Intra- and interlimb effects of gait retraining in individuals with knee hyperextension.

BACKGROUND: Gait retraining, which typically focuses on the most severely affected limb or joint, has shown promising results in treating faulty running and walking patterns. The closed-chain nature of gait during the stance phase may influence kinematic changes in the adjacent joints of the trained leg. In addition, the coupled nature of the lower extremity motion of gait suggests that changes in one leg may transfer to the other. This study aimed to assess the intra- and inter-limb transfer of kinematic changes following gait retraining to reduce knee extension in individuals with hyperextension walking patterns. METHODS: Seventeen women with knee hyperextension gait patterns participated in six treadmill retraining sessions. All participants received verbal and real-time visual kinematic feedback in the form of knowledge of results. This intervention study took place at the Gait Analysis Laboratory at the University of Iowa. Mean peak sagittal-plane lower extremity joint kinematics during overground walking at pretraining, post-training, and 1- and 8-month follow-ups were calculated for analysis and comparisons. FINDINGS: The post-training changes in ankle range of motion returned to baseline values by the 8-month follow-up. There was a significant transfer effect of kinematic changes to the untrained knee following gait retraining. INTERPRETATION: Training one knee did not result in long-term compensatory kinematic changes in the other joints. In addition, the improvements in knee extension range of motion were transferred to the untrained knee and retained at the 8-month follow-up. This study supports the use of gait retraining as an effective clinical intervention.

Unilateral high-load resistance training induced a similar cross-education of strength between the dominant and non-dominant arm.

It was previously hypothesized that the cross-education of strength is asymmetrical, where a greater transfer of strength is observed from the dominant to the non-dominant limb. The purpose of this study was to examine if the magnitude of cross-education of strength differed between dominant and non-dominant limbs following unilateral high-load resistance training. One hundred and twenty-two participants were randomized to one of the three groups: 1) training on the dominant arm (D-Only), 2) training on the non-dominant arm (ND-Only) and 3) a time-matched non-exercise control (Control). The training groups completed 6 weeks (18 sessions) of unilateral elbow flexion exercise. Each training session started with one-repetition maximum (1RM) training (≤ five attempts), followed by four sets of high-load exercise (i.e. 8-12RM). Strength changes of the untrained arm were compared between groups. Changes in the strength of the untrained arm were greater in D-Only (1.5 kg) and ND-Only (1.3 kg) compared to Control (-0.2 kg), without differences between D-Only and ND-Only. Unilateral resistance training increased strength in the opposite untrained arm, and the magnitude of this effect was similar regardless of which arm was trained. However, there is still considerable uncertainty on this topic and additional research is warranted to confirm the current findings.

Lateralization of acquisition and consolidation in direction but not amplitude of a motor skill task.

Previous research suggests that the neural processes underlying specification of movement direction and amplitude are independently represented in the nervous system. However, our understanding of acquisition and consolidation processes in the direction and distance learning remains limited. We designed a virtual air hockey task, in which the puck direction is determined by the hand direction at impact, while the puck distance is determined by the amplitude of the velocity. In two versions of this task, participants were required to either specify the direction or the distance of the puck, while the alternate variable did not contribute to task success. Separate groups of right-handed participants were recruited for each task. Each participant was randomly assigned to one of two groups with a counter-balanced arm practice sequence (right to left, or left to right). We examined acquisition and, after 24 h, we examined two aspects of consolidation: 1) same hand performance to test the durability and 2) the opposite hand to test the effector-independent consolidation (interlimb transfer) of learning. The distance task showed symmetry between hands in the extent of acquisition as well as in both aspects of consolidation. In contrast, the direction task showed asymmetry in both acquisition and consolidation: the dominant right arm showed faster and greater acquisition and greater transfer from the opposite arm training. The asymmetric acquisition and consolidation processes shown in the direction task might be explained by lateralized control and mapping of direction, an interpretation consistent with previous findings on motor adaptation paradigms.

Interlimb transfer of sequential motor learning between upper and lower effectors.

