Task-specific strength increases after lower-limb compound resistance training occurred in the absence of corticospinal changes in vastus lateralis.

NEW FINDINGS: What is the central question of the study? Are corticospinal responses to acute and short-term squat resistance training task-specific? What is the main finding and its importance? A single bout of resistance training increased spinal excitability, but no changes in corticospinal responses were noted following 4 weeks of squat training despite task-specific increases in strength. The present data suggest that processes along the corticospinal pathway of the knee extensors play a limited role in the task-specific increase in strength following resistance training. ABSTRACT: Neural adaptations subserving strength increases have been shown to be task-specific, but responses and adaptation to lower-limb compound exercises such as the squat are commonly assessed in a single-limb isometric task. This two-part study assessed neuromuscular responses to an acute bout (Study A) and 4 weeks (Study B) of squat resistance training at 80% of one-repetition-maximum, with measures taken during a task-specific isometric squat (IS) and non-specific isometric knee extension (KE). Eighteen healthy volunteers (25 ± 5 years) were randomised into either a training (n = 10) or a control (n = 8) group. Neural responses were evoked at the intracortical, corticospinal and spinal levels, and muscle thickness was assessed using ultrasound. The results of Study A showed that the acute bout of squat resistance training decreased maximum voluntary contraction (MVC) for up to 45 min post-exercise (-23%, P < 0.001). From 15-45 min post-exercise, spinally evoked responses were increased in both tasks (P = 0.008); however, no other evoked responses were affected (P ≥ 0.240). Study B demonstrated that following short-term resistance training, participants improved their one repetition maximum squat (+35%, P < 0.001), which was reflected by a task-specific increase in IS MVC (+49%, P = 0.001), but not KE (+1%, P = 0.882). However, no training-induced changes were observed in muscle thickness (P = 0.468) or any evoked responses (P = 0.141). Adjustments in spinal motoneuronal excitability are evident after acute resistance training. After a period of short-term training, there were no changes in the responses to central nervous system stimulation, which suggests that alterations in corticospinal properties of the vastus lateralis might not contribute to increases in strength.

Corticospinal responses during passive shortening and lengthening of tibialis anterior and soleus in older compared to younger adults.

NEW FINDINGS: What is the central question of this study? Are there age-related differences in corticospinal responses whilst passively changing muscle length? What is the main finding and its importance? In contrast to young, older adults exhibited no modulation of corticospinal excitability in tibialis anterior during passive ankle movement. These data show impaired sensorimotor response in older adults during length changes of tibialis anterior, thus contributing to our understanding of age-related changes in sensorimotor control. ABSTRACT: Corticospinal responses have been shown to increase and decrease with passive muscle shortening and lengthening, respectively, as a result of changes in muscle spindle afferent feedback. The ageing sensory system is accompanied by a number of alterations that might influence the processing and integration of sensory information. Consequently, corticospinal excitability might be modulated differently whilst changing muscle length. In 10 older adults (66 ± 4 years), corticospinal responses (MEP/Mmax ) were evoked in a static position, and during passive shortening and lengthening of soleus (SOL) and tibialis anterior (TA), and these data were compared to the re-analysed data pool of 18 younger adults (25 ± 4 years) published previously. Resting motor threshold was greater in SOL compared to TA (P < 0.001), but did not differ between young and older (P = 0.405). No differences were observed in MEP/Mmax between the static position, passive shortening or lengthening in SOL (young: all 0.02 ± 0.01; older: 0.05 ± 0.04, 0.03 ± 0.02 and 0.04 ± 0.01, respectively; P = 0.298), and responses were not dependent on age (P = 0.090). Conversely, corticospinal responses in TA were modulated differently between the age groups (P = 0.002), with greater MEP/Mmax during passive shortening (0.22 ± 0.12) compared to passive lengthening (0.13 ± 0.10) and static position (0.10 ± 0.05) in young (P < 0.001), but unchanged in older adults (0.19 ± 0.11, 0.22 ± 0.11 and 0.18 ± 0.07, respectively; P ≥ 0.867). The present experiment shows that length-dependent changes in corticospinal excitability in TA of the young are not evident in older adults. This suggests impaired sensorimotor response during muscle length changes in older age that might only be present in ankle flexors, but not extensors.

Compound maximal motor unit response is modulated by contraction intensity, but not contraction type in tibialis anterior.

