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This is the first study to examine the influence of activity in one limb on corticospinal excitability to the contralateral limb during a locomotor output. Corticospinal and spinal excitability to the biceps brachii of the ipsilateral arm were assessed using transcranial magnetic stimulation (TMS) of the motor cortex and transmastoid electrical stimulation (TMES) of corticospinal axons, respectively. Responses were evoked during the mid-elbow extension position of arm cycling across three different cycling tasks: (1) bilateral arm cycling (BL), (2) unilateral, contralateral cycling with the ipsilateral arm moving passively (IP), and (3) unilateral, contralateral cycling with the ipsilateral arm at rest (IR). Each of these three tasks were performed at two cadences: 60 and 90 rpm. TMS-induced motor evoked potential (MEPs) amplitudes were significantly smaller during BL compared to the IP and IR conditions; however, MEP amplitudes were not significantly different between IP and IR. TMES-evoked cervicomedullary MEP (CMEPs) amplitudes followed a similar pattern of task-dependent modulation, with BL having the smallest CMEPs and IR having the largest. In line with our previous findings, MEP amplitudes increased and CMEP amplitudes decreased as the cadence increased from 60 to 90 rpm. We suggest that the higher corticospinal excitability to the ipsilateral limb during the IP and IR conditions was predominantly due to disinhibition at both the cortical and spinal levels.

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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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A patient with bilateral diffuse uveal melanocytic proliferation (BDUMP) associated with endometrial cancer was treated with plasmapheresis, but failed therapy with progressive serous retinal detachment. We collected plasma before and after plasmapheresis therapy. Our goal was to determine if the cultured melanocyte elongation and proliferation (CMEP) factor and hepatocyte growth factor (HGF) was present in the IgG enriched fraction and understand why our patient failed plasmapheresis therapy. Melanocytes were cultured for 3-5 days in the presence of control medium, unfractionated pre-plasmapheresis BDUMP medium, IgG enriched or IgG depleted BDUMP medium, or unfractionated post-plasmapheresis BDUMP medium. Subretinal fluid was collected from patients with BDUMP and control retinal detachments and analyzed by electropheresis with immunoblotting. Medium with unfractionated BDUMP plasma stimulated melanocyte growth 1.4-1.5 fold compared to control medium on days 3-5 (p < 0.001 for all). Both IgG enriched and IgG depleted BDUMP medium mildly increased melanocyte growth 1.3 fold (p < 0.05 for enriched, p < 0.01 for depleted) compared to control. In comparison, unfractionated BDUMP medium caused a 1.7-fold increase in melanocyte growth, which was significantly more than the enriched (p < 0.01) and depleted (p < 0.05) fractions. Pre-plasmapheresis and post-plasmapheresis unfractionated BDUMP medium equally stimulated melanocyte growth 1.7-fold (p < 0.05) compared to control. HGF was present in IgG depleted, pre-plasmapheresis, and post-plasmapheresis samples, but absent in the IgG enriched fraction. There was no enrichment of IgG in the subretinal fluid from eyes with BDUMP. In conclusion, CMEP factor is not concentrated in the IgG enriched plasma fraction in our patient who failed plasmapheresis therapy. HGF levels have no correlation with melanocyte growth. Because plasmapheresis preferentially removes immunoglobulins from the plasma, our patient responded poorly to plasmapheresis treatment with worsening retinal detachment.

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Torque depression (TD) is the reduction in steady-state isometric torque following active muscle shortening when compared to an isometric reference contraction at the same muscle length and activation level. Central nervous system excitability differs in the TD state. While torque production about a joint is influenced by both agonist and antagonist muscle activation, investigations of corticospinal excitability have focused on agonist muscle groups. Hence, it is unknown how the TD state affects spinal and supraspinal excitability of an antagonist muscle. Eight participants (~ 24y, three female) performed 14 submaximal dorsiflexion contractions at the intensity needed to maintain a level of integrated electromyographic activity in the soleus equivalent to 15% of that recorded during a maximum plantar flexion contraction. The seven contractions of the TD protocol included a 2 s isometric phase at an ankle angle of 140°, a 1 s shortening phase at 40°/s, and a 7 s isometric phase at an angle of 100°. The seven isometric reference contractions were performed at an ankle angle of 100° for 10 s. Motor evoked potentials (MEPs), cervicomedullary motor evoked potentials (CMEPs), and maximal M-waves (Mmax) were recorded from the soleus in both conditions. In the TD compared to isometric reference state, a 13% reduction in dorsiflexor torque was accompanied by 10% lower spinal excitability (normalized CMEP amplitude; CMEP/Mmax), and 17% greater supraspinal excitability (normalized MEP amplitude; MEP/CMEP) for the soleus muscle. These findings demonstrate a neuromechanical coupling following active muscle shortening and indicate that the underlying mechanisms of TD influence antagonist activation during voluntary force production.

