Motor Equivalence Research Articles

Anticipatory Planning and Control of Grasp Positions and Forces for Dexterous Two-Digit Manipulation

Dexterous object manipulation requires anticipatory control of digit positions and forces. Despite extensive studies on sensorimotor learning of digit forces, how humans learn to coordinate digit positions and forces has never been addressed. Furthermore, the functional role of anticipatory modulation of digit placement to object properties remains to be investigated. We addressed these questions by asking human subjects (12 females, 12 males) to grasp and lift an inverted T-shaped object using precision grip at constrained or self-chosen locations. The task requirement was to minimize object roll during lift. When digit position was not constrained, subjects could have implemented many equally valid digit position-force coordination patterns. However, choice of digit placement might also have resulted in large trial-to-trial variability of digit position, hence challenging the extent to which the CNS could have relied on sensorimotor memories for anticipatory control of digit forces. We hypothesized that subjects would modulate digit placement for optimal force distribution and digit forces as a function of variable digit positions. All subjects learned to minimize object roll within the first three trials, and the unconstrained device was associated with significantly smaller grip forces but larger variability of digit positions. Importantly, however, digit load force modulation compensated for position variability, thus ensuring consistent object roll minimization on each trial. This indicates that subjects learned object manipulation by integrating sensorimotor memories with sensory feedback about digit positions. These results are discussed in the context of motor equivalence and sensorimotor integration of grasp kinematics and kinetics.

Immune Algorithm Based Non-Intrusive Efficiency Determination for In-Situ Induction Motor

On average, industrial motors operate at less than 60% of their rated load, and thus at reduced efficiency, which results in wasted energy. The evaluation of motor efficiency enables energy savings in industry. However, because of the uninterrupted characteristic of industrial process, traditional methods cannot be used for in-situ motors. This paper proposes an immune algorithm based non-intrusive method for in-situ motor efficiency determination based on a modified induction motor equivalence circuit. Rotor stray load loss is considered in the equivalence circuit by adding an equivalent resister in series with the rotor circuit. The proposed method is verified by testing a 5 HP (3.75 kW) squirrel cage induction motor, and the results are compared with the torque-gauge method and the conventional equivalent circuit parameters estimation method using no-load and locked-rotor tests.

Shared neural resources between left and right interlimb coordination skills: The neural substrate of abstract motor representations

Functional magnetic resonance imaging was used to reveal the shared neural resources between movements performed with effectors of the left versus right body side. Prior to scanning, subjects extensively practiced a complex coordination pattern involving cyclical motions of the ipsilateral hand and foot according to a 90° out-of-phase coordination mode. Brain activity associated with this (nonpreferred) coordination pattern was contrasted with pre-existing isodirectional (preferred) coordination to extract the learning-related brain networks. To identify the principal candidates for effector-independent movement encoding, the conjunction of training-related activity for left and right limb coordination was determined. A dominantly left-lateralized parietal-to-(pre)motor activation network was identified, with activation in inferior and superior parietal cortex extending into intraparietal sulcus and activation in the premotor areas, including inferior frontal gyrus (pars opercularis). Similar areas were previously identified during observation of complex coordination skills by expert performers. These parietal-premotor areas are principal candidates for abstract (effector-independent) movement encoding, promoting motor equivalence, and they form the highest level in the action representation hierarchy.

Inter-speaker variability and the articulatory-acoustic relations in German and English /sh/

The aim of this study is to investigate speaker-specific articulatory-acoustic relations and their potential causes in the realisation of /sh/. By means of electropalatographic and acoustic data for 12 German and 12 English native speakers (6 males and 6 females for each language), we obtained the following results: 1) relatively invariant COG values of the frication noise among the subjects, 2) huge inter-speaker variability in the articulatory place of articulation, especially for the males. We suggest that in the realisation of /sh/ motor equivalence principles can be used speaker-dependently. In particular, the length of palatal channel, the size of the front cavity (including its sublingual portion) as well as lip protrusion are the relevant articulatory parameters involved in the adjustment of the frication noise. Speaker-specific articulatory strategies will be discussed in the light of the formant transition characteristics, palatal morphology, and the prototypical articulatory patterns will be illustrated by means of acoustic modelling.