Determining a single compound maximal motor response (MMAX ) or an average superimposed MMAX response (MSUP ) are commonly used reference values in experiments eliciting raw electromyographic, motor evoked potentials, H-reflexes, and V-waves. However, existing literature is limited in detailing the most appropriate method to normalize these electrophysiological measures. Due to the accessibility of assessment from a cortical and spinal perspective, the tibialis anterior is increasingly used in literature and hence investigated in this study. The aims of the present study were to examine the differences and level of agreement in MMAX /MSUP under different muscle actions and contraction intensities. Following a familiarization session, 22 males visited the laboratory on a single occasion. MMAX was recorded under 10% isometric and 25% and 100% shortening and lengthening maximal voluntary contractions (MVC) at an angular velocity of 15° sec-1 . MSUP was also recorded during 100% shortening and lengthening with an average of five responses recorded. There were no differences in MMAX or MSUP between contraction types. All variables showed large, positive correlations (P < 0.001, r2  ≥ 0.64). MMAX amplitude was larger (P < 0.001) at 100% shortening and lengthening intensity compared to MMAX amplitude at 10% isometric and 25% lengthening MVC. Bland-Altman plots revealed a bias toward higher MMAX at the higher contraction intensities. Despite MSUP being significantly smaller than MMAX (P < 0.001) at 100% MVC, MSUP showed a large positive correlation (P < 0.001, r2  ≥ 0.64) with all variables. It is our recommendation that MMAX should be recorded at specific contraction intensity but not necessarily a specific contraction type.

Neurophysiological responses and adaptation following repeated bouts of maximal lengthening contractions in young and older adults.

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.

Corticospinal excitability of tibialis anterior and soleus differs during passive ankle movement.

The purpose of this study was to assess corticospinal excitability of soleus (SOL) and tibialis anterior (TA) at a segmental level during passive ankle movement. Four experimental components were performed to assess the effects of passive ankle movement and muscle length on corticospinal excitability (MEP/Mmax) at different muscle lengths, subcortical excitability at the level of lumbar spinal segments (LEP/Mmax), intracortical inhibition (SICI) and facilitation (ICF), and H-reflex in SOL and TA. In addition, the degree of fascicle length changes between SOL and TA was assessed in a subpopulation during passive ankle movement. Fascicles shortened and lengthened with joint movement during passive shortening and lengthening of SOL and TA to a similar degree (p < 0.001). Resting motor threshold was greater in SOL compared to TA (p ≤ 0.014). MEP/Mmax was facilitated in TA during passive shortening relative to the static position (p ≤ 0.023) and passive lengthening (p ≤ 0.001), but remained similar during passive ankle movement in SOL (p ≥ 0.497), regardless of muscle length at the point of stimulus (p = 0.922). LEP/Mmax (SOL: p = 0.075, TA: p = 0.071), SICI (SOL: p = 0.427, TA: p = 0.540), and ICF (SOL: p = 0.177, TA: p = 0.777) remained similar during passive ankle movement. H-reflex was not different across conditions in TA (p = 0.258), but was reduced during passive lengthening compared to shortening in SOL (p = 0.048). These results suggest a differential modulation of corticospinal excitability between plantar and dorsiflexors during passive movement. The corticospinal behaviour observed might be mediated by an increase in corticospinal drive as a result of reduced afferent input during muscle shortening and appears to be flexor-biased.

Reduced corticospinal responses in older compared with younger adults during submaximal isometric, shortening, and lengthening contractions.

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.

Electrical stimulation of human corticospinal axons at the level of the lumbar spinal segments.

Electrical stimulation over the mastoids or thoracic spinous processes has been used to assess subcortical contribution to corticospinal excitability, but responses are difficult to evoke in the resting lower limbs or are limited to only a few muscle groups. This might be mitigated by delivering the stimuli lower on the spinal column, where the descending tracts contain a greater relative density of motoneurons projecting to lower limb muscles. We investigated activation of the corticospinal axons innervating tibialis anterior (TA) and rectus femoris (RF) by applying a single electrical stimulus over the first lumbar spinous process (LS). LS was paired with transcranial magnetic stimulation (TMS) at interstimulus intervals (ISIs) of -16 (TMS before LS) to 14 ms (LS before TMS). The relationship between muscle contraction strength (10%-100% maximal) and the amplitude of single-pulse TMS and LS responses was also investigated. Compared to the responses to TMS alone, responses to paired stimulation were significantly occluded in both muscles for ISIs ≥-8 ms (p ≤ 0.035), consistent with collision of descending volleys from TMS with antidromic volleys originating from LS. This suggests that TMS and LS activate some of the same corticospinal axons. Additionally, the amplitude of TMS and LS responses increased with increasing contraction strengths with no change in onset latency, suggesting responses to LS are evoked transsynaptically and have a monosynaptic component. Taken together, these experiments provide evidence that LS is an alternative method that could be used to discern segmental changes in the corticospinal tract when targeting lower limb muscles.

