Memory consolidation of sequence learning and dynamic adaptation during wakefulness.

Motor learning involves acquiring new movement sequences and adapting motor commands to novel conditions. Labile motor memories, acquired through sequence learning and dynamic adaptation, undergo a consolidation process during wakefulness after initial training. This process stabilizes the new memories, leading to long-term memory formation. However, it remains unclear if the consolidation processes underlying sequence learning and dynamic adaptation are independent and if distinct neural regions underpin memory consolidation associated with sequence learning and dynamic adaptation. Here, we first demonstrated that the initially labile memories formed during sequence learning and dynamic adaptation were stabilized against interference through time-dependent consolidation processes occurring during wakefulness. Furthermore, we found that sequence learning memory was not disrupted when immediately followed by dynamic adaptation and vice versa, indicating distinct mechanisms for sequence learning and dynamic adaptation consolidation. Finally, by applying patterned transcranial magnetic stimulation to selectively disrupt the activity in the primary motor (M1) or sensory (S1) cortices immediately after sequence learning or dynamic adaptation, we found that sequence learning consolidation depended on M1 but not S1, while dynamic adaptation consolidation relied on S1 but not M1. For the first time in a single experimental framework, this study revealed distinct neural underpinnings for sequence learning and dynamic adaptation consolidation during wakefulness, with significant implications for motor skill enhancement and rehabilitation.

Off-line learning in a rhythmic bimanual task: early feedback dependency is reduced over wakefulness.

Research has supported two distinct forms of motor skill consolidation that can occur between practice sessions: (1) off-line learning, and (2) memory stabilization. Off-line learning describes performance improvement between practice sessions that is above the gain observed at the end of practice, while memory stabilization describes a gain in performance that is maintained between practice sessions. This study used a Lissajous plot to provide concurrent feedback to train participants to produce a 90° relative phase between the index fingers (flexion/extension motion). Significant improvements in performance emerged after ten trials (5 min) of practice. At the end of training, participants were divided into two delay interval groups before retesting, 2-h and 6-h. The retesting session started with participants performing an interference task (10 trials, 5 min) that required training on a 45° relative phase between the fingers with concurrent feedback from the Lissajous plot. When training with the interference task was completed participants were retested with the 90° relative phase without the Lissajous plot feedback. In the retest of the 90° pattern, a performance loss was found in the 2-h delay group, whereas the 6-h delay group maintained the end of practice performance level. Maintenance of the same level of performance without the Lissajous plot represents memory stabilization of the initially trained 90° pattern. The findings are discussed within the context of current positions regarding procedural consolidation and the coordination dynamics framework wherein action and perception are linked through the informational nature of relative phase.

Motor and spatial representations of action: corticospinal excitability in M1 after training with a bimanual skill.

The purpose of the study was twofold: (1) determine if different time delays (30 min or 6 h) between training and a post-training test with a rhythmic bimanual pattern (90° relative phase) would be associated with different levels of consolidation for the motor and spatial representations of the pattern; and (2) determine if training with the rhythmic bimanual pattern would lead to enhanced corticospinal excitability in M1 linked to changes in motor and spatial performance measures. Coordination accuracy and stability of the 90° pattern improved over practice. Coordination accuracy and stability were the same after a 30-min or 6-h delay between training and the post-training test, indicating equivalent levels of consolidation in the motor representation. The 6-h delay interval resulted in shorter visual recognition times compared to the 30-min delay and was centered on the trained 90° pattern. These findings indicate the consolidation of the spatial representation was more time sensitive compared to the motor representation in the current task. Motor evoked potentials (MEPs) from the first dorsal interosseous muscle (FDI) generated by single-pulse transcranial magnetic stimulation (TMS) were measured at baseline (before training) and at 6-min and 21-min intervals post-training with the 90° pattern. Increased corticospinal excitability in M1 was evidenced by larger MEPs of the FDI muscle at the 6-min interval. This increased excitability after training is a novel finding after training with a difficult and initially unstable rhythmic bimanual pattern. No significant correlations were found between the MEP data and behavioral data; thus, the increased excitability in M1 may have been linked to the difficulty in performing the pattern, consolidation processes, or both.

