Christine K. Thomas

Contractile properties of human thenar muscles paralyzed by spinal cord injury

The electrical and mechanical properties of paralyzed human thenar muscles were measured in response to supramaximal stimulation of the median nerve in individuals with chronic cervical spinal cord injury. These data were compared to those recorded from control muscles. Spontaneous motor unit activity was common in paralyzed muscles. There was significantly more variance in the twitch and tetanic forces, twitch/tetanus force ratios, twitch and tetanic half‐relaxation times, and the stimulus frequencies which generated half‐maximal force in paralyzed versus control muscles. Approximately half the paralyzed thenar muscles were significantly weaker than control muscles and their compound action potential amplitudes were reduced significantly. Paralyzed muscles had significantly higher twitch/tetanus force ratios. The mean stimulus frequency which generated half‐maximal force was also reduced significantly. Thus for rehabilitation purposes, lower stimulation rates are required to elicit any given submaximal force from chronically paralyzed thenar muscles.

Muscle fatigue induced by stimulation with and without doublets.

Muscles are usually stimulated by shocks delivered at some constant rate. However, human thenar motor units generate optimum force per pulse when excited by impulse trains that begin with one or two short interpulse intervals (“doublets”), followed by longer intervals. Our aim was to determine whether the rate of force and force–time integral reduction during fatigue of thenar muscles is influenced by an initial doublet, and/or the number of pulses per train. We first matched thenar force–time integral using two different pulse patterns, one of which began with a doublet. Fatigue induced by trains that contained a doublet resulted in slower rates of force and force–time integral reduction and smaller increases in half‐relaxation time than that evoked by bursts of 40‐HZ stimulation. When the force was measured in each protocol after equal numbers of pulses had been delivered, the force loss was still significantly less for pulse trains containing a doublet. These results have useful implications when designing stimulation to strengthen weak muscles or to drive paralyzed muscles.

Motor unit activation order during electrically evoked contractions of paralyzed or partially paralyzed muscles

The activation order of motor units during electrically evoked contractions of paralyzed or partially paralyzed thenar muscles was determined in seven subjects with chronic cervical spinal cord injury. The median nerve was stimulated percutaneously with pulses of graded intensity to produce increments in the compound electromyogram (EMG) and force. Each increment corresponded to the activation of another unit. The evoked unit EMG and force was obtained by digital subtraction. The thenar muscles had between 15 and 83 units (26 ± 19) that produced 114.3 ± 127.1 mN force (n = 290). In six subjects, a significant positive correlation was found between activation order and unit force indicating that weaker units were excited before stronger units. These data are contrary to the notion that a reversal of unit activation order occurs during evoked versus voluntary contractions.

Fatigue of muscles weakened by death of motoneurons

Weakness is a characteristic of muscles influenced by the postpolio syndrome (PPS), amyotrophic lateral sclerosis (ALS), and spinal cord injury (SCI). The strength deficits relate to changes in muscle use and to the chronic denervation that can follow the spinal motoneuron death common to these disorders. PPS, ALS, and SCI also involve variable amounts of supraspinal neuron death, the effects of which on muscle weakness remains unclear. Nevertheless, weakness of muscle itself defines the functional consequences of these disorders. A weaker muscle requires an individual to work that muscle at higher than usual intensities relative to its maximal capacity, inducing progressive fatigue and an increased sense of effort. Little evidence is available to suggest that the fatigue commonly experienced by individuals with these disorders relates to an increase in the intrinsic fatigability of the muscle fibers. The only exception is when SCI induces chronic muscle paralysis. To reduce long‐term functional deficits in these disorders, studies must identify the signaling pathways that influence neuron survival and determine the factors that encourage and limit sprouting of motor axons. This may ensure that a greater proportion of the fibers in each muscle remain innervated and available for use. Muscle Nerve, 2005

Motor unit forces and recruitment patterns after cervical spinal cord injury.