BACKGROUND: Interlimb transfer of sequential motor learning (SML) refers to the positive influence of prior experiences in performing the same sequential movements using different effectors. Despite evidence from intermanual SML, and while most daily living activities involve interlimb cooperation and coordination between the four limbs, nothing is known about bilateral SML transfer between the upper and lower limbs. RESEARCH QUESTION: We examined the transfer of bilateral SML from the upper to the lower limbs and vice versa. METHODS: Twenty-four participants had to learn an initial bilateral SML task using the upper limbs and then performed the same sequence using the lower limbs during a transfer SML task. They performed the reversed situation 1 month apart. The performance was evaluated at the beginning and the end of both initial and transfer SML practice phases. RESULTS: Significant and reciprocal transfer gains in performance were observed regardless of the effectors. Greater transfer gains in performance were observed at the beginning of the transfer SML from the lower to the upper limbs (44 %) but these gains vanished after practice with the transfer effectors (5 %). Although smaller gains were initially achieved in the transfer of SML from the upper to the lower limbs (15 %), these gains persisted and remained significant (9 %) after practice with the transfer effectors. SIGNIFICANCE: Our results provide evidence of a reciprocal and asymmetrical interlimb transfer of bilateral SML between the upper and lower limbs. These findings could be leveraged as a relevant strategy in the context of sports and functional rehabilitation.

Structural Learning Benefits in a Visuomotor Adaptation Task Generalize to a Contralateral Effector.

Structural learning is characterized by facilitated adaptation following training on a set of sensory perturbations all belonging to the same structure (e.g., 'visuomotor rotations'). This generalization of learning is a core feature of the motor system and is often studied in the context of interlimb transfer. However, such transfer has only been demonstrated when participants learn to counter a specific perturbation in the sensory feedback of their movements; we determined whether structural learning in one limb generalized to the contralateral limb. We trained 13 participants to counter random visual feedback rotations between +/-90 degrees with the right hand and subsequently tested the left hand on a fixed rotation. The structural training group showed faster adaptation in the left hand in both feedforward and feedback components of reaching compared to 13 participants who trained with veridical reaching, with lower initial reaching error, and straighter, faster, and smoother movements than in the control group. The transfer was ephemeral - benefits were confined to roughly the first 20 trials. The results demonstrate that the motor system can extract invariant properties of seemingly random environments in one limb, and that this information can be accessed by the contralateral limb.

Bilateral transfer of motor performance as a function of motor imagery training: a systematic review and meta-analysis.

Objective: The objective of this review was to evaluate the efficacy of mental imagery training (MIT) in promoting bilateral transfer (BT) of motor performance for healthy subjects. Data sources: We searched 6 online-databases (Jul-Dec 2022) using terms: "mental practice," "motor imagery training," "motor imagery practice," "mental training," "movement imagery," "cognitive training," "bilateral transfer," "interlimb transfer," "cross education," "motor learning," "strength," "force" and "motor performance." Study selection and data extraction: We selected randomized-controlled studies that examined the effect of MIT on BT. Two reviewers independently determined if each study met the inclusion criteria for the review. Disagreements were resolved through discussion and, if necessary, by a third reviewer. A total of 9 articles out of 728 initially identified studies were chosen for the meta-analysis. Data synthesis: The meta-analysis included 14 studies for the comparison between MIT and no-exercise control (CTR) and 15 studies for the comparison between MIT and physical training (PT). Results: MIT showed significant benefit in inducing BT compared to CTR (ES = 0.78, 95% CI = 0.57-0.98). The effect of MIT on BT was similar to that of PT (ES = -0.02, 95% CI = -0.15-0.17). Subgroup analyses showed that internal MIT (IMIT) was more effective (ES = 2.17, 95% CI = 1.57-2.76) than external MIT (EMIT) (ES = 0.95, 95% CI = 0.74-1.17), and mixed-task (ES = 1.68, 95% CI = 1.26-2.11) was more effective than mirror-task (ES = 0.46, 95% CI = 0.14-0.78) and normal-task (ES = 0.56, 95% CI = 0.23-0.90). No significant difference was found between transfer from dominant limb (DL) to non-dominant limb (NDL) (ES = 0.67, 95% CI = 0.37-0.97) and NDL to DL (ES = 0.87, 95% CI = 0.59-1.15). Conclusion: This review concludes that MIT can serve as a valuable alternative or supplement to PT in facilitating BT effects. Notably, IMIT is preferable to EMIT, and interventions incorporating tasks that have access to both intrinsic and extrinsic coordinates (mixed-task) are preferred over those that involve only one of the two coordinates (mirror-task or normal-task). These findings have implications for rehabilitation of patients such as stroke survivors.

