A teleological explanation of Weber's law and the motor unit size law

It is shown that the Weber-Fechner law. which relates the response of a sensory biosystem to the intensity of the input stimulus, can be derived from a teleological principle of minimum transentropy (maximal noise reduction) provided the relative mean fluctuation (coefficient of variation) of the input intensity can be assumed to be (approximately) constant for all feasible mean input intensities. A law is then deduced from experimental results which quantifies the relationship existing between the relative amount of activated muscle mass and the “size” (which term is clearly defined) of a newly recruited motor unit. This law is found to be formally equivalent to the Weber-Fechner law when applied to motor unit recruitment. It is then shown that, in general, the ratio of the force increment upon recruitment, to the present force output does not obey Weber's law. Finally, it is proved that the “motor unit size law” as derived in this paper implies a fixed sequential order in the recruitment of motor units and that it may be viewed as the realization, by the mammalian neuromuscular system, of a general principle of maximum grading sensitivity.     

Role of reflex gain and reflex delay in spinal stability--a dynamic simulation

The goal of this study was to investigate the role of reflex and reflex time delay in muscle recruitment and spinal stability. A dynamic biomechanical model of the musculoskeletal spine with reflex response was implemented to investigate the relationship between reflex gain, co-contraction, and stability in the spine. The first aim of the study was to investigate how reflex gain affected co-contraction predicted in the model. It was found that reflexes allowed the model to stabilize with less antagonistic co-contraction and hence lower metabolic power than when limited to intrinsic stiffness alone. In fact, without reflexes there was no feasible recruitment pattern that could maintain spinal stability when the torso was loaded with 200N external load. Reflex delay is manifest in the paraspinal muscles and represents the time from a perturbation to the onset of reflex activation. The second aim of the study was to investigate the relationship between reflex delay and the maximum tolerable reflex gain. The maximum acceptable upper bound on reflex gain decreased logarithmically with reflex delay. Thus, increased reflex delay and reduced reflex gain requires greater antagonistic co-contraction to maintain spinal stability. Results of this study may help understanding of how patients with retarded reflex delay utilize reflex for stability, and may explain why some patients preferentially recruit more intrinsic stiffness than healthy subjects.     

Abdominal motor unit activity during respiratory and nonrespiratory tasks

Abdominalmuscles serve multiple roles, but the functional organization of theirmotoneurons remains unclear. To gain insight, we recorded single motorunit potentials from the internal oblique (IO) and transversusabdominis (TA) muscles of three standing subjects during quietbreathing, a leg lift, and an expiratory threshold load. Inspiratoryairflow, recorded from a pneumotachometer, provided tidal volumes andrespiratory cycle timing. Fine wires, implanted under ultrasonicimaging, detected single motor unit potentials that were visuallydistinguished by their spike morphology. From the number of spikes,firing profiles, times of occurrence in the respiratory cycle, andtheir onset, instantaneous, mean, and peak firing frequencies wededuced that 1) breathing patterns varied across tasks, 2) differentmotor units were recruited for each task with essentially no overlap,3) their firing displayed prominentexpiratory activity during each task, and4) the recruitment levels anddischarge patterns of IO and TA were different. We conclude that the IOand TA motor pools receive a strong central respiratory drive, yet eachpool receives its own distinct, task-dependent synapticinput.

     

The proprioceptive activation of stretch tension in pretibial flexor muscles induced by stimulation of antagonistic muscle afferents.

1. The effect of tetanic stimulation of ipsilateral group I afferents from the GS muscle on a synchronous stretch of the flexor EDL/TA muscles has been investigated in precollicular decerebrate cats. 2. The stretch-induced tension of the EDL/TA muscles increases remarkably during simultaneous stimulation of the GS nerve with maximal intensities corresponding to 1.5 times the threshold for the group 1 afferents. This increas appears above all in the dynamic part of stretch. 3. Under our experimental conditions there is no activation fo flexor-alpha-motoneurones during tetanic stimulation of the GS afferents without muscle stretch, as measured by the resting tension of the EDL/TA muscles. 4. Desptie an increase in the stretch-induced tension during fusimotor stimulation of antagonistic group I afferents, a transmission loss in the excitation via the psi-loop to the flexor-alpha-motoneurones occurs. This could be demonstrated by the ratio: increase in the tension T /increase in the number of Ia spikes. This is explained by snychronous convergence of the discharges of Ia inhibitory interneurones to the flexor-alpha-motoneurones. 5. The system studied demonstrates an input-output relation of the stretch reflex during conditions in which both reciprocal inhibition and autogenetic excitation via the psi-loop occur. It appears however, that the reciprocal inhibition is partly overwhelmed by the autogenetic excitation which results from the increase in the Ia discharge rate during fusimotor reflex. 6. It is postulated that static rather than dynamic psi-moto-neurones are involved in the investigated reflex arc.     

