Recruitment Of Motoneurons Research Articles

Principles governing recruitment of motoneurons during swimming in zebrafish

Locomotor movements are coordinated by a network of neurons that produces sequential muscle activation. Different motoneurons need to be recruited in an orderly manner to generate movement with appropriate speed and force. However, the mechanisms governing recruitment order have not been fully clarified. Using an in vitro juvenile/adult zebrafish brainstem-spinal cord preparation, we found that motoneurons were organized into four pools with specific topographic locations and were incrementally recruited to produce swimming at different frequencies. The threshold of recruitment was not dictated by the input resistance of motoneurons, but was instead set by a combination of specific biophysical properties and the strength of the synaptic currents. Our results provide insights into the cellular and synaptic computations governing recruitment of motoneurons during locomotion.

Monosynaptic Rabies Virus Reveals Premotor Network Organization and Synaptic Specificity of Cholinergic Partition Cells

Movement is the behavioral output of neuronal activity in the spinal cord. Motor neurons are grouped into motor neuron pools, the functional units innervating individual muscles. Here we establish an anatomical rabies virus-based connectivity assay in early postnatal mice. We employ it to study the connectivity scheme of premotor neurons, the neuronal cohorts monosynaptically connected to motor neurons, unveiling three aspects of organization. First, motor neuron pools are connected to segmentally widely distributed yet stereotypic interneuron populations, differing for pools innervating functionally distinct muscles. Second, depending on subpopulation identity, interneurons take on local or segmentally distributed positions. Third, cholinergic partition cells involved in the regulation of motor neuron excitability segregate into ipsilaterally and bilaterally projecting populations, the latter exhibiting preferential connections to functionally equivalent motor neuron pools bilaterally. Our study visualizes the widespread yet precise nature of the connectivity matrix for premotor interneurons and reveals exquisite synaptic specificity for bilaterally projecting cholinergic partition cells.

Influence of stimulus pulse width on M‐waves, H‐reflexes, and torque during tetanic low‐intensity neuromuscular stimulation

Neuromuscular electrical stimulation (NMES) has been shown to generate contractions that include a central recruitment of motoneurons; however, the effect of pulse width on electromyographic (EMG) and torque responses during NMES are not well documented. Soleus EMG and isometric plantarflexion torque were recorded from 14 subjects with NMES delivered to the tibial nerve using 50, 200, 500, and 1000 μs pulse widths. M-waves were significantly smaller during 20 Hz NMES compared with responses evoked by single pulses of 200, 500, and 1000 μs, but not 50 μs pulse widths. At all pulse widths, stimulation at 20 Hz depressed soleus H-reflexes compared with single pulses. Two seconds of 100 Hz NMES significantly increased H-reflexes and torque during the subsequent 20 Hz NMES with 200, 500, and 1000 μs, but not 50 μs, pulse widths. NMES delivered using wide pulses generated larger contractions with a relatively greater central contribution than narrow pulses. This may help reduce atrophy and produce fatigue-resistant contractions for rehabilitation.

*Poster 181: Motor Control by Shifting Spatial Reference Frames for Motoneuronal Recruitment

*Poster 181: Motor Control by Shifting Spatial Reference Frames for Motoneuronal Recruitment

Assessment of the effect of L-glutamine supplementation on DOMS

Glutamine is an amino acid and is not considered one of the eight essential aminos. Amino acid supplementation outspreaded to enhancing athletic performance, preparation, removal fatigue and minimising risk of...

