Journal articles on the topic 'Corticospinal responses'
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Maier, Marc A., Peter A. Kirkwood, Thomas Brochier, and Roger N. Lemon. "Responses of single corticospinal neurons to intracortical stimulation of primary motor and premotor cortex in the anesthetized macaque monkey." Journal of Neurophysiology 109, no. 12 (June 15, 2013): 2982–98. http://dx.doi.org/10.1152/jn.01080.2012.
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The responses of individual primate corticospinal neurons to localized electrical stimulation of primary motor (M1) and of ventral premotor cortex (area F5) are poorly documented. To rectify this and to study interactions between responses from these areas, we recorded corticospinal axons, identified by pyramidal tract stimulation, in the cervical spinal cord of three chloralose-anesthetized macaque monkeys. Single stimuli (≤400 μA) were delivered to the hand area of M1 or F5 through intracortical microwire arrays. Only 14/112 (13%) axons showed responses to M1 stimuli that indicated direct intracortical activation of corticospinal neurons (D-responses); no D-responses were seen from F5. In contrast, 62 axons (55%) exhibited consistent later responses to M1 stimulation, corresponding to indirect activation (I-responses), showing that single-pulse intracortical stimulation of motor areas can result in trans-synaptic activation of a high proportion of the corticospinal output. A combined latency histogram of all axon responses was nonperiodic, clearly different from the periodic surface-recorded corticospinal volleys. This was readily explained by correcting for conduction velocities of individual axons. D-responding axons, taken as originating in neurons close to the M1 stimulating electrodes, showed more I-responses from M1 than those without a D-response, and 8/10 of these axons also responded to F5 stimulation. Altogether, 33% of tested axons responded to F5 stimulation, most of which also showed I-responses from M1. These excitatory effects are in keeping with facilitation of hand muscles evoked from F5 being relayed via M1. This was further demonstrated by facilitation of test responses from M1 by conditioning F5 stimuli.
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Taylor, J. L., and S. C. Gandevia. "Noninvasive stimulation of the human corticospinal tract." Journal of Applied Physiology 96, no. 4 (April 2004): 1496–503. http://dx.doi.org/10.1152/japplphysiol.01116.2003.
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Spinal tracts can be stimulated noninvasively in human subjects by passing a high-voltage stimulus between the mastoids or by magnetic stimulation over the back of the head. The stimulus probably activates the corticospinal tract at the cervicomedullary junction (pyramidal decussation) and evokes large, short-latency motor responses in the arm muscles. These responses have a large monosynaptic component. Responses in leg muscles can be elicited by cervicomedullary junction stimulation or by stimulation over the cervical or thoracic spine. Because nerve roots are more easily activated than spinal tracts, stimulus spread to motor axons can occur. Facilitation of responses by voluntary activity confirms that the responses are evoked synaptically. Stimulation of the corticospinal tract is useful in studies of central conduction and studies of the behavior of motoneurons during different tasks. It also provides an important comparison to allow interpretation of changes in responses to stimulation of the motor cortex. The major drawback to the use of electrical stimulation of the corticospinal tract is that each stimulus is transiently painful.
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Mason, Joel, Ashlyn K. Frazer, Janne Avela, Alan J. Pearce, Glyn Howatson, and Dawson J. Kidgell. "Tracking the corticospinal responses to strength training." European Journal of Applied Physiology 120, no. 4 (February 14, 2020): 783–98. http://dx.doi.org/10.1007/s00421-020-04316-6.
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Boyne, Pierce, Colleen Meyrose, Jennifer Westover, Dustyn Whitesel, Kristal Hatter, Darcy S. Reisman, David Cunningham, et al. "Exercise intensity affects acute neurotrophic and neurophysiological responses poststroke." Journal of Applied Physiology 126, no. 2 (February 1, 2019): 431–43. http://dx.doi.org/10.1152/japplphysiol.00594.2018.
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Aerobic exercise may acutely prime the brain to be more responsive to rehabilitation, thus facilitating neurologic recovery from conditions like stroke. This aerobic priming effect could occur through multiple mechanisms, including upregulation of circulating brain-derived neurotrophic factor (BDNF), increased corticospinal excitability, and decreased intracortical inhibition. However, optimal exercise parameters for targeting these mechanisms are poorly understood. This study tested the effects of exercise intensity on acute BDNF and neurophysiological responses. Sixteen ambulatory persons >6 mo poststroke performed three different 20-min exercise protocols in random order, approximately 1 wk apart, including the following: 1) treadmill high-intensity interval training (HIT-treadmill); 2) seated-stepper HIT (HIT-stepper); and 3) treadmill moderate-intensity continuous exercise (MCT-treadmill). Serum BDNF and transcranial magnetic stimulation measures of paretic lower limb excitability and inhibition were assessed at multiple time points during each session. Compared with MCT-treadmill, HIT-treadmill elicited significantly greater acute increases in circulating BDNF and corticospinal excitability. HIT-stepper initially showed BDNF responses similar to HIT-treadmill but was no longer significantly different from MCT-treadmill after decreasing the intensity in reaction to two hypotensive events. Additional regression analyses showed that an intensity sufficient to accumulate blood lactate appeared to be important for eliciting BDNF responses, that the interval training approach may have facilitated the corticospinal excitability increases, and that the circulating BDNF response was (negatively) related to intracortical inhibition. These findings further elucidate neurologic mechanisms of aerobic exercise and inform selection of optimal exercise-dosing parameters for enhancing acute neurologic effects. NEW & NOTEWORTHY Acute exercise-related increases in circulating BDNF and corticospinal excitability are thought to prime the brain for learning. Our data suggest that these responses can be obtained among persons with stroke using short-interval treadmill high-intensity interval training, that a vigorous aerobic intensity sufficient to generate lactate accumulation is needed to increase BDNF, that interval training facilitates increases in paretic quadriceps corticospinal excitability, and that greater BDNF response is associated with lesser intracortical inhibition response.
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Mason, Joel, Ashlyn K. Frazer, Alan J. Pearce, Alicia M. Goodwill, Glyn Howatson, Shapour Jaberzadeh, and Dawson J. Kidgell. "Determining the early corticospinal-motoneuronal responses to strength training: a systematic review and meta-analysis." Reviews in the Neurosciences 30, no. 5 (July 26, 2019): 463–76. http://dx.doi.org/10.1515/revneuro-2018-0054.
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Abstract Several studies have used transcranial magnetic stimulation to probe the corticospinal-motoneuronal responses to a single session of strength training; however, the findings are inconsistent. This systematic review and meta-analysis examined whether a single bout of strength training affects the excitability and inhibition of intracortical circuits of the primary motor cortex (M1) and the corticospinal-motoneuronal pathway. A systematic review was completed, tracking studies between January 1990 and May 2018. The methodological quality of studies was determined using the Downs and Black quality index. Data were synthesised and interpreted from meta-analysis. Nine studies (n=107) investigating the acute corticospinal-motoneuronal responses to strength training met the inclusion criteria. Meta-analyses detected that after strength training compared to control, corticospinal excitability [standardised mean difference (SMD), 1.26; 95% confidence interval (CI), 0.88, 1.63; p<0.0001] and intracortical facilitation (ICF) (SMD, 1.60; 95% CI, 0.18, 3.02; p=0.003) were increased. The duration of the corticospinal silent period was reduced (SMD, −17.57; 95% CI, −21.12, −14.01; p=0.00001), but strength training had no effect on the excitability of the intracortical inhibitory circuits [short-interval intracortical inhibition (SICI) SMD, 1.01; 95% CI, −1.67, 3.69; p=0.46; long-interval intracortical inhibition (LICI) SMD, 0.50; 95% CI, −1.13, 2.13; p=0.55]. Strength training increased the excitability of corticospinal axons (SMD, 4.47; 95% CI, 3.45, 5.49; p<0.0001). This systematic review and meta-analyses revealed that the acute neural changes to strength training involve subtle changes along the entire neuroaxis from the M1 to the spinal cord. These findings suggest that strength training is a clinically useful tool to modulate intracortical circuits involved in motor control.
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Cherni, Yosra, Alexia Tremblay, Margaux Simon, Floriane Bretheau, Andréanne K. Blanchette, and Catherine Mercier. "Corticospinal Responses Following Gait-Specific Training in Stroke Survivors: A Systematic Review." International Journal of Environmental Research and Public Health 19, no. 23 (November 24, 2022): 15585. http://dx.doi.org/10.3390/ijerph192315585.
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Corticospinal excitability is subject to alterations after stroke. While the reversal of these alterations has been proposed as an underlying mechanism for improved walking capacity after gait-specific training, this has not yet been clearly demonstrated. Therefore, the objective of this review is to evaluate the effect of gait-specific training on corticospinal excitability in stroke survivors. We conducted an electronic database search in four databases (i.e., Medline, Embase, CINAHL and Web of Science) in June 2022. Two authors screened in an independent way all the studies and selected those that investigated the effect of gait-specific training on variables such as motor-evoked potential amplitude, motor threshold, map size, latency, and corticospinal silent period in stroke survivors. Nineteen studies investigating the effect of gait-specific training on corticospinal excitability were included. Some studies showed an increased MEP amplitude (7/16 studies), a decreased latency (5/7studies), a decreased motor threshold (4/8 studies), an increased map size (2/3 studies) and a decreased cortical silent period (1/2 study) after gait-specific training. No change has been reported in terms of short interval intracortical inhibition after training. Five studies did not report any significant effect after gait-specific training on corticospinal excitability. The results of this systematic review suggest that gait-specific training modalities can drive neuroplastic adaptation among stroke survivors. However, given the methodological disparity of the included studies, additional clinical trials of better methodological quality are needed to establish conclusions. The results of this review can therefore be used to develop future studies to better understand the effects of gait-specific training on the central nervous system.