Action-dependent perceptual invariants: From ecological to sensorimotor approaches

Ecological and sensorimotor theories of perception build on the notion of action-dependent invariants as the basic structures underlying perceptual capacities. In this paper we contrast the assumptions these theories make on the nature of perceptual information modulated by action. By focusing on the question, how movement specifies perceptual information, we show that ecological and sensorimotor theories endorse substantially different views about the role of action in perception. In particular we argue that ecological invariants are characterized with reference to transformations produced in the sensory array by movement: such invariants are transformation-specific but do not imply motor-specificity. In contrast, sensorimotor theories assume that perceptual invariants are intrinsically tied to specific movements. We show that this difference leads to different empirical predictions and we submit that the distinction between motor equivalence and motor-specificity needs further clarification in order to provide a more constrained account of action/perception relations.

Vestibular System May Provide Equivalent Motor Actions Regardless of the Number of Body Segments Involved in the Task

The vestibulospinal system likely plays an essential role in motor equivalence--the ability to reach the desired motor goal despite intentional or imposed changes in the number of body segments involved in the task. To test this hypothesis, we compared the ability of healthy subjects and patients with unilateral vestibular lesions (surgical acoustic neuroma resection 0.6 to 6.7 yr before the study) to maintain either the same hand position or the same trajectory of within arm reach movements while flexing the trunk, in the absence of vision. In randomly selected trials, the trunk motion was prevented by an electromagnetic device. Healthy subjects were able to preserve the hand position or trajectory by modifying the elbow and shoulder joint rotations in a condition-dependent way, at a minimal latency of about 60 ms after the trunk movement onset. In contrast, six of seven patients showed deficits in the compensatory angular modifications at least in one of two tasks so that 30-100% of the trunk displacement was not compensated and thus influenced the hand position or trajectory. Results suggest that vestibular influences evoked by the head motion during trunk flexion play a major role in maintaining the consistency of arm motor actions in external space despite changes in the number of body segments involved. Our findings also suggest that despite long-term plasticity in the vestibular system and related neural structures, unilateral vestibular lesion may reduce the capacity of the nervous system to achieve motor equivalence.

Mental rotation of anthropoid hands: a chronometric study

It has been shown that mental rotation of objects and human body parts is processed differently in the human brain. But what about body parts belonging to other primates? Does our brain process this information like any other object or does it instead maximize the structural similarities with our homologous body parts? We tried to answer this question by measuring the manual reaction time (MRT) of human participants discriminating the handedness of drawings representing the hands of four anthropoid primates (orangutan, chimpanzee, gorilla, and human). Twenty-four right-handed volunteers (13 males and 11 females) were instructed to judge the handedness of a hand drawing in palm view by pressing a left/right key. The orientation of hand drawings varied from 0 masculine (fingers upwards) to 90 masculine lateral (fingers pointing away from the midline), 180 masculine (fingers downwards) and 90 masculine medial (finger towards the midline). The results showed an effect of rotation angle (F(3, 69) = 19.57, P < 0.001), but not of hand identity, on MRTs. Moreover, for all hand drawings, a medial rotation elicited shorter MRTs than a lateral rotation (960 and 1169 ms, respectively, P < 0.05). This result has been previously observed for drawings of the human hand and related to biomechanical constraints of movement performance. Our findings indicate that anthropoid hands are essentially equivalent stimuli for handedness recognition. Since the task involves mentally simulating the posture and rotation of the hands, we wondered if "mirror neurons" could be involved in establishing the motor equivalence between the stimuli and the participants' own hands.

