous unified electronic (CUE) diary method, a longitudinal, event-based, electronic parent report method that allows real-time recording of infant and child behavior in natural contexts. Thirty-nine expectant mothers were trained to identify and record target behaviors into programmed handheld computers. From birth to 18?months, maternal reporters recorded the initial, second, and third occurrences of seven target motor behaviors: palmar grasp, rolls from side to back, reaching when sitting, pincer grip, crawling, walking, and climbing stairs. Compliance was assessed as two valid entries per behavior: 97?% of maternal reporters met compliance criteria. Reliability was assessed by comparing diary entries with researcher assessments for three of the motor behaviors: palmar grasp, pincer grip and walking. A total of 81?% of maternal reporters met reliability criteria. For those three target behaviors, age of emergence was compared across data from the CUE diary method and researcher assessments. The CUE diary method was found to detect behaviors earlier and with greater sensitivity to individual differences. The CUE diary method is shown to be a reliable methodological tool for studying processes of change in human development.     

Conflict in object affordance revealed by grip force

Viewing objects can result in automatic, partial activation of motor plans associated with them-"object affordance". Here, we recorded grip force simultaneously from both hands in an object affordance task to investigate the effects of conflict between coactivated responses. Participants classified pictures of objects by squeezing force transducers with their left or right hand. Responses were faster on trials where the object afforded an action with the same hand that was required to make the response (congruent trials) compared to the opposite hand (incongruent trials). In addition, conflict between coactivated responses was reduced if it was experienced on the preceding trial, just like Gratton adaptation effects reported in "conflict" tasks (e.g., Eriksen flanker). This finding suggests that object affordance demonstrates conflict effects similar to those shown in other stimulus-response mapping tasks and thus could be integrated into the wider conceptual framework on overlearnt stimulus-response associations. Corrected erroneous responses occurred more frequently when there was conflict between the afforded response and the response required by the task, providing direct evidence that viewing an object activates motor plans appropriate for interacting with that object. Recording continuous grip force, as here, provides a sensitive way to measure coactivated responses in affordance tasks.     

Conflict in object affordance revealed by grip force

Viewing objects can result in automatic, partial activation of motor plans associated with them—“object affordance”. Here, we recorded grip force simultaneously from both hands in an object affordance task to investigate the effects of conflict between coactivated responses. Participants classified pictures of objects by squeezing force transducers with their left or right hand. Responses were faster on trials where the object afforded an action with the same hand that was required to make the response (congruent trials) compared to the opposite hand (incongruent trials). In addition, conflict between coactivated responses was reduced if it was experienced on the preceding trial, just like Gratton adaptation effects reported in “conflict” tasks (e.g., Eriksen flanker). This finding suggests that object affordance demonstrates conflict effects similar to those shown in other stimulus–response mapping tasks and thus could be integrated into the wider conceptual framework on overlearnt stimulus–response associations. Corrected erroneous responses occurred more frequently when there was conflict between the afforded response and the response required by the task, providing direct evidence that viewing an object activates motor plans appropriate for interacting with that object. Recording continuous grip force, as here, provides a sensitive way to measure coactivated responses in affordance tasks.     

Effects of Sensory Deficit on Phalanx Force Deviation During Power Grip Post Stroke

The effect of sensory deficits on power grip force from individual phalanges was examined. The authors found that stroke survivors with sensory deficits (determined by the Semmes-Weinstein monofilament test) gripped with phalanx force directed more tangential to the object surface, than those without, although both groups had similar motor deficits (Chedoke-McMaster and Fugl-Meyer), grip strength, and skin friction. Altered grip force direction elevates risk of finger slippage against the object thus grip loss/object dropping, hindering activities of daily living. Altered grip force direction was associated with altered muscle activation patterns. In summary, the motor impairment level alone may not describe hand motor control in detail. Information about sensory deficits helps elucidate patients' hand motor control with functional relevance.     

Viewing objects and planning actions: on the potentiation of grasping behaviours by visual objects

How do humans interact with tools? Gibson (1979) suggested that humans perceive directly what tools afford in terms of meaningful actions. This “affordances” hypothesis implies that visual objects can potentiate motor responses even in the absence of an intention to act. Here we explore the temporal evolution of motor plans afforded by common objects. We presented objects that have a strong significance for action (pinching and grasping) and objects with no such significance. Two experimental tasks involved participants viewing objects presented on a computer screen. For the first task, they were instructed to respond rapidly to changes in background colour by using an apparatus mimicking precision and power grip responses. For the second task, they received stimulation of their primary motor cortex using transcranial magnetic stimulation (TMS) while passively viewing the objects. Muscular responses (motor evoked potentials: MEPs) were recorded from two intrinsic hand muscles (associated with either a precision or power grip). The data showed an interaction between type of response (or muscle) and type of object, with both reaction time and MEP measures implying the generation of a congruent motor plan in the period immediately after object presentation. The results provide further support for the notion that the physical properties of objects automatically activate specific motor codes, but also demonstrate that this influence is rapid and relatively short lived.     

