The type 1 submovement conundrum: an investigation into the function of velocity zero-crossings within two-component aiming movements.

In rapid manual aiming, traditional wisdom would have it that two components manifest from feedback-based processes, where error accumulated within the primary submovement can be corrected within the secondary submovement courtesy of online sensory feedback. In some aiming contexts, there are more type 1 submovements (overshooting) compared to types 2 and 3 submovements (undershooting), particularly for more rapid movements. These particular submovements have also been attributed to a mechanical artefact involving movement termination and stabilisation. Hence, the goal of our study was to more closely examine the function of type 1 submovements by revisiting some of our previous datasets. We categorised these submovements according to whether the secondary submovement moved the limb closer (functional), or not (non-functional), to the target. Overall, there were both functional and non-functional submovements with a significantly higher proportion for the former. The displacement at the primary and secondary submovements, and negative velocity peak were significantly greater in the functional compared to non-functional. The influence of submovement type on other movement characteristics, including movement time, was somewhat less clear. These findings indicate that the majority of type 1 submovements are related to intended feedforward- and/or feedback-based processes, although there are a portion that can be attributed an indirect manifestation of a mechanical artefact. As a result, we suggest that submovements should be further categorised by their error-reducing function.

The multiple process model of goal-directed aiming/reaching: insights on limb control from various special populations.

Several years ago, our research group forwarded a model of goal-directed reaching and aiming that describes the processes involved in the optimization of speed, accuracy, and energy expenditure Elliott et al. (Psychol Bull 136:1023-1044, 2010). One of the main features of the model is the distinction between early impulse control, which is based on a comparison of expected to perceived sensory consequences, and late limb-target control that involves a spatial comparison of limb and target position. Our model also emphasizes the importance of strategic behaviors that limit the opportunity for worst-case or inefficient outcomes. In the 2010 paper, we included a section on how our model can be used to understand atypical aiming/reaching movements in a number of special populations. In light of a recent empirical and theoretical update of our model Elliott et al. (Neurosci Biobehav Rev 72:95-110, 2017), here we consider contemporary motor control work involving typical aging, Down syndrome, autism spectrum disorder, and tetraplegia with tendon-transfer surgery. We outline how atypical limb control can be viewed within the context of the multiple-process model of goal-directed reaching and aiming, and discuss the underlying perceptual-motor impairment that results in the adaptive solution developed by the specific group.

Effects of wrist tendon vibration and eye movements on manual aiming.

In the present study, we investigated whether visual information mediates a proprioceptive illusion effect induced by muscle tendon vibration in manual aiming. Visual information was gradually degraded from a situation in which the targets were present and participants (n = 20; 22.3 ± 2.7 years) were permitted to make saccadic eye movements to designated target positions, to a condition in which the targets were not visible and participants were required to perform cyclical aiming while fixating a point between the two target positions. Local tendon vibration applied to the right wrist extensor muscles induced an illusory reduction of 15% in hand movement amplitude. This effect was greater in the fixation than in the saccade condition. Both anticipatory control and proprioceptive feedback are proposed to contribute to the observed effects. The primary saccade amplitude was also reduced by almost 4% when muscle tendon vibration was locally applied to the wrist. These results confirm a tight link between eye movements and manual perception and action. Moreover, the impact of the proprioceptive illusion on the ocular system indicates that the interaction between systems is bidirectional.

The violation of Fitts' Law: an examination of displacement biases and corrective submovements.

Fitts' Law holds that, to maintain accuracy, movement times of aiming movements must change as a result of varying degrees of movement difficulty. Recent evidence has emerged that aiming to a target located last in an array of placeholders results in a shorter movement time than would be expected by the Fitts' equation-a violation of Fitts' Law. It has been suggested that the violation emerges because the performer adopts an optimized movement strategy in which they partially pre-plan an action to the closest placeholder (undershoot the last placeholder) and rely on a secondary acceleration to propel the limb toward the last location when it is selected as the target (Glazebrook et al. in Hum Mov Sci 39:163-176, 2015). In the current study, we examine this proposal and further elucidate the processes underlying the violation by examining limb displacement and corrective submovements that occur when performers aim to different target locations. For our Main Study, participants executed discrete aiming movements in a five-placeholder array. We also reanalyzed data from a previously reported study in which participants aimed in placeholder and no-placeholder conditions (Blinch et al. in Exp Brain Res 223:505-515, 2012). The results showed the violation of Fitts' Law unfolded following peak velocity (online control). Further, the analysis showed that movements to the last target tended to overshoot and had a higher proportion of secondary submovements featuring a reversal than other categories of submovement (secondary accelerations, discontinuities). These findings indicate that the violation of Fitts' Law may, in fact, result from a strategic bias toward planning farther initial displacements of the limb which accommodates a shorter time in online control.