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Forman, Davis A., Garrick N. Forman, Bernadette A. Murphy, and Michael W. R. Holmes. "Sustained Isometric Wrist Flexion and Extension Maximal Voluntary Contractions on Corticospinal Excitability to Forearm Muscles during Low-Intensity Hand-Gripping." Brain Sciences 10, no. 7 (July 13, 2020): 445. http://dx.doi.org/10.3390/brainsci10070445.
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The wrist extensors demonstrate an earlier fatigue onset than the wrist flexors. However, it is currently unclear whether fatigue induces unique changes in muscle activity or corticospinal excitability between these muscle groups. The purpose of this study was to examine how sustained isometric wrist extension/flexion maximal voluntary contractions (MVCs) influence muscle activity and corticospinal excitability of the forearm. Corticospinal excitability to three wrist flexors and three wrist extensors were measured using motor evoked potentials (MEPs) elicited via transcranial magnetic stimulation. Responses were elicited while participants exerted 10% of their maximal handgrip force, before and after a sustained wrist flexion or extension MVC (performed on separate sessions). Post-fatigue measures were collected up to 10-min post-fatigue. Immediately post-fatigue, extensor muscle activity was significantly greater following the wrist flexion fatigue session, although corticospinal excitability (normalized to muscle activity) was greater on the wrist extension day. Responses were largely unchanged in the wrist flexors. However, for the flexor carpi ulnaris, normalized MEP amplitudes were significantly larger following wrist extension fatigue. These findings demonstrate that sustained isometric flexion/extension MVCs result in a complex reorganization of forearm muscle recruitment strategies during hand-gripping. Based on these findings, previously observed corticospinal behaviour following fatigue may not apply when the fatiguing task and measurement task are different.
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Škarabot, Jakob, Paul Ansdell, Callum G. Brownstein, Kirsty M. Hicks, Glyn Howatson, Stuart Goodall, and Rade Durbaba. "Reduced corticospinal responses in older compared with younger adults during submaximal isometric, shortening, and lengthening contractions." Journal of Applied Physiology 126, no. 4 (April 1, 2019): 1015–31. http://dx.doi.org/10.1152/japplphysiol.00987.2018.
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The aim of this study was to assess differences in motor performance, as well as corticospinal and spinal responses to transcranial magnetic and percutaneous nerve stimulation, respectively, during submaximal isometric, shortening, and lengthening contractions between younger and older adults. Fifteen younger [26 yr (SD 4); 7 women, 8 men] and 14 older [64 yr (SD 3); 5 women, 9 men] adults performed isometric and shortening and lengthening dorsiflexion on an isokinetic dynamometer (5°/s) at 25% and 50% of contraction type-specific maximums. Motor evoked potentials (MEPs) and H reflexes were recorded at anatomical zero. Maximal dorsiflexor torque was greater during lengthening compared with shortening and isometric contractions ( P < 0.001) but was not age dependent ( P = 0.158). However, torque variability was greater in older compared with young adults ( P < 0.001). Background electromyographic (EMG) activity was greater in older compared with younger adults ( P < 0.005) and was contraction type dependent ( P < 0.001). As evoked responses are influenced by both the maximal level of excitation and background EMG activity, the responses were additionally normalized {[MEP/maximum M wave (Mmax)]/root-mean-square EMG activity (RMS) and [H reflex (H)/Mmax]/RMS}. (MEP/Mmax)/RMS and (H/Mmax)/RMS were similar across contraction types but were greater in young compared with older adults ( P < 0.001). Peripheral motor conduction times were prolonged in older adults ( P = 0.003), whereas peripheral sensory conduction times and central motor conduction times were not age dependent ( P ≥ 0.356). These data suggest that age-related changes throughout the central nervous system serve to accommodate contraction type-specific motor control. Moreover, a reduction in corticospinal responses and increased torque variability seem to occur without a significant reduction in maximal torque-producing capacity during older age. NEW & NOTEWORTHY This is the first study to have explored corticospinal and spinal responses with aging during submaximal contractions of different types (isometric, shortening, and lengthening) in lower limb musculature. It is demonstrated that despite preserved maximal torque production capacity corticospinal responses are reduced in older compared with younger adults across contraction types along with increased torque variability during dynamic contractions. This suggests that the age-related corticospinal changes serve to accommodate contraction type-specific motor control.
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Krishnan, Chandramouli, Edward P. Washabaugh, Aviroop Dutt-Mazumder, Scott R. Brown, Edward M. Wojtys, and Riann M. Palmieri-Smith. "Conditioning Brain Responses to Improve Quadriceps Function in an Individual With Anterior Cruciate Ligament Reconstruction." Sports Health: A Multidisciplinary Approach 11, no. 4 (April 5, 2019): 306–15. http://dx.doi.org/10.1177/1941738119835163.
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Background: Persistent quadriceps weakness and activation failure are common in individuals with anterior cruciate ligament (ACL) reconstruction. A growing body of evidence indicates that this chronic quadriceps dysfunction could be partly mediated due to reduced corticospinal excitability. However, current rehabilitation approaches do not directly target corticospinal deficits, which may be critical for restoring optimal clinical outcomes after the surgery. This case study tested the feasibility of operant conditioning of torque responses evoked by transcranial magnetic stimulation (TMS) to improve quadriceps function after ACL reconstruction. Hypothesis: Operant conditioning of motor evoked torque responses would improve quadriceps strength, voluntary activation, and corticospinal excitability. Study Design: Case study and research report. Level of Evidence: Level 5. Methods: A 24-year-old male with an ACL reconstruction (6 months postsurgery) trained for 20 sessions (2-3 times per week for 8 weeks) to increase his TMS-induced motor evoked torque response (MEP torque) of the quadriceps muscles using operant conditioning principles. Knee extensor strength, voluntary quadriceps muscle activation, and quadriceps corticospinal excitability were evaluated at 3 time points: preintervention (pre), 4 weeks (mid), and immediately after the intervention (post). Results: The participant was able to successfully condition (ie, increase) the quadriceps MEP torque after 1 training session, and the conditioned MEP torque gradually increased over the course of 20 training sessions to reach about 500% of the initial value at the end of training. The participant’s control MEP torque values and corticospinal excitability, which were measured outside of the conditioning paradigm, also increased with training. These changes were paralleled by improvements in knee extensor strength and voluntary quadriceps muscle activation. Conclusion: This study shows that operant conditioning of MEP torque is a feasible approach to improving quadriceps corticospinal excitability and quadriceps function after ACL reconstruction and encourages further testing in a larger cohort of ACL-reconstructed individuals. Clinical Relevance: Operant conditioning may serve as a potential therapeutic adjuvant for ACL rehabilitation.
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Sidhu, Simranjit K., Ben W. Hoffman, Andrew G. Cresswell, and Timothy J. Carroll. "Corticospinal contributions to lower limb muscle activity during cycling in humans." Journal of Neurophysiology 107, no. 1 (January 2012): 306–14. http://dx.doi.org/10.1152/jn.00212.2011.
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The purpose of the current study was to investigate corticospinal contributions to locomotor drive to leg muscles involved in cycling. We studied 1) if activation of inhibitory interneurons in the cortex via subthreshold transcranial magnetic stimulation (TMS) caused a suppression of EMG and 2) how the responses to stimulation of the motor cortex via TMS and cervicomedullary stimulation (CMS) were modulated across the locomotor cycle. TMS at intensities subthreshold for activation of the corticospinal tract elicited suppression of EMG for approximately one-half of the subjects and muscles during cycling, and in matched static contractions in vastus lateralis. There was also significant modulation in the size of motor-evoked potentials (MEPs) elicited by TMS across the locomotor cycle ( P < 0.001) that was strongly related to variation in background EMG in all muscles ( r > 0.86; P < 0.05). When MEP and CMEP amplitudes were normalized to background EMG, they were relatively larger prior to the main EMG burst and smaller when background EMG was maximum. Since the pattern of modulation of normalized MEP and CMEP responses was similar, the data suggest that phase-dependent modulation of corticospinal responses during cycling in humans is driven mainly by spinal mechanisms. However, there were subtle differences in the degree to which normalized MEP and CMEP responses were facilitated prior to EMG burst, which might reflect small increases in cortical excitability prior to maximum muscle activation. The data demonstrate that the motor cortex contributes actively to locomotor drive, and that spinal factors dominate phase-dependent modulation of corticospinal excitability during cycling in humans.
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Martin, P. G., J. E. Butler, S. C. Gandevia, and J. L. Taylor. "Noninvasive Stimulation of Human Corticospinal Axons Innervating Leg Muscles." Journal of Neurophysiology 100, no. 2 (August 2008): 1080–86. http://dx.doi.org/10.1152/jn.90380.2008.
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These studies investigated whether a single electrical stimulus over the thoracic spine activates corticospinal axons projecting to human leg muscles. Transcranial magnetic stimulation of the motor cortex and electrical stimulation over the thoracic spine were paired at seven interstimulus intervals, and surface electromyographic responses were recorded from rectus femoris, tibialis anterior, and soleus. The interstimulus intervals (ISIs) were set so that the first descending volley evoked by cortical stimulation had not arrived at (positive ISIs), was at the same level as (0 ISI) or had passed (negative ISIs) the site of activation of descending axons by the thoracic stimulation at the moment of its delivery. Compared with the responses to motor cortical stimulation alone, responses to paired stimuli were larger at negative ISIs but reduced at positive ISIs in all three leg muscles. This depression of responses at positive ISIs is consistent with an occlusive interaction in which an antidromic volley evoked by the thoracic stimulation collides with descending volleys evoked by cortical stimulation. The cortical and spinal stimuli activate some of the same corticospinal axons. Thus it is possible to examine the excitability of lower limb motoneuron pools to corticospinal inputs without the confounding effects of changes occurring within the motor cortex.