Age-related changes of the stretch reflex excitability in human ankle muscles

The purpose of this study was to characterize the effects of aging on the stretch reflex in the ankle muscles, and in particular to compare the effects on the ankle dorsi-flexor (tibialis anterior: TA) and the plantar-flexor (soleus: SOL). Stretch reflex responses were elicited in the TA and SOL at rest and during weak voluntary contractions in 20 elderly and 23 young volunteers. The results indicated that, in the TA muscle, the elderly group had a remarkably larger long-latency reflex (LLR), whereas no aging effect was found in the short latency reflex (SLR). These results were very different from those in the SOL muscle, which showed significant aging effects in the SLR and medium latency reflex (MLR), but not in the LLR. Given the fact that the LLR of the TA stretch reflex includes the cortical pathway, it is probable that the effects of aging on the TA stretch reflex involve alterations not only at the spinal level but also at the cortical level. The present results indicate that the stretch reflexes of each of the ankle antagonistic muscles are affected differently by aging, which might have relevance to the neural properties of each muscle.     

Torso flexion modulates stiffness and reflex response.

Neuromuscular factors that contribute to spinal stability include trunk stiffness from passive and active tissues as well as active feedback from reflex response in the paraspinal muscles. Trunk flexion postures are a recognized risk factor for occupational low-back pain and may influence these stabilizing control factors. Sixteen healthy adult subjects participated in an experiment to record trunk stiffness and paraspinal muscle reflex gain during voluntary isometric trunk extension exertions. The protocol was designed to achieve trunk flexion without concomitant influences of external gravitational moment, i.e., decouple the effects of trunk flexion posture from trunk moment. Systems identification analyses identified reflex gain by quantifying the relation between applied force disturbances and time-dependent EMG response in the lumbar paraspinal muscles. Trunk stiffness was characterized from a second order model describing the dynamic relation between the force disturbances versus the kinematic response of the torso. Trunk stiffness increased significantly with flexion angle and exertion level. This was attributed to passive tissue contributions to stiffness. Reflex gain declined significantly with trunk flexion angle but increased with exertion level. These trends were attributed to correlated changes in baseline EMG recruitment in the lumbar paraspinal muscles. Female subjects demonstrated greater reflex gain than males and the decline in reflex gain with flexion angle was greater in females than in males. Results reveal that torso flexion influences neuromuscular factors that control spinal stability and suggest that posture may contribute to the risk of instability injury.     

Proprioceptive proximo-distal relationships between the quadriceps and soleus muscles in man

Electrical stimulation of femoral nerve modulates voluntary tonic activity o of ipsilateral soleus muscle. Stimulus time-locked inhibitory and facilitatory phases can be distinguished. EMG temporal analysis suggests that early perturbations are correlated with spinal effects of centripetal electrical activity. The inhibitory effects which momentarily abolish voluntary soleus activity are thought to result from quadriceps Ib fibres recruitment. While no heteronymous activity is induced at rest, femoral nerve Ia fibres activation can produce soleus muscle reflex when soleus motor nucleus excitability is increased by voluntary command. Recurrent discharge resulting from soleus reflex response enhances inhibition initially due to quadriceps Ib volley. Secondary effects of isometric quadriceps contraction (and soleus contraction when the femoral stimulus elicits a reflex in this muscle) have their own effects later. These findings suggest that proprioceptive relationships of the two muscular groups are efficient during tonic isometric voluntary command.     