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The present study investigated the effects of cadence and power output on corticospinal excitability to the biceps (BB) and triceps brachii (TB) during arm cycling. Supraspinal and spinal excitability were assessed using transcranial magnetic stimulation (TMS) of the motor cortex and transmastoid electrical stimulation (TMES) of the corticospinal tract, respectively. Motor-evoked potentials (MEPs) elicited by TMS and cervicomedullary motor-evoked potentials (CMEPs) elicited by TMES were recorded at two positions during arm cycling corresponding to mid-elbow flexion and mid-elbow extension (i.e., 6 and 12 o'clock made relative to a clock face, respectively). Arm cycling was performed at combinations of two cadences (60 and 90 rpm) at three relative power outputs (20, 40, and 60% peak power output). At the 6 o'clock position, BB MEPs increased ~11.5% as cadence increased and up to ~57.2% as power output increased ( P < 0.05). In the TB, MEPs increased ~15.2% with cadence ( P = 0.013) but were not affected by power output, while CMEPs increased with cadence (~16.3%) and power output (up to ~19.1%, P < 0.05). At the 12 o'clock position, BB MEPs increased ~26.8% as cadence increased and up to ~96.1% as power output increased ( P < 0.05), while CMEPs decreased ~29.7% with cadence ( P = 0.013) and did not change with power output ( P = 0.851). In contrast, TB MEPs were not different with cadence or power output, while CMEPs increased ~12.8% with cadence and up to ~23.1% with power output ( P < 0.05). These data suggest that the "type" of intensity differentially modulates supraspinal and spinal excitability in a manner that is phase- and muscle dependent. NEW & NOTEWORTHY There is currently little information available on how changes in locomotor intensity influence excitability within the corticospinal pathway. This study investigated the effects of arm cycling intensity (i.e., alterations in cadence and power output) on corticospinal excitability projecting to the biceps and triceps brachii during arm cycling. We demonstrate that corticospinal excitability is modulated differentially by cadence and power output and that these modulations are dependent on the phase and the muscle examined.

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In quadrupeds, special circuity located within the spinal cord, referred to as central pattern generators (CPGs), is capable of producing complex patterns of activity such as locomotion in the absence of descending input. During these motor outputs, the electrical properties of spinal motoneurones are modulated such that the motoneurone is more easily activated. Indirect evidence suggests that like quadrupeds, humans also have spinally located CPGs capable of producing locomotor outputs, albeit descending input is considered to be of greater importance. Whether motoneurone properties are reconfigured in a similar manner to those of quadrupeds is unclear. The purpose of this review is to summarize our current state of knowledge regarding the modulation of motoneurone excitability during CPG-mediated motor outputs using animal models. This will be followed by more recent work initially aimed at understanding changes in motoneurone excitability during CPG-mediated motor outputs in humans, which quickly expanded to also include supraspinal excitability.