Corticospinal excitability during shortening and lengthening actions with incremental torque output.

NEW FINDINGS: What is the central question of this study? The relationship between motor unit recruitment and firing rate has been related to the size of the corticospinal output with variations in the nervous system gain during isometric contractions. However, corticospinal behaviour with incremental torque output might differ during anisometric contractions owing to differences in neural control of anisometric contraction types. What is the main finding and its importance? Corticospinal excitability during lengthening contractions was smaller compared with shortening but increased with incremental torque output in a similar manner between contraction types. The relationship between motor unit recruitment and firing rates is probably the main determinant of the size of an evoked response with variations in system gain. ABSTRACT: The modulation of motor evoked potentials (MEPs), an index of corticospinal excitability, has been shown to increase during isometric contractions with incremental torque output in accordance with the contribution between motor unit recruitment and firing rate of the muscle to increases in required torque output. However, the motor unit strategy of the muscle might not be the only factor influencing this behaviour, because differences in pre- and postsynaptic control have been reported between lengthening and shortening or isometric contractions. In 30 healthy adults, MEPs were elicited in tibialis anterior during shortening and lengthening contractions at 15, 25, 50 and 80% of contraction-type-specific maximal voluntary contraction torque. Background EMG activity increased progressively with greater torque output (P < 0.001) but was similar between contraction types (P = 0.162). When normalized to the maximal muscle response, MEPs were greater during shortening compared with lengthening contractions (P = 0.004) and increased stepwise with increased contraction intensities (P = 0.001). These data show an increase in corticospinal excitability with torque output from lower to higher contraction intensities, suggesting a greater contribution of motor unit recruitment to increased nervous system gain in the tibialis anterior. Despite differences in corticospinal control of shortening and lengthening contractions, the data suggest that the corticospinal responses to increases in torque output are not dependent on contraction type, because corticospinal excitability increased to a similar extent during shortening and lengthening actions. Thus, it is likely that the relationship between motor unit recruitment and firing rate of the muscle is the main determinant of corticospinal output with variations in nervous system gain.

Differences in force normalising procedures during submaximal anisometric contractions.

Eccentric contractions are thought to require a unique neural activation strategy. However, due to greater intrinsic force generating capacity of muscle fibres during eccentric contraction, the understanding of neural modulation of different contraction types during submaximal contractions may be impeded by the force normalisation procedure employed. In the present experiment, subjects performed maximal isometric dorsiflexion at shorter (80°), intermediate (90°) and longer (100°) muscle lengths, and maximal concentric and eccentric contractions. Thereafter, submaximal concentric and eccentric contractions were performed normalised to either isometric maximum at 90° (ISO), contraction type specific maximum (CTS) or muscle length specific maximum (MLS). When using ISO or MLS for normalisation, mean submaximal eccentric torque levels were significantly lower when compared to CTS normalisation (11 and 7% lower compared to CTS; p = 0.003 and p = 0.018 for ISO and MLS, respectively). These experimentally observed differences closely matched those expected from the predictive model. During submaximal concentric contraction, mean torque levels were similar between ISO and CTS normalisation with similar discrepancies noted in EMG activity. These findings suggest that normalising to ISO and MLS might not be accurate for assessment and prescription of submaximal eccentric contractions.

Muscle spindle and fusimotor activity in locomotion.