Motor unit firing patterns of lower leg muscles during isometric plantar flexion with flexed knee joint position

The ankle flexor and extensor muscles are essential for pedal movements associated with car driving. Neuromuscular activation of lower leg muscles is influenced by the posture during a given task, such as the flexed knee joint angle during car driving. This study aimed to investigate the influence of flexion of the knee joint on recruitment threshold-dependent motor unit activity in lower leg muscles during isometric contraction. Twenty healthy participants performed plantar flexor and dorsiflexor isometric ramp contractions at 30 % of the maximal voluntary contraction (MVC) with extended (0°) and flexed (130°) knee joint angles. High-density surface electromyograms were recorded from medial gastrocnemius (MG), soleus (SOL), and tibialis anterior (TA) muscles and decomposed to extract individual motor units. The torque-dependent change (Δpps /Δ%MVC) of the motor unit activity of MG (recruited at 15 %MVC) and SOL (recruited at 5 %MVC) muscles was higher with a flexed compared with an extended knee joint (p < 0.05). The torque-dependent change of TA MU did not different between the knee joint angles. The motor units within certain limited recruitment thresholds recruited to exert plantar flexion torque can be excited to compensate for the loss of MG muscle torque output with a flexed knee joint.     

Gravitational force modulates muscle activity during mechanical oscillation of the tibia in humans

Mechanical oscillation (vibration) is an osteogenic stimulus for bone in animal models and may hold promise as an anti-osteoporosis measure in humans with spinal cord injury (SCI). However, the level of reflex induced muscle contractions associated with various loads (g force) during limb segment oscillation is uncertain. The purpose of this study was to determine whether certain gravitational loads (g forces) at a fixed oscillation frequency (30 Hz) increases muscle reflex activity in individuals with and without SCI. Nine healthy subjects and two individuals with SCI sat with their hip and knee joints at 90° and the foot secured on an oscillation platform. Vertical mechanical oscillations were introduced at 0.3, 0.6, 1.2, 3 and 5g force for 20 s at 30 Hz. Non-SCI subjects received the oscillation with and without a 5% MVC background contraction. Peak soleus and tibialis anterior (TA) EMG were normalized to M-max. Soleus and TA EMG were <2.5% of M-max in both SCI and non-SCI subjects. The greatest EMG occurred at the highest acceleration (5g). Low magnitude mechanical oscillation, shown to enhance bone anabolism in animal models, did not elicit high levels of reflex muscle activity in individuals with and without SCI. These findings support the g force modulated background muscle activity during fixed frequency vibration. The magnitude of muscle activity was low and likely does not influence the load during fixed frequency oscillation of the tibia.     

Paraspinal muscle reflex dynamics

Neuromuscular control of spinal stability may be represented as a control system wherein the paraspinal muscle reflex acts as feedback response to kinetic and kinematic disturbances of the trunk. The influence of preparatory muscle recruitment for the control of spinal stability has been previously examined, but there are few reported studies that characterize paraspinal reflex gain as feedback response. In the current study, the input-output dynamics of paraspinal reflexes were quantified by means of the impulse response function (IRF), with trunk perturbation force representing the input signal and EMG the output signal. Surface EMGs were collected from the trunk muscles in response to a brief anteriorly directed impact force applied to the trunk of healthy participants. Reflex behavior was measured in response to three levels of force impulse, 6.1, 9.2 and 12.0 Ns, and two different levels of external trunk flexion preload, 0 and 110 N anterior force. Reflex EMG was quantifiable in response to 91% of the perturbations. Mean reflex onset latency was 30.7+/-21.3 ms and reflex amplitude increased with perturbation amplitude. Impulse response function gain, G(IRF), was defined as the peak amplitude of the measured IRF and provided a consistent measure of response behavior. EMG reflex amplitude and G(IRF) increased with force impulse. Mean G(IRF) was 2.27+/-1.31% MVC/Ns and demonstrated declining trend with flexion preload. Results agree with a simple systems model of the neuromechanical feedback behavior. The relative contribution of the reflex dynamics to spinal stability must be investigated in future research.     