Role of Intrinsic Properties in Drosophila Motoneuron Recruitment During Fictive Crawling

Motoneurons in most organisms conserve a division into low-threshold and high-threshold types that are responsible for generating powerful and precise movements. Drosophila 1b and 1s motoneurons may be analogous to low-threshold and high-threshold neurons, respectively, based on data obtained at the neuromuscular junction, although there is little information available on intrinsic properties or recruitment during behavior. Therefore in situ whole cell patch-clamp recordings were used to compare parameters of 1b and 1s motoneurons in Drosophila larvae. We find that resting membrane potential, voltage threshold, and delay-to-spike distinguish 1b from 1s motoneurons. The longer delay-to-spike in 1s motoneurons is a result of the shal-encoded A-type K(+) current. Functional differences between 1b and 1s motoneurons are behaviorally relevant because a higher threshold and longer delay-to-spike are observed in MNISN-1s in pairwise whole cell recordings of synaptically evoked activity during bouts of fictive locomotion.

Vibration-induced extra torque during electrically-evoked contractions of the human calf muscles

BackgroundHigh-frequency trains of electrical stimulation applied over the lower limb muscles can generate forces higher than would be expected from a peripheral mechanism (i.e. by direct activation of motor axons). This phenomenon is presumably originated within the central nervous system by synaptic input from Ia afferents to motoneurons and is consistent with the development of plateau potentials. The first objective of this work was to investigate if vibration (sinusoidal or random) applied to the Achilles tendon is also able to generate large magnitude extra torques in the triceps surae muscle group. The second objective was to verify if the extra torques that were found were accompanied by increases in motoneuron excitability.MethodsSubjects (n = 6) were seated on a chair and the right foot was strapped to a pedal attached to a torque meter. The isometric ankle torque was measured in response to different patterns of coupled electrical (20-Hz, rectangular 1-ms pulses) and mechanical stimuli (either 100-Hz sinusoid or gaussian white noise) applied to the triceps surae muscle group. In an additional investigation, Mmax and F-waves were elicited at different times before or after the vibratory stimulation.ResultsThe vibratory bursts could generate substantial self-sustained extra torques, either with or without the background 20-Hz electrical stimulation applied simultaneously with the vibration. The extra torque generation was accompanied by increased motoneuron excitability, since an increase in the peak-to-peak amplitude of soleus F waves was observed. The delivery of electrical stimulation following the vibration was essential to keep the maintained extra torques and increased F-waves.ConclusionsThese results show that vibratory stimuli applied with a background electrical stimulation generate considerable force levels (up to about 50% MVC) due to the spinal recruitment of motoneurons. The association of vibration and electrical stimulation could be beneficial for many therapeutic interventions and vibration-based exercise programs. The command for the vibration-induced extra torques presumably activates spinal motoneurons following the size principle, which is a desirable feature for stimulation paradigms.

Control of wrist position and muscle relaxation by shifting spatial frames of reference for motoneuronal recruitment: possible involvement of corticospinal pathways.

It has previously been established that muscles become active in response to deviations from a threshold (referent) position of the body or its segments, and that intentional motor actions result from central shifts in the referent position. We tested the hypothesis that corticospinal pathways are involved in threshold position control during intentional changes in the wrist position in humans. Subjects moved the wrist from an initial extended to a final flexed position (and vice versa). Passive wrist muscle forces were compensated with a torque motor such that wrist muscle activity was equalized at the two positions. It appeared that motoneuronal excitability tested by brief muscle stretches was also similar at these positions. Responses to mechanical perturbations before and after movement showed that the wrist threshold position was reset when voluntary changes in the joint angle were made. Although the excitability of motoneurons was similar at the two positions, the same transcranial magnetic stimulus (TMS) elicited a wrist extensor jerk in the extension position and a flexor jerk in the flexion position. Extensor motor-evoked potentials (MEPs) elicited by TMS at the wrist extension position were substantially bigger compared to those at the flexion position and vice versa for flexor MEPs. MEPs were substantially reduced when subjects fully relaxed wrist muscles and the wrist was held passively in each position. Results suggest that the corticospinal pathway, possibly with other descending pathways, participates in threshold position control, a process that pre-determines the spatial frame of reference in which the neuromuscular periphery is constrained to work. This control strategy would underlie not only intentional changes in the joint position, but also muscle relaxation. The notion that the motor cortex may control motor actions by shifting spatial frames of reference opens a new avenue in the analysis and understanding of brain function.

