Coding of on-line and pre-planned movement sequences

Recent experiments have demonstrated that complex multi-element movement sequences were coded in visual-spatial coordinates even after extensive practice, while relatively simple spatial-temporal movement sequences are coded in motor coordinates after a single practice session. The purpose of the present experiment was to determine if the control process rather than the difficulty of the sequence played a role in determining the pattern of effector transfer. To accomplish this, different concurrent feedback conditions were provided to two groups of participants during practice of the same movement sequence. The results indicated that when concurrent visual feedback was provided during the production of the movement, which was thought to encourage on-line control, the participants performed transfer tests with the contra-lateral limb better when the visual-spatial coordinates were reinstated than when the motor coordinates were reinstated. When concurrent visual feedback was not provided, which was thought to encourage pre-planned control, the opposite was observed. The data are consistent with the hypothesis that the mode of control dictates the coordinate system used to code the movement sequence rather than sequence difficulty or stage of practice as has been proposed.     

Transfer of movement sequences: bigger is better

Experiment 1 was conducted to determine if proportional transfer from "small to large" scale movements is as effective as transferring from "large to small." We hypothesize that the learning of larger scale movement will require the participant to learn to manage the generation, storage, and dissipation of forces better than when practicing smaller scale movements. Thus, we predict an advantage for transfer of larger scale movements to smaller scale movements relative to transfer from smaller to larger scale movements. Experiment 2 was conducted to determine if adding a load to a smaller scale movement would enhance later transfer to a larger scale movement sequence. It was hypothesized that the added load would require the participants to consider the dynamics of the movement to a greater extent than without the load. The results replicated earlier findings of effective transfer from large to small movements, but consistent with our hypothesis, transfer was less effective from small to large (Experiment 1). However, when a load was added during acquisition transfer from small to large was enhanced even though the load was removed during the transfer test. These results are consistent with the notion that the transfer asymmetry noted in Experiment 1 was due to factors related to movement dynamics that were enhanced during practice of the larger scale movement sequence, but not during the practice of the smaller scale movement sequence. The findings that the movement structure is unaffected by transfer direction but the movement dynamics are influenced by transfer direction is consistent with hierarchal models of sequence production.     

Inter-manual transfer and practice: Coding of simple motor sequences

Previous research suggests that movements are represented early in practice in visual-spatial coordinates/codes, which are effector independent, and later in practice in motor coordinates/codes (e.g., joint angles, activation patterns), which are effector dependent. In the present experiments, the task was to reproduce 1.3 s patterns of elbow flexions and extensions. An inter-manual transfer paradigm was used in Experiment 1 and an inter-manual practice paradigm was used in Experiment 2. The present results clearly indicated a strong advantage of effector transfer when the motor coordinates available during acquisition were reinstated (Experiment 1) and demonstrate that inter-manual practice with the same motor coordinates results in enhanced retention performance relative to transfer and practice where the same visual-spatial coordinates are used. These results demonstrate that the more effective movement code (motor or visual-spatial) is dependent on the movement sequence characteristics (e.g., difficulty, number of elements, and mode of control [preplanned or on-line]). These results are also interesting because they indicate, contrary to previous findings with more complex movement sequences, that an effective motor code can be developed relatively early in practice for rapid movement sequences.     

Parkinson's disease differentially affects adaptation to gradual as compared to sudden visuomotor distortions

Patients with Parkinson’s disease (PD) have difficulties in movement adaptation to optimize performance in novel environmental contexts such as altered screen cursor-hand relationships. Prior studies have shown that the time course of the distortion differentially affects visuomotor adaptation to screen cursor rotations, suggesting separate mechanisms for gradual and sudden adaptation. Moreover, studies in human and non-human primates suggest that adaptation to sudden kinematic distortions may engage the basal ganglia, whereas adaptation to gradual kinematic distortions involves cerebellar structures. In the present studies, participants were patients with PD, who performed center-out pointing movements, using either a digitizer tablet and pen or a computer trackball, under normal or rotated screen cursor feedback conditions. The initial study tested patients with PD using a cross-over experimental design for adaptation to gradual as compared with sudden rotated hand-screen cursor relationships and revealed significant after-effects for the gradual adaptation task only. Consistent with these results, findings from a follow-up experiment using a trackball that required only small finger movements showed that patients with PD adapt better to gradual as against sudden perturbations, when compared to age-matched healthy controls. We conclude that Parkinson’s disease affects adaptation to sudden visuomotor distortions but spares adaptation to gradual distortions.     