Hand Grip and Load Force Coordination of the Ipsilesional Hand of Chronic Stroke Individuals

Object manipulation depends on a refined control of grip force (GF) and load force (LF). After a brain injury, the GF control is altered in the paretic hand but what happens with the non-paretic hand is still unclear. In this study, we compared the GF control and GF–LF coordination of the non-paretic hand of 10 stroke individuals who suffered right brain damage (RBD) and 10 who suffered left brain damage (LBD), with 20 healthy individuals during lifting and oscillation task, using an instrumented object. GF was recorded with a force transducer, and LF was estimated from the object weight and acceleration. Overall, the ipsilesional hand of stroke individuals, independent of the lesion side, presented similar GF control and GF–LF coordination. However, LBD individuals took longer to start lifting the object, which may be due to the need of more time to obtain somatosensory information from the contact with the object. The findings indicate that stroke individuals preserve their ability to control and coordinate GF and LF when using their ipsilesional hand for object manipulation and the left hemisphere may play an essential role in the processing of somatosensory information needed for the GF control.     

计算机时代的书写心理学

计算机的普及使人们阅读及书写的能力降低, 这引发了社会对书写的关注。书写涉及大量高级神经活动, 与认知、人格、疾病等均有关联, 对个体发展也具有重要作用。现代书写心理学通过使用书写板等设备, 实现了对书写过程及结果的量化研究, 发现了书写速度、压力、握笔等指标与智力、认知负荷等的一些关联。未来对书写过程的研究可以进一步深入; 测量方面的研究成果可以应用到日常人机交互及教学、司法等实践中; 还可以研究用书写促进个体认知及人格发展的具体途径。     

Bimanual Lifting: Do Fingertip Forces Work Independently or Interactively?

Bimanual coordination is a commonplace activity, but the consequences of using both hands simultaneously are not well understood. The authors examined fingertip forces across 4 experiments in which participants undertook a range of bimanual tasks. They first measured fingertip forces during simultaneous lifts of 2 identical objects, noting that individuals held the objects with more force bimanually than unimanually. They then varied the mass of the objects held by each hand, noting that when both hands lifted together performance was equivalent to unimanual lifts. The authors next measured one hand's static grip force while the other hand lifted an object. They found a gradual reduction of grip force throughout the trial, but once again no evidence of one hand influencing the other. In the final experiment the authors tested whether tapping with one hand could influence the static grip force of its counterpart. Although the authors found no changes in static grip force as a direct consequence of the other hand's actions, they found clear differences from one task to the other, suggesting an effect of task instruction. Overall, these results suggest that fingertip forces are largely independent between hands in a bimanual lifting context, but are sensitive to different task requirements.     

The stability of precision grip force in older adults

The exceedingly large grip forces that many older adults employ when lifting objects with a precision pinch grip (Cole, 1991) may compensate for a reduced capability to produce a stable isometric force. That is, their grip force may fluctuate enough from moment to moment to yield grip forces that approach the force at which the object would slip from grasp. We examined the within-trial variability of isometric force in old (68-85 years, n = 13) and young (n = 11) human subjects (a) when they were asked to produce a constant pinch force at three target levels (0.49, 2.25, and 10.5 N) with external support of the arm, hand, and force transducer and (b) when they were asked to grasp, lift, and hold a small test object with a precision grip. Pinch force produced in the first task was equally stable across the two subject groups during analysis intervals that lasted 4 s. The elderly subjects produced grip forces when lifting objects that averaged twice as much as those produced by the young subjects. The force variability during the static (hold) phase of the lift for the old subjects was comparable with that used by the young subjects, after adjusting for the difference in grip force. The failure to observe less stable grip force in older adults contradicts a similar recent study. Differences in task (isometric grip force versus isometric abduction torque of a single digit) may account for this conflict, however. Thumb and finger forces for grip are produced through coactivation of many muscles and thus promote smooth force output through temporal summation of twitches. We conclude that peripheral reorganization of muscle in older adults does not yield increased instability of precision grip force and therefore does not contribute directly to increased grip forces in this population. However, force instability may affect other grip configurations (e.g., lateral pinch) or manipulation involving digit abduction or adduction forces.     

Motor prediction at the edge of instability: alteration of grip force control during changes in bimanual coordination

Predicting the consequences of actions is fundamental for skilled motor behavior. We investigated whether motor prediction is influenced by the fact that some movements are easier to perform and stabilize than others. Twelve subjects performed a bimanual rhythmical task either symmetrically or asymmetrically (the latter being more difficult and less stable) while oscillating in each hand an object attached to an elastic cord. Motor prediction was monitored through the adequacy of anticipatory grip force adjustments with respect to the elastic resisting force. Results showed less adequate predictive control during asymmetrical movements (compared with symmetrical ones). Furthermore, switching between modes of coordination induced even larger alterations. An interesting finding was that grip force control did not always stabilize around the expected value after voluntary transition. We conclude that motor prediction is affected by the degree of coordination between the upper limbs and by phase transitions and is prone to carryover effects.     