Changes in motor unit recruitment strategy during pain alters force direction

Motor unit (MU) recruitment is altered (decreased discharge rate and cessation of discharge in some units, and recruitment of new units) in force-matched contractions during pain compared to contractions performed before pain. As MU’s within a motoneurone pool have different force direction properties we hypothesised that altered MU recruitment during experimental knee pain would change the force vector (total force ( F T): amplitude and angle) generated by the quadriceps. Force was produced at two levels during 1 × 60-s and 3 × 10-s isometric contractions of knee extensors, and recorded by two force transducers at right angles. This enabled calculation of both F E (extension force) and F T. MU recruitment was recorded from the medial and lateral vastii with four fine-wire electrodes. Pain was induced by hypertonic saline injection in the infra-patella fat pad. Nine subjects matched F E and six subjects also matched both medial and lateral forces ( F T) before and during pain. Changes in MU discharge pattern (decreased discharge rate ( P < 0.001), complete cessation of firing, and recruitment of new units) during pain were associated with a ∼5° change in absolute force angle. As force angle changed in both directions (left/right) for individual subjects with pain there was no change in average F T amplitude between conditions. When both medial and lateral forces were matched MU discharge rate decreased ( P < 0.001) with pain, but, fewer units ceased firing or were newly recruited during pain. Change in motoneurone recruitment during pain alters direction of muscle force. This may be a strategy to avoid pain or protect the painful part.

The estimation of short intra-cortical inhibition depends on the proportion of spinal motoneurones activated by corticospinal inputs

The high variability of SICI limits its utility and by extension that of TMS in clinical neurophysiology. Non-linear summation of descending volleys due to heterogeneous motoneurone properties, on which MEP size depends, has not previously been implicated as an issue in SICI evaluation. MEP size and SICI were normalised to the test MEP (mV), and as a percentage of M(max) to take account of the proportion of motoneurone pool activated by TMS. Two EMG systems, producing large and small MEPs, were used to determine how the normalisation affected MEPs of different amplitude. M(max) normalisation (i) counteracted the influence of recording conditions on the MEP size, (ii) revealed a significant influence of the test size on SICI (between medium and large MEPs), and of test size on the conditioning intensity (the larger the MEP, the stronger the SICI), and (iii) decreased the variability. Data normalised to M(max) better reflect the motoneurone recruitment after SICI. To enhance reproducibility, MEP should be normalised to M(max). This adjusts for some of the non-linear properties at the spinal, and possibly, at cortical levels. To reduce variability is important because TMS is becoming widely adopted and is being used in patients.

Amphetamine Increases Persistent Inward Currents in Human Motoneurons Estimated From Paired Motor-Unit Activity

Recruitment and repetitive firing of spinal motoneurons depend on the activation of persistent inward calcium and sodium currents (PICs) that are in turn facilitated by serotonin and norepinephrine that arise primarily from the brain stem. Considering that in rats motoneuron PICs are greatly facilitated by increasing the presynaptic release of norepinephrine with amphetamine, we sought similar evidence for the modulation of PICs in human motoneurons. Pairs of motor units were recorded during a gradually increasing and then decreasing voluntary contraction. The firing frequency (F) of the lower-threshold (control) motor unit was used as an estimate of the synaptic input to the higher-threshold (test) motor unit. Generally, PICs are initiated during the recruitment of a motoneuron and subsequently provide a fixed depolarizing current that helps the synaptic input maintain firing until derecruitment. Thus the amplitude of the PIC in the test motor unit was estimated from the difference in synaptic input (DeltaF) needed to maintain minimal firing once the PIC was fully activated (measured at the time of test unit derecruitment) compared with the larger synaptic input required to initiate firing prior to full PIC activation (measured at the time of test unit recruitment; DeltaF = F(recruit) - F(derecruit)). Moreover, the activation time of the PIC was estimated as the minimal contraction duration needed to produce a maximal PIC (DeltaF). In five subjects, oral administration of amphetamine, but not placebo, increased the DeltaF by 62% [from 3.7 +/- 0.6 to 6.0 +/- 0.8 (SD) imp/s, P = 0.001] and decreased the time needed to activate a maximal DeltaF from approximately 2 to 0.5 s. Both findings suggest that the endogenous facilitation of PICs from brain stem derived norepinephrine plays an important role in modulating human motoneuron excitability, readying motoneurons for rapid and sustained activity during periods of high arousal such as stress or fear.