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Maslovat, Dana, Faven Teku, Victoria Smith, Neil M. Drummond, and Anthony N. Carlsen. "Bimanual but not unimanual finger movements are triggered by a startling acoustic stimulus: evidence for increased reticulospinal drive for bimanual responses." Journal of Neurophysiology 124, no. 6 (December 1, 2020): 1832–38. http://dx.doi.org/10.1152/jn.00309.2020.
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The relative contributions of reticulospinal and corticospinal pathways to movement initiation are relatively unknown but appear to depend on the involved musculature. Here, we show that unimanual finger movements, which are predominantly initiated via corticospinal pathways, are not triggered at short latency by a startling acoustic stimulus (SAS), while bimanual finger movements are triggered by the SAS. This distinction is attributed to increased reticulospinal drive for bilateral responses.
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Taube, Wolfgang, Martin Schubert, Markus Gruber, Sandra Beck, Michael Faist, and Albert Gollhofer. "Direct corticospinal pathways contribute to neuromuscular control of perturbed stance." Journal of Applied Physiology 101, no. 2 (August 2006): 420–29. http://dx.doi.org/10.1152/japplphysiol.01447.2005.
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The antigravity soleus muscle (Sol) is crucial for compensation of stance perturbation. A corticospinal contribution to the compensatory response of the Sol is under debate. The present study assessed spinal, corticospinal, and cortical excitability at the peaks of short- (SLR), medium- (MLR), and long-latency responses (LLR) after posterior translation of the feet. Transcranial magnetic stimulation (TMS) and peripheral nerve stimulation were individually adjusted so that the peaks of either motor evoked potential (MEP) or H reflex coincided with peaks of SLR, MLR, and LLR, respectively. The influence of specific, presumably direct, corticospinal pathways was investigated by H-reflex conditioning. When TMS was triggered so that the MEP arrived in the Sol at the same time as the peaks of SLR and MLR, EMG remained unaffected. Enhanced EMG was observed when the MEP coincided with the LLR peak ( P < 0.001). Similarly, conditioning of the H reflex by subthreshold TMS facilitated H reflexes only at LLR ( P < 0.001). The earliest facilitation after perturbation occurred after 86 ms. The TMS-induced H-reflex facilitation at LLR suggests that increased cortical excitability contributes to the augmentation of the LLR peaks. This provides evidence that the LLR in the Sol muscle is at least partly transcortical, involving direct corticospinal pathways. Additionally, these results demonstrate that ∼86 ms after perturbation, postural compensatory responses are cortically mediated.
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Dongés, Siobhan C., Jessica M. D’Amico, Jane E. Butler, and Janet L. Taylor. "Involvement of N-methyl-d-aspartate receptors in plasticity induced by paired corticospinal-motoneuronal stimulation in humans." Journal of Neurophysiology 119, no. 2 (February 1, 2018): 652–61. http://dx.doi.org/10.1152/jn.00457.2017.
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Plasticity can be induced at human corticospinal-motoneuronal synapses by delivery of repeated, paired stimuli to corticospinal axons and motoneurons in a technique called paired corticospinal-motoneuronal stimulation (PCMS). To date, the mechanisms of the induced plasticity are unknown. To determine whether PCMS-induced plasticity is dependent on N-methyl-d-aspartate receptors (NMDARs), the effect of the noncompetitive NMDAR antagonist dextromethorphan on PCMS-induced facilitation was assessed in a 2-day, double-blind, placebo-controlled experiment. PCMS consisted of 100 pairs of stimuli, delivered at an interstimulus interval that produces facilitation at corticospinal-motoneuronal synapses that excite biceps brachii motoneurons. Transcranial magnetic stimulation elicited corticospinal volleys, which were timed to arrive at corticospinal-motoneuronal synapses just before antidromic potentials elicited in motoneurons with electrical brachial plexus stimulation. To measure changes in the corticospinal pathway at a spinal level, biceps responses to cervicomedullary stimulation (cervicomedullary motor evoked potentials, CMEPs) were measured before and for 30 min after PCMS. Individuals who displayed a ≥10% increase in CMEP size after PCMS on screening were eligible to take part in the 2-day experiment. After PCMS, there was a significant difference in CMEP area between placebo and dextromethorphan days ( P = 0.014). On the placebo day PCMS increased average CMEP areas to 127 ± 46% of baseline, whereas on the dextromethorphan day CMEP area was decreased to 86 ± 33% of baseline (mean ± SD; placebo: n = 11, dextromethorphan: n = 10). Therefore, dextromethorphan suppressed the facilitation of CMEPs after PCMS. This indicates that plasticity induced at synapses in the human spinal cord by PCMS may be dependent on NMDARs. NEW & NOTEWORTHY Paired corticospinal-motoneuronal stimulation can strengthen the synaptic connections between corticospinal axons and motoneurons at a spinal level in humans. The mechanism of the induced plasticity is unknown. In our 2-day, double-blind, placebo-controlled study we show that the N-methyl-d-aspartate receptor (NMDAR) antagonist dextromethorphan suppressed plasticity induced by paired corticospinal-motoneuronal stimulation, suggesting that an NMDAR-dependent mechanism is involved.
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Giesebrecht, Sabine, Peter G. Martin, Simon C. Gandevia, and Janet L. Taylor. "Facilitation and Inhibition of Tibialis Anterior Responses to Corticospinal Stimulation After Maximal Voluntary Contractions." Journal of Neurophysiology 103, no. 3 (March 2010): 1350–56. http://dx.doi.org/10.1152/jn.00879.2009.
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The corticospinal pathway is the major pathway controlling human voluntary movements. After strong voluntary contractions, the efficacy of corticospinal transmission to elbow flexors is reduced for ∼90 s, and this limits motoneuronal output. This reduction may reflect activity-dependent changes at cortico-motoneuronal synapses. We investigated whether similar changes occur in a leg muscle, tibialis anterior (TA). Electrical stimuli over high thoracic vertebrae activated corticospinal axons to evoke an EMG response in TA (TMEP). Stimuli were delivered before and after short 10-s and prolonged 1-min maximal contractions (MVCs) of ankle dorsiflexors. In two studies, stimuli were given with the muscle relaxed. In other studies, stimuli were given during weak contraction. After a 10-s MVC ( study 1, n = 10), TMEPs increased immediately to 349 ± 335% (mean ± SD) of control values. By 1 min after contraction, TMEPs decreased to 38 ± 28% of control and remained depressed for >10 min. Facilitation (191 ± 133% control) and depression (18 ± 22% control) occurred over the same time course after the 1-min MVC ( study 2, n = 10). When tested during weak contraction ( study 3, n = 10), TMEPs showed less facilitation (131 ± 41% control) and less depression (67 ± 21% control) and responses returned to baseline over ∼15 min. In contrast to TMEPs, H-reflexes in TA were little changed after a 10-s MVC ( study 4, n = 7). Our findings reveal an immediate facilitation and subsequent longer-lasting depression in corticospinal transmission to TA, which originate at a premotoneuronal site. This behavior differs markedly from that in elbow flexor muscles and suggests that activity-dependent changes in the motor pathway may be muscle specific.
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Smith, Victoria, Dana Maslovat, Neil M. Drummond, Joëlle Hajj, Alexandra Leguerrier, and Anthony N. Carlsen. "High-intensity transcranial magnetic stimulation reveals differential cortical contributions to prepared responses." Journal of Neurophysiology 121, no. 5 (May 1, 2019): 1809–21. http://dx.doi.org/10.1152/jn.00510.2018.
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Corticospinal output pathways have typically been considered to be the primary driver for voluntary movements of the hand/forearm; however, more recently, reticulospinal drive has also been implicated in the production of these movements. Although both pathways may play a role, the reticulospinal tract is thought to have stronger connections to flexor muscles than to extensors. Similarly, movements involuntarily triggered via a startling acoustic stimulus (SAS) are believed to receive greater reticular input than voluntary movements. To investigate a differential role of reticulospinal drive depending on movement type or acoustic stimulus, corticospinal drive was transiently interrupted using high-intensity transcranial magnetic stimulation (TMS) applied during the reaction time (RT) interval. This TMS-induced suppression of cortical drive leads to RT delays that can be used to assess cortical contributions to movement. Participants completed targeted flexion and extension movements of the wrist in a simple RT paradigm in response to a control auditory go signal or SAS. Occasionally, suprathreshold TMS was applied over the motor cortical representation for the prime mover. Results revealed that TMS significantly increased RT in all conditions. There was a significantly longer TMS-induced RT delay seen in extension movements than in flexion movements and a greater RT delay in movements initiated in response to control stimuli compared with SAS. These results suggest that the contribution of reticulospinal drive is larger for wrist flexion than for extension. Similarly, movements triggered involuntarily by an SAS appear to involve greater reticulospinal drive, and relatively less corticospinal drive, than those that are voluntarily initiated.NEW & NOTEWORTHY Through the use of the transcranial magnetic stimulation-induced silent period, we provide novel evidence for a greater contribution of reticulospinal drive, and a relative decrease in corticospinal drive, to movements involuntarily triggered by a startle compared with voluntary movements. These results also provide support for the notion that both cortical and reticular structures are involved in the neural pathway underlying startle-triggered movements. Furthermore, our results indicate greater reticulospinal contribution to wrist flexion than extension movements.