Age-related effects in interlimb practice on coding complex movement sequences

Hikosaka et al. (1999) proposed that sequential movements are acquired in independent visual-spatial and motor coordinate systems with coding initially represented in visual-spatial coordinates, and later after extended practice in motor coordinates. One aspect of sequence learning that has not been systematically studied, however, is the question of whether or not older adults show the same pattern of coding in inter-limb practice as younger learners. In the present experiment an inter-limb practice paradigm was designed to determine the role that visual-spatial (Cartesian) and motor (joint angles, activation patterns) coordinates play in the coding and learning of a complex movement sequence. Younger and older adults practiced a 16-element movement sequence with one limb on Day 1 and the contra-lateral limb on Day 2. Practice involved the same sequence with either the same visual-spatial or motor coordinates on the two days. Retention tests were conducted on Day 3. Results indicated that keeping the visual-spatial coordinates the same during acquisition resulted in superior retention only for younger adults. Results also indicated the overall slowing of sequential movement production for older adults which appears to result from these participants inability to impose a structure on the sequence. This provides strong evidence that the visual-spatial code plays a dominant role in complex movement sequences and this code is represented in an effector-independent manner for younger adults, but not for older adults.     

Goal-directed imitation: the means to an end

The effects of goal-directed imitation and observational learning were examined whilst learning a goal-directed motor skill (three-ball cascade juggling). An observational learning (OL) group observed a model and a control (CON) group received minimal verbal instructions regarding how to hold and release the juggling balls. The OL group performed more juggling cycles across practice and retention than the CON group. In addition, the OL group's upper limb coordination and ball flight trajectory pattern were more similar to the model's movements than the CON group. These data show that when the to-be-learnt movement pattern and end-goal are not specified by the task's mechanical constraints, or can be achieved by modifying a pre-existing motor skill, individuals have difficulty learning on the basis of discovery processes alone. Under these circumstances, observational learning is effective because it conveys to the individual the specific means by which the end-goal can be achieved. These findings lead us to suggest that when the end-goal and the means to achieve the end-goal are directly linked, the means are given sufficient weight in the goal hierarchy such that the model's movement is imitated.     

The impact of concurrent visual feedback on coding of on-line and pre-planned movement sequences

The purpose of this study was to determine the extent to which participants could effectively switch from on-line (OL) to pre-planned (PP) control (or vice versa) depending on previous practice conditions and whether concurrent visual feedback was available during transfer testing. The task was to reproduce a 2000 ms spatial–temporal pattern of a sequence of elbow flexions and extensions. Participants were randomly assigned to one of two practice conditions termed OL or PP. In the OL condition the criterion waveform and the cursor were provided during movement production while this information was withheld during movement production for the PP condition. A retention test and two effector transfer tests were administered to half of the participants in each acquisition conditions under OL conditions and the other half under PP conditions. The mirror effector transfer test required the same pattern of muscle activation and limb joint angles as required during acquisition. The non-mirror transfer test required movements to the same visual–spatial locations as experienced during acquisition. The results indicated that when visual information was available during the transfer tests performers could switch from PP to OL. When visual information was withdrawn, they shifted from the OL to the PP-control mode. This finding suggests that performers adopt a mode of control consistent with the feedback conditions provided during testing.     

Solution strategies as possible explanations of individual and sex differences in a dynamic spatial task

When individuals perform spatial tasks, individual differences emerge in accuracy and speed as well as in the response patterns used to cope with the task. The purpose of this study is to identify, through empirical criteria, the different response patterns or strategies used by individuals when performing the dynamic spatial task presented in the Spatial Orientation Dynamic Test-Revised (SODT-R). Results show that participants can be classified according to their response patterns. Three different ways of solving a task are described, and their relation to (a) performance factors (response latency, response frequency, and invested time) and (b) ability tests (analytical reasoning, verbal reasoning, and spatial estimation) are investigated. Sex differences in response patterns and performance are also analyzed. It is found that the frequency with which men and women employ each one of the strategies described here, is different and statistically significant. Thus, employed strategy plays an important role when interpreting sex differences on dynamic spatial tasks.     

