Representation of movement sequences is related to task characteristics

Recent experiments have produced mixed results in terms of performance when, after learning a sequential task, the same visual-spatial coordinates or the same motor coordinates were reinstated on a subsequent effector transfer test. Given the diversity of tasks and especially sequence characteristics used in previous experiments, the cross-experimental comparison makes inferences and unambiguous interpretations difficult. The purpose of the present experiment was to determine in a principled manner how the spatio-temporal structure of a sequence influences the way the sequence is represented. The results indicated that after limited amount of practice relatively more simple sequences (S1) are coded more efficiently in a mirror (motor) representation which requires the same pattern of homologous muscle activation. Conversely, relatively more complex sequences (S2) are more efficiently coded in a visual-spatial coordinate system which requires movements to the same spatial locations as during acquisition. The data are also consistent with the notion that sequences with different spatio-temporal structures rely to a different degree on distinct control mechanisms (pre-planned vs. on-line, respectively).     

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.     

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.     

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.     

The influence of writing practice on letter recognition in preschool children: a comparison between handwriting and typing

A large body of data supports the view that movement plays a crucial role in letter representation and suggests that handwriting contributes to the visual recognition of letters. If so, changing the motor conditions while children are learning to write by using a method based on typing instead of handwriting should affect their subsequent letter recognition performances. In order to test this hypothesis, we trained two groups of 38 children (aged 3-5 years) to copy letters of the alphabet either by hand or by typing them. After three weeks of learning, we ran two recognition tests, one week apart, to compare the letter recognition performances of the two groups. The results showed that in the older children, the handwriting training gave rise to a better letter recognition than the typing training.     

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.     

Consistently modeling the same movement strategy is more important than model skill level in observational learning contexts

The experiment undertaken was designed to elucidate the impact of model skill level on observational learning processes. The task was bimanual circle tracing with a 90° relative phase lead of one hand over the other hand. Observer groups watched videos of either an instruction model, a discovery model, or a skilled model. The instruction and skilled model always performed the task with the same movement strategy, the right-arm traced clockwise and the left-arm counterclockwise around circle templates with the right-arm leading. The discovery model used several movement strategies (tracing-direction/hand-lead) during practice. Observation of the instruction and skilled model provided a significant benefit compared to the discovery model when performing the 90° relative phase pattern in a post-observation test. The observers of the discovery model had significant room for improvement and benefited from post-observation practice of the 90° pattern. The benefit of a model is found in the consistency with which that model uses the same movement strategy, and not within the skill level of the model. It is the consistency in strategy modeled that allows observers to develop an abstract perceptual representation of the task that can be implemented into a coordinated action. Theoretically, the results show that movement strategy information (relative motion direction, hand lead) and relative phase information can be detected through visual perception processes and be successfully mapped to outgoing motor commands within an observational learning context.     

Offline improvement occurs for temporal stability but not accuracy following practice of integer and non-integer rhythms

Procedural learning benefits from memory processes occurring outside practice resulting in offline learning. Offline gains have been demonstrated almost exclusively for the ordinal structure of sequential motor tasks. Many skills also demand that the correct serial order of events be appropriately timed. Evidence indicates that the temporal aspect of a procedural skill can be encoded independent of serial order knowledge and governed by at least two distinct neural circuits. The present experiment determined if (a) offline gains emerge for temporal learning, and (b) if such gains occur for timing supervised by distinct timing systems. Participants experienced 216 practice trials of a 7-key press sequence that involved integer- or non-integer timing rhythms. Twenty-four hours after training 30 test trials were administered. Results revealed robust offline enhancement for timing performance of the non-integer based temporal sequences. This improvement was localized to stabilization of the required relative but not absolute time profiles. The neural circuitry central to supporting the performance of non-integer timing sequences is also a principal constituent of what is described as the "cognitive" timing system. Timing governed by this system appears most susceptible to offline gains via consolidation.     

The direction of bilateral transfer depends on the performance parameter

To acquire a more comprehensive understanding of the learning benefits associated with bilateral transfer and to gain knowledge of possible mechanisms behind bilateral transfer, we investigated the transfer direction of several parameters which are assumed to represent important features of movement control in a visuo-motor task. During the study, participants learned a multidirectional point-to-point drawing task in which the visual feedback was rotated 45° and the gain was increased. Performance changes of the untrained hand in movement time, trajectory length, normalized jerk, initial direction error, ratio of the primary sub-movement time to the total movement time, and the accuracy of the aiming movement after the primary sub-movement were investigated as indices of learning from bilateral transfer. The results showed that performance parameters related to the initial production of the movement, such as the initial direction, ratio of primary sub-movement to the total movement time, and movement accuracy after the primary sub-movement, only transferred to the non-dominant, while hand performance variables related to the overall outcome, such as movement duration, movement smoothness, and trajectory length, transferred in both directions. The findings of the current study support the basic principle of the “dynamic dominance model” because it is suggested that overall improvements in the non-dominant system are controlled by trajectory parameters in visuo-motor tasks, which resulted in transference of the afore mentioned production parameters to rather occur to the non-dominant hand as opposed to transference to the dominant hand.     