Motor equivalent control of the center of mass in response to support surface perturbations

To claim that the center of mass (CM) of the body is a controlled variable of the postural system is difficult to verify experimentally. In this report, a new variant of the method of the uncontrolled manifold (UCM) hypothesis was used to evaluate CM control in response to an abrupt surface perturbation during stance. Subjects stood upright on a support surface that was displaced in the posterior direction. Support surface translations between 0.03 and 0.12 m, each lasting for 275 ms, were presented randomly. The UCM corresponding to all possible combinations of joints that are equivalent with respect to producing the average pre-perturbation anterior-posterior position of the center of mass (CM(AP)) were linearly estimated for each trial. At each point in time thereafter, the difference between the current joint configuration and the average pre-perturbation joint configuration was computed. This joint difference vector was then projected onto the pre-perturbation UCM as a measure of motor equivalence, and onto its complementary subspace, which represents joint combinations that lead to a different CM(AP) position. A similar analysis was performed related to control of the trunk's spatial orientation. The extent to which the joint velocity vector acted to stabilize the CM(AP) position was also examined. Excursions of the hip and ankle joints both increased linearly with perturbation magnitude. The configuration of joints at each instance during the perturbation differed from the mean configuration prior to the perturbation, as evidenced by the joint difference vector. Most of this joint difference vector was consistent, however, with the average pre-perturbation CM(AP) position rather than leading to a different CM(AP )position. This was not the case, however, when performing this analysis with respect to the UCM corresponding to the control of the pre-perturbation trunk orientation. The projection of the instantaneous joint velocity vector also was found to lie primarily in the UCM corresponding to the pre-perturbation CM(AP) position, indicating that joint motion was damped in directions leading to a change away from the pre-perturbation CM(AP) position. These results provide quantitative support for the argument that the CM position is a planned variable of the postural system and that its control is achieved through selective, motor equivalent changes in the joint configuration in response to support surface perturbations. The results suggest that the nervous system accomplishes postural control by a control strategy that considers all DOFs. This strategy presumably resists combinations of DOFs that affect the stability of important task-relevant variables (CM(AP) position) while, to a large extent, freeing from control combinations of those DOFs that have no effect on the task-relevant variables (Schöner in Ecol Psychol 8:291-314, 1995).

Hand path priming in manual obstacle avoidance: Evidence for abstract spatiotemporal forms in human motor control.

Previous research suggests that motor equivalence is achieved through reliance on effector-independent spatiotemporal forms. Here the authors report a series of experiments investigating the role of such forms in the production of movement sequences. Participants were asked to complete series of arm movements in time with a metronome and, on some trials, with an obstacle between 1 or more of the target pairs. In moves following an obstacle, participants only gradually reduced the peak heights of their manual jumping movements. This hand path priming effect, scaled with obstacle height, was preserved when participants cleared the obstacle with 1 hand and continued with the other, and it was modulated by future task demands. The results are consistent with the hypothesis that the control of movement sequences relies on abstract spatiotemporal forms. The data also support the view that motor programming is largely achieved by changing just those features that distinguish the next movement to be made from the movement that was just made.

From Parallel Sequence Representations to Calligraphic Control: A Conspiracy of Neural Circuits

Calligraphic writing presents many challenges for motor control, including: learning and recall of stroke sequences; critical timing of stroke onsets and durations; fine control of grip and contact forces; and letterform invariance under size scaling, which entails fine control of stroke directions and amplitudes during recruitment and derecruitment of musculoskeletal degrees of freedom. Experimental and computational studies in behavioral neuroscience have progressed toward explaining the learning, planning, and control exercised in tasks that share features with calligraphic writing and drawing. This article highlights component operations ranging from parallel sequence representations to fine force control. Treated in succession are: competitive queuing models of sequence representation, performance, learning, and recall; letter size scaling and motor equivalence; cursive handwriting models in which sensory-motor transformations are performed by circuits that learn inverse differential kinematic mappings; and fine-grained control of timing and transient forces by circuit models that learn to solve inverse dynamics problems.

Amplitude and direction errors in kinesthetic pointing.

We investigated the accuracy with which, in the absence of vision, one can reach again a 2D target location that had been previously identified by a guided movement. A robotic arm guided the participant's hand to a target (locating motion) and away from it (homing motion). Then, the participant pointed freely toward the remembered target position. Two experiments manipulated separately the kinematics of the locating and homing motions. Some robot motions followed a straight path with the bell-shaped velocity profile that is typical of natural movements. Other motions followed curved paths, or had strong acceleration and deceleration peaks. Current motor theories of perception suggest that pointing should be more accurate when the homing and locating motion mimics natural movements. This expectation was not borne out by the results, because amplitude and direction errors were almost independent of the kinematics of the locating and homing phases. In both experiments, participants tended to overshoot the target positions along the lateral directions. In addition, pointing movements towards oblique targets were attracted by the closest diagonal (oblique effect). This error pattern was robust not only with respect to the manner in which participants located the target position (perceptual equivalence), but also with respect to the manner in which they executed the pointing movements (motor equivalence). Because of the similarity of the results with those of previous studies on visual pointing, it is argued that the observed error pattern is basically determined by the idiosyncratic properties of the mechanisms whereby space is represented internally.