Force was measured from triceps brachii motor units in individuals with chronic cervical spinal cord injury (SCI) and in able‐bodied (A‐B) control subjects using spike‐triggered averaging (175 and 48 units, respectively). Eleven percent of units from the SCI population generated normal electromyograms (EMGs) but exerted no measurable force, 65% generated force comparable to the control data, while 24% were stronger than usual. Weak units probably reflect disuse. Muscle shortening, densely innervated territories, and polyphasic EMG potentials suggested strong units resulted from intact axons sprouting to reinnervate denervated muscle. Many units from SCI subjects had faster than normal contraction times (CTs). The force and CT distributions from the SCI and A‐B populations differed significantly. Motor units of SCI subjects were recruited in order of increasing force output and increasing contraction time. Chronic cervical SCI therefore seems to alter the expected triceps brachii motor unit force‐speed relations.

The role of motor unit rate modulation versus recruitment in repeated submaximal voluntary contractions performed by control and spinal cord injured subjects

The relative roles of motor unit firing rate modulation and recruitment were evaluated when individuals with cervical spinal cord injury (SCI) and able-bodied controls performed a brief (6 s), 50% maximal voluntary contraction (50% MVC; target contraction) of triceps brachii every 10 s until it required maximal effort to achieve the target force. Mean (+/-SD) endurance times for SCI and control subjects were 34+/-26 and 15+/-5 min, respectively, at which point significant reductions in maximal triceps force had occurred. Twitch occlusion analysis in controls indicated that force declines resulted largely from peripheral contractile failure. In SCI subjects, triceps surface EMG and motor unit potential amplitude declined in parallel suggesting failure at axon branch points and/or alterations in muscle membrane properties. The force of low threshold units, measured by spike-triggered averaging, declined in SCI but not control subjects, suggesting that higher threshold units fatigued in controls. Central fatigue was also obvious after SCI. Mean (+/-SD) MVC motor unit firing rates declined significantly with fatigue for control (24.6+/-7.1 to 17.3+/-5.1Hz), but not SCI subjects (25.9+/-12.7 to 20.1+/-9.7Hz). Unit firing rates were unchanged during target contractions for each subject group, but with the MVC rate decreases, units of SCI and control subjects were activated intensely at endurance time (88% and 99% MVC rates, respectively). New unit recruitment also maintained the target contractions although it was limited after SCI because many descending inputs to triceps motoneurons were disrupted. This resulted in sparse EMG, even during MVCs, but allowed the same unit to be recorded throughout. These EMG data showed that both unit recruitment and rate modulation were important for maintaining force during repeated submaximal intermittent contractions of triceps brachii muscles performed by SCI subjects. Similar results were found for control subjects. Muscles weakened by SCI may therefore provide a useful model in which to directly study motor unit rate modulation and recruitment during weak or strong voluntary contractions.

Firing rates of motor units during strong dynamic contractions.

Muscle behavior is usually studied during isometric contractions but many tasks include contractions that involve changes in muscle length. Our aim was to record motor unit action potentials and surface electromyograms (EMGs) from triceps brachii muscles during rhythmic dynamic contractions (3‐s concentric, 3‐s eccentric; 40°/s; four subjects) performed at the highest voluntary forces subjects could exert (maximal concentric contraction) and at various submaximal intensities. Mean unit firing rates and surface EMG increased significantly with contraction intensity in both concentric and eccentric contractions, but at each intensity mean concentric values were significantly higher than eccentric values. In contrast, mean unit firing rates and surface EMGs were similar during maximal concentric and maximal isometric contractions. These data suggest muscles were activated maximally during the strongest concentric contractions but submaximally during the strongest eccentric efforts. After estimated eccentric contraction intensity was adjusted using surface EMG data, mean unit firing rates during eccentric contractions were still lower than the concentric values. Thus, protective mechanisms may limit motor unit firing rates during forceful lengthening contractions to minimize damage. Muscle Nerve, 2005

