Neural pathways mediating cross education of motor function

Cross education is the process whereby training of one limb gives rise to enhancements in the performance of the opposite, untrained limb. Despite interest in this phenomenon having been sustained for more than a century, a comprehensive explanation of the mediating neural mechanisms remains elusive. With new evidence emerging that cross education may have therapeutic utility, the need to provide a principled evidential basis upon which to design interventions becomes ever more pressing. Generally, mechanistic accounts of cross education align with one of two explanatory frameworks. Models of the 'cross activation' variety encapsulate the observation that unilateral execution of a movement task gives rise to bilateral increases in corticospinal excitability. The related conjecture is that such distributed activity, when present during unilateral practice, leads to simultaneous adaptations in neural circuits that project to the muscles of the untrained limb, thus facilitating subsequent performance of the task. Alternatively, 'bilateral access' models entail that motor engrams formed during unilateral practise, may subsequently be utilised bilaterally - that is, by the neural circuitry that constitutes the control centres for movements of both limbs. At present there is a paucity of direct evidence that allows the corresponding neural processes to be delineated, or their relative contributions in different task contexts to be ascertained. In the current review we seek to synthesise and assimilate the fragmentary information that is available, including consideration of knowledge that has emerged as a result of technological advances in structural and functional brain imaging. An emphasis upon task dependency is maintained throughout, the conviction being that the neural mechanisms that mediate cross education may only be understood in this context. © 2013 Ruddy and Carson.

Anodal tDCS Prolongs the Cross-education of Strength and Corticomotor Plasticity

To assess the efficacy of applying anodal transcranial direct current stimulation (a-tDCS) to the ipsilateral motor cortex (iM1) during unilateral strength training to enhance the neurophysiological and functional effects of cross-education.

Increased cross-education of muscle strength and reduced corticospinal inhibition following eccentric strength training

AIM: Strength training of one limb results in a substantial increase in the strength of the untrained limb, however, it remains unknown what the corticospinal responses are following either eccentric or concentric strength training and how this relates to the cross-education of strength. The aim of this study was to determine if eccentric or concentric unilateral strength training differentially modulates corticospinal excitability, inhibition and the cross-transfer of strength. METHODS: Changes in contralateral (left limb) concentric strength, eccentric strength, motor-evoked potentials, short-interval intracortical inhibition and silent period durations were analyzed in groups of young adults who exercised the right wrist flexors with either eccentric (N=9) or concentric (N=9) contractions for 12 sessions over 4weeks. Control subjects (N=9) did not train. RESULTS: Following training, both groups exhibited a significant strength gain in the trained limb (concentric group increased concentric strength by 64% and eccentric group increased eccentric strength by 62%) and the extent of the cross-transfer of strength was 28% and 47% for the concentric and eccentric group, respectively, which was different between groups (P=0.031). Transcranial magnetic stimulation revealed that eccentric training reduced intracortical inhibition (37%), silent period duration (15-27%) and increased corticospinal excitability (51%) compared to concentric training for the untrained limb (P=0.033). There was no change in the control group. CONCLUSION: The results show that eccentric training uniquely modulates corticospinal excitability and inhibition of the untrained limb to a greater extent than concentric training. These findings suggest that unilateral eccentric contractions provide a greater stimulus in cross-education paradigms and should be an integral part of the rehabilitative process following unilateral injury to maximize the response.

The effects of whole-body vibration on the cross-transfer of strength

This study investigated whether the use of superimposed whole-body vibration (WBV) during cross-education strength training would optimise strength transfer compared to conventional cross-education strength training. Twenty-one healthy, dominant right leg volunteers (21±3 years) were allocated to a strength training (ST, m = 3, f = 4), a strength training with WBV (ST + V, m = 3, f = 4), or a control group (no training, m = 3, f = 4). Training groups performed 9 sessions over 3 weeks, involving unilateral squats for the right leg, with or without WBV (35 Hz; 2.5mm amplitude). All groups underwent dynamic single leg maximum strength testing (1RM) and single and paired pulse transcranial magnetic stimulation (TMS) prior to and following training. Strength increased in the trained limb for the ST (41%; ES = 1.14) and ST + V (55%; ES = 1.03) groups, which resulted in a 35% (ES = 0.99) strength transfer to the untrained left leg for the ST group and a 52% (ES = 0.97) strength transfer to the untrained leg for the ST + V group, when compared to the control group. No differences in strength transfer between training groups were observed (P = 0.15). For the untrained leg, no differences in the peak height of recruitment curves or SICI were observed between ST and ST + V groups (P = 1.00). Strength training with WBV does not appear to modulate the cross-transfer of strength to a greater magnitude when compared to conventional cross-education strength training.

Neural adaptations associated with interlimb transfer in a ballistic wrist flexion task.

Cross education is the process whereby training of one limb gives rise to increases in the subsequent performance of its opposite counterpart. The execution of many unilateral tasks is associated with increased excitability of corticospinal projections from primary motor cortex (M1) to the opposite limb. It has been proposed that these effects are causally related. Our aim was to establish whether changes in corticospinal excitability arising from prior training of the opposite limb determine levels of interlimb transfer.