Modulation of Motor Cortical Excitability Induced by Combined Visual Attention to Guide Finger Movement

Objective: To evaluate the effect of multiple sensory stimulation on cortical excitability by using transcranial magnetic stimulation (TMS). Method: Thirteen right handed young adults without neurological deficit were enrolled. Cortical excitability was tested by measuring recruitment of motor evoked potentials [recruitment curve (RC)], intracortical inhibition (ICI), and intracortical facilitation (ICF) at the abductor pollicis brevis of the dominant hand in two different conditions: (1) group A: active thumb and index finger pinch movement while observing the congruent finger movement in a screen with 0.2 Hz bell sound for 30 minutes, (2) group B: the same active finger movement in a dark screen with the same bell sound for 30 minutes. All of these procedures were done with a randomized crossover design. Results: The amplitude of MEP and the slope of all RC (140%,160% of the resting motor threshold) of group A showed increment after visually-guided finger movement and the level of ICI showed decrement after visually guided finger movement (p 0.05) in group A. In group B, the amplitude of MEP and the levels of ICI, ICF showed no significant changes following finger movement with no visual guidance (p>0.05), but the slope of RC with 140% showed increment (p<0.05). Conclusion: These findings suggest that cortical excitability can be enhanced by simple repetitive motor practice. The congruent sensori-motor stimulations lead to corresponding additional effect on cortical excitability, presumably by recruitment of remote motor neurons. (Brain & NeuroRehabilitation 2010; 3: 106-110)

Rank-ordered Recruitment of Masseter Motoneurons during Slow Closing Phase of Mastication Cycle Presumably by the Activity of Mesencephalic Trigeminal Neurons

Rank-ordered Recruitment of Masseter Motoneurons during Slow Closing Phase of Mastication Cycle Presumably by the Activity of Mesencephalic Trigeminal Neurons

Proliferative activity and motoneurone recruitment persist at the spinal cord central canal during larval and some postlarval stages in the rainbow trout (Oncorhynchus mykiss).

We previously found a linear relationship between the cross sectional myotomal area and the motoneurone number in the growing trout during postlarval stages. These neurones increased in number until a fish length of 150 mm, which prompted us to examine how motor neurones are recruited afterwards to meet the growth of their target myotomal muscle. Young adult (260 mm in length), fingerlings (F, 120-170 mm), fry (Fr, 70 mm) and eleutherembryos (Es, 20-30 mm) of rainbow trout (Oncorhyncus mykiss) were employed in this study. PCNA immunohistochemistry was used for monitoring the proliferative activity in the epithelium of the spinal cord central canal. This activity was quantified as the number of PCNA labelled cells for each spinal cord section. In Es and Fry, a mean value of 3-5 labelled cells for each section was found with a sharp decrease in young F (120 mm long). After this fish length, it was not possible to quantitatively evaluate the proliferative activity at the central canal. However, labelled cells were seldom found in the spinal cord sections until a fish length of 260 mm. From these data it is possible to conclude that motoneurone recruitment in the trout spinal cord is down-regulated at the F stage. Afterwards, we found that motoneurones increase in size to meet the growth of their target myotomal muscle.

