Heteronymous reflex connections in human upper limb muscles in response to stretch of forearm muscles.

Torque motor produced stretch of upper limb muscles results in two distinct reflex peaks in the electromyographic activity. Whereas the short-latency reflex (SLR) response is mediated largely by the spinal monosynaptic reflex pathway, the longer-latency reflex (LLR) is suggested to involve a transcortical loop. For the SLRs, patterns of heteronymous monosynaptic Ia connections have been well-studied for a large number of muscles in the cat and in humans. For LLRs, information is available for perturbations to proximal joints, although the protocols for most of these studies did not focus on heteronymous connections. The main objective of the present study was to elicit both SLRs and LLRs in wrist flexors and extensors and to examine heteronymous connections from these muscles to elbow flexors (biceps brachii; BiBr) and extensors (triceps brachii; TriBr) and to selected distal muscles, including abductor pollicis longus (APL), first dorsal interosseous (FDI), abductor digiti minimi (ADM), and Thenars. The stretch of wrist flexors produced SLR and LLR peaks in APL, FDI, ADM, Thenars, and BiBr while simultaneously inducing inhibition of wrist extensors and TriBr. When wrist extensors were stretched, SLR and LLR peaks were observed in TriBr, whereas the primary wrist flexors, APL and BiBr, were inhibited; response patterns of FDI, ADM, and Thenars were less consistent. The main conclusions from the observed data are that: 1) as in the cat, afferents from wrist flexors and extensors make heteronymous connections with proximal and distal upper limb muscles; and 2) the strength of heteronymous connections is greater for LLRs than SLRs in the distal muscles, whereas the opposite is true for the proximal muscles. In the majority of observations, SLR and LLR excitatory peaks were observed together. However, on occasion, LLRs were observed without the SLR response in hand muscles when wrist extensors were stretched.

Motor unit rotation in a variety of human muscles.

The phenomena of substitution and rotation among motor units of a muscle were examined in seven different muscles. Intramuscular motor unit activity and surface electromyographic (EMG) activity were recorded from one of the following muscles: abductor digiti minimi, first dorsal interosseous, extensor digitorum communis, flexor and extensor carpi radialis, tibialis anterior, and soleus. The subject was asked to discharge a discernible unit at a comfortable constant or rhythmically (pseudosinusoidally) modulated rate with audio and visual feedback. Results are reported from a total of 42 sets of motor units from all seven muscles. We observed that when a subject fired a motor unit for a long period, an additional motor unit frequently started to discharge after a few minutes. When the subject was asked to keep activity down to one unit, very often it was Unit 1 that dropped and Unit 2 continued to fire. Whereas Unit 2 had fired for a few minutes, Unit 1 resumed firing without any conscious effort by the subject. If the subject was then asked to retain just one unit, it was Unit 2 that dropped. Rhythmic modulation of firing rate of a tonically firing unit showed that whereas the threshold of this unit increased, the threshold of a phasically discharging unit decreased substantially. The increase in threshold of a tonically discharging unit is suggested to arise from inactivation of Na(+) and Ca(2+) channels and the decrease in threshold of higher-threshold units is suggested to arise from an increase in persistent inward currents that may occur during prolonged contractions. Whether a unit stops or starts to fire is suggested to depend on a balance between the strength of the central motor command, persistent inward currents, and inactivation of voltage-gated channels. Such rotations among low-threshold motoneurons would ensure low-level sustained contractions to be viable not only in small hand muscles but also in larger limb muscles.

Responses of human motoneurons to high-frequency stimulation of Ia afferents.

This study was designed to extend to humans the findings of classical studies on anesthetized cats, which have examined the discharge of spinal motoneurons in response to high-frequency stimulus trains delivered to Ia afferents. Experiments were conducted on the monosynaptic pathway in the flexor carpi radialis (FCR) and soleus muscles. Subjects maintained a rhythmic discharge of a single motor unit (SMU) in either the FCR or soleus while homonymous Ia afferents were stimulated with either a single- or multipulse train. An n@IPI stimulus train had n pulses (n = 2-4) and an interpulse interval (IPI) of 1-8 ms. For each condition and motor unit, surface electromyographic (EMG) activity was averaged, and peristimulus-time histograms (PSTHs) were constructed for the SMU. The magnitude of the EMG was high for IPI = 1 ms, low for IPI = 2-3 ms, and high for IPI = 4-8 ms. SMU responses showed a similar pattern, which indicated that the increased EMG response was due to the presence of multiple peaks in a PSTH. The key results indicate that: (1) a short, high-frequency stimulus train enhances the discharge probability of a motoneuron above that observed with a single pulse; and (2) the increased motoneuron responses are significantly greater for the FCR than for the soleus muscle.