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Rahman, Simin, Ummutal Siddique, Ashlyn Frazer, Alan Pearce, and Dawson Kidgell. "tDCS Anodal tDCS increases bilateral corticospinal excitability irrespective of hemispheric dominance." Journal of Science and Medicine 2, no. 2 (June 3, 2020): 1–17. http://dx.doi.org/10.37714/josam.v2i2.40.
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Background: Transcranial direct current stimulation (tDCS) is a non-invasive brain stimulation technique that utilizes weak direct currents to induce polarity-dependent modulation of corticospinal excitability. Although tDCS exerts a modulatory effect over the stimulation region, several studies have also demonstrated that distal areas of the brain connected to the region of stimulation may also be affected, as well as the contralateral hemisphere. Objective: We examined the effect of a single session of anodal tDCS on corticospinal excitability and inhibition of both the stimulated and non-stimulated hemisphere and examined the influence of these responses by the brain-derived neurotrophic factor (BDNF) polymorphism. Methods: In a randomized cross-over design, changes in corticospinal excitability and inhibition of the stimulated and non-stimulated hemispheres were analysed in 13 participants in both the dominant and non-dominant primary motor cortex (M1). Participants were exposed to 20 min of anodal and sham tDCS and also undertook a blood sample for BDNF genotyping. Results: TMS revealed a bilateral increase in corticospinal excitability irrespective of which hemisphere (dominant vs non-dominant) was stimulated (all P < 0.05). Furthermore, the induction of corticospinal excitability was influenced by the BDNF polymorphism. Conclusion: This finding shows that anodal tDCS induces bilateral effects in corticospinal excitability irrespective of hemispheric dominance. This finding provides scientists and medical practitioners with a greater understanding as to how this technique may be used as a therapeutic tool for clinical populations.
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Carroll, Timothy J., Michael Lee, Marlene Hsu, and Janel Sayde. "Unilateral practice of a ballistic movement causes bilateral increases in performance and corticospinal excitability." Journal of Applied Physiology 104, no. 6 (June 2008): 1656–64. http://dx.doi.org/10.1152/japplphysiol.01351.2007.
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It has long been known that practicing a task with one limb can result in performance improvements with the opposite, untrained limb. Hypotheses to account for cross-limb transfer of performance state that the effect is mediated either by neural adaptations in higher order control centers that are accessible to both limbs, or that there is a “spillover” of neural drive to the opposite hemisphere that results in bilateral adaptation. Here we address these hypotheses by assessing performance and corticospinal excitability in both hands after unilateral practice of a ballistic finger movement. Participants ( n = 9) completed 300 practice trials of a ballistic task with the right hand, the aim of which was to maximize the peak abduction acceleration of the index finger. Practice caused a 140% improvement in right-hand performance and an 82% improvement for the untrained left hand. There were bilateral increases in the amplitude of responses to transcranial magnetic stimulation, but increased corticospinal excitability was not correlated with improved performance. There were no significant changes in corticospinal excitability or task performance for a control group that did not train ( n = 9), indicating that performance testing for the left hand alone did not induce performance or corticospinal effects. Although the data do not provide conclusive evidence whether increased corticospinal excitability in the untrained hand is causally related to the cross-transfer of ballistic performance, the finding that ballistic practice can induce bilateral corticospinal adaptations may have important clinical implications for movement rehabilitation.
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Thompson, Aiko K., Rachel H. Cote, Janice M. Sniffen, and Jodi A. Brangaccio. "Operant conditioning of the tibialis anterior motor evoked potential in people with and without chronic incomplete spinal cord injury." Journal of Neurophysiology 120, no. 6 (December 1, 2018): 2745–60. http://dx.doi.org/10.1152/jn.00362.2018.
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The activity of corticospinal pathways is important in movement control, and its plasticity is essential for motor skill learning and re-learning after central nervous system (CNS) injuries. Therefore, enhancing the corticospinal function may improve motor function recovery after CNS injuries. Operant conditioning of stimulus-induced muscle responses (e.g., reflexes) is known to induce the targeted plasticity in a targeted pathway. Thus, an operant conditioning protocol to target the corticospinal pathways may be able to enhance the corticospinal function. To test this possibility, we investigated whether operant conditioning of the tibialis anterior (TA) motor evoked potential (MEP) to transcranial magnetic stimulation can enhance corticospinal excitability in people with and without chronic incomplete spinal cord injury (SCI). The protocol consisted of 6 baseline and 24 up-conditioning/control sessions over 10 wk. In all sessions, TA MEPs were elicited at 10% above active MEP threshold while the sitting participant provided a fixed preset level of TA background electromyographic activity. During baseline sessions, MEPs were simply measured. During conditioning trials of the conditioning sessions, the participant was encouraged to increase MEP and was given immediate feedback indicating whether MEP size was above a criterion. In 5/8 participants without SCI and 9/10 with SCI, over 24 up-conditioning sessions, MEP size increased significantly to ~150% of the baseline value, whereas the silent period (SP) duration decreased by ~20%. In a control group of participants without SCI, neither MEP nor SP changed. These results indicate that MEP up-conditioning can facilitate corticospinal excitation, which is essential for enhancing motor function recovery after SCI. NEW & NOTEWORTHY We investigated whether operant conditioning of the motor evoked potential (MEP) to transcranial magnetic stimulation can systematically increase corticospinal excitability for the ankle dorsiflexor tibialis anterior (TA) in people with and without chronic incomplete spinal cord injury. We found that up-conditioning can increase the TA MEP while reducing the accompanying silent period (SP) duration. These findings suggest that MEP up-conditioning produces the facilitation of corticospinal excitation as targeted, whereas it suppresses inhibitory mechanisms reflected in SP.
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Carroll, Timothy J., Stephan Riek, and Richard G. Carson. "Corticospinal Responses to Motor Training Revealed by Transcranial Magnetic Stimulation." Exercise and Sport Sciences Reviews 29, no. 2 (April 2001): 54–59. http://dx.doi.org/10.1249/00003677-200104000-00003.
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NUZZO, JAMES L., BENJAMIN K. BARRY, SIMON C. GANDEVIA, and JANET L. TAYLOR. "Acute Strength Training Increases Responses to Stimulation of Corticospinal Axons." Medicine & Science in Sports & Exercise 48, no. 1 (January 2016): 139–50. http://dx.doi.org/10.1249/mss.0000000000000733.
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Carroll, Timothy J., Stephan Riek, and Richard G. Carson. "Corticospinal Responses to Motor Training Revealed by Transcranial Magnetic Stimulation." Exercise and Sport Sciences Reviews 29, no. 2 (April 2001): 54–59. http://dx.doi.org/10.1097/00003677-200104000-00003.
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Roeder, Luisa, Tjeerd W. Boonstra, Simon S. Smith, and Graham K. Kerr. "Dynamics of corticospinal motor control during overground and treadmill walking in humans." Journal of Neurophysiology 120, no. 3 (September 1, 2018): 1017–31. http://dx.doi.org/10.1152/jn.00613.2017.
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Increasing evidence suggests cortical involvement in the control of human gait. However, the nature of corticospinal interactions remains poorly understood. We performed time-frequency analysis of electrophysiological activity acquired during treadmill and overground walking in 22 healthy, young adults. Participants walked at their preferred speed (4.2, SD 0.4 km/h), which was matched across both gait conditions. Event-related power, corticomuscular coherence (CMC), and intertrial coherence (ITC) were assessed for EEG from bilateral sensorimotor cortices and EMG from the bilateral tibialis anterior (TA) muscles. Cortical power, CMC, and ITC at theta, alpha, beta, and gamma frequencies (4–45 Hz) increased during the double support phase of the gait cycle for both overground and treadmill walking. High beta (21–30 Hz) CMC and ITC of EMG was significantly increased during overground compared with treadmill walking, as well as EEG power in theta band (4–7 Hz). The phase spectra revealed positive time lags at alpha, beta, and gamma frequencies, indicating that the EEG response preceded the EMG response. The parallel increases in power, CMC, and ITC during double support suggest evoked responses at spinal and cortical populations rather than a modulation of ongoing corticospinal oscillatory interactions. The evoked responses are not consistent with the idea of synchronization of ongoing corticospinal oscillations but instead suggest coordinated cortical and spinal inputs during the double support phase. Frequency-band dependent differences in power, CMC, and ITC between overground and treadmill walking suggest differing neural control for the two gait modalities, emphasizing the task-dependent nature of neural processes during human walking. NEW & NOTEWORTHY We investigated cortical and spinal activity during overground and treadmill walking in healthy adults. Parallel increases in power, corticomuscular coherence, and intertrial coherence during double support suggest evoked responses at spinal and cortical populations rather than a modulation of ongoing corticospinal oscillatory interactions. These findings identify neurophysiological mechanisms that are important for understanding cortical control of human gait in health and disease.
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Painter, Rhys, Alan Pearce, Mohamad Rostami, Ashlyn Frazer, and Dawson Kidgell. "Determining the Corticospinal and Neuromuscular Responses Following a Warm-Up Protocol." Journal of Science and Medicine 2, no. 2 (June 10, 2020): 1–12. http://dx.doi.org/10.37714/josam.v2i2.45.