Motor unit recruitment patterns during reflex compensation of muscle yield investigated by computer simulations

 An important function of the stretch reflex in the soleus muscle in the decerebrate cat preparation is to compensate for the tendency of muscle suddenly to yield during ramp increases in length. As the level of background (i.e. pre-stretch) force increases, there is a systematic change in the curvature of the force trajectory during this reflex compensation, from concave to convex with respect to increasing force. The hypothesis that this change in curvature was due to background force-dependent changes in the recruitment pattern of motor units was investigated with a combined computer simulation/ experimental technique. The simulation consisted of 20 model motor units for the soleus muscle, each based on a distributed moment muscle model. The timing of recruitment of the motor units was optimized to allow the simulation outputs to fit a set of experimental data records on the reflex response to stretch initiated at five different levels of pre-stretch force. The resulting recruitment patterns showed that a tendency for recruitment to be concentrated progressively in the early portion of the stretch as pre-stretch force increased could account for the changes in reflex force curvature. These results are consistent with the skewed distribution of intrinsic electrical thresholds of motoneurons, in which low-threshold units are much more frequent than high-threshold ones. Therefore the changes in recruitment pattern and reflex force curvature may be due primarily to the intrinsic properties of motoneurons. Received: 18 September 1995/Accepted in revised form: 21 May 1996     

"Donor" muscle structure and function after end-to-side neurorrhaphy

End-to-end nerve coaptation is the preferred surgical technique for peripheral nerve reconstruction after injury or tumor extirpation. However, if the proximal nerve stump is not available for primary repair, then end-to-side neurorrhaphy may be a reasonable alternative. Numerous studies have demonstrated the effectiveness of this technique for muscle reinnervation. However, very little information is available regarding the potential adverse sequelae of end-to-side neurorrhaphy on the innervation and function of muscles innervated by the "donor" nerve. End-to-side neurorrhaphy is hypothesized to (1) acutely produce partial donor muscle denervation and (2) chronically produce no structural or functional deficits in muscles innervated by the donor nerve. Adult Lewis rats were allocated to one of two studies to determine the acute (2 weeks) and chronic (6 months) effects of end-to-side neurorrhaphy on donor muscle structure and function. In the acute study, animals underwent either sham exposure of the peroneal nerve (n = 13) or end-to-side neurorrhaphy between the end of the tibial nerve and the side of the peroneal nerve (n = 7). After a 2-week recovery period, isometric force (F(0) was measured, and specific force (sF(0) was calculated for the extensor digitorum longus muscle ("donor" muscle) for each animal. Immunohistochemical staining for neural cell adhesion molecule (NCAM) was performed to identify populations of denervated muscle fibers. In the chronic study, animals underwent either end-to-side neurorrhaphy between the end of the peroneal nerve and the side of the tibial nerve (n = 6) or sham exposure of the tibial nerve with performance of a peroneal nerve end-to-end nerve coaptation approximately 6), to match the period of anterior compartment muscle denervation in the end-to-side neurorrhaphy group. After a 6-month recovery period, contractile properties of the medial gastrocnemius muscle ("donor" muscle) were measured. Acutely, a fivefold increase in the percentage of denervated muscle fibers (1 +/0 0.7 percent to 5.4 +/-2.7 percent) was identified in the donor muscles of the animals with end-to-side neurorrhaphy (p < 0.001). However, no skeletal muscle force deficits were identified in these donor muscles. Chronically, the contractile properties of the medial gastrocnemius muscles were identical in the sham and end-to-side neurorrhaphy groups. These data support our two hypotheses that end-to-side neurorrhaphy causes acute donor muscle denervation, suggesting that there is physical disruption of axons at the time of nerve coaptation. However, end-to-side neurorrhaphy does not affect the long-term structure or function of muscles innervated by the donor nerve.     