Refractoriness of sural nerve in humans.

High-frequency stimulation of peripheral nerve bundles is frequently used in clinical tests and physiologic experiments to study presynaptic and postsynaptic effects. To understand the postsynaptic effects, it is important to ensure that each pulse in the train is equally effective in stimulating the presynaptic nerve bundle; however, the optimal interpulse interval (IPI) and the stimulus intensity at which each pulse is equally effective in stimulating the same number of axons are not known. The magnitude of the compound action potential produced by each pulse in a train was tested on the sural nerve of 4 healthy human subjects. The stimulus train (2-4 pulses) was applied to the sural nerve at the lateral malleolus, and neural responses were recorded from just below the knee. With 2-pulse trains, families of curves between IPIs (1-6 ms) and normalized amplitudes of the second response were plotted for different stimulus intensities. Visual inspection of the data showed that the curves fell into 2 groups: with stimulus intensities <2.5x perception threshold (Th), the test response appeared partially at longer IPIs, whereas with stimulus intensities > or = 3x Th, partial recovery of the test response was earlier. The interval for complete recovery was statistically the same for low- and high-intensity stimulation. With more than 2 pulses in a stimulus train (IPI = 5 ms), the amplitude of the compound action potential (CAP) was not affected significantly. These results are important in understanding both the presynaptic and postsynaptic responses when presynaptic axon bundles are stimulated at high frequencies.

Rotation of motoneurons during prolonged isometric contractions in humans.

Prolonged and weak isometric contractions can result in neuromuscular fatigue. Alternation of discharge of motor units with similar thresholds (termed rotation) could be useful to minimize neuromuscular fatigue by providing periods for metabolic recovery of the contractile elements. In the present study, we investigated the prevalence of motoneuron rotation during prolonged contractions of distal limb muscles. Electromyographic (EMG; needle and surface) was recorded from muscles of the forearm and distal leg. The subject made a slowly increasing isometric contraction to recruit and discharge a motor unit (1) for a prolonged period of time (> 30 min). Sometimes an additional motor unit (2) was recruited in which case the subject relaxed the contraction slightly so that only one motor unit remained tonic. Often it was this newly recruited motor unit (i.e., unit 2) that continued discharging, while motor unit 1 fell silent. Continued contraction would then lead to the resumption of tonic discharge of unit 1 and silence of unit 2. This would complete a rotation between motor units 1 and 2. During prolonged contractions, rotation was observed in approximately 80% of the motor-unit pairs examined. There was no difference in rotation incidence by muscle type. For the unit pairs showing rotation, surface EMG values were significantly higher immediately prior to rotation than after rotation had occurred. Our findings show that rotation of motor units with similar recruitment thresholds during such contractions is common in distal muscles of the arm and leg and may help offset neuromuscular fatigue.

Rate-coding of spinal motoneurons with high-frequency magnetic stimulation of human motor cortex.

Rate-coding in spinal motoneurons was studied using high-frequency magnetic stimulation of the human motor cortex. The subject made a weak contraction to cause rhythmic (i.e., tonic) discharge of a single motor unit in flexor (or extensor) carpi radialis or tibialis anterior, while the motor cortical representation of that muscle was stimulated with brief trains of pulses from a Pyramid stimulator (4 Magstim units connected by 3 BiStim modules). An "m@n" stimulus train consisted of m number of pulses (1-4), with an interpulse interval (IPI) of n ms (1-6). Peristimulus time histograms were constructed for each stimulus condition of a given motor unit, and related to the average rectified surface electromyography (EMG) from that muscle. Surface EMG responses showed markedly more facilitation than single-pulse stimulation, with increasing numbers of pulses in the train; responses also tended to increase in magnitude for the longer IPI values (4 and 6 ms) tested. Motor-unit response probability increased in a manner comparable to that of surface EMG. In particular, motoneurons frequently responded twice to a given stimulus train. In addition to recruitment of new motor units, the increased surface EMG responses were, in part, a direct consequence of short-term rate-coding within the tonically discharging motoneuron. Our results suggest that human corticomotoneurons are capable of reliably following high-frequency magnetic stimulation rates, and that this activity pattern is carried over to the spinal motoneuron, enabling it to discharge at extremely high rates for brief periods of time, a pattern known to be optimal for force generation at the onset of a muscle contraction.

Neural control of motor output: can training change it?

Henneman's size principle of motor unit recruitment and rate coding reduces fatigue, minimizes error in transfer of information from the nervous system, and produces smooth force output. Plasticity present at various sites of the motor system may change endurance, force, speed, or precision with training, but not the recruitment order.