Application of anodal tDCS at primary motor cortex immediately after practice of a motor sequence does not improve offline gain.

Tecchio et al. (J Neurophysiology 104: 1134-1140, 2010) reported that the application of anodal tDCS at primary motor cortex (M1) immediately after practice of a procedural motor skill enhanced consolidation, which in turn improved offline gain. Tecchio et al. noted, however, that this study did not account for known after-effects associated with this form of non-invasive stimulation. The present study was designed to explicitly reevaluate Tecchio et al.'s claim. As in the original study, individuals experienced either anodal or sham stimulation at M1 after practice of a serial reaction time task (SRTT) followed by test trials 15-min later. Two additional novel conditions experienced the test trials after 120-min rather than 15-min thus allowing potential stimulation after-effects to dissipate. The expectation was that if anodal stimulation influences post-practice consolidation leading to offline gain, this effect would be present not only at 15-min but also after 120-min. In agreement with the working hypothesis, findings revealed offline gain at both 15-min and the longer 2-h time period. Unexpectedly, we found no interaction between real and sham conditions. The lack of difference between Real and Sham effects weakens confidence in the potential of post-practice tDCS for consolidation enhancement, while it is more consistent with other claims that decoupling practice and anodal tDCS stimulation in time can reduce the effectiveness of exogenous stimulation for procedural skill gain.

Mirror-hand selection is influenced by training perspective and model skill level in a motor-learning task.

This study examined mirror and non-mirror arm selection processes in an observational learning context. Observer groups watched either a novice (instruction or discovery) or skilled model performing a bimanual task with the right arm leading the left arm. The models were viewed from a third-person perspective. Observers of the skilled model more often selected a mirror-image (left-hand) hand-lead in post-observations tests, while observers of the novice models more often selected a non-mirror image (right hand) hand-lead in post-observation tests. This is a novel finding regarding arm selection processes in a learning context, yet it is consistent with imaging data that has revealed specific neural areas linked to the selection of mirror and non-mirror imitation processes for first- and third-person viewing perspectives. The skilled model also supported more accurate and stable performance of the bimanual task in observers compared to the instruction and novice models. It is concluded that a skilled model supports attention focus being directed at pattern analysis, while novice models support attention focus being allocated to strategy identification first, followed by pattern analysis.

Bimanual coordination patterns are stabilized under monitoring-pressure.

The influence of monitoring-pressure on the performance of anti-phase and in-phase bimanual coordination was examined. The two bimanual patterns were produced under no-monitoring and monitoring-pressure conditions at self-paced frequencies. Anti-phase coordination was always less stable than in-phase coordination, with or without monitoring. When performed under monitoring-pressure, the coordination patterns were performed with less variability in relative phase for both patterns across a range of self-paced movement frequencies compared to performance without monitoring. Thus, while monitoring-pressure did induce a behavioral change, it consisted of performance stabilization rather than degradation, a finding inconsistent with explicit-monitoring theory. However, the findings are consistent with the theory of coordination dynamics and studies that have revealed increased stability for the system's intrinsic dynamics as a result of attentional focus and intentional control.

Observation and physical practice: different practice contexts lead to similar outcomes for the acquisition of kinematic information.

This study differentiated the contributions of physical and observational practice to the learning of a single-limb multi-joint coordination pattern. Three groups (physical-practice, observation-practice, observation-physical) practiced for 2 days and were given two performance tests 24 h after the second practice session. The performance tests revealed that physical and observational practice contributed similarly to identifying and using kinematic information related to the relative motion direction between joints (lead/lag relationship) and to the to-be-learned relative phase pattern (ϕ = 90°). Physical practice resulted in more stable coordination during performance tests and in the ability to produce different joint amplitudes with less variability. A serendipitous finding was that maximum elbow flexion (point of movement reversal) emerged as a kinematic event around which elbow and wrist coordination were organized. Movement reversals often serve to anchor the movement dynamics, and this anchoring effect was evident following both physical and observational practice, yet physical practice resulted in an advantage with regard to this anchor point on several kinematic measures. The results are discussed within the context of contemporary behavioral theories (coordination dynamics, visual perception perspective) of observational learning.