Role of action observation and action in sequence learning and coding

A complex sequence learning task was used to determine if the type of coding acquired through physical practice (PP), observation of the stimulus (Obs-S), or observation of stimulus and action (Obs-SA) differs between conditions and whether the type of observation influences subsequent learning of the task when physical practice was permitted. Participants in the Obs-S group were permitted to watch the sequentially illuminated stimuli on the screen. In the Obs-SA group participants could see both flexion-extension movements of the model's arm performing the sequence and the sequentially illuminated stimuli on the screen. Participants in the PP group actually performed the 16-element sequence with their dominant right arm. Delayed retention tests and two inter-manual transfer tests were completed following each of two acquisition sessions. First, the data indicated that learning the sequence structure, as revealed by response time per element, occurred similarly irrespective of the initial practice condition. Secondly, the movement sequence appeared to be coded in abstract visual-spatial coordinates resulting in effector-independent performance. Finally, observing the model's action and sequential stimuli allows participants to transfer the perceived aspects of the movement sequence into efficient coordination patterns when additional physical practice is permitted.     

Cognitive underpinnings of contextual interference during motor learning

The reported study examined the cognitive processes underlying contextual interference (CI) in motor learning. This experiment was designed to assess the combined influence of practice schedule (blocked or random) and task similarity (similar or dissimilar) on acquisition and retention performance. Participants (N = 60) learned a set of three variations of a timing task according to a similar (900, 1000 and 1100 ms) or dissimilar parameter condition (700, 1000 and 1300 ms) with either a blocked or random practice order: this resulted in 4 experimental groups. Performance in delayed retention demonstrated a typical CI effect due to the schedule of practice for the dissimilar parameter condition with the random practice group outperforming the blocked practice group. Conversely, no blocked-random difference was found for the similar parameter condition. These findings lend support for the reconstruction hypothesis by showing that supplementing random practice with additional intertask elaboration (i.e., similar parameter condition) did not facilitate subsequent retention performance.     

Interference within hands: Retrieval-induced forgetting of left and right hand movements

We examined retrieval-induced forgetting of motor sequences that were categorized by the effectors (left or right hand) involved in their execution. This left–right categorization was independent from input locations or input devices. In addition, the acquired motor sequences were arbitrarily assigned to left and right. Participants learned twelve sequential joystick movements as responses to letter stimuli. Half of the sequences pertained to the left, half to the right hand. Subsequent retrieval-practice of half the items of one hand induced forgetting for the non-retrieved rest of the items of that hand in a final recall test. This finding demonstrates that the hands were used to organize the memory storage of motor sequences in a way that gave rise to later interference between commonly stored items, that is, linked to the same hand.     

Retrieval practice in motor learning

In this study we sought to determine whether testing promotes the generalization of motor skills during the process of encoding and/or consolidation. We used a dynamic arm movement task that required participants to reproduce a spatial-temporal pattern of elbow extensions and flexions with their dominant right arm. Generalization of motor learning was tested by the ability to transfer the original pattern (extrinsic transformation) or the mirrored pattern (intrinsic transformation) to the unpractised left arm. To investigate the testing effects during both encoding and consolidation processing, participants were administered an initial testing session during early practice before being evaluated on a post-practice testing session administered either 10 min (Testing-Encoding group) or 24 hr apart (Testing-Consolidation group), respectively. Control groups were required to perform a post-practice testing session administered after either a 10-min (Control-Encoding group) or 24-hr delay (Control-Consolidation group). The findings revealed that testing produced rapid, within-practice skill improvements, yielding better effector transfer at the 10-min testing for the Testing-Encoding group on both extrinsic and intrinsic transformation tests when compared with the Control-Encoding group. Furthermore, we found better performance for the Testing-Consolidation group at the 24-hr testing for extrinsic and intrinsic transformations of the movement pattern when compared with the Control-Consolidation group. However, our results did not indicate any significant testing advantage on the latent, between-session development of the motor skill representation (i.e., from the 10-min to the 24-hr testing). The testing benefits expressed at the 10-min testing were stabilised but did not extend during the period of consolidation. This indicates that testing contributes to the generalisation of motor skills during encoding but not consolidation.     