Complimentary lower-level and higher-order systems underpin imitation learning.

We examined whether the temporal representation developed during motor training with reduced-frequency knowledge of results (KR; feedback available on every other trial) was transferred to an imitation learning task. To this end, four groups first practised a three-segment motor sequence task with different KR protocols. Two experimental groups received reduced-frequency KR, one group received high-frequency KR (feedback available on every trial), and one received no-KR. Compared to the no-KR group, the groups that received KR learned the temporal goal of the movement sequence, as evidenced by increased accuracy and consistency across training. Next, all groups learned a single-segment movement that had the same temporal goal as the motor sequence task but required the imitation of biological and nonbiological motion kinematics. Kinematic data showed that whilst all groups imitated biological motion kinematics, the two experimental reduced-frequency KR groups were on average ∼ 800 ms more accurate at imitating movement time than the high-frequency KR and no-KR groups. The interplay between learning biological motion kinematics and the transfer of temporal representation indicates imitation involves distinct, but complementary lower-level sensorimotor and higher-level cognitive processing systems.

Effector mass and trajectory optimization in the online regulation of goal-directed movement.

Goal-directed aiming movements are planned and executed so that they optimize speed, accuracy and energy expenditure. In particular, the primary submovements involved in manual aiming attempts typically undershoot targets in order to avoid costly time and energy overshoot errors. Furthermore, in aiming movements performed over a series of trials, the movement planning process considers the sensory information associated with the most recent aiming attempt. The goal of the current study was to gain further insight into how the sensory consequences associated with the recent and forthcoming aiming attempts impact performance. We first examined whether performers are more conservative in their aiming movements with a heavy, as opposed to a light, stylus by determining whether primary submovements undershot the target to a greater extent in the former due to an anticipated increase in spatial variability. Our results show that movements with the heavy stylus demonstrated greater undershoot biases in the primary submovements, as well as greater trial-to-trial spatial variability at specific trajectory kinematic landmarks. In addition, we also sought to determine whether the sensory information experienced on a previous aiming movement affected movement planning and/or online control on the subsequent aiming attempt. To vary the type sensory consequences experienced on a trial-to-trial basis, participants performed aiming movements with light and heavy styli in either blocked or random orderings of trials. In the random-order conditions, some participants were provided advance information about stylus mass for the upcoming trial, while others were not. The blocked and random trial orders had minimal impacts on end point aiming performance. Furthermore, similarities in the times to key kinematic landmarks in the trajectories of the random-order groups suggest that recent trial experience had a greater effect on the upcoming aiming movement compared with advance task knowledge.

Factors underlying age-related changes in discrete aiming.

Age has a clear impact on one's ability to make accurate goal-directed aiming movements. Older adults seem to plan slower and shorter-ranged initial pulses towards the target, and rely more on sensory feedback to ensure endpoint accuracy. Despite the fact that these age-related changes in manual aiming have been observed consistently, the underlying mechanism remains speculative. In an attempt to isolate four commonly suggested underlying factors, young and older adults were instructed to make discrete aiming movements under varying speed and accuracy constraints. Results showed that older adults were physically able to produce fast primary submovements and that they demonstrated similar movement-programming capacities as young adults. On the other hand, considerable evidence was found supporting a decreased visual feedback-processing efficiency and the implementation of a play-it-safe strategy in older age. In conclusion, a combination of the latter two factors seems to underlie the age-related changes in manual aiming behaviour.

The impact of age and physical activity level on manual aiming performance.