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Background: The effect of warming-up prior to exercise on increased neuromuscular transmission speed remains largely untested. Objective: This study used transcranial magnetic stimulation (TMS) and peripheral nerve stimulation (PNS) to quantify neuromuscular transmission along the corticospinal tract (CST) before and after a warm-up protocol of the elbow flexors. Method: Using a single-group, pre-test-post-test design, 30 participants (20 male; 10 female; mean age 26.3 ± 7.4 years) completed four sets of bicep curls that aimed to increase heart rate (HR) and biceps brachii (BB) muscle temperature by a minimum of 40 beats per minute (bpm) and 1°C, respectively. Single-pulse TMS was applied to the primary motor cortex, and over the cervical and thoracic (C7-T1) areas of the spine to quantify motor evoked potentials (MEPs) and spinal evoked potentials (SEPs), respectively. Central motor conduction time (CMCT) was determined by calculating the difference in latency time of the onset of MEPs and SEPs. Peripheral motor conduction time (PMCT) was calculated following stimuli from Erb’s point to the onset of the maximal compound muscle action potential twitch (MMAX latency). MMAX time to peak twitch was also measured. MMAX amplitude was used to normalize the MEP to quantify corticospinal excitability. Results: Following the warm-up, significant increases in mean heart rate (44.8 ± 11.7 bpm; P < 0.001) and muscle temperature (1.4 ± 0.6°C; P < 0.001) were observed. No changes were seen in corticospinal excitability (P = 0.39), CMCT (P = 0.09), or MMAX latency (P = 0.24). However, MMAX time to peak twitch was significantly reduced (P = 0.003). Conclusion: This study has shown that exercise-based warm-ups improve neuromuscular conduction velocity via thermoregulatory processes that result in the onset of muscle contraction being more rapid, but not as a result of changes in the efficacy of neural transmission along the CST.
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Sheets, Patrick L., Benjamin A. Suter, Taro Kiritani, C. Savio Chan, D. James Surmeier, and Gordon M. G. Shepherd. "Corticospinal-specific HCN expression in mouse motor cortex: Ih-dependent synaptic integration as a candidate microcircuit mechanism involved in motor control." Journal of Neurophysiology 106, no. 5 (November 2011): 2216–31. http://dx.doi.org/10.1152/jn.00232.2011.
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Motor cortex is a key brain center involved in motor control in rodents and other mammals, but specific intracortical mechanisms at the microcircuit level are largely unknown. Neuronal expression of hyperpolarization-activated current ( Ih) is cell class specific throughout the nervous system, but in neocortex, where pyramidal neurons are classified in various ways, a systematic pattern of expression has not been identified. We tested whether Ih is differentially expressed among projection classes of pyramidal neurons in mouse motor cortex. Ih expression was high in corticospinal neurons and low in corticostriatal and corticocortical neurons, a pattern mirrored by mRNA levels for HCN1 and Trip8b subunits. Optical mapping experiments showed that Ih attenuated glutamatergic responses evoked across the apical and basal dendritic arbors of corticospinal but not corticostriatal neurons. Due to Ih, corticospinal neurons resonated, with a broad peak at ∼4 Hz, and were selectively modulated by α-adrenergic stimulation. Ih reduced the summation of short trains of artificial excitatory postsynaptic potentials (EPSPs) injected at the soma, and similar effects were observed for short trains of actual EPSPs evoked from layer 2/3 neurons. Ih narrowed the coincidence detection window for EPSPs arriving from separate layer 2/3 inputs, indicating that the dampening effect of Ih extended to spatially disperse inputs. To test the role of corticospinal Ih in transforming EPSPs into action potentials, we transfected layer 2/3 pyramidal neurons with channelrhodopsin-2 and used rapid photostimulation across multiple sites to synaptically drive spiking activity in postsynaptic neurons. Blocking Ih increased layer 2/3-driven spiking in corticospinal but not corticostriatal neurons. Our results imply that Ih-dependent synaptic integration in corticospinal neurons constitutes an intracortical control mechanism, regulating the efficacy with which local activity in motor cortex is transferred to downstream circuits in the spinal cord. We speculate that modulation of Ih in corticospinal neurons could provide a microcircuit-level mechanism involved in translating action planning into action execution.
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Aboodarda, Saied Jalal, Cindy Xin Yu Zhang, Ruva Sharara, Madeleine Cline, and Guillaume Y. Millet. "Exercise-Induced Fatigue in One Leg Does Not Impair the Neuromuscular Performance in the Contralateral Leg but Improves the Excitability of the Ipsilateral Corticospinal Pathway." Brain Sciences 9, no. 10 (September 25, 2019): 250. http://dx.doi.org/10.3390/brainsci9100250.
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To investigate the influence of pre-induced fatigue in one leg on neuromuscular performance and corticospinal responses of the contralateral homologous muscles, three experiments were conducted with different exercise protocols; A (n = 12): a 60 s rest vs. time-matched sustained left leg knee extension maximum voluntary contraction (MVC), B (n = 12): a 60 s rest vs. time-matched left leg MVC immediately followed by 60 s right leg MVC, and C (n = 9): a similar protocol to experiment B, but with blood flow occluded in the left leg while the right leg was performing the 60 s MVC. The neuromuscular assessment included 5 s knee extensions at 100%, 75%, and 50% of MVC. At each force level, transcranial magnetic and peripheral nerve stimuli were elicited to investigate the influence of different protocols on the right (tested) knee extensors’ maximal force output, voluntary activation, corticospinal excitability, and inhibition. The pre-induced fatigue in the left leg did not alter the performance nor the neuromuscular responses recorded from the right leg in the three experiments (all p > 0.3). However, enhanced corticospinal pathway excitability was evident in the tested knee extensors (p = 0.002). These results suggest that the pre-induced fatigue and muscle ischemia in one leg did not compromise the central and peripheral components of the neuromuscular function in the tested contralateral leg.
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Sidhu, Simranjit K., Andrew G. Cresswell, and Timothy J. Carroll. "Motor cortex excitability does not increase during sustained cycling exercise to volitional exhaustion." Journal of Applied Physiology 113, no. 3 (August 1, 2012): 401–9. http://dx.doi.org/10.1152/japplphysiol.00486.2012.
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The excitability of the motor cortex increases as fatigue develops during sustained single-joint contractions, but there are no previous reports on how corticospinal excitability is affected by sustained locomotor exercise. Here we addressed this issue by measuring spinal and cortical excitability changes during sustained cycling exercise. Vastus lateralis (VL) and rectus femoris (RF) muscle responses to transcranial magnetic stimulation of the motor cortex (motor evoked potentials, MEPs) and electrical stimulation of the descending tracts (cervicomedullary evoked potentials, CMEPs) were recorded every 3 min from nine subjects during 30 min of cycling at 75% of maximum workload (Wmax), and every minute during subsequent exercise at 105% of Wmax until subjective task failure. Responses were also measured during nonfatiguing control bouts at 80% and 110% of Wmax prior to sustained exercise. There were no significant changes in MEPs or CMEPs ( P > 0.05) during the sustained cycling exercise. These results suggest that, in contrast to sustained single-joint contractions, sustained cycling exercise does not increase the excitability of motor cortical neurons. The contrasting corticospinal responses to the two modes of exercise may be due to differences in their associated systemic physiological consequences.
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Proessl, F., M. C. Canino, M. E. Beckner, A. M. Sinnott, S. R. Eagle, A. D. LaGoy, W. R. Conkright, et al. "Characterizing off-target corticospinal responses to double-cone transcranial magnetic stimulation." Experimental Brain Research 239, no. 4 (February 6, 2021): 1099–110. http://dx.doi.org/10.1007/s00221-021-06044-5.
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Kidgell, Dawson J., Daniel R. Bonanno, Ashlyn K. Frazer, Glyn Howatson, and Alan J. Pearce. "Corticospinal responses following strength training: a systematic review and meta-analysis." European Journal of Neuroscience 46, no. 11 (October 9, 2017): 2648–61. http://dx.doi.org/10.1111/ejn.13710.
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Schubert, M., A. Curt, L. Jensen, and V. Dietz. "Corticospinal input in human gait: modulation of magnetically evoked motor responses." Experimental Brain Research 115, no. 2 (June 16, 1997): 234–46. http://dx.doi.org/10.1007/pl00005693.
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Stewart, Mark, Gregory J. Quirk, and VahéE Amassian. "Corticospinal responses to electrical stimulation of motor cortex in the rat." Brain Research 508, no. 2 (February 1990): 341–44. http://dx.doi.org/10.1016/0006-8993(90)90421-7.
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Martin, Peter G., Anna L. Hudson, Simon C. Gandevia, and Janet L. Taylor. "Reproducible Measurement of Human Motoneuron Excitability With Magnetic Stimulation of the Corticospinal Tract." Journal of Neurophysiology 102, no. 1 (July 2009): 606–13. http://dx.doi.org/10.1152/jn.91348.2008.
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It is difficult to test responses of human motoneurons in a controlled way or to make longitudinal assessments of adaptive changes at the motoneuron level. These studies assessed the reliability of responses produced by magnetic stimulation of the corticospinal tract. Cervicomedullary motor evoked potentials (CMEPs) were recorded in the first dorsal interosseus (FDI) on 2 separate days. On each day, four sets of stimuli were delivered at the maximal output of the stimulator, with the final two sets ≥10 min after the initial sets. Sets of stimuli were also delivered at different stimulus intensities to obtain stimulus-response curves. In addition, on the second day, responses at different stimulus intensities were evoked during weak voluntary contractions. Responses were normalized to the maximal muscle compound action potential ( Mmax). CMEPs evoked in the relaxed FDI were small, even when stimulus intensity was maximal (3.6 ± 2.5% Mmax) but much larger during a weak contraction (e.g., 26.2 ± 10.2% Mmax). CMEPs evoked in the relaxed muscle at the maximal output of the stimulator were highly reproducible both within (ICC = 0.83, session 1; ICC = 0.87, session 2) and between sessions (ICC = 0.87). ICCs for parameters of the input-output curves, which included measures of motor threshold, slope, and maximal response size, ranged between 0.87 and 0.62. These results suggest that responses to magnetic stimulation of the corticospinal tract can be assessed in relaxation and contraction and can be reliably obtained for longitudinal studies of motoneuronal excitability.