Relationship between the interlimb transfer of a visuomotor learning task and interhemispheric inhibition in healthy humans.

The "interlimb transfer" phenomenon consists of improved performance of the trained and untrained contralateral limbs after unilateral motor practice. We here assessed whether a visuomotor learning task can be transferred from one hemisphere to the other, whether this occurs symmetrically, and the cortical neurophysiological correlates of this phenomenon, focusing on interhemispheric connectivity measures. We enrolled 33 healthy subjects (age range: 24-73 years). Participants underwent two randomized sessions, which investigated the transfer from the dominant to the nondominant hand and vice versa. Measures of cortical and intracortical excitability and interhemispheric inhibition were assessed through transcranial magnetic stimulation before and after a visuomotor task. The execution of the visuomotor task led to an improvement in motor performance with the dominant and nondominant hands and induced a decrease in intracortical inhibition in the trained hemisphere. Participants were also able to transfer the visuomotor learned skill. The interlimb transfer, however, only occurred from the dominant to the nondominant hand and positively correlated with individual learning-related changes in interhemispheric inhibition. We here demonstrated that the "interlimb transfer" of a visuomotor task occurs asymmetrically and relates to the modulation of specific inhibitory interhemispheric connections. The study results have pathophysiological, clinical, and neuro-rehabilitative implications.

To assess the effects of cross-education on strength and motor function in post stroke rehabilitation: a systematic literature review and meta-analysis.

BACKGROUND: Cross-education refers to the increase in motor output of the untrained limb following unilateral training of the opposite limb. Cross education has been shown to be beneficial in clinical settings. OBJECTIVES: This systematic literature and meta-analysis aims to assess the effects of cross-education on strength and motor function in post stroke rehabilitation. DATA SOURCES: MEDLINE, CINAHL, Cochrane Library, PubMed, PEDro, Web of Science, ClinicalTrails.gov and Cochrane Central registers were searched up to 1st October 2022. STUDY SELECTION: Controlled trials using unilateral training of the less affected limb in individuals diagnosed with stroke and English language. DATA SYNTHESIS: Methodological quality was assessed using Cochrane Risk-of-Bias tools. Quality of evidence was evaluated using Grading of Recommendations Assessment, Development and Evaluation. Meta-analyses were performed using RevMan 5.4.1. RESULTS: Five studies capturing 131 participants were included in the review and three studies capturing 95 participants were included in the meta-analysis. Cross education was shown to have a statistically and clinically significant effect on upper limb strength (p < 0.003; SMD 0.58; 95% CI 0.20-0.97; n = 117) and upper limb function (p = 0.04; SMD 0.40; 95% CI 0.02-0.77; n = 119). LIMITATIONS: Small number of studies, with all studies identified as having some risk of bias. Quality of evidence graded 'low' due to limitations and imprecision. CONCLUSION: Cross education may be beneficial in improving strength and motor function in the more affected upper limb post stroke. Further studies are needed as the research into the benefits of cross education in stroke rehabilitation is still limited. Systematic Review Registration Number: PROSPERO (CRD42020219058).

Neural mechanisms mediating cross education: With additional considerations for the ageing brain.