Older adults traditionally adapt their discrete aiming movements, thereby traveling a larger proportion of the movement under closed-loop control. As the beneficial impact of a physically active lifestyle in older age has been described for several aspects of motor control, we compared the aiming performance of young controls to active and sedentary older adults. To additionally determine the contribution of visual feedback, aiming movements were executed with and without saccades. Results showed only sedentary older adults adopted the typical movement changes, highlighting the impact of a physically active lifestyle on manual aiming in older age. In an attempt to reveal the mechanism underlying the movement changes, evidence for an age-related decline in force control was found, which in turn resulted in an adapted aiming strategy. Finally, prohibiting saccades did not affect older adults' performance to a greater extent, suggesting they do not rely more on visual feedback than young controls.

Primary and submovement control of aiming in C6 tetraplegics following posterior deltoid transfer.

BACKGROUND: Upper limb motor control in fast, goal-directed aiming is altered in tetraplegics following posterior-deltoid musculotendinous transfer. Specifically, movements have similar end-point accuracy but longer duration and lower peak velocity than those of age-matched, neurotypical controls. Here, we examine in detail the interplay between primary movement and submovement phases in five C6 tetraplegic and five control participants. METHODS: Aiming movements were performed in two directions (20 cm away or toward), with or without vision. Trials that contained a submovement phase (i.e., discontinuity in velocity, acceleration or jerk) were identified. Discrete kinematic variables were then extracted on the primary and submovements phases. RESULTS: The presence of submovements did not differ between the tetraplegic (68%) and control (57%) groups, and almost all submovements resulted from acceleration and jerk discontinuities. Tetraplegics tended to make a smaller amplitude primary movement, which had lower peak velocity and greater spatial variability at peak velocity. This was followed by a larger amplitude and longer duration secondary submovement. Peak velocity of primary movement was not related to submovement incidence. Together, the primary and submovement phases of both groups were equally effective in reducing end-point error. CONCLUSIONS: C6 tetraplegic participants exhibit some subtle differences in measures of motor behaviour compared to control participants, but importantly feedforward and feedback processes work effectively in combination to achieve accurate goal-directed aiming.

The influence of target context and early and late vision on goal-directed reaching.

The online visual control of movement involves contributions from 2 processes: a process early in the trajectory concerned with comparisons between actual and expected sensory consequences and another process late in the trajectory that reduces the discrepancy between the position of the hand and the target. This experiment was designed to determine how early and late visual controls are impacted by the illusory characteristics of the target in a rapid reaching task. Participants performed 500 ms movements to the vertices of Müller-Lyer figures with the availability of full vision on the majority of trials. However, on a fraction of the trials, movements to the targets were performed with either early vision (first 200 ms of movement), late vision (last 200 ms of movement) or no vision. Although participants undershoot the targets under all target and visual conditions, the impact of the target configuration was greatest when vision was available during only the final portion of the movement trajectory and least when only early vision was available for limb regulation. Aiming bias under full-vision and no-vision conditions was intermediate. These findings indicate that visual context has a greater impact on late discrete limb regulation than on early dynamic control of the limb trajectory.

Movement strategies in vertical aiming of older adults.

The current study examined the movement kinematics of older adults when aiming to vertically oriented targets. Late middle-age and early old-age participants completed 20 trials to a small target located downward or upward by 0.16 m from a home position at shoulder height. Aiming direction had a significant effect, resulting in more submovements, which were mostly reflective of undershooting when aiming to the downward compared to upward target. In trials containing a submovement, both groups exhibited shorter total movement time, concomitant with a decrease in duration of the primary movement and an increase in submovement amplitude, when aiming to the downward target. Measures of dispersion also differed in accord with the amplitude of submovements, such that there was greater spatial and temporal variability in the primary movement when aiming in the downward direction. While there was limited evidence of a difference between the groups, there were significant correlations between age and several dependent measures when aiming to the downward target. Of note, in trials containing submovements, older participants exhibited larger amplitude and longer duration submovements, as well as shorter amplitude primary movement. Spatial variability at peak velocity also increased as a function of age when aiming downward, but not in trials without submovements. An explanation related to physical limits on movement production is discounted given the lack of consistent findings between trial types. Instead, we suggest older participants' exhibit strategic differences in movement kinematics when aiming to vertically located targets, and that these change progressively with age in order to maintain speed-accuracy relations.

Dissociable contributions of motor-execution and action-observation to intramanual transfer.