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Nuzzo, James L., Gabriel S. Trajano, Benjamin K. Barry, Simon C. Gandevia, and Janet L. Taylor. "Arm posture-dependent changes in corticospinal excitability are largely spinal in origin." Journal of Neurophysiology 115, no. 4 (April 1, 2016): 2076–82. http://dx.doi.org/10.1152/jn.00885.2015.
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Biceps brachii motor evoked potentials (MEPs) from cortical stimulation are influenced by arm posture. We used subcortical stimulation of corticospinal axons to determine whether this postural effect is spinal in origin. While seated at rest, 12 subjects assumed several static arm postures, which varied in upper-arm (shoulder flexed, shoulder abducted, arm hanging to side) and forearm orientation (pronated, neutral, supinated). Transcranial magnetic stimulation over the contralateral motor cortex elicited MEPs in resting biceps and triceps brachii, and electrical stimulation of corticospinal tract axons at the cervicomedullary junction elicited cervicomedullary motor evoked potentials (CMEPs). MEPs and CMEPs were normalized to the maximal compound muscle action potential (Mmax). Responses in biceps were influenced by upper-arm and forearm orientation. For upper-arm orientation, biceps CMEPs were 68% smaller ( P = 0.001), and biceps MEPs 31% smaller ( P = 0.012), with the arm hanging to the side compared with when the shoulder was flexed. For forearm orientation, both biceps CMEPs and MEPs were 34% smaller (both P < 0.046) in pronation compared with supination. Responses in triceps were influenced by upper-arm, but not forearm, orientation. Triceps CMEPs were 46% smaller ( P = 0.007) with the arm hanging to the side compared with when the shoulder was flexed. Triceps MEPs and biceps and triceps MEP/CMEP ratios were unaffected by arm posture. The novel finding is that arm posture-dependent changes in corticospinal excitability in humans are largely spinal in origin. An interplay of multiple reflex inputs to motoneurons likely explains the results.
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Riddle, C. Nicholas, and Stuart N. Baker. "Convergence of Pyramidal and Medial Brain Stem Descending Pathways Onto Macaque Cervical Spinal Interneurons." Journal of Neurophysiology 103, no. 5 (May 2010): 2821–32. http://dx.doi.org/10.1152/jn.00491.2009.
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We investigated the control of spinal interneurons by corticospinal and medial brain stem descending tracts in two macaque monkeys. Stimulating electrodes were implanted in the left pyramidal tract (PT), and the right medial longitudinal fasciculus (MLF), which contains reticulospinal, vestibulospinal, and some tectospinal fibers. Single unit discharge was recorded from 163 interneurons in the intermediate zone of the right spinal cord (segmental levels C6–C8) in the awake state; inputs from descending pathways were assessed from the responses to stimulation through the PT and MLF electrodes. Convergent input from both pathways was the most common finding (71/163 cells); responses to PT and MLF stimulation were of similar amplitude. Interneuron discharge was also recorded while the animal performed a reach and grasp task with the right hand; the output connections of the recorded cells were determined by delivering intraspinal microstimulation (ISMS) at the recording sites. Convergent input from MLF/PT stimulation was also common when analysis was restricted to cells that increased their rate during grasp (14/23 cells) or to cells recorded at sites where ISMS elicited finger or wrist movements (23/57 cells). We conclude that medial brain stem and corticospinal descending pathways have largely overlapping effects on spinal interneurons, including those involved in the control of the hand. This may imply a more important role for the brain stem in coordinating hand movements in primates than commonly assumed; brain stem pathways could contribute to the restoration of function seen after lesions to the corticospinal tract.
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Tuszynski, Mark H., Keith Murai, Armin Blesch, Ray Grill, and Ian Miller. "Functional Characterization of Ngf-Secreting Cell Grafts to the Acutely Injured Spinal Cord." Cell Transplantation 6, no. 3 (May 1997): 361–68. http://dx.doi.org/10.1177/096368979700600318.
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Previously we reported that grafts of cells genetically modified to produce human nerve growth factor (hNGF) promoted specific and robust sprouting of spinal sensory, motor, and noradrenergic axons. In the present study we extend these investigations to assess NGF effects on corticospinal motor axons and on functional outcomes after spinal cord injury. Fibroblasts from adult rats were transduced to express human NGF; control cells were not genetically modified. Fibroblasts were then grafted to sites of midthoracic spinal cord dorsal hemisection lesions. Three months later, recipients of NGF-secreting grafts showed deficits on conditioned locomotion over a wire mesh that did not differ in extent from control-lesioned animals. On histological examination, NGF-secreting grafts elicited specific sprouting from spinal primary sensory afferent axons, local motor axons, and putative cerulospinal axons as previously reported, but no specific responses from corticospinal axons. Axons responding to NGF robustly penetrated the grafts but did not exit the grafts to extend to normal innervation territories distal to grafts. Grafted cells continued to express NGF protein through the experimental period of the study. These findings indicate that 1) spinal cord axons show directionally sensitive growth responses to neurotrophic factors, 2) growth of axons responding to a neurotrophic factor beyond an injury site and back to their natural target regions will likely require delivery of concentration gradients of neurotrophic factors toward the target, 3) corticospinal axons do not grow toward a cellular source of NGF, and 4) functional impairments are not improved by strictly local sprouting response of nonmotor systems.
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Latella, Christopher, Matheus D. Pinto, James L. Nuzzo, and Janet L. Taylor. "Effects of postexercise blood flow occlusion on quadriceps responses to transcranial magnetic stimulation." Journal of Applied Physiology 130, no. 5 (May 1, 2021): 1326–36. http://dx.doi.org/10.1152/japplphysiol.01082.2020.
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Maintained exercise-induced firing of group III/IV quadriceps muscle afferents counteracts known reductions in corticospinal excitability that occur with fatigue. However, the results suggest that this increased excitability is not underpinned by changes in intracortical facilitatory or inhibitory networks. These findings are not consistent with previous findings for hand muscle, which reported preserved intracortical facilitation with fatigue-related sustained group III/IV muscle afferent firing.
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Bretzner, Frédéric, and Trevor Drew. "Motor Cortical Modulation of Cutaneous Reflex Responses in the Hindlimb of the Intact Cat." Journal of Neurophysiology 94, no. 1 (July 2005): 673–87. http://dx.doi.org/10.1152/jn.01247.2004.
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We have used the technique of spatial facilitation to examine the interactions between the signals conveyed by the corticospinal tract and those of cutaneous afferents in the hindlimb of the intact, walking cat. Microstimulation was applied to 20 cortical sites in the hindlimb representation of the motor cortex and to three different cutaneous nerves innervating the hindpaw in four cats. Conditioning stimuli to the motor cortex induced both facilitation and depression of cutaneous reflexes evoked by stimulation of nerves in the hindlimb contralateral to the cortical stimulation site. Facilitation was most frequently evoked by conditioning stimuli in the range of 10–30 ms before the cutaneous stimulation; depression was normally evoked by shorter and longer conditioning delays. Similar changes were observed after conditioning stimuli to the pyramidal tract, suggesting that the changes were independent of any changes in cortical excitability. Modulation of reflex activity varied according to the muscle under study, the cutaneous nerve used to evoke the reflex and the cortical site used to condition the reflex. Together, these results suggest that there is spatial convergence of corticospinal and cutaneous afferent activity and that this convergence is mediated by distinct subpopulations of spinal interneurons.
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Oya, T., B. W. Hoffman, and A. G. Cresswell. "Corticospinal-evoked responses in lower limb muscles during voluntary contractions at varying strengths." Journal of Applied Physiology 105, no. 5 (November 2008): 1527–32. http://dx.doi.org/10.1152/japplphysiol.90586.2008.
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This study investigated corticospinal-evoked responses in lower limb muscles during voluntary contractions at varying strengths. Similar investigations have been made on upper limb muscles, where evoked responses have been shown to increase up to ∼50% of maximal force and then decline. We elicited motor-evoked potentials (MEPs) and cervicomedullary motor-evoked potentials (CMEPs) in the soleus (Sol) and medial gastrocnemius (MG) muscles using magnetic stimulation over the motor cortex and cervicomedullary junction during voluntary plantar flexions with the torque ranging from 0 to 100% of a maximal voluntary contraction. Differences between the MEP and CMEP were also investigated to assess whether any changes were occurring at the cortical or spinal levels. In both Sol and MG, MEP and CMEP amplitudes [normalized to maximal M wave (Mmax)] showed an increase, followed by a plateau, over the greater part of the contraction range with responses increasing from ∼0.2 to ∼6% of Mmax for Sol and from ∼0.3 to ∼10% of Mmax for MG. Because both MEPs and CMEPs changed in a similar manner, the observed increase and lack of decrease at high force levels are likely related to underlying changes occurring at the spinal level. The evoked responses in the Sol and MG increase over a greater range of contraction strengths than for upper limb muscles, probably due to differences in the pattern of motor unit recruitment and rate coding for these muscles and the strength of the corticospinal input.
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Olivier, E., S. N. Baker, K. Nakajima, T. Brochier, and R. N. Lemon. "Investigation Into Non-Monosynaptic Corticospinal Excitation of Macaque Upper Limb Single Motor Units." Journal of Neurophysiology 86, no. 4 (October 1, 2001): 1573–86. http://dx.doi.org/10.1152/jn.2001.86.4.1573.