CALVERT, G.H.M., and CARSON, R.G. Neural mechanisms mediating cross education: With additional considerations for the ageing brain. NEUROSCI BIOBEHAV REV 21(1) XXX-XXX, 2021. - Cross education (CE) is the process whereby a regimen of unilateral limb training engenders bilateral improvements in motor function. The contralateral gains thus derived may impart therapeutic benefits for patients with unilateral deficits arising from orthopaedic injury or stroke. Despite this prospective therapeutic utility, there is little consensus concerning its mechanistic basis. The precise means through which the neuroanatomical structures and cellular processes that mediate CE may be influenced by age-related neurodegeneration are also almost entirely unknown. Notwithstanding the increased incidence of unilateral impairment in later life, age-related variations in the expression of CE have been examined only infrequently. In this narrative review, we consider several mechanisms which may mediate the expression of CE with specific reference to the ageing CNS. We focus on the adaptive potential of cellular processes that are subserved by a specific set of neuroanatomical pathways including: the corticospinal tract, corticoreticulospinal projections, transcallosal fibres, and thalamocortical radiations. This analysis may inform the development of interventions that exploit the therapeutic utility of CE training in older persons.

Low-Load Unilateral and Bilateral Resistance Training to Restore Lower Limb Function in the Early Rehabilitation After Total Knee Arthroplasty: A Randomized Active-Controlled Clinical Trial.

Background: Continuous passive motion (CPM) is frequently used during rehabilitation following total knee arthroplasty (TKA). Low-load resistance training (LLRT) using continuous active motion (CAM) devices is a promising alternative. We investigated the effectiveness of CPM compared to LLRT using the affected leg (CAMuni) and both legs (CAMbi) in the early post-operative rehabilitation. HYPOTHESES: (I) LLRT (CAMuni and CAMbi) is superior to CPM, (II) additional training of the unaffected leg (CAMbi) is more effective than unilateral training (CAMuni). Materials and Methods: Eighty-five TKA patients were randomly assigned to three groups, respectively: (i) unilateral CPM of the operated leg; (ii) unilateral CAM of the operated leg (CAMuni); (iii) bilateral alternating CAM (CAMbi). Patients were assessed 1 day before TKA (pre-test), 1 day before discharge (post-test), and 3 months post-operatively (follow-up). PRIMARY OUTCOME: active knee flexion range of motion (ROMFlex). SECONDARY OUTCOMES: active knee extension ROM (ROMExt), swelling, pain, C-reactive protein, quality of life (Qol), physical activity, timed-up-and-go performance, stair-climbing performance, quadriceps muscle strength. Analyses of covariances were performed (modified intention-to-treat and per-protocol). Results: Hypothesis I: Primary outcome: CAMbi resulted in a higher ROMFlex of 9.0° (95%CI -18.03-0.04°, d = 0.76) and 6.3° (95%CI -14.31-0.99°, d = 0.61) compared to CPM at post-test and follow-up, respectively. SECONDARY OUTCOMES: At post-test, C-reactive protein was lower in both CAM groups compared with CPM. Knee pain was lower in CAMuni compared to CPM. Improved ROMExt, reduced swelling, better stair-climbing and timed-up-and-go performance were observed for CAMbi compared to CPM. At follow-up, both CAM groups reported higher Qol and CAMbi showed a better timed-up-and-go performance. Hypothesis II: Primary outcome: CAMbi resulted in a higher knee ROMFlex of 6.5° (95%CI -2.16-15.21°, d = 0.56) compared to CAMuni at post-test. SECONDARY OUTCOMES: At post-test, improved ROMExt, reduced swelling, and better timed-up-and-go performance were observed in CAMbi compared to CAMuni. Conclusions: Additional LLRT of the unaffected leg (CAMbi) seems to be more effective for recovery of function than training of the affected leg only (CAMuni), which may be mediated by positive transfer effects from the unaffected to the affected limb (cross education) and/or preserved neuromuscular function of the trained, unaffected leg. Trial Registration: ClinicalTrials.gov Identifier: NCT02062138.

Training load but not fatigue affects cross-education of maximal voluntary force.