Creating number semantics through finger movement perception

Communication, language and conceptual knowledge related to concrete objects may rely on the sensory–motor systems from which they emerge. How abstract concepts can emerge from these systems is however still unknown. Here we report a functional interaction between a specific meaningful finger movement, such as a finger grip closing, and a concept as abstract as numerical magnitude. Participants were presented with Arabic digits to recall before or after they perceived a biological or non-biological hand movement. The results show that perceiving a grip closing slows down the processing of large magnitude numbers. Importantly, we show that this motor-to-semantic interaction differs from the reverse semantic-to-motor interaction, and that it does not result from a general movement amplitude processing as it is only observed for biological hand movements. These results demonstrate the functional link between number meaning and goal-directed finger movements, and show how abstract concept semantics can emerge from the sensory–motor circuits of the brain.     

Relations between the inflection point on the force-time curve and force-time parameters during static explosive grip

Individual differences in muscle contractile speed during static explosive muscle contraction are reflected in the developmental phase of the force-time curve. The purposes of this study were to clarify the properties and reliability of the inflection point of force-time, statistically dividing speed during static explosive grip into two phases and to assess the relations between that inflection point and others. Static explosive grip data were measured two times with a 5-min. rest (sampling frequency; 100 Hz). 32 healthy, young men (age: 15.5 +/- 0.8 yr., height: 173.9 +/- 7.3 cm, body mass: 71.5 +/- 11.2 kg) participated. 8 static explosive grip parameters were selected: time of reaching, integrated area, and quotient values of the integrated areas up to 0.25, 0.5, and 1.0 sec. divided by maximal grip force. The inflection point was calculated statistically from two regression lines fitted to a developmental phase and the almost steady-state phase of reaching maximal grip force by applying a two-phase regression model. The reliabilities of maximal grip force, time of reaching 90% of maximal grip force, and the integrated area until 0.5 sec. and 1.0 sec. after the onset of grip were good (ICC=.77 to .93). The time of reaching an inflection force value appeared at 0.3 sec. after the onset of grip, corresponding to 80% of maximal grip force, and the reliabilities of the parameters regarding inflection point were good (ICC=.77 to .95). The time determined by boundary data between the former and the latter regression data set and the regression coefficient during the developmental phase correlated significantly with the time of reaching 90% of maximal grip force, the integrated area, and the quotient values of the integrated areas up to 0.25, 0.5, and 1.0 sec. divided by maximal grip force (rs=-.78 to -.96 and -.75 to 0.88, respectively, p<.05). However, these parameters did not correlate with maximal grip force. A force during the developmental phase and maximal grip force can depend on different physiological factors. The time determined by boundary data between the former and the latter regression data set and the regression coefficient during the developmental phase are useful parameters for evaluating static explosive grip.     

Right, left or both? Brain hemispheres and apraxia of naturalistic actions

Neuropsychology allows us to see more clearly how brain processes are organized by looking at the way they fall apart. For example, a hundred years of research on brain-damaged individuals with deficits of higher-level motor abilities have shed light on the organization of normal motor abilities. A consistent finding has been the dominant role of the left hemisphere in motor control in right-handers. However, the recent paper of Hartmann et al. challenges this tradition by suggesting that both hemispheres can contribute to complex object and tool use.     

Grip morphology and hand use in chimpanzees (Pan troglodytes): evidence of a left hemisphere specialization in motor skill

Three experiments on grip morphology and hand use were conducted in a sample of chimpanzees. In Experiment 1, grip morphology when grasping food items was recorded, and it was found that subjects who adopted a precision grip were more right-handed than chimpanzees using other grips. In Experiment 2, the effect of food type on grasping was assessed. Smaller food items elicited significantly more precision grips for the right hand. In Experiment 3, error rates in grasping foods were compared between the left and right hands. Significantly more errors were made for the left compared with the right hand. The cumulative results indicate that chimpanzees show a left-hemisphere asymmetry in motor skill that is associated with the use of precision grips.     

A study of a bimanual synergy associated with holding an object

The task of supporting an object with one or two hands was used to test the applicability of the notion of synergy. Subjects sat with their dominant forearm supported up to the wrist while holding a cylindrical “cup” between their thumb and fingers. Force transducers recorded the grip force applied normal to the cup's side by the thumb and the force applied normal to the cup bottom. On different series, a supporting force was added to and released from the bottom of the cup by the subject's non-dominant hand or by the experimenter. As predicted, the results indicated feedforward adjustments of the grip force, and of the EMGs, and significant correlations between grip force and supporting force when they were produced by two hands of one person, and the lack of such closely tied changes when the two forces were produced by two different persons. In the latter case, different subjects could demonstrate grip force changes in different directions. The findings suggest that grip force adjustments represented peripheral patterns of a single central process (a single synergy) rather than being separately controlled focal and postural components of the action.PsycINFO classification: 2330