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There has been considerable recent debate as to relative importance, in the primate, of propriospinal transmission of corticospinal excitation to upper limb motoneurons. Previous studies in the anesthetized macaque monkey suggested that, compared with the cat, the transmission of such excitation via a system of C3–C4 propriospinal neurons may be relatively weak. However, it is possible that in the anesthetized preparation, propriospinal transmission of cortical inputs to motoneurons may be depressed. To address this issue, the current study investigated the responses of single motor units (SMUs) to corticospinal inputs in either awake ( n = 1) or lightly sedated ( n = 3) macaque monkeys. Recordings in the awake state were made during performance of a precision grip task. The responses of spontaneously discharging SMUs to electrical stimulation of the pyramidal tract (PT) via chronically implanted electrodes were examined for evidence of non-monosynaptic, presumed propriospinal, effects. Single PT stimuli (up to 250 μA; duration, 0.2 ms, 2 Hz) were delivered during steady discharge of the SMU (10–30 imp/s). SMUs were recorded from muscles acting on the thumb (adductor pollicis and abductor pollicis brevis, n = 18), wrist (extensor carpi radialis, n = 29) and elbow (biceps, n = 9). In all SMUs, the poststimulus time histograms to PT stimulation consisted of a single peak at a fixed latency and with a brief duration [0.74 ± 0.25 (SD) ms, n = 56], consistent with the responses being mediated by monosynaptic action of cortico-motoneuronal (CM) impulses. Later peaks, indicating non-monosynaptic action, were not present even when the probability of the initial peak response was low and when there was no evidence for suppression of ongoing SMU activity following this peak ( n = 20 SMUs). Even when repetitive (double-pulse) PT stimuli were used to facilitate transmission through oligosynaptic linkages, no later peaks were observed (16 SMUs). In some thumb muscle SMUs ( n = 8), responses to PT stimulation were compared with those evoked by transcranial magnetic stimulation, using a figure-eight coil held over the motor cortex. Responses varied according the orientation of the coil: in the latero-medial position, single peak responses similar to those from the PT were obtained; their latencies confirmed direct excitation of CM cells, and there were no later peaks. In the posterio-anterior orientation, responses had longer latencies and consisted of two to three subpeaks. At least under the conditions that we have tested, the results provide no positive evidence for transmission of cortical excitation to upper limb motoneurons by non-monosynaptic pathways in the macaque monkey.
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Mason, Joel, Ashlyn K. Frazer, Shapour Jaberzadeh, Juha P. Ahtiainen, Janne Avela, Timo Rantalainen, Michael Leung, and Dawson J. Kidgell. "Determining the Corticospinal Responses to Single Bouts of Skill and Strength Training." Journal of Strength and Conditioning Research 33, no. 9 (September 2019): 2299–307. http://dx.doi.org/10.1519/jsc.0000000000003266.
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Škarabot, Jakob, Paul Ansdell, John Temesi, Glyn Howatson, Stuart Goodall, and Rade Durbaba. "Neurophysiological responses and adaptation following repeated bouts of maximal lengthening contractions in young and older adults." Journal of Applied Physiology 127, no. 5 (November 1, 2019): 1224–37. http://dx.doi.org/10.1152/japplphysiol.00494.2019.
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A bout of maximal lengthening contractions is known to produce muscle damage, but confers protection against subsequent damaging bouts, with both tending to be lower in older adults. Neural factors contribute to this adaptation, but the role of the corticospinal pathway remains unclear. Twelve young (27 ± 5 yr) and 11 older adults (66 ± 4 yr) performed two bouts of 60 maximal lengthening dorsiflexions 2 weeks apart. Neuromuscular responses were measured preexercise, immediately postexercise, and at 24 and 72 h following both bouts. The initial bout resulted in prolonged reductions in maximal voluntary torque (MVC; immediately postexercise onward, P < 0.001) and increased creatine kinase (from 24 h onward, P = 0.001), with both responses being attenuated following the second bout ( P < 0.015), demonstrating adaptation. Smaller reductions in MVC following both bouts occurred in older adults ( P = 0.005). Intracortical facilitation showed no changes ( P ≥ 0.245). Motor-evoked potentials increased 24 and 72 h postexercise in young ( P ≤ 0.038). Torque variability ( P ≤ 0.041) and H-reflex size ( P = 0.024) increased, while short-interval intracortical inhibition (SICI; P = 0.019) and the silent period duration (SP) decreased ( P = 0.001) in both groups immediately postexercise. The SP decrease was smaller following the second bout ( P = 0.021), and there was an association between the change in SICI and reduction in MVC 24 h postexercise in young adults ( R = −0.47, P = 0.036). Changes in neurophysiological responses were mostly limited to immediately postexercise, suggesting a modest role in adaptation. In young adults, neural inhibitory changes are linked to the extent of MVC reduction, possibly mediated by the muscle damage–related afferent feedback. Older adults incurred less muscle damage, which has implications for exercise prescription. NEW & NOTEWORTHY This is the first study to have collectively assessed the role of corticospinal, spinal, and intracortical activity in muscle damage attenuation following repeated bouts of exercise in young and older adults. Lower levels of muscle damage in older adults are not related to their neurophysiological responses. Neural inhibition transiently changed, which might be related to the extent of muscle damage; however, the role of processes along the corticospinal pathway in the adaptive response is limited.
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Coates, Kyla D., Saied Jalal Aboodarda, Renata L. Krüger, Tristan Martin, Luanne M. Metz, Scott E. Jarvis, and Guillaume Y. Millet. "Multiple sclerosis-related fatigue: the role of impaired corticospinal responses and heightened exercise fatigability." Journal of Neurophysiology 124, no. 4 (October 1, 2020): 1131–43. http://dx.doi.org/10.1152/jn.00165.2020.
Full textAbstract:
The etiology of fatigability from whole body exercise was examined for the first time to accurately elucidate the relationship between fatigue and fatigability in multiple sclerosis (MS). Compromised corticospinal responsiveness predicted fatigue severity, providing a novel, objective indicator of fatigue in MS. Although the impaired corticomotor transmission did not aggravate muscle activation in this group of people with multiple sclerosis (PwMS) of lower disability, heightened muscle fatigability was seen to contribute to perceptions of fatigue in PwMS.
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Lockyer, Evan J., Katarina Hosel, Anna P. Nippard, Duane C. Button, and Kevin E. Power. "Corticospinal-Evoked Responses from the Biceps Brachii during Arm Cycling across Multiple Power Outputs." Brain Sciences 9, no. 8 (August 19, 2019): 205. http://dx.doi.org/10.3390/brainsci9080205.
Full textAbstract:
Background: We examined corticospinal and spinal excitability across multiple power outputs during arm cycling using a weak and strong stimulus intensity. Methods: We elicited motor evoked potentials (MEPs) and cervicomedullary motor evoked potentials (CMEPs) in the biceps brachii using magnetic stimulation over the motor cortex and electrical stimulation of corticospinal axons during arm cycling at six different power outputs (i.e., 25, 50, 100, 150, 200 and 250 W) and two stimulation intensities (i.e., weak vs. strong). Results: In general, biceps brachii MEP and CMEP amplitudes (normalized to maximal M-wave (Mmax)) followed a similar pattern of modulation with increases in cycling intensity at both stimulation strengths. Specifically, MEP and CMEP amplitudes increased up until ~150 W and ~100 W when the weak and strong stimulations were used, respectively. Further increases in cycling intensity revealed no changes on MEP or CMEP amplitudes for either stimulation strength. Conclusions: In general, MEPs and CMEPs changed in a similar manner, suggesting that increases and subsequent plateaus in overall excitability are likely mediated by spinal factors. Interestingly, however, MEP amplitudes were disproportionately larger than CMEP amplitudes as power output increased, despite being initially matched in amplitude, particularly with strong stimulation. This suggests that supraspinal excitability is enhanced to a larger degree than spinal excitability as the power output of arm cycling increases.
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Thorstensen, Jacob R., Janet L. Taylor, and Justin J. Kavanagh. "Human corticospinal-motoneuronal output is reduced with 5-HT2 receptor antagonism." Journal of Neurophysiology 125, no. 4 (April 1, 2021): 1279–88. http://dx.doi.org/10.1152/jn.00698.2020.
Full textAbstract:
Voluntary contractions and responses to magnetic stimulation of the motor cortex are dependent on serotonin activity in the central nervous system. 5-HT2 antagonism decreased evoked potential size to high-intensity stimulation, and reduced torque and lengthened inhibitory silent periods during maximal contractions. We provide novel evidence that 5-HT2 receptors are involved in muscle activation, where 5-HT effects are strongest when a large number of descending inputs activate motoneurons.
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Martin, P. G., S. C. Gandevia, and J. L. Taylor. "Output of Human Motoneuron Pools to Corticospinal Inputs During Voluntary Contractions." Journal of Neurophysiology 95, no. 6 (June 2006): 3512–18. http://dx.doi.org/10.1152/jn.01230.2005.
Full textAbstract:
This study investigated transmission of corticospinal output through motoneurons over a wide range of voluntary contraction strengths in humans. During voluntary contraction of biceps brachii, motor evoked potentials (MEPs) to transcranial magnetic stimulation of the motor cortex grow up to about 50% maximal force and then decrease. To determine whether the decrease reflects events at a cortical or spinal level, responses to stimulation of the cortex and corticospinal tract (cervicomedullary motor evoked potentials, CMEPs) as well as maximal M-waves (Mmax) were recorded during strong contractions at 50 to 100% maximum. In biceps and brachioradialis, MEPs and CMEPs (normalized to Mmax) evoked by strong stimuli decreased during strong elbow flexions. Responses were largest during contractions at 75% maximum and both potentials decreased by about 25% Mmax during maximal efforts ( P < 0.001). Reductions were smaller with weaker stimuli, but again similar for MEPs and CMEPs. Thus the reduction in MEPs during strong voluntary contractions can be accounted for by reduced responsiveness of the motoneuron pool to stimulation. During strong contractions of the first dorsal interosseous, a muscle that increases voluntary force largely by frequency modulation, MEPs declined more than in either elbow flexor muscle (35% Mmax, P < 0.001). This suggests that motoneuron firing rates are important determinants of evoked output from the motoneuron pool. However, motor cortical output does not appear to be limited at high contraction strengths.