The purpose of this study was to determine the effects of training load (25% vs. 75% of one repetition maximum [1RM]) and fatigue (failure vs. non-failure) during four weeks of unilateral knee extension resistance training (RT) on maximal voluntary force in the trained and the untrained knee extensors. Healthy young adults (n = 42) were randomly assigned to control (CON, n = 9, 24 ± 4.3 years), low-load RT to failure (LLF, n = 11, 21 ± 1.3 years, three sets to failure at 25% of 1RM), high-load RT to failure (HLF, n = 11, 21 ± 1.4 years, three sets to failure at 75% of 1RM), and high-load RT without failure (HLNF, n = 11, 22 ± 1.5 years, six sets of five repetitions at 75% of 1RM) groups. Before and after the four weeks of training, 1RM, maximal voluntary isometric force, and corticospinal excitability (CSE) were measured. 1RM in the trained (20%, d = 0.70, 15%, d = 0.61) and the untrained knee extensors (5%, d = 0.27, 6%, d = 0.26) increased only in the HLF and HLNF groups, respectively. MVIC force increased only in the trained leg of the HLF (5%, d = 0.35) and HLNF groups (12%, d = 0.67). CSE decreased in the VL of both legs in the HLNF group (-19%, d = 0.44) and no changes occurred in the RF. In conclusion, high- but not low-load RT improves maximal voluntary force in the trained and the untrained knee extensors and fatigue did not further enhance these adaptations. Voluntary force improvements were unrelated to CSE changes in both legs.

Contralateral effects of eccentric resistance training on immobilized arm.

This study compared the effects of contralateral eccentric-only (ECC) and concentric-/eccentric-coupled resistance training (CON-ECC) of the elbow flexors on immobilized arm. Thirty healthy participants (18-34 y) were randomly allocated to immobilization only (CTRL; n = 10), immobilization and ECC (n = 10), or immobilization and CON-ECC group (n = 10). The non-dominant arms of all participants were immobilized (8 h·day-1 ) for 4 weeks, during which ECC and CON-ECC were performed by the dominant (non-immobilized) arm 3 times a week (3-6 sets of 10 repetitions per session) with an 80%-120% and 60%-90% of one concentric repetition maximum (1-RM) load, respectively, matching the total training volume. Arm circumference, 1-RM and maximal voluntary isometric contraction (MVIC) strength, biceps brachii surface electromyogram amplitude (sEMGRMS ), rate of force development (RFD), and joint position sense (JPS) were measured for both arms before and after immobilization. CTRL showed decreases (P < .05) in MVIC (-21.7%), sEMGRMS (-35.2%), RFD (-26.0%), 1-RM (-14.4%), JPS (-87.4%), and arm circumference (-5.1%) of the immobilized arm. These deficits were attenuated or eliminated by ECC and CON-ECC, with greater effect sizes for ECC than CON-ECC in MVIC (0.29: +12.1%, vs -0.18: -0.1%) and sEMGRMS (0.31:17.5% vs -0.15: -5.9%). For the trained arm, ECC showed greater effect size for MVIC than CON-ECC (0.47 vs 0.29), and increased arm circumference (+2.9%), sEMGRMS (+77.9%), and RDF (+31.8%) greater (P < .05) than CON-ECC (+0.6%, +15.1%, and + 15.8%, respectively). The eccentric-only resistance training of the contralateral arm was more effective to counteract the negative immobilization effects than the concentric-eccentric training.

How does cross-education affects muscles of paretic upper extremity in subacute stroke survivors?