Are correlated streams of information necessary for implicit sequence learning?

We investigated whether the existence of correlated streams of information is necessary for incidental sequence learning to occur. We ran three separate experiments with a total of 201 undergraduate students. Each experiment had at least one condition with two streams of sequenced information that were correlated. The correlations differed in terms of the kind of responses that were required, the kind of tasks and stimuli, the on-screen locations at which they occurred and how they were combined. Only in conditions with correlated sequences was implicit sequence learning found. Our results suggest that the presence of correlated streams of information may be the main pre-requisite for implicit sequence learning.     

Moving and memorizing: Motor planning modulates the recency effect in serial and free recall

Motor planning has generally been studied in situations where participants carry out physical actions without a particular purpose. Yet in everyday life physical actions are usually carried out for higher-order goals. We asked whether two previously discovered motor planning phenomena - the end-state comfort effect and motor hysteresis - would hold up if the actions were carried out in the service of higher-order goals. The higher-order goal we chose to study was memorization. By focusing on memorization, we asked not only how and whether motor planning is affected by the need to memorize, but also how memory performance might depend on the cognitive demands of motor planning. We asked university-student participants to retrieve cups from a column of drawers and memorize as many letters as possible from the inside of the cups. The drawers were opened either in a random order (Experiment 1) or in a regular order (Experiments 2 and 3). The end-state comfort effect and motor hysteresis were replicated in these conditions, indicating that the effects hold up when physical actions are carried out for the sake of a higher-order goal. Surprisingly, one of the most reliable effects in memory research was eliminated, namely, the tendency of recent items to be recalled better than earlier items - the recency effect. This outcome was not an artifact of memory being uniformly poor, because the tendency of initial items to be recalled better than later items - the primacy effect - was obtained. Elimination of the recency effect was not due to the requirement that participants recall items in their correct order, for the recency effect was also eliminated when the items could be recalled in any order (Experiment 3). These and other aspects of the results support recent claims for tighter links between perceptual-motor control and intellectual (symbolic) processing than have been assumed in the past.     

Grasp posture planning during multi-segment object manipulation tasks — Interaction between cognitive and biomechanical factors

The present study examined adaptations in the planning of initial grasp postures during a multi-segment object manipulation task. Participants performed a grasping and placing task that consisted of one, two, or three movement segments. The position of the targets was manipulated such that the degree of object rotation between the home and temporally proximal positions, and between the temporally proximal and distal target positions, varied. Participants selected initial grasp postures based on the specific requirements of the temporally proximal and temporally distal action segments, and adjustments in initial grasp posture depended on the temporal order of target location. In addition, during the initial stages of the experimental session initial grasp postures were influenced to a larger extent by the demands of the temporally proximal segment. However, over time, participants overcame these cognitive limitations and adjusted their initial grasp postures more strongly to the requirements of the temporally distal segment. Taken together, these results indicate that grasp posture planning is influenced by cognitive and biomechanical factors, and that participants learn to anticipate the task demands of temporally distal task demands, which we hypothesize, reduce the burden on the central nervous system.     

Manipulability impairs association-memory: Revisiting effects of incidental motor processing on verbal paired-associates

Imageability is known to enhance association-memory for verbal paired-associates. High-imageability words can be further subdivided by manipulability, the ease by which the named object can be functionally interacted with. Prior studies suggest that motor processing enhances item-memory, but impairs association-memory. However, these studies used action verbs and concrete nouns as the high- and low-manipulability words, respectively, confounding manipulability with word class. Recent findings demonstrated that nouns can serve as both high- and low-manipulability words (e.g., CAMERA and TABLE, respectively), allowing us to avoid this confound. Here participants studied pairs of words that consisted of all possible pairings of high- and low-manipulability words and were tested with immediate cued recall. Recall was worse for pairs that contained high-manipulability words. In free recall, participants recalled more high- than low-manipulability words. Our results provide further evidence that manipulability influences memory, likely occurring through automatic motor imagery.     