The Coordination Dynamics of Observational Learning: Relative Motion Direction and Relative Phase as Informational Content Linking Action-Perception to Action-Production.

The primary goal of this chapter is to merge together the visual perception perspective of observational learning and the coordination dynamics theory of pattern formation in perception and action. Emphasis is placed on identifying movement features that constrain and inform action-perception and action-production processes. Two sources of visual information are examined, relative motion direction and relative phase. The visual perception perspective states that the topological features of relative motion between limbs and joints remains invariant across an actor's motion and therefore are available for pickup by an observer. Relative phase has been put forth as an informational variable that links perception to action within the coordination dynamics theory. A primary assumption of the coordination dynamics approach is that environmental information is meaningful only in terms of the behavior it modifies. Across a series of single limb tasks and bimanual tasks it is shown that the relative motion and relative phase between limbs and joints is picked up through visual processes and supports observational learning of motor skills. Moreover, internal estimations of motor skill proficiency and competency are linked to the informational content found in relative motion and relative phase. Thus, the chapter links action to perception and vice versa and also links cognitive evaluations to the coordination dynamics that support action-perception and action-production processes.

The perception-action dynamics of action competency are altered by both physical and observational training.

Action competency is defined as the ability of an individual to self-evaluate their own performance capabilities. The current experiment demonstrated that physical and observational training with a motor skill alters action competency ratings in a similar manner. Using a pre-test and post-test protocol, the results revealed that action competency is constrained prior to training by the intrinsic dynamics of relative phase (ϕ), with in-phase (ϕ = 0°) and anti-phase (ϕ = 180°) patterns receiving higher competency ratings than other relative phase patterns. After 2 days of training, action competency ratings for two trained relative phase patterns, +60° and +120°, increased following physical practice or observational practice. A transfer test revealed that both physical performance ability and action competency ability transferred to the symmetry partners (-60° and -120°) of the two trained relative phase patterns following physical or observational training. The findings also revealed that relative motion direction acts as categorical information that helps to organize action production and facilitate action competency. The results are interpreted based on the coordination dynamics theory of perception-action coupling, and extend this theory by showing that visual perception, action production, and action competency are all constrained in a consistent manner by the dynamics of the order parameter relative phase. As a whole, the findings revealed that relative motion, relative phase, and possibly relative amplitude information are all distinct sources of information that contribute to the emergence of a kinematic understanding of action in the nervous system.

Spontaneity competes with intention to influence the coordination dynamics of interpersonal performance tendencies.

Research has shown that spontaneous visual coupling supports frequency entrainment, phase attraction, and intermittent interpersonal coordination when co-actors are switched from a no-vision (NV) to vision (V) context. In two experiments, co-actors started in a NV context while producing the same or different amplitude movements. The same amplitude resulted in similar self-paced frequencies, while different amplitudes resulted in disparate frequencies. In experiment 1, co-actors were instructed to maintain amplitude while receiving no instructions to coordinate their actions. Frequency and phase entrainment was limited in the V context even when co-actors started the NV context with the same amplitude. In experiment 2, co-actors were instructed to maintain amplitude and intentionally coordinate together, but not at a specific pattern. Significant frequency modulations occurred to maintain amplitude as the co-actors sought to coordinate their actions. With the open-ended instructions, co-actors produced in-phase and anti-phase coordination along with intermittent performance exhibited by shifts between a variety of stable relative phase patterns. The proposed hypotheses and findings are discussed within the context of a shared manifold representation for joint action contexts, with the coordination dynamics expressed by the HKB model of relative phase serving to conceptualization the representations in the shared manifold.