INTRODUCTION: This study aimed to evaluate the benefits of adding electromuscular stimulation (EMS) to the flexors of wrist muscles on the nonparetic limb in conventional stroke training to strengthen homologous agonist and antagonist muscles on the paretic side in patients with subacute stroke. METHODS: The EMS group patients (n = 15) received conventional therapy for 30 sessions for 6 weeks (60 min/session) with 30 min of electrical stimulation to their nonparetic forearm using wrist flexors, with 5 min of pre- and post-warm-up. The transcutaneous electrical nerve stimulation (TENS) group patients (n = 15) received the same conventional rehabilitation training with 30 min of conventional antalgic TENS at a barely sensible level to their nonparetic forearm. The Fugl-Meyer motor function assessment for upper extremity (FMA-UE), functional independence measure (FIM), Brunnstrom staging of recovery for hand, maximum and mean wrist flexion force (flexionmax and flexionmean), and wrist extension force (extensionmax and extensionmean) of paretic untrained limb were evaluated before and after the treatment. RESULTS: EMS and TENS group patients improved similarly in terms of FMA-UE, FIM, and Brunnstrom staging for hand recovery. However, flexionmax and flexionmean of the paretic limb increased more in the EMS group than in the TENS group. Extensionmax and extensionmean on the paretic side increased in the EMS group but did not differ in the TENS group. CONCLUSION: Cross-education via EMS may have a beneficial effect as an adjunct to conventional treatment methods. This study is retrospectively registered and is available at www.clinicaltrials.gov (ID: NCT04113369).

Contralateral Transfer of the Phenomenon of Repeated Bout Rate Enhancement in Unilateral Index Finger Tapping.

These hypotheses were tested: (1) Freely chosen frequency in unilateral index finger tapping is correlated between the two index fingers, and (2) A 3-min bout of unilateral index finger tapping followed by 10 min rest results in an increase of the freely chosen tapping frequency performed by the contralateral index finger in a second bout. Thirty-two adults participated. Freely chosen tapping frequencies from first bouts were 167.2 ± 79.0 and 161.5 ± 69.4 taps/min for the dominant and non-dominant hand, respectively (p=.434). These variables correlated (R=.86, p<.001). When bout one and two were performed with the dominant and non-dominant hand, respectively, the frequency increased by 8.1%±17.2% in bout two (p=.011). In opposite order, the frequency increased by 14.1%±17.5% (p<.001), which was not different from the ∼8% (p=.157).

Effects of acute and chronic unilateral resistance training variables on ipsilateral motor cortical excitability and cross-education: A systematic review.

OBJECTIVE: The increase in voluntary force of an untrained limb (i.e. Cross-education) after unilateral resistance training (RT) is believed to be a consequence of cortical adaptations. However, studies measuring neurophysiological adaptations with transcranial magnetic stimulation (TMS) found inconsistent results. One unexamined factor contributing to the conflicting data is the variation in the type and intensity of muscle contractions, fatigue, and the strategies of pacing the movement. Therefore, the purpose was to analyse how those unilateral RT variables affect the adaptations in ipsilateral M1 (iM1) and cross-education. METHODS: We performed a systematic literature review, with the following search terms with Boolean conjunctions: "Transcranial magnetic stimulation" AND "Ipsilateral cortex" AND "Resistance training". RESULTS: The 11 acute and 12 chronic studies included partially support the idea of increased cortical excitability and reduced intracortical inhibition in iM1, but the inconsistency between studies was high. CONCLUSIONS: Differences in type and intensity of contraction, fatigue, and strategies of pacing the movement contributed to the inconsistencies. The tentative conclusion is that high intensity eccentric or externally paced contractions are effective to increase iM1 excitability but cross-education can occur in the absence of such changes. Thus, the mechanism of the cross-education examined with TMS remains unclear.

Interference between competing motor memories developed through learning with different limbs.

Learning from motor errors that occur across different limbs is essential for effective tool use, sports training, and rehabilitation. To probe the neural organization of error-driven learning across limbs, we asked whether learning opposing visuomotor mappings with the two arms would interfere. Young right-handers first adapted to opposite visuomotor rotations A and B with different arms and were then reexposed to A 24 h later. We observed that relearning of A was never faster nor were initial errors smaller than prior A learning, which would be expected if there was no interference from B. Rather, errors were greater than or similar to, and learning rate was slower than or comparable to, previous A learning depending on the order in which the arms learned. This indicated robust interference between the motor memories of A and B when they were learned with different arms in close succession. We then proceeded to uncover that the order-dependent asymmetry in performance upon reexposure resulted from asymmetric transfer of learning from the left arm to the right but not vice versa and that the observed interference was retrograde in nature. Such retrograde interference likely occurs because the two arms require the same neural resources for learning, a suggestion consistent with that of our past work showing impaired learning following left inferior parietal damage regardless of the arm used. These results thus point to a common neural basis for formation of new motor memories with different limbs and hold significant implications for how newly formed motor memories interact. NEW & NOTEWORTHY In a series of experiments, we demonstrate robust retrograde interference between competing motor memories developed through error-based learning with different arms. These results provide evidence for shared neural resources for the acquisition of motor memories across different limbs and also suggest that practice with two effectors in close succession may not be a sound approach in either sports or rehabilitation. Such training may not allow newly acquired motor memories to be stabilized.