We examined the hypothesis that different processes and representations are associated with the learning of a movement sequence through motor-execution and action-observation. Following a pre-test in which participants attempted to achieve an absolute, and relative, time goal in a sequential goal-directed aiming movement, participants received either physical or observational practice with feedback. Post-test performance indicated that motor-execution and action-observation participants learned equally well. Participants then transferred to conditions where the gain between the limb movements and their visual consequences were manipulated. Under both bigger and smaller transfer conditions, motor-execution and action-observation participants exhibited similar intramanual transfer of absolute timing. However, participants in the action-observation group exhibited superior transfer of relative timing than the motor-execution group. These findings suggest that learning via action-observation is underpinned by a visual-spatial representation, while learning via motor-execution depends more on specific force-time planning (feed forward) and afferent processing associated with sensorimotor feedback. These behavioural effects are discussed with reference to neural processes associated with striatum, cerebellum and motor cortical regions (pre-motor cortex; SMA; pre-SMA).

125 years of perceptual-motor skill research.

This article celebrates the contribution that the American Journal of Psychology (AJP) has made to the area of perceptual-motor skill over its 125-year history. We highlight the articles published in AJP and trace the technical and theoretical developments that stem from this groundbreaking work. Included in our overview are AJP articles on the excitability of the motor system, motor learning, adaptation to visual rearrangement, the ecological approach to perception and action, and the measurement of human handedness. We conclude by identifying a number of areas associated with perceptual-motor skill where AJP continues to make an important contribution.

Optimization in Manual Aiming: Relating Inherent Variability and Target Size, and Its Influence on Tendency.

For manual aiming, the optimized submovement model predicts a tendency toward target-center of primary movement endpoints (probabilistic strategy), while the minimization model predicts target undershooting ("play-it-safe" strategy). The spatial variability of primary movement endpoints directed toward a cross-hair (400-500 ms) (Session 1) were scaled by a multiplicative factor (x1 - 4) to form circular targets of different sizes (Session 2). In recognition of both models, it was predicted that the more that inherent variability exceeded the target size, the greater the tendency to shift from target-center aiming to target undershooting. The central tendency of primary movement endpoints was not influenced by the targets, while it neared target-center. These findings concur with a probabilistic strategy, although we speculate on factors that might otherwise foster a "play-it-safe" strategy.

Action representations in perception, motor control and learning: implications for medical education.

OBJECTIVES: the motor behaviours or 'actions' that provide the basis for precision limb control, including the performance of complex medical procedures, are represented at different levels in the central nervous system. This review focuses on how these representations influence the way people perceive, execute and learn goal-directed movements. PERCEPTION AND ATTENTION: the neural processes associated with paying attention to an object are part and particle of the same processes engaged to physically interact with that object. The automatic way in which specific actions are engaged makes it important that we structure perceptual motor environments in a manner that facilitates goal actions and minimises the likelihood of unwanted actions. MOTOR CONTROL: most actions are organised to optimise speed, accuracy and energy expenditure while avoiding worst-case outcomes. To achieve a good outcome on movements, the performer must have the opportunity to experiment with the way specific actions are executed. Early in the discovery process, errors are necessary if the performer is to determine his or her performance boundaries. motor learning: as learning progresses, representations of action become predictive. For example, if rapid corrective processes are to operate, the performer needs to anticipate sensorimotor consequences of movement. Thus, practice should be specific to the conditions under which actions are performed, and the performer. Although nothing can replace physical practice, complex representations of action can develop by observing both expert performers and learners. In many cases, practice scenarios that include both physical practice and observations of other learners can be the most efficient use of time and resources. CONCLUSIONS: although most of the experiments reviewed here involved laboratory tasks such as rapid aiming and movement sequencing, the majority of the principles apply to motor control and learning in more complex situations. Thus, they should be considered when developing methods to train medical personnel to perform perceptual motor procedures with precision.

Intermittent Vision and Goal-Directed Movement: A Review.