Contributions from associative and explicit sequence knowledge to the execution of discrete keying sequences

Research has provided many indications that highly practiced 6-key sequences are carried out in a chunking mode in which key-specific stimuli past the first are largely ignored. When in such sequences a deviating stimulus occasionally occurs at an unpredictable location, participants fall back to responding to individual stimuli (Verwey & Abrahamse, 2012). The observation that in such a situation execution still benefits from prior practice has been attributed to the possibility to operate in an associative mode. To better understand the contribution to the execution of keying sequences of motor chunks, associative sequence knowledge and also of explicit sequence knowledge, the present study tested three alternative accounts for the earlier finding of an execution rate increase at the end of 6-key sequences performed in the associative mode. The results provide evidence that the earlier observed execution rate increase can be attributed to the use of explicit sequence knowledge. In the present experiment this benefit was limited to sequences that are executed at the moderately fast rates of the associative mode, and occurred at both the earlier and final elements of the sequences.     

Distinct modes of executing movement sequences: reacting, associating, and chunking

Responding to individual key-specific stimuli in entirely unfamiliar keying sequences is said to involve a reaction mode. With practice, short keying sequences can be executed in the chunking mode. This is indicated by the first stimulus sufficing for rapid execution of the entire sequence. The present study explored whether an associative mode develops also in participants who practice short keying sequences. This associative mode would involve priming by earlier events of responses to external stimuli, and is believed to be responsible for skill in the Serial Reaction Time task. In the present study participants practiced two discrete 6-key sequences. In the ensuing test phase, participants were prevented from using the chunking mode by including two deviant stimuli in most sequences. The results from the remaining - unchanged - familiar sequences confirmed that participants no longer used the chunking mode, but as predicted by associative learning these sequences were executed faster than unfamiliar sequences.     

Individual differences in Sternberg’s memory scanning task

This study provides a new perspective on both the cognitive processes actually implemented and the effect of a simple experimental control - the recall constraint - in Sternberg’s memory scanning task. These findings were highlighted by adopting a new approach based on the comparison of qualitative and quantitative results.The analysis of individual processing, on 72 adults, each participating in one of two experimental conditions (with or without sequence recall), highlighted a large variability in quantitative results as well as qualitative procedures. Based on the participants’ retrospective verbalisations, two categories of strategies were identified: (1) the procedures used to memorize the sequence of digits, and (2) the procedures used to compare this sequence with the test digit, which includes strategies for coding the items and processes for searching them in memory. The analysis of the strategies shows that their frequencies of use depend not only on the experimental condition, but also on the participants, the level of task difficulty and the interaction between participants and level of difficulty. This variability questions the accuracy of Sternberg’s mean model. Furthermore, this approach suggests some answers to the old debate concerning the exhaustive search pattern for the yes response. Indeed, our results show three types of strategies that can be identified according to the different models of search suggested in the literature. The “exhaustive” search, that would only be involved in the recall condition and only for some of the participants, the “self-terminating” search and the “immediate” strategy, which can be identified with a model of parallel search with limited resources. Thus our study suggests that the different search models are appropriate but depend on both the specific experimental conditions and participant’s strategy. Our results should help to improve the interpretation of data collected with this paradigm in cognitive and neuroscientific studies of memory.     

Eye movements are not a prerequisite for learning movement sequence timing through observation

The present experiment examined learning of a three-segment movement sequence using physical or observational practice, and whether permitting eye movements to be made during observation is a prerequisite for learning such a movement sequence. Specifically, participants were required to move a mouse cursor through a three-segment movement sequence in order to satisfy one of three movement time goals (800, 1000, 1200 ms). A yoked-participant design was used in which a physical practice group acted as a learning model, which was viewed simultaneously by two groups that carried out different observational practice procedures. An observation group was permitted to move their eyes whilst observing the model, whereas the fixation group was instructed to maintain fixation on a central target. The difference between pre-test and post-test data indicated that all the three experimental groups significantly altered their timing accuracy, variability and movement kinematics over practice, while the control group’s behaviour was unchanged. These data indicate that movement time as well as the underlying movement control was learned following observation of a movement with or without an explicit contribution from eye movements, albeit to a lesser extent during the final segment of the sequence when compared to the physical practice group. The implication is that while similar processes might normally be involved in physical and observational practice, information afforded by eye movements during observation (e.g., efference copy and eye proprioception) is not necessary for movement sequence learning.