The quest to understand bimanual coordination

Many skillful manipulations engage both hands for goal achievement. Whereas the goal is planned consciously and achieved quasi-invariantly, the articulators are mobilized automatically, but in a flexible manner (Lashley's principle of motor equivalence). In brain disorders affecting hand functions, adaptive mechanisms are mobilized to improve goal achievement. Thus, chronic cerebellar patients were found to initiate a bimanual drawer task with marked intermanual desynchronization as compared to control subjects. This was partly compensated for, however, by adjusting the kinematics as the individual limbs move toward the goal, thereby improving the initial desynchronization. Adaptive strategies rarely correct deficits completely, however. Bimanual movement patterns, either in-phase or anti-phase are relatively stable in healthy human subjects, whereas brain pathology may preferentially impair the anti-phase pattern. This is the case in patients with acquired pathology of the corpus callosum, thereby suggesting that this structure is important for maintaining temporally independent limb and hand movements.

Learning a single limb multijoint coordination pattern: the impact of a mechanical constraint on the coordination dynamics of learning and transfer.

The coordination dynamics of learning and transfer were studied in a single limb multijoint task requiring rhythmic elbow and wrist motions. Participants were required to learn a continuous 90 degrees relative phase pattern between the elbow and wrist such that an angle-angle plot of elbow and wrist motion produced a circle with a diameter of 80 degrees. Joint motion was restricted to elbow and wrist flexion-extension on the sagittal plane and the to-be-learned 90 degrees relative phase pattern was always practiced with the learning arm supine. Cycling frequency was controlled by a pacing metronome set at 0.75 Hz. Issues regarding effector-independent and effector-specific transfer were addressed with three transfer conditions: (1). learning arm prone (LP), (2). non-learning arm supine (NS), and (3). non-learning arm prone (NP). Four subjects learned the required relative phase (90 degrees ) and amplitude (80 degrees ) pattern with their dominant arm and four with their non-dominant arm. The experiment produced three main findings with regard to elbow-wrist control processes: First, seven of eight participants spontaneously produced a wrist-lagging coordination pattern (wrist motion lagged elbow motion) in learning to produce a continuous relative phase pattern of 90 degrees between the elbow and wrist. The wrist-lagging pattern may emerge as a result of the central nervous system exploiting the transfer of angular momentum from the elbow to the wrist as the elbow rotates up and down. The influence of interactive torque on elbow-wrist coordination represents an important mechanical constraint on the selection of intralimb coordination strategies during learning. The transfer conditions revealed that this mechanical constraint was effector-independent with regard to ipsilateral limb transfer (LP) and contralateral limb transfer (NS and NP). Second, consistent transfer of the learned relative phase pattern across ipsilateral and contralateral conditions demonstrates an effector-independent representation for this control variable. The effector-independent and effector-specific nature of joint amplitude transfer was dependent to some degree on learning arm, dominant or non-dominant, and the amount of practice, 1 day versus 5 days. Third, learning of the required 90 degrees relative phase pattern may be characterized as a phase transition leading to the formation of a stable attractor in the elbow-wrist coordination landscape. The above findings are discussed with respect to motor programming and coordination dynamic viewpoints on effector-independent and effector-specific aspects of motor equivalence.

Learning to juggle: on the assembly of functional subsystems into a task-specific dynamical organization.