It is well known that vision makes an important contribution to the control of goal-directed movements. However, task performance can be maintained when vision is interrupted, such as when a goalkeeper faces a free kick in soccer and the ball moves behind teammates and opposing players. To maintain behavior, it is necessary to process the visual information available from intermittent samples. In this review, we consider the performance and learning effects of intermittent vision in tasks such as aiming, reaching and grasping, goal-directed locomotion and ball-catching. We review research that finds both interocular and intraocular integration contribute to continuous upper limb control with intermittent visual pickup/sampling. Recent work using intermittent visual presentation (i.e., stroboscopic vision) to facilitate learning of general and task-specific visual-motor skills indicates that training/learning protocols that challenge, but don't alter, the visual-motor processing associated with a specific visual-motor task can be effective. In this theoretical context, we discuss methodological and design factors that could impact the effectiveness of future training studies.

General motor representations are developed during action-observation.

This study was designed to examine the generality of motor learning by action-observation. During practice, action-observation participants watched a learning model (e.g., physical practice participants) perform a motor sequence-timing task involving mouse/cursor movements on a computer screen; control participants watched a blank screen. Participants transferred to either a congruent (same mouse-cursor gain), or an incongruent (different mouse-cursor gain) condition. As predicted, motor sequence timing was learned through action-observation as well as physical practice. Moreover, transfer of learning to an incongruent set of task demands indicates that the motor representation developed through observation includes generalised visual-motor procedures associated with the use of feedback utilization.

Sensory-motor equivalence: manual aiming in C6 tetraplegics following musculotendinous transfer surgery at the elbow.

Cervical spinal lesions at C6 result in paralysis of the triceps brachii while leaving deltoid and elbow flexor function intact. We examined the spatial-temporal characteristics of goal-directed aiming movements performed by C6 tetraplegics who had undergone musculotendinous transfer surgery in which the posterior deltoid replaces the triceps as the elbow extensor. On some trials, liquid crystal goggles were used to eliminate vision of the limb and target upon movement initiation. Although tetraplegic participants achieved the same degree of movement accuracy/consistency as control participants, their movement times were longer regardless of whether the movements were made away from (elbow extension) or towards the body (elbow flexion). Longer movement times were related to lower peak velocities, and not the symmetry of the aiming profiles. The tetraplegic participants were no more dependent on visual feedback for limb regulation than control participants. Although the characteristics of the movement trajectories were surprisingly similar, in both vision conditions, tetraplegics required more real and proportional time to reduce spatial variability in the limb's trajectory for elbow extensions. Our results indicate that the sensorimotor system is adaptable and that the representations governing limb control are not muscle specific.

Between-person effects on attention and action: Joe and Fred revisited.

Previous study indicates that target-target inhibition of return (IOR) is not restricted to a single nervous system. Specifically, watching another person perform a goal-directed aiming movement engages similar inhibitory processes on a subsequent aiming attempt as if having performed the preceding movement oneself. This between-person effect has been attributed to the mirror neuron system. In the study reported here, we replicated this finding and examined the relative importance of automatic stimulus alerting events and action-observation by dissociating these two influences. This was done by having two people alternately perform sets of two aiming trials to the same equally probable targets. Under some experimental conditions, one or both of the performers moved to a non-illuminated target. In this way, we dissociated the stimulus and observed event under some between-person conditions. Although IOR was greatest when the stimulus and observed events were compatible, both contributed to the between-person inhibitory processes slowing the responses (Experiment 1). The impact of observing another person perform an aiming movement appears to have more to do with realizing a particular spatial goal than seeing the biological motion associated with achieving that goal (Experiment 2). Findings that both the illumination of a visual target signal and the observation of another person's action engage similar attention-action processes are consistent with action-based accounts of visual selective attention.

Visual regulation of manual aiming: a comparison of methods.

Visual regulation of upper limb movements occurs throughout the trajectory and is not confined to discrete control in the target area. Early control is based on the dynamic relationship between the limb, the target, and the environment. Despite robust outcome differences between protocols involving visual manipulations, it remains difficult to identify the kinematic events that characterize these differences. In this study, participants performed manual aiming movements with and without vision. We compared several traditional approaches to movement analysis with two new methods of quantifying online limb regulation. As expected, participants undershot the target and their movement endpoints were more variable when vision was not available. Although traditional measures such as reaction time, time after peak velocity, and the presence of discontinuities in acceleration were sensitive to the visual manipulation, measures quantifying the trial-to-trial spatial variability throughout the trajectory were the most effective in isolating the time course of online regulation.