Neuroprotective effect of testosterone treatment on motoneuron recruitment following the death of nearby motoneurons

Motoneuron loss is a significant medical problem, capable of causing severe movement disorders or even death. We have previously shown that motoneuron death induces marked dendritic atrophy in surviving nearby motoneurons. Additionally, in quadriceps motoneurons, this atrophy is accompanied by decreases in motor nerve activity. However, treatment with testosterone partially attenuates changes in both the morphology and activation of quadriceps motoneurons. Testosterone has an even larger neuroprotective effect on the morphology of motoneurons of the spinal nucleus of the bulbocavernosus (SNB), in which testosterone treatment can completely prevent dendritic atrophy. The present experiment was performed to determine whether the greater neuroprotective effect of testosterone on SNB motoneuron morphology was accompanied by a greater neuroprotective effect on motor activation. Right side SNB motoneurons were killed by intramuscular injection of cholera toxin-conjugated saporin in adult male Sprague-Dawley rats. Animals were either given Silastic testosterone implants or left untreated. Four weeks later, left side SNB motor activation was assessed with peripheral nerve recording. The death of right side SNB motoneurons resulted in several changes in the electrophysiological response properties of surviving left side SNB motoneurons, including decreased background activity, increased response latency, increased activity duration, and decreased motoneuron recruitment. Treatment with exogenous testosterone attenuated the increase in activity duration and completely prevented the decrease in motoneuron recruitment. These data provide a functional correlate to the known protective effects of testosterone treatment on the morphology of these motoneurons, and further support a role for testosterone as a therapeutic agent in the injured nervous system.

Spike synchronization is population specific in the respiratory pattern generator

Breathing accounts for only a small fraction of basalmetabolism and its motor pattern is highly efficientmechanically [1]. While this view is widely accepted, noone understands whether this property is malleable andwhether it can be utilized to attenuate respiratory failureas well as to facilitate weaning from artificial ventilation.Several factors influence muscle force including firing rateand recruitment of pre-motor and motoneurons (MNs),which increase energy consumption. Another factor is thesynchronization of spikes of MNs, which increases muscleforce in an energy efficient manner, preserving rather thanincreasing the firing frequency of active motor units. Inline with Otis's findings, we hypothesized that the respira-tory control network uses synchrony to generate an effi-cient inspiratory pattern, minimizing the work ofdiaphragm contraction, chest wall expansion and lunginflation.

Effect of a peripheral nerve block on torque produced by repetitive electrical stimulation

Neuromuscular electrical stimulation (NMES) generates contractions by activation of motor axons (peripheral mechanism), but the afferent volley also contributes by recruiting spinal motoneurons synaptically (central mechanism), which recruits motoneurons according to Henneman's size principle. Thus, we hypothesized that contractions that develop due to a combination of peripheral and central mechanisms will fatigue less rapidly than when electrically evoked contractions are generated by the activation of motor axons alone. Plantar-flexion torque evoked by NMES over the triceps surae was compared in five able-bodied subjects before (Intact) and during (Blocked) a complete anesthetic block of the tibial and common peroneal nerves. In the Blocked condition, plantar-flexion torque could only develop from the direct activation of motor axons beneath the stimulating electrodes. NMES was delivered using three protocols: protocol A, constant 100 Hz for 30 s; protocol B, four 2-s bursts of 100 Hz alternating with 20-Hz stimulation; and protocol C, alternating 100 Hz bursts (1 s on, 1 s off) for 30 s. The percent change in evoked plantar flexion torque from the beginning to the end of the stimulation differed (P < 0.05) between Intact and Blocked conditions for all protocols (Intact: protocol A = +125%, B = +230%, C = +78%; Blocked: protocol A = -79%, B = -15%, C = -35%). These results corroborate previous evidence that NMES can evoke contractions via the recruitment of spinal motoneurons in addition to the direct recruitment of motor axons. We now show that NMES delivered for periods of up to 30 s generates plantar-flexion torque which decreases when only motor axons are recruited and increases when the central nervous system can contribute.