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OBJECTIVE The aim of this study was to determine the most effective electrode montage to elicit lower-extremity transcranial motor evoked potentials (LE-tMEPs) using a minimum stimulation current. METHODS A realistic 3D head model was created from T1-weighted images. Finite element methods were used to visualize the electric field in the brain, which was generated by transcranial electrical stimulation via 4 electrode montage models. The stimulation threshold level of LE-tMEPs in 52 patients was also studied in a practical clinical setting to determine the effects of each electrode montage. RESULTS The electric field in the brain radially diffused from the brain surface at a maximum just below the electrodes in the finite element models. The Cz-inion electrode montage generated a centrally distributed high electric field with a current direction longitudinal and parallel to most of the pyramidal tract fibers of the lower extremity. These features seemed to be effective in igniting LE-tMEPs. Threshold level recordings of LE-tMEPs revealed that the Cz-inion electrode montage had a lower threshold on average than the C3-C4 montage, 76.5 ± 20.6 mA and 86.2 ± 20.6 mA, respectively (31 patients, t = 4.045, p < 0.001, paired t-test). In 23 (74.2%) of 31 cases, the Cz-inion montage could elicit LE-tMEPs at a lower threshold than C3-C4. CONCLUSIONS The C3-C4 and C1-C2 electrode montages are the standard for tMEP monitoring in neurosurgery, but the Cz-inion montage showed lower thresholds for the generation of LE-tMEPs. The Cz-inion electrode montage should be a good alternative for LE-tMEP monitoring when the C3-C4 has trouble igniting LE-tMEPs.

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OBJECTIVE Transcranial motor evoked potential (tMEP) monitoring is popular in neurosurgery; however, the accuracy of tMEP can be impaired by craniotomy. Each craniotomy procedure and changes in the CSF levels affects the current spread. The aim of this study was to investigate the influence of several craniotomies on tMEP monitoring by using C3-4 transcranial electrical stimulation (TES). METHODS The authors used the finite element method to visualize the electric field in the brain, which was generated by TES, using realistic 3D head models developed from T1-weighted MR images. Surfaces of 5 layers of the head (brain, CSF, skull, subcutaneous fat, and skin layer) were separated as accurately as possible. The authors created 5 models of the head, as follows: normal head; frontotemporal craniotomy; parietal craniotomy; temporal craniotomy; and occipital craniotomy. The computer simulation was investigated by finite element methods, and clinical recordings of the stimulation threshold level of upper-extremity tMEP (UE-tMEP) during neurosurgery were also studied in 30 patients to validate the simulation study. RESULTS Bone removal during the craniotomy positively affected the generation of the electric field in the motor cortex if the motor cortex was just under the bone at the margin of the craniotomy window. This finding from the authors' simulation study was consistent with clinical reports of frontotemporal craniotomy cases. A major decrease in CSF levels during an operation had a significantly negative impact on the electric field when the motor cortex was exposed to air. The CSF surface level during neurosurgery depends on the body position and location of the craniotomy. The parietal craniotomy and temporal craniotomy were susceptible to the effect of the changing CSF level, based on the simulation study. A marked increase in the threshold following a decrease in CSF was actually recorded in clinical reports of the UE-tMEP threshold from a temporal craniotomy. However, most frontotemporal craniotomy cases were minimally affected by a small decrease in CSF. CONCLUSIONS Bone removal during a craniotomy positively affects the generation of the electric field in the motor cortex if the motor cortex is just under the bone at the margin of the craniotomy window. The CSF decrease and the shifting brain can negatively affect tMEP ignition. These changes should be minimized to maintain the original conductivity between the motor cortex and the skull, and the operation team must remember the fluctuation of the tMEP threshold.

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This is the first study to examine corticospinal excitability (CSE) to antagonistic muscle groups during arm cycling. Transcranial magnetic stimulation (TMS) of the motor cortex and transmastoid electrical stimulation (TMES) of the corticospinal tract were used to assess changes in supraspinal and spinal excitability, respectively. TMS induced motor evoked potentials (MEPs) and TMES induced cervicomedullary evoked potentials (CMEPs) were recorded from the biceps and triceps brachii at two positions, mid-elbow flexion and extension, while cycling at 5% and 15% of peak power output. While phase-dependent modulation of MEP and CMEP amplitudes occurred in the biceps brachii, there was no difference between flexion and extension for MEP amplitudes in the triceps brachii and CMEP amplitudes were higher during flexion than extension. Furthermore, MEP amplitudes in both biceps and triceps brachii increased with increased workload. CMEP amplitudes increased with higher workloads in the triceps brachii, but not biceps brachii, though the pattern of change in CMEPs was similar to MEPs. Differences between changes in CSE between the biceps and triceps brachii suggest that these antagonistic muscles may be under different neural control during arm cycling. Putative mechanisms are discussed.