Fatigue in human thenar muscles paralysed by spinal cord injury

Muscle fatigue (force loss) induced by constant frequency stimulation (36 Hz) and variable rate stimulation (36 Hz to 18 Hz over 60 s) were compared in six individuals with thenar muscles which were paralysed by chronic cervical spinal cord injury (SCI), and in six volunteers with no known neurological disorder. The variable stimulation rate pattern represented the general decline in thenar motor unit firing rates recorded during 60 s maximum voluntary contractions performed by the able-bodied (AB) subjects. Constant and variable rate stimulation produced similar resultant force declines, as measured from abduction and flexion force components. However, significant force loss always occurred earlier and was of greater magnitude in SCI subjects, irrespective of the stimulation pattern (all, P < 0.01). Because more force was generally lost in one force component versus the other, the direction of the resultant force could also change with fatigue. The recordings from SCI participants were also contaminated by spontaneous motor unit activity, spasms and F responses. The stimulation frequency needed to produce half-maximum tetanic force increased for SCI subjects after fatigue, so higher, not lower stimulation frequencies were needed to produce any given submaximal force. Therefore, to match stimulation rate to changes in muscle contractile properties, these parameters have to be monitored and controlled on-line. The fatigue during each stimulation protocol, and for each subject population, was attributed primarily to contractile failure because any decrements in M-wave amplitude or area recovered completely within the first minute whereas twitch and tetanic forces remained somewhat depressed.

Understanding Therapeutic Benefits of Overground Bionic Ambulation: Exploratory Case Series in Persons With Chronic, Complete Spinal Cord Injury

OBJECTIVE To explore responses to overground bionic ambulation (OBA) training from an interdisciplinary perspective including key components of neuromuscular activation, exercise conditioning, mobility capacity, and neuropathic pain. DESIGN Case series. SETTING Academic research center. PARTICIPANTS Persons (N=3; 2 men, 1 woman) aged 26 to 38 years with complete spinal cord injury (SCI) (American Spinal Injury Association Impairment Scale grade A) between the levels of T1 and T10 for ≥1 year. INTERVENTION OBA 3d/wk for 6 weeks. MAIN OUTCOME MEASURES To obtain a comprehensive understanding of responses to OBA, an array of measures were obtained while walking in the device, including walking speeds and distances, energy expenditure, exercise conditioning effects, and neuromuscular and cortical activity patterns. Changes in spasticity and pain severity related to OBA use were also assessed. RESULTS With training, participants were able to achieve walking speeds and distances in the OBA device similar to those observed in persons with motor-incomplete SCI (10-m walk speed, .11-.33m/s; 2-min walk distance, 11-33m). The energy expenditure required for OBA was similar to walking in persons without disability (ie, 25%-41% of peak oxygen consumption). Subjects with lower soleus reflex excitability walked longer during training, but there was no change in the level or amount of muscle activity with training. There was no change in cortical activity patterns. Exercise conditioning effects were small or nonexistent. However, all participants reported an average reduction in pain severity over the study period ranging between -1.3 and 1.7 on a 0-to-6 numeric rating scale. CONCLUSIONS OBA training improved mobility in the OBA device without significant changes in exercise conditioning or in neuromuscular or cortical activity. However, pain severity was reduced and no severe adverse events were encountered during training. OBA therefore opens the possibility to reduce the common consequences of chronic, complete SCI such as reduced functional mobility and neuropathic pain.

Properties of medial gastrocnemius motor units and muscle fibers reinnervated by embryonic ventral spinal cord cells

Severe muscle atrophy occurs after complete denervation. Here, Embryonic Day 14-15 ventral spinal cord cells were transplanted into the distal tibial nerve stump of adult female Fischer rats to provide a source of neurons for muscle reinnervation. Our aim was to characterize the properties of the reinnervated motor units and muscle fibers. Some reinnervated motor units contracted spontaneously. Electrical stimulation of the transplants at increasing intensity produced an average (+/- SE) of 7 +/- 1 electromyographic and force steps. Each signal increment represented the excitation of another motor unit. These reinnervated units exerted an average force of 12.0 +/- 1.5 mN, strength similar to that of control fatigue-resistant units. Repeated transplant stimulation depleted 17% of the muscle fibers of glycogen, an indication of some functional reinnervation. Reinnervated (glycogen-depleted), denervated (no cells transplanted), and control fibers were of histochemical type I, IIA, or IIB. Fibers of the same type were grouped after reinnervation. The proportion of fiber types also changed. Reinnervated fibers were primarily type IIA, whereas most fibers in denervated and control muscles were type IIB. Reinnervated fibers of each type had significantly larger cross-sectional areas than the corresponding fiber types in denervated muscles. These data suggest that neurons with different properties can reside in the unusual environment of the adult rat peripheral nerve, make functional connections with muscle, specify muscle fiber type, and reduce the amount that each type atrophies.