We used three vision conditions shown previously to modulate the excitability of corticospinal projections to the inactive (right) limb during wrist flexion movements performed by the training (left) limb. These were: mirrored visual feedback of the training limb; no visual feedback of either limb; and visual feedback of the inactive limb. Training comprised 300 discrete, ballistic wrist flexion movements executed as rapidly as possible. Performance of the right limb on the same task was assessed prior to, at the mid point of, and following left limb training.

There was no evidence that variations in the excitability of corticospinal projections (assessed by transcranial magnetic stimulation (TMS)) to the inactive limb were associated with, or predictive of, the extent of interlimb transfer that was expressed. There were however associations between alterations in muscle activation dynamics observed for the untrained limb, and the degree of positive transfer that arose from training of the opposite limb.

The results suggest that the acute adaptations that mediate the bilateral performance gains realised through unilateral practice of this ballistic wrist flexion task are mediated by neural elements other than those within M1 that are recruited at rest by single-pulse TMS.

CHANGES OF GENE EXPRESSION IN ELECTRICALLY STIMULATED AND CONTRALATERAL RAT SOLEUS MUSCLES

In this study we investigate the effect of a single session of high-intensity contractions on expression of pleiotropic genes and, in particular, those genes associated with metabolism in soleus muscle from electrically stimulated (ES) and contralateral (CL) limbs. The right limbs of male Wistar rats were submitted to contractions by 200-ms trains of electrical stimulation at 100-Hz frequency with pulses of 0.1 ms (voltage 24 3 V) delivered each second for 1 hour. Soleus muscles were isolated 1 hour after contraction, and gene expression was analyzed by a macroarray technique (Atlas Toxicology 1.2 Array; Clontech Laboratories). Electrical stimulation increased expression in 92 genes (16% of the genes present in the membrane). Sixty-six genes were upregulated in both ES and CL soleus muscles, and expression of 26 genes was upregulated in the ES muscle only. The most altered genes were those related to stress response and metabolism. Electrical stimulation also raised expression of transcription factors, translation and posttranslational modification of proteins, ribosomal proteins, and intracellular transducers/effectors/modulators. The results indicate that a single session of electrical stimulation upregulated expression of genes related to metabolism and oxidative stress in soleus muscle from both ES and CL limbs. These findings may indicate an association with tissue hypertrophy and metabolic adaptations induced by physical exercise training not only in the ES but also in the CL non-stimulated muscle, suggesting a cross-education phenomenon. Muscle Nerve 40: 838-846, 2009

Reduction in corticospinal inhibition in the trained and untrained limb following unilateral leg strength training

This study used transcranial magnetic stimulation to measure the corticospinal responses following 8 weeks of unilateral leg strength training. Eighteen healthy, non-strength trained participants (14 male, 4 female; 18–35 years of age) were matched for age, gender, and pre-training strength; and assigned to a training or control group. The trained group participated in unilateral horizontal leg press strength training, progressively overloaded and wave periodised, thrice per week for 8 weeks. Testing occurred prior to the intervention, at the end of 4 weeks and at the completion of training at 8 weeks. Participants were tested in both legs for one repetition maximum strength, muscle thickness, maximal electromyography (EMG) activity, and corticospinal excitability and inhibition. No changes were observed in muscle thickness in either leg. The trained leg showed an increase in strength of 21.2% (P = 0.001) and 29.0% (P = 0.007, compared to pre-testing) whilst the untrained contralateral leg showed 17.4% (P = 0.01) and 20.4% (P = 0.004, compared to pre-testing) increases in strength at 4 and 8 weeks, respectively. EMG and corticospinal excitability did not change; however, corticospinal inhibition was significantly reduced by 17.7 ms (P = 0.003) and 17.3 ms (P = 0.001) at 4 and 8 weeks, respectively, in the trained leg, and 25.1 ms (P = 0.001) and 20.8 ms (P = 0.001) at 4 and 8 weeks, respectively, in the contralateral untrained leg. This data support the theory of corticospinal adaptations underpinning cross-education gains in the lower limbs following unilateral strength training.

Corticospinal adaptations and strength maintenance in the immobilized arm following 3 weeks unilateral strength training

Cross-education strength training has being shown to retain strength and muscle thickness in the immobilized contralateral limb. Corticospinal mechanisms have been proposed to underpin this phenomenon; however, no transcranial magnetic stimulation (TMS) data has yet been presented. This study used TMS to measure corticospinal responses following 3 weeks of unilateral arm training on the contralateral, immobilize arm. Participants (n = 28) were randomly divided into either immobilized strength training (Immob + train) immobilized no training (Immob) or control. Participants in the immobilized groups had their nondominant arm rested in a sling, 15 h/day for 3 weeks. The Immob + train group completed unilateral arm curl strength training, while the Immob and control groups did not undertake training. All participants were tested for corticospinal excitability, strength, and muscle thickness of both arms. Immobilization resulted in a group x time significant reduction in strength, muscle thickness and corticospinal excitability for the untrained limb of the Immob group. Conversely, no significant change in strength, muscle thickness, or corticospinal excitability occurred in the untrained limb of the Immob + train group. These results provide the first evidence of corticospinal mechanisms, assessed by TMS, underpinning the use of unilateral strength training to retain strength and muscle thickness following immobilization of the contralateral limb.