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The purpose of this study was to examine the influence of neutral and pronated handgrip positions on corticospinal excitability to the biceps brachii during arm cycling. Corticospinal and spinal excitability were assessed using motor evoked potentials (MEPs) elicited via transcranial magnetic stimulation (TMS) and cervicomedullary-evoked potentials (CMEPs) elicited via transmastoid electrical stimulation (TMES), respectively. Participants were seated upright in front on arm cycle ergometer. Responses were recorded from the biceps brachii at two different crank positions (6 and 12 o'clock positions relative to a clock face) while arm cycling with neutral and pronated handgrip positions. Responses were also elicited during tonic elbow flexion to compare/contrast the results to a non-rhythmic motor output. MEP and CMEP amplitudes were significantly larger at the 6 o'clock position while arm cycling with a neutral handgrip position compared to pronated (45.6 and 29.9%, respectively). There were no differences in MEP and CMEP amplitudes at the 12 o'clock position for either handgrip position. For the tonic contractions, MEPs were significantly larger with a neutral vs. pronated handgrip position (32.6% greater) while there were no difference in CMEPs. Corticospinal excitability was higher with a neutral handgrip position for both arm cycling and tonic elbow flexion. While spinal excitability was also higher with a neutral handgrip position during arm cycling, no difference was observed during tonic elbow flexion. These findings suggest that not only is corticospinal excitability to the biceps brachii modulated at both the supraspinal and spinal level, but that it is influenced differently between rhythmic arm cycling and tonic elbow flexion.

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Based on H-reflex data, spinal mechanisms are proposed to be responsible for the first 50-80 ms of the transcranial magnetic stimulation (TMS)-induced silent period. As several methodological issues can compromise H-reflex validity as a measure of motoneuron excitability, this study used transmastoid stimulation to elicit cervicomedullary motor evoked potentials (CMEPs) during the silent period. Eleven subjects made 1-3 visits which involved 32 or 44 brief (~3 s) isometric elbow flexor contractions at 25 % of maximal torque. During each contraction, transmastoid stimulation was delivered in isolation to elicit an unconditioned CMEP and at interstimulus intervals (ISIs) ranging from 50 to 150 ms after TMS to elicit a conditioned CMEP. Stimulus intensities for TMS and transmastoid stimulation were set to elicit a silent period of ~200 ms and an unconditioned CMEP of 15, 50, or 85 % of the maximal compound muscle action potential (M max), respectively. At all ISIs and intensities of transmastoid stimulation, the conditioned CMEP was significantly smaller than the unconditioned CMEP (p < 0.001). However, suppression of the conditioned CMEP was significantly less at 85 % compared to 15 or 50 % M max (p = 0.001). Contrary to published H-reflex data, the conditioned CMEP did not recover within 50-80 ms, remaining significantly suppressed at the longest ISI tested (150 ms). These data suggest the spinal portion of the TMS-evoked silent period is considerably longer than reported previously. Transmastoid stimulation, unlike peripheral nerve stimulation, does not impact proprioceptive inflow to motoneurons. Hence, relative to the H-reflex, the CMEP will be subjected to greater afferent-mediated disfacilitation and inhibition due to the TMS-induced muscle twitch.

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This is the first study to examine changes in corticospinal excitability to the biceps brachii during the onset of arm cycling from a resting position to a point when steady-state arm cycling was obtained. Supraspinal and spinal excitability were assessed using motor-evoked potentials (MEPs) elicited via transcranial magnetic stimulation and cervicomedullary evoked potentials (CMEPs) elicited via transmastoid electrical stimulation, respectively. Evoked responses were recorded from the biceps brachii during elbow flexion (6 o'clock relative to a clock face) for both arm cycling and an intensity-matched tonic contraction at three separate periods: (1) immediately at the onset of motor output and after completion of the (2) 4th revolution and (3) 9th revolution. There was no difference during initiation between tasks for MEP (P = 0.79) or CMEP amplitudes (P = 0.57). However, MEP amplitudes were significantly larger during arm cycling than an intensity-matched tonic contraction after the completion of the 4th (Cycling 76.48 ± 17.35 % of M max, Tonic 63.45 ± 18.45 % of M max, P < 0.05) and 9th revolutions (Cycling 72.37 ± 15.96 % of M max, Tonic 58.1 ± 24.23 % of M max, P < 0.05). There were no differences between conditions in CMEP amplitudes at the 4th (Cycling 49.6 ± 25.4 % of M max, Tonic 41.6 ± 11.2 % of M max, P = 0.31) or the 9th revolution (Cycling 47.2 ± 17.0 % of M max, Tonic 40.8 ± 13.6 % of M max, P = 0.29). These results demonstrate that corticospinal excitability is not different between arm cycling and a tonic contraction at motor output onset, but supraspinal excitability is enhanced during steady-state arm cycling. This suggests a similarity in the way the corticospinal tract initiates motor outputs in humans, regardless of the differences that present themselves in the later, steady-state stages.