Better functional outcome of compression spinal cord injury in mice is associated with enhanced H-reflex responses

Alterations in spinal reflexes and functional improvements occur after incomplete spinal cord injury but the relationship between these phenomena is not understood. Here we show that spontaneous functional recovery after compression injury of the spinal cord at low-thoracic level (Th10–12) in C57BL/6J mice is associated with a progressively increasing, over 3 months, excitability of the plantar H-reflex. The stimulation rate-sensitive H-reflex depression, already strongly reduced at 1 week after injury, when compared with non-injured mice, decreased further during the observation time period. Twelve weeks after injury, the degree of motor recovery estimated by single-frame motion analysis in individual animals correlated positively with their H-reflex responses at 2-Hz stimulation. Functional recovery and reflex alterations were accompanied by an increase in glycine/GABAergic and glutamatergic terminals around motoneuron cell bodies between 6 and 12 weeks after injury. Enhanced H-reflex responses at frequencies between 0.1 and 5 Hz were also observed in mice deficient in the extracellular matrix glycoprotein tenascin-R and the adhesion molecule close homolog of L1, mice previously shown to have better motor recovery after spinal cord injury than wild-type littermates. These results indicate that better functional outcome of compression spinal cord injury in mice is associated with alterations of the monosynaptic reflex pathway which facilitate motoneuron recruitment. Our observations support the view that plasticity of spinal circuitries underlies specific aspects of motor recovery and demonstrate the usefulness of H-reflex analyses in studies on spinal cord injury in mice.

Motoneurone recruitment is altered with pain induced in non-muscular tissue

Motoneurone discharge rate is reduced despite the maintenance of force when pain is induced via injection of hypertonic saline into muscle. Two aspects require consideration. First, hypertonic saline may have direct effects on axons other than small diameter pain fibres including the motoneurones that innervate the painful muscle. Second, it is unclear how force is maintained, when motoneurone discharge rate is decreased. We aimed to determine; (1) if motoneurone discharge rate is reduced during force-matched tasks when pain is induced in non-muscle tissue (to exclude direct effects on motoneurones) and (2) if the reduction of discharge rate is associated with additional changes in motoneurone recruitment over multiple muscle regions. Motoneurone discharge was recorded in the quadriceps with eight pairs of fine-wire electrodes. Seven subjects performed 30-s low-level, force-matched contractions before and during anterior knee pain, which was induced by a bolus (0.25ml) injection of 5% hypertonic saline into the infra-patellar fat pad. In total, 119 motor units were identified. Of these, 34 were identified both before and during pain. The discharge rate of these units decreased during pain from 8.9(1.5) to 7.2(1.4)Hz (P<0.0001). In addition, 31 units were recruited in the no-pain condition but not during pain, when 53 new units were recruited. These changes coincided with a large variability in gross muscle activity measures between muscle regions. These data confirm that motoneurone recruitment is altered when direct effects of saline on motoneurones are excluded. Recruitment of additional motor units may explain force maintenance despite reduced discharge rate of some units.

Continuous shifts in the active set of spinal interneurons during changes in locomotor speed

The classic ‘size principle’ of motor control describes how increasingly forceful movements arise by the recruitment of motoneurons of progressively larger size and force output into the active pool. Here, we explore the activity of pools of spinal interneurons in larval zebrafish and find that increases in swimming speed are not associated with the simple addition of cells to the active pool. Instead, the recruitment of interneurons at faster speeds is accompanied by the silencing of those driving movements at slower speeds. This silencing occurs both between and within classes of rhythmically-active premotor excitatory interneurons. Thus, unlike motoneurons, there is a continuous shift in the set of cells driving the behavior, even though changes in the speed of the movements and the frequency of the motor pattern appear smoothly graded. We conclude that fundamentally different principles may underlie the recruitment of motoneuron and interneuron pools.