Mammals may exhibit different forms of locomotion even within a species. A particular form of locomotion (e.g. walk, run, bound) appears to be selected by supraspinal commands, but the precise pattern, i.e. phasing of limbs and muscles, is generated within the spinal cord by so-called central pattern generators. Peripheral sense organs, particularly the muscle spindle, play a crucial role in modulating the central pattern generator output. In turn, the feedback from muscle spindles is itself modulated by static and dynamic fusimotor (gamma) neurons. The activity of muscle spindle afferents and fusimotor neurons during locomotion in the cat is reviewed here. There is evidence for some alpha-gamma co-activation during locomotion involving static gamma motoneurons. However, both static and dynamic gamma motoneurons show patterns of modulation that are distinct from alpha motoneuron activity. It has been proposed that static gamma activity may drive muscle spindle secondary endings to signal the intended movement to the central nervous system. Dynamic gamma motoneuron drive appears to prime muscle spindle primary endings to signal transitions in phase of the locomotor cycle. These findings come largely from reduced animal preparations (decerebrate) and require confirmation in freely moving intact animals.

Effect of mental fatigue on induced tremor in human knee extensors.

In this study, the effects of mental fatigue on mechanically induced tremor at both a low (3-6Hz) and high (8-12Hz) frequency were investigated. The two distinct tremor frequencies were evoked using two springs of different stiffness, during 20s sustained contractions of the knee extensor muscles at 30% maximum voluntary contraction (MVC) before and after 100min of a mental fatigue task, in 12 healthy (29±3.7years) participants. Mental fatigue resulted in a 6.9% decrease in MVC and in a 9.4% decrease in the amplitude of the agonist muscle EMG during sustained 30% MVC contractions in the induced high frequency only. Following the mental fatigue task, the coefficient of variation and standard deviation of the force signal decreased at 8-12Hz induced tremor by 31.7% and 35.2% respectively, but not at 3-6Hz induced tremor. Similarly, the maximum value and area underneath the peak in the power spectrum of the force signal decreased by 55.5% and 53.1% respectively in the 8-12Hz range only. In conclusion, mental fatigue decreased mechanically induced 8-12Hz tremor and had no effect on induced 3-6Hz tremor. We suggest that the reduction could be attributed to the decreased activation of the agonist muscles.

The effects of isometric resistance training on stretch reflex induced tremor in the knee extensor muscles.

This study examines the effect of 4 wk of high-intensity isometric resistance training on induced tremor in knee extensor muscles. Fourteen healthy volunteers were assigned to either the training group (n = 7) or the nontraining control group (n = 7). Induced tremor was assessed by measuring force fluctuations during anisometric contractions against spring loading, whose compliance was varied to allow for preferential activation of the short or long latency stretch reflex components. Effects of high-intensity isometric resistance training on induced tremor was assessed under two contraction conditions: relative force matching, where the relative level of activity was equal for both pre- and post-training sessions, set at 30% maximum voluntary contraction (MVC), and absolute force matching, where the level of activity was set to 30% pretrained MVC. The training group experienced a 26.5% increase in MVC in contrast to the 0.8% for the control group. For relative force-matching contractions, induced tremor amplitude and frequency did not change in either the training or control group. During absolute force-matching contractions, induced tremor amplitude was decreased by 37.5% and 31.6% for the short and long components, respectively, with no accompanying change in frequency, for the training group. No change in either measure was observed in the control group for absolute force-matching contractions. The results are consistent with high-intensity isometric resistance training induced neural changes leading to increased strength, coupled with realignment of stretch reflex automatic gain compensation to the new maximal force output. Also, previous reported reductions in anisometric tremor following strength training may partly be due to changed stretch reflex behavior.

Role of the fusimotor system in locomotion.

The contribution of muscle spindles to the control of locomotion depends on the patterns of discharge that occur in static and dynamic gamma motoneurones (gammaS and gammaD). Discharges of gamma-axons to the MG muscle were studied during treadmill locomotion in pre-mammillary, decerebrated cats. All gammaS-efferents increased their rate of discharge at onset of locomotion and were modulated with movement. Type-1 increased their rate with muscle shortening whereas type-2 gammaS efferents increased their discharge rate during muscle lengthening. The type-1 gammaS pattern appears to be a temporal template of the intended movement. The type-2 gammaS pattern may be appropriate for afferent biasing through bag2 intrafusal fibres. The gammaD axons showed an interrupted discharge pattern with sudden onset of firing at the start of muscle shortening and falling quiet shortly after the beginning of lengthening. The gammaD discharge would prepare primary endings to respond with high sensitivity at the start of muscle lengthening.