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Carroll, Timothy J., Evan R. L. Baldwin, David F. Collins, and E. Paul Zehr. "Corticospinal Excitability Is Lower During Rhythmic Arm Movement Than During Tonic Contraction." Journal of Neurophysiology 95, no. 2 (February 2006): 914–21. http://dx.doi.org/10.1152/jn.00684.2005.
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Humans perform rhythmic, locomotor movements with the arms and legs every day. Studies using reflexes to probe the functional role of the CNS suggest that spinal circuits are an important part of the neural control system for rhythmic arm cycling and walking. Here, by studying motor-evoked potentials (MEPs) in response to transcranial magnetic stimulation (TMS) of the motor cortex, and H-reflexes induced by electrical stimulation of peripheral nerves, we show a reduction in corticospinal excitability during rhythmic arm movement compared with tonic, voluntary contraction. Responses were compared between arm cycling and tonic contraction at four positions, while participants generated similar levels of muscle activity. Both H-reflexes and MEPs were significantly smaller during arm cycling than during tonic contraction at the midpoint of arm flexion ( F = 13.51, P = 0.006; F = 11.83, P = 0.009). Subthreshold TMS significantly facilitated the FCR H-reflex during tonic contractions, but did not significantly modulate H-reflex amplitude during arm cycling. The data indicate a reduction in the responsiveness of cells constituting the fast, monosynaptic, corticospinal pathway during arm cycling and suggest that the motor cortex may contribute less to motor drive during rhythmic arm movement than during tonic, voluntary contraction. Our results are consistent with the idea that subcortical regions contribute to the control of rhythmic arm movements despite highly developed corticospinal projections to the human upper limb.
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Jo, Hang Jin, and Monica A. Perez. "Corticospinal-motor neuronal plasticity promotes exercise-mediated recovery in humans with spinal cord injury." Brain 143, no. 5 (May 1, 2020): 1368–82. http://dx.doi.org/10.1093/brain/awaa052.
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Abstract Rehabilitative exercise in humans with spinal cord injury aims to engage residual neural networks to improve functional recovery. We hypothesized that exercise combined with non-invasive stimulation targeting spinal synapses further promotes functional recovery. Twenty-five individuals with chronic incomplete cervical, thoracic, and lumbar spinal cord injury were randomly assigned to 10 sessions of exercise combined with paired corticospinal-motor neuronal stimulation (PCMS) or sham-PCMS. In an additional experiment, we tested the effect of PCMS without exercise in 13 individuals with spinal cord injury with similar characteristics. During PCMS, 180 pairs of stimuli were timed to have corticospinal volleys evoked by transcranial magnetic stimulation over the primary motor cortex arrive at corticospinal-motor neuronal synapses of upper- or lower-limb muscles (depending on the injury level), 1–2 ms before antidromic potentials were elicited in motor neurons by electrical stimulation of a peripheral nerve. Participants exercised for 45 min after all protocols. We found that the time to complete subcomponents of the Graded and Redefined Assessment of Strength, Sensibility and Prehension (GRASSP) and the 10-m walk test decreased on average by 20% after all protocols. However, the amplitude of corticospinal responses elicited by transcranial magnetic stimulation and the magnitude of maximal voluntary contractions in targeted muscles increased on overage by 40–50% after PCMS combined or not with exercise but not after sham-PCMS combined with exercise. Notably, behavioural and physiological effects were preserved 6 months after the intervention in the group receiving exercise with PCMS but not in the group receiving exercise combined with sham-PCMS, suggesting that the stimulation contributed to preserve exercise gains. Our findings indicate that targeted non-invasive stimulation of spinal synapses might represent an effective strategy to facilitate exercise-mediated recovery in humans with different degrees of paralysis and levels of spinal cord injury.
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Jo, Hang Jin, and Monica A. Perez. "Changes in motor-evoked potential latency during grasping after tetraplegia." Journal of Neurophysiology 122, no. 4 (October 1, 2019): 1675–84. http://dx.doi.org/10.1152/jn.00671.2018.
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The corticospinal pathway contributes to the control of grasping in intact humans. After spinal cord injury (SCI), there is an extensive reorganization in the corticospinal pathway; however, its contribution to the control of grasping after the injury remains poorly understood. We addressed this question by using transcranial magnetic stimulation (TMS) over the hand representation of the motor cortex to elicit motor-evoked potentials (MEPs) in an intrinsic finger muscle during precision grip and power grip with the TMS coil oriented to induce currents in the brain in the latero-medial (LM) direction to activate corticospinal axons directly and in the posterior-anterior (PA) and anterior-posterior (AP) directions to activate the axon indirectly through synaptic inputs in humans with and without cervical incomplete SCI. We found prolonged MEP latencies in all coil orientations in both tasks in SCI compared with control subjects. The latencies of MEPs elicited by AP relative to LM stimuli were consistently longer during power compared with precision grip in controls and SCI subjects. In contrast, PA relative to LM MEP latencies were similar between tasks across groups. Central conduction time of AP MEPs was prolonged during power compared with precision grip in controls and SCI participants. Our results support evidence indicating that inputs activated by AP and PA currents are engaged to a different extent during fine and gross grasping in humans with and without SCI. NEW & NOTEWORTHY The mechanisms contributing to the control of hand function in humans with spinal cord injury (SCI) remain poorly understood. Here, we demonstrate for the first time that the latency of corticospinal responses elicited by transcranial magnetic stimulation anterior-posterior induced currents, relative to latero-medial currents, was prolonged during power compared with precision grip in humans with and without SCI. Gross grasping might represent a stragegy to engage networks activated by anterior-posterior currents after SCI.
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Thomas, Sarah L., and Monica A. Gorassini. "Increases in Corticospinal Tract Function by Treadmill Training After Incomplete Spinal Cord Injury." Journal of Neurophysiology 94, no. 4 (October 2005): 2844–55. http://dx.doi.org/10.1152/jn.00532.2005.
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In this study, we examined if several months of intensive locomotor training increases the function of spared corticospinal tract pathways after chronic spinal cord injury (SCI) in association with the recovery of locomotor function. Transcranial magnetic stimulation (TMS) at incrementing levels of intensity was applied over the motor cortex supplying either the tibialis anterior or vastus lateralis muscles, and the resulting peak-to-peak amplitude of the motor-evoked potentials (MEPs) were measured to obtain a recruitment curve both before and after training. In the majority of subjects (7/8), 3–5 mo of daily intensive training increased the responses to TMS in at least one of the leg muscles tested (9/13). On average, across all muscles tested MEPmax, which was evoked at high stimulation intensities, increased by 46% and MEPh, which was evoked at intermediate stimulation intensities, increased by 45% (both significantly different from 0), indicating an increase in corticospinal tract connectivity from training. The slope of the sigmoid function fit to the recruitment curve increased by 24% after training (significantly different), indicating an expansion and/or increased excitability of corticospinal circuits supplying muscles to the lower leg. We also observed that the average duration of the silent period measured at MEPmax increased after training from 130 to 178 ms, suggesting that training had effects on cortical circuits thought to mediate this long-lasting inhibition. The percentage increase in MEPmax was positively and significantly correlated to the degree of locomotor recovery as assessed by the WISCI II score, the distance a subject could walk in 6 min, and the amplitude of the locomotor EMG activity, suggesting that the corticospinal tract, in part, mediated the functional locomotor recovery produced from training.
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Sohn, Young H., Nguyet Dang, and Mark Hallett. "Suppression of Corticospinal Excitability During Negative Motor Imagery." Journal of Neurophysiology 90, no. 4 (October 2003): 2303–9. http://dx.doi.org/10.1152/jn.00206.2003.
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To investigate the effect of negative motor imagery on corticospinal excitability, we performed transcranial magnetic stimulation (TMS) studies in seven healthy subjects during imagination of suppressing movements. Subjects were asked to imagine suppression of TMS-induced twitching movement of their nondominant left hands by attempting to increase the amount of relaxation after receiving an auditory NoGo cue (negative motor imagery), but to imagine squeezing hands after a Go cue (positive motor imagery). Single- and paired-pulse TMS were triggered at 2 s after Go or NoGo cues. Motor-evoked potentials (MEPs) were recorded in the first dorsal interosseus (FDI), abductor pollicis brevis (APB), and abductor digiti minimi (ADM) muscles of the left hand. Paired-pulse TMS with subthreshold conditioning stimuli at interstimulus intervals of 2 (short intracortical inhibition) and 15 ms (intracortical facilitation) and that with suprathreshold conditioning stimuli at interstimulus interval of 80 ms (long intracortical inhibition) were performed in both negative motor imagery and control conditions. Compared with the control state (no imagination), MEP amplitudes of FDI (but not APB and ADM) were significantly suppressed in negative motor imagery, but those from all three muscles were unchanged during positive motor imagery. F-wave responses (amplitudes and persistence) were unchanged during both negative and positive motor imagery. During negative motor imagery, resting motor threshold was significantly increased, but short and long intracortical inhibition and intracortical facilitation were unchanged. The present results demonstrate that excitatory corticospinal drive is suppressed during imagination of suppressing movements.