We examined the development of task-specific couplings among functional subsystems (i.e., ball circulation, respiration, and body sway) when learning to juggle a three-ball cascade, with a focus on learning-induced changes in the coupling between ball movements and respiration and the coupling between ball movements and body sway. Six novices practiced to juggle three balls in cascade fashion for one hour per day for twenty days. On specific days (7 in total), ball movements, center-of-pressure (CoP) trajectories and respiration traces were measured simultaneously. Discrete, time-continuous and spectral analyses revealed that the spatio-temporal variability of the juggling patterns decreased with practice and that the degree to which the task constraints were satisfied increased gradually. No conclusive evidence was found for ball movement-respiration coupling. In contrast, clear-cut evidence was found for the presence of 1:3 and 2:3 frequency locking between the vertical component of the ball trajectories and both the anterior-posterior and the medio-lateral components of the CoP. Incidence and expression of these mode locks varied across individuals and altered in the course of learning. Gradual changes in locking strength, appearances and disappearances of mode locks, as well as abrupt transitions between coupled states were observed. These results indicate that dissimilar learning dynamics may arise in the functional embedding of subsystems into a task-specific organization and that motor equivalence is an inherent property of such emerging task-specific organizations.

Directional invariance during loading-related modulations of muscle activity: evidence for motor equivalence.

In the present study, we investigated the influence of external force manipulations on movements in different directions, while keeping the amplitude invariant. Subjects ( n=10) performed a series of cyclical anteroposterior, mediolateral, and oblique line-drawing movements (star drawing task) with their dominant limb in the horizontal plane. To dissociate kinematics from the underlying patterns of muscle activation, spring loading was applied to the forearm of the moving limb. Whereas spring loading of the arm resulted in considerable changes in the overall amount of muscle activation in the elbow and shoulder muscles, invariance was largely maintained at the kinematic level. Subjects produced the required movement directions and amplitudes of the star drawing largely successfully, irrespective of the force bias induced by the spring. These observations demonstrate motor equivalence and strengthen the notion that the spatial representation of drawing movements is encoded in the higher brain regions in a rather abstract form that is dissociated from the concrete muscle activation patterns underlying a particular movement direction. To achieve this goal, the central nervous system shifted between two or more muscle grouping strategies to overcome modulations in the interaction among posture-dependent (joint stiffness), dynamic (inertial), and elastic (spring) torque components in the joints. Spring loading induced general changes in the overall amount of EMG activity, which was largely muscle but not direction specific, presumably to represent the posture-dependent biasing force of the spring. Loading was mainly shown to increase muscle coactivation in the elbow joint. This indicates that the subjects tended to increase stiffness in the elbow to compensate for changes in the spring bias forces in order to minimize trajectory errors. Changes in muscle grouping of the shoulder antagonists were mainly a consequence of movement direction but were also affected partly by loading, presumably reflecting the influence of dynamic force components. Taken together, the results confirmed the hypothesis that changes of movement direction and direction of force in the end-effector generated specific sets of muscle grouping to overcome the dynamic requirements in the joints while keeping the kinematics largely unchanged. This suggests that directional tuning in muscle activity and changes in muscle grouping reflects the formation of appropriate internal models in the CNS that give rise to motor equivalence.

Kinematics of a coordinated goal-directed bimanual task

The experiments address the problem of bimanual coordination in a familiar task of everyday life. A goal-directed drawer-pulling task, with asymmetrical assignments among hands, was analyzed with the objective to detect discrete kinematic events (‘anchors’) that potentially could serve in proper goal synchronization. The left hand reached out for the drawer and opened it while the right hand performed a prehension movement to pick up a peg from the drawer. The task was smoothly performed, independently of vision. Typically, trajectories and velocity profiles of the leading pull-hand were more stereotypical than the more variable ones of the pick-hand. The pull-hand had a large velocity peak during reaching, followed by a small peak during pulling. Velocity profiles of the pick-hand were not bell-shaped and exhibited one or two broad waves, often with an irregular and probing evolution. Velocity profiles of both hands were aligned with the first or the second velocity peak of the leading pull-hand. In the majority of cases, temporal associations of events in the kinematics of the two limbs could thus be identified, which could serve to synchronize the hands at the goal. The nearly straight biphasic reach-and-pull trajectory of the leading hand contrasted with the more curved trajectory of the right pick-hand whereas, in the same unimanual action, the latter trajectories were quasi-rectilinear. Changing constraints (no vision, cutaneous anesthesia of pulling fingers) could change the coordination pattern. We argue that bimanual coordination relies on two interacting mechanisms: (1) feedforward control on the basis of sensorimotor memory; (2) temporal adjustments during the evolving bimanual synergy. Multiple strategies, imposed by the leading pull-hand, appeared to be responsible for feedback-induced corrections in the pick-hand and were found to contribute to the goal-invariance and to the principle of motor equivalence.