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This is the first study to report the influence of different cadences on the modulation of supraspinal and spinal excitability during arm cycling. Supraspinal and spinal excitability were assessed using transcranial magnetic stimulation of the motor cortex and transmastoid electrical stimulation of the corticospinal tract, respectively. Transcranial magnetic stimulation-induced motor evoked potentials and transmastoid electrical stimulation-induced cervicomedullary evoked potentials (CMEPs) were recorded from the biceps brachii at two separate positions corresponding to elbow flexion and extension (6 and 12 o'clock relative to a clock face, respectively) while arm cycling at 30, 60 and 90 rpm. Motor evoked potential amplitudes increased significantly as cadence increased during both elbow flexion (P < 0.001) and extension (P = 0.027). CMEP amplitudes also increased with cadence during elbow flexion (P < 0.01); however, the opposite occurred during elbow extension (i.e., decreased CMEP amplitude; P = 0.01). The data indicate an overall increase in the excitability of corticospinal neurons which ultimately project to biceps brachii throughout arm cycling as cadence increased. Conversely, changes in spinal excitability as cadence increased were phase dependent (i.e., increased during elbow flexion and decreased during elbow extension). Phase- and cadence-dependent changes in spinal excitability are suggested to be mediated via changes in the balance of excitatory and inhibitory synaptic input to the motor pool, as opposed to changes in the intrinsic properties of spinal motoneurons.

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It is well known that the H-reflex amplitude decreases during passive muscle lengthening in comparison with passive shortening. However, this decrease in spinal synaptic efficacy observed during passive lengthening seems to be lesser during eccentric voluntary contraction. The aim of the present study was to examine whether spinal excitability during lengthening condition could be modulated by magnetic brain stimulation. H reflexes of the triceps surae muscles were elicited on 10 young healthy subjects, and conditioned by a sub-threshold transcranial magnetic stimulation (TMS). The conditioning stimulation was applied over the M1 area of triceps surae muscles at an intensity below motor threshold with a conditioning-test interval of 5ms. Conditioned and non-conditioned H-reflexes were elicited at rest, during passive lengthening and shortening, and during submaximal contractions (concentric, eccentric and isometric). During passive and active lengthening, H reflexes conditioned by a sub-threshold TMS pulse increased on average by 50% compared with non-conditioned responses. No significant effect was found during isometric and concentric conditions. Activation of the corticospinal pathway would partially cancel inhibitions caused by muscle stretch, and according to the time-delayed effect, this result suggested the existence of a specific polysynaptic pathway. In additional experiments, H responses were conditioned by cervico-medullary stimulations, showing that the modulation described by the previous results involves subcortical mechanisms. This study provides further evidences that the modulation of the final cortico-spinal command reaching the muscle depends on a central mechanism that controls peripheral input, such as Ia afference discharge during lengthening.

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The responsiveness of the human central nervous system can change profoundly with exercise, injury, disuse, or disease. Changes occur at both cortical and spinal levels but in most cases excitability of the motoneuron pool must be assessed to localize accurately the site of adaptation. Hence, it is critical to understand, and employ correctly, the methods to test motoneuron excitability in humans. Several techniques exist and each has its advantages and disadvantages. This review examines the most common techniques that use evoked compound muscle action potentials to test the excitability of the motoneuron pool and describes the merits and limitations of each. The techniques discussed are the H-reflex, F-wave, tendon jerk, V-wave, cervicomedullary motor evoked potential (CMEP), and motor evoked potential (MEP). A number of limitations with these techniques are presented.