Changes in mitochondrial function and mitochondria associated protein expression in response to 2-weeks of high intensity interval training

PURPOSE: High-intensity short-duration interval training (HIT) stimulates functional and metabolic adaptation in skeletal muscle, but the influence of HIT on mitochondrial function remains poorly studied in humans. Mitochondrial metabolism as well as mitochondrial-associated protein expression were tested in untrained participants performing HIT over a 2-week period. METHODS: Eight males performed a single-leg cycling protocol (12 × 1 min intervals at 120% peak power output, 90 s recovery, 4 days/week). Muscle biopsies (vastus lateralis) were taken pre- and post-HIT. Mitochondrial respiration in permeabilized fibers, citrate synthase (CS) activity and protein expression of peroxisome proliferator-activated receptor gamma coactivator (PGC-1α) and respiratory complex components were measured. RESULTS: HIT training improved peak power and time to fatigue. Increases in absolute oxidative phosphorylation (OXPHOS) capacities and CS activity were observed, but not in the ratio of CCO to the electron transport system (CCO/ETS), the respiratory control ratios (RCR-1 and RCR-2) or mitochondrial-associated protein expression. Specific increases in OXPHOS flux were not apparent after normalization to CS, indicating that gross changes mainly resulted from increased mitochondrial mass. CONCLUSION: Over only 2 weeks HIT significantly increased mitochondrial function in skeletal muscle independently of detectable changes in mitochondrial-associated and mitogenic protein expression.

Motor cortex excitability is not differentially modulated following skill and strength training

AIM: A single session of skill or strength training can modulate the primary motor cortex (M1), which manifests as increased corticospinal excitability (CSE) and decreased short-latency intra-cortical inhibition (SICI). We tested the hypothesis that both skill and strength training can propagate the neural mechanisms mediating cross-transfer and modulate the ipsilateral M1 (iM1). METHODS: Transcranial magnetic stimulation (TMS) measured baseline CSE and SICI in the contralateral motor cortex (cM1) and iM1. Participants completed 4 sets of unilateral training with their dominant arm, either visuomotor tracking, metronome-paced strength training (MPST), self-paced strength training (SPST) or control. Immediately post training, TMS was repeated in both M1s. RESULTS: Motor-evoked potentials (MEPs) increased and inhibition was reduced for skill and MPST training from baseline in both M1s. Self-paced strength training and control did not produce changes in CSE and SICI when compared to baseline in both M1s. After training, skill and MPST increased CSE and decreased SICI in cM1 compared to SPST and control. Skill and MPST training decreased SICI in iM1 compared to SPST and control post intervention; however, CSE in iM1 was not different across groups post training. CONCLUSION: Both skill training and MPST facilitated an increase in CSE and released SICI in iM1 and cM1 compared to baseline. Our results suggest that synchronizing to an auditory or a visual cue promotes neural adaptations within the iM1, which is thought to mediate cross transfer.

Treinamento de força unilateral: efeitos sobre a taxa de desenvolvimento de força do membro contralateral

The aim of this study was to analyze the effects of unilateral isokinetic strength training at high speed (180°.s-1) of the dominant leg on the rate of force development (RFD) of the contralateral limb. The study included 18 male individuals, apparently healthy, with an average of 23 years, and without regular practice of physical activities. The subjects were divided randomly into two groups: control group (GC) and isokinetic group (GISOC). Assessments were made of the values of peak isometric torque (PT isom) and RFD of both lower limbs in the pre-training. So, the GISOC underwent an isokinetic strength training with high speed (180°.s-1) for 6 weeks, with the dominant leg only and then reassessed. It was found that there was no significant improvement in value of PT isom (p> 0.05), and the RFD for the dominant limb had a significant improvement (p< 0.05) compared to the pre-training and that there was no transfer of values to the contralateral side (p> 0.05). It can be concluded that the isokinetic training at high speed was not enough for significant cross education

A model for engaging and evaluating innovative cross-sectoral education reform : case studies from Queensland, Australia

This paper will explore how a general education can contribute successfully to vocational outcomes using both Participatory Action Research (PAR) and Program Theory methodology. The paper will focus on the development aspects of ‘marrying’ vocational and general education including engagement processes, student, teacher, institute and employer preparation and the pathway possibilities that emerge. Successful cases presented include the: Healthy Futures program (pathways into the Health and Allied industries); Accounting Pathways program (simultaneously studying a general Accounting subject and a Certificate III vocational qualification); and Sustainable Sciences initiative (development of a vocational qualification that focuses on the emerging renewable energy industry and is linked to school science programs). The case studies have been selected because they are unique in character and application and can be used as a basis for future program development in other settings or curriculum areas.