Preliminary studies on respiratory activity in speech

Work at Edinburgh University in the 1950s, summarized by Ladefoged [Three Area of Experimental Phonetics (Oxford University Press, Oxford, 1967)] claimed that (1) the mean power for speech was provided by actions of the respiratory muscles. The external intercostals check the outgoing airflow when speaking with high lung volumes, and the internal intercostals, rectus abdominis, and other muscles become more active as lung volume decreases. (2) Stressed syllables are produced by increases of respiratory power above the mean, usually by further activity of the internal intercostals. Hixon and Weismer (1995) pointed out technical limitations of these early studies, but wrongly indicated that they were of little value. These studies have been replicated with modern methods to measure intramuscular EMG and airflow and pressure. The two subjects studied to date follow the pattern previously described. In addition the improved techniques indicate that speakers exercise further control by bringing in stabilizing antagonistic muscles. These results are consistent with the general hypothesis that the central nervous system is able to control subglottal pressure by using various combinations of passive elastic and active muscle forces, thereby achieving motor equivalence and acoustic invariance over a wide range of lung volumes.

Contributions of gait and trunk movements to prehension: perspectives from world- and body-centered coordinates.

The present paper reviews a series of prehension experiments recently conducted at Simon Fraser University's Human Motor Systems Laboratory, and attempts to place them into the larger context of multi-segmental control theory. Two related lines of experiments are reported: (a) experiments involving prehension during walking, and (b) experiments involving trunk-assisted reaching. Three-dimensional analyses of movements were performed via both world- and body-centered coordinates. Our results are supportive of the idea that both types of tasks are carried out using task-specific synergies. Furthermore, we assert that the actions of these synergies are comprised of variable contributions of different movement systems and result in smooth, world-centered end-point trajectories. We show evidence that this "motor equivalence" is the result of increasing the complexity of a given task. Finally, the implications of the present findings on prevailing motor control theory are discussed in terms of the theoretical mechanisms underlying the coordination of the transport and grasp components of prehension.

Human inferior parietal cortex ‘programs’ the action class of grasping

If one writes with a pen grasped between the toes, or a pencil held in the mouth, the handwriting style may be of poor quality but can be identified as belonging to a particular individual. Like other actions, such as grasping or pointing, different body parts can be used to produce the movement. These findings, of reasonably consistent spatial and temporal productions by different effectors, have been used to argue for the concept of motor equivalence and the existence of motor programs abstracted from particular effectors. In this study subjects were required to perform an action (grasping a sweet) with different effectors (the mouth or the hand) while the brain was scanned. Activation of the inferior parietal lobe during real and imagined mouth grasping, and during real hand grasping actions was demonstrated. Primate neurophysiological research has implicated this region in a movement-planning role. Our results confirm the importance of the inferior parietal lobe in integrating converging multimodal sensory information for coding of general action patterns in humans.

Evidence of motor equivalence in a pointing task involving locomotion.

A pointing task was performed both while subjects stood beside and while subjects walked past targets that involved differing movement amplitudes and differing sizes. The hand kinematics were considered relative both to a fixed frame of reference in the movement environment (end effector kinematics) and to the subject's body (kinematics of the hand alone). From the former view, there were few differences between standing and walking versions of the task, indicating similarity of the kinematics of the hand. However, when the hand was considered alone, marked differences in the kinematics and spatial trajectories between standing and walking were achieved. Furthermore, kinematic analyses of the trunk showed that subjects used differing amounts of both flexion-extension and rotation movements at the waist depending on whether they were standing or walking as well as on the constraints imposed by target width and movement amplitude. The present results demonstrate the existence of motor equivalence in a combined upper and lower extremity task and that this motor equivalence is a control strategy to cope with increasing task demands. Given the complexity involved in controlling the arm, the torso, and the legs (during locomotion), the movements involved in the present tasks appear to be planned and controlled by considering the whole body as a single unit.