Somatosensory Electrical Stimulation Does Not Augment Motor Skill Acquisition and Intermanual Transfer in Healthy Young Adults-A Pilot Study.

Sensory input can modify motor function and magnify interlimb transfer. We examined the effects of low-intensity somatosensory electrical stimulation (SES) on motor practice-induced skill acquisition and intermanual transfer. Participants practiced a visuomotor skill for 25 min and received SES to the practice or the transfer arm. Responses to single- and double-pulse transcranial magnetic stimulation were measured in both extensor carpi radialis. SES did not further increase skill acquisition (motor practice with right hand [RMP]: 30.8% and motor practice with right hand + somatosensory electrical stimulation to the right arm [RMP + RSES]: 27.8%) and intermanual transfer (RMP: 13.6% and RMP + RSES: 9.8%) when delivered to the left arm (motor practice with right hand + somatosensory electrical stimulation to the left arm [RMP + LSES]: 44.8% and 18.6%, respectively). Furthermore, transcranial magnetic stimulation measures revealed no changes in either hand. Future studies should systematically manipulate SES parameters to better understand the mechanisms of how SES affords motor learning benefits documented but not studied in patients.

Enhancing Generalization of Visuomotor Adaptation by Inducing Use-dependent Learning.

Learning a motor task in one condition typically generalizes to another, although it is unclear why it generalizes substantially in certain situations, but only partially in other situations (e.g., across movement directions and motor effectors). Here, we demonstrate that generalization of motor learning across directions and effectors can be enhanced substantially by inducing use-dependent learning, that is, by having subjects experience motor actions associated with a desired trajectory repeatedly during reaching movements. In Experiments 1 and 2, healthy human adults adapted to a visuomotor rotation while concurrently experiencing repetitive passive movements guided by a robot. This manipulation increased the extent of generalization across movement directions (Expt. 1) and across the arms (Expt. 2) by up to 50% and 42%, respectively, indicating crucial contribution of use-dependent learning to motor generalization. In Experiment 3, we applied repetitive transcranial magnetic stimulation (rTMS) to the left primary motor cortex (M1) of the human subjects prior to passive training with the right arm to increase cortical excitability. This intervention resulted in increased motor-evoked potentials (MEPs) and decreased short-interval intracortical inhibition (SICI) in the rTMS group, but not in the sham group. These changes observed in the rTMS group were accompanied by enhanced generalization of visuomotor adaptation across the arms, which was not the case in the sham group. Collectively, these findings confirm the involvement of M1 in use-dependent learning, and suggest that use-dependent learning can contribute not only to motor learning, but also to motor generalization.

What Do TMS-Evoked Motor Potentials Tell Us About Motor Learning?

Thirty years ago, the first magnetic device capable of stimulating the human brain without discomfort through the intact skull was unveiled in Sheffield, England (Barker et al. in Lancet 1:1106-1107, 1985). Since that time, transcranial magnetic stimulation (TMS) has become the tool of choice for many scientists investigating human motor control and learning. In light of the fact that there are limits to the information that can be provided by any experimental technique, we first make the case that the necessarily restricted explanatory scope of the TMS technique-and the motor-evoked potentials to which it gives rise, is not yet reflected adequately in the research literature. We also argue that this inattention, coupled with the pervasive adoption of TMS as an investigative tool, may be restricting the elaboration of knowledge concerning the neural processes that mediate human motor learning. In order to make these points, we use as an exemplar the study of cross-education-the interlimb transfer of functional capacity.