Adaptation to direction-dependent visuo-motor rotations and its decay in younger and older adults

We studied adaptation to a direction-dependent visuo-motor rotation in adults of early and late working age. For hand movements to the right, visual motion of the cursor on a monitor was rotated clockwise, for forward movements rotation of the cursor motion was zero, and for directions in-between rotation was intermediate. In contrast to previous studies, in which adaptation was more difficult (larger visuo-motor rotation, larger number of targets during practice) and the older age group was of higher age, we found no age-related deficit of adaptation. However, consistent with previous studies we found an age-related impairment of explicit knowledge of the visuo-motor rotation and no age-related differences of aftereffects. Across periods of not performing the task for 24 h and of performing the task for a prolonged period of time without visual feedback, we observed a decay of adaptation which did not depend on age. The present findings cast doubts on the prevalent interpretation of age-related impairments of adaptation in the absence of age-related changes of aftereffects as resulting from intentional strategic corrections, which become less efficient at higher age.     

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.     

When the mask falls: The role of facial motor resonance in memory for emotional language

The recognition and interpretation of emotional information (e.g., about happiness) has been shown to elicit, amongst other bodily reactions, spontaneous facial expressions occurring in accordance to the relevant emotion (e.g. a smile). Theories of embodied cognition act on the assumption that such embodied simulations are not only an accessorial, but a crucial factor in the processing of emotional information. While several studies have confirmed the importance of facial motor resonance during the initial recognition of emotional information, its role at later stages of processing, such as during memory for emotional content, remains unexplored. The present study bridges this gap by exploring the impact of facial motor resonance on the retrieval of emotional stimuli. In a novel approach, the specific effects of embodied simulations were investigated at different stages of emotional memory processing (during encoding and/or retrieval). Eighty participants underwent a memory task involving emotional and neutral words consisting of an encoding and retrieval phase. Depending on the experimental condition, facial muscles were blocked by a hardening facial mask either during encoding, during retrieval, during both encoding and retrieval, or were left free to resonate (control). The results demonstrate that not only initial recognition but also memory of emotional items benefits from embodied simulations occurring during their encoding and retrieval.     

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.     

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.     

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.     

Increased dynamic regulation of postural tone through Alexander Technique training

Gurfinkel and colleagues (2006) recently found that healthy adults dynamically modulate postural muscle tone in the body axis during anti-gravity postural maintenance and that this modulation is inversely correlated with axial stiffness. Our objective in the present study was to investigate whether dynamic modulation of axial postural tone can change through training. We examined whether teachers of the Alexander Technique (AT), who undergo “long-term” (3-year) training, have greater modulation of axial postural tone than matched control subjects. In addition, we performed a longitudinal study on the effect of “short-term” (10-week) AT training on the axial postural tone of individuals with low back pain (LBP), since short term AT training has previously been shown to reduce LBP. Axial postural tone was quantified by measuring the resistance of the neck, trunk and hips to small (±10°), slow (1°/s) torsional rotation during stance. Modulation of tone was determined by the torsional resistance to rotation (peak-to-peak, phase-advance, and variability of torque) and axial muscle activity (EMG). Peak-to-peak torque was lower (∼50%), while phase-advance and cycle-to-cycle variability were enhanced for AT teachers compared to matched control subjects at all levels of the axis. In addition, LBP subjects decreased trunk and hip stiffness following short-term AT training compared to a control intervention. While changes in static levels of postural tone may have contributed to the reduced stiffness observed with the AT, our results suggest that dynamic modulation of postural tone can be enhanced through long-term training in the AT, which may constitute an important direction for therapeutic intervention.     

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.     

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.     

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.     

Covariation of attentional cost and stability provides further evidence for two routes to learning new coordination patterns

This study investigated how learning a new bimanual coordination pattern affects the attentional resources allotted by the CNS to maintain it throughout the acquisition process. The repertoire of the existing stable coordination patterns was individually evaluated before and after practice in order to detect expected changes with learning. Bistable participants, who initially exhibited stable and accurate coordination patterns at 0° and 180° of relative phase, practiced a 90° pattern, whereas multistable participants, who already mastered the 90° pattern, practiced 135° pattern instead. In a typical dual-task paradigm, all participants had to simultaneously perform a reaction time task that assessed the associated attentional cost. Beyond an overall increase in accuracy, the results revealed a significant decrease in the attentional cost for bistable participants, accompanying the stabilization of the 90° pattern with learning, but not for multistable participants, as the 135° pattern barely stabilized. Pattern stability and attentional cost co-evolve during learning and the process follows two different routes depending on the interplay between the task and the learner’s coordination abilities before practice.