Comparative response of 2-week and 6-month old rat myocardium to hypoxia

The effect of hypoxia on contractile performance of left ventricular papillary muscles from 2-week old and 6-month old rats was studied. Left ventricular glycogen concentration was not significantly different between 2-week and 6-month old hearts; however, the lactate dehydrogenase isoenzyme pattern of the left ventricular myocardium from the 2-week old rats demonstrated a shift toward M-type (anaerobic) distribution relative to 6-month-old animals. Papillary muscle performance was studied at 28 degrees C while contracting 12 times/min at the peak of the active tension curve in oxygenated (95% oxygen, 5% carbon dioxide) Krebs-Henseleit solution containing glucose concentrations from 5.5 to 22 mM. In a bath containing 5.5 mM glucose, the decline in developed tension during hypoxia (95% nitrogen, 5% carbon dioxide) of preparations from the 2-week old rats was significantly less (P less than 0.05) than the 6-month old group only at 15 min. Raising the glucose concentrations of the bath to 11 and 16.5 mM resulted in improved performance throughout 60 min of hypoxia in preparations from 2-week compared with 6-month old animals. Practolol (5 X 10(-5)M) did not prevent the increased tolerance to hypoxia observed in preparations from 2-week animals. Thus, increasing anaerobic substrate has differential effects on the performance of 2-week and 6-month myocardium during hypoxia. The data suggests that in contrast to 6-month old heart muscle, myocardium from 2-week old rats is capable of modulating its active mechanical activity during hypoxia in accordance with the availability of glycolyptic substrate.     

Separation and estimation of muscle spindle and tension receptor populations by vibration of the biceps muscle in the frog

Frog spinal cord reflex behaviors have been used to test the idea of spinal primitives. We have suggested a significant role for proprioception in regulation of primitives. However the in vivo behavior of spindle and golgi tendon receptors in frogs in response to vibration are not well described and the proportions of these proprioceptors are not established. In this study, we examine the selectivity of muscle vibration in the spinal frog. The aim of the study was (1) to examine how hindlimb muscle spindles and GTO receptors are activated by muscle vibration and (2) to estimate the relative numbers of GTO receptors and spindle afferents in a selected muscle, for comparison with the mammal. Single muscle afferents from the biceps muscle were identified in the dorsal roots. These were tested in response to biceps vibration, intramuscular stimulation and biceps nerve stimulation. Biceps units were categorized into two types: First, spindle afferents which had a high conduction velocity (approximately 20-30 m/s), responded reliably (were entrained 1:1) to muscle vibration, and exhibited distinct pauses to shortening muscle contractions. Second, golgi tendon organ afferents, which had a lower conduction velocity (approximately 10-20 m/s), responded less reliably to muscle vibration at physiologic muscle lengths, but responded more reliably at extended lengths or with background muscle contraction, and exhibited distinct bursts to shortening muscle contractions. Vibration responses of these units were tested with and without muscle curarization. Ensemble (suction electrode) recordings from the dorsal roots were used to provide rough estimates of the proportions of the two muscle afferent types.     

Orderly recruitment among motoneurons supplying different muscles.

Virtually all movements involve the recruitment of motor units from multiple muscles. Given the functional diversity of motor units (motoneurons and the muscle fibers they supply), the effective production of specific movements undoubtedly depends upon some principle(s) to organize the ensemble of active motor units. The principle acting to organize the recruitment of motor units within muscles is the size principle, whereby the first motor units to be recruited have the smallest values for axonal conduction velocity and contractile force, and are the slowest to contract and fatigue. Here we consider the possibility that the size principle applies in the recruitment of motor units across muscles, i.e., that regardless of their muscles of origin, active motor units are recruited in rank order, for example, from low to high conduction velocity. The benefits of orderly recruitment across muscles could be similar to the acknowledged advantages of orderly recruitment within muscles. One benefit is that the neural process involved in organizing active motor units would be simplified. In a muscle-based scheme, the size principle would organize only those motor units within individual muscles, leaving the nervous system with the additional task of coordinating the relative activities of motor units from different muscles. By contrast, in an ensemble-based scheme, orderly recruitment of all motor units according to the size principle would automatically coordinate motor units both within and across motor nuclei. Another potential benefit is the provision for movements with smooth trajectory, the result of interleaving the divergent torque contributions made by motor units from muscles that differ in their orientations about joints. Otherwise, if order were restricted within muscles, the torque trajectory of a joint would change unevenly as participating muscles begin contracting at different times and grade activity at different rates. These considerations support speculation that motor units recruited from co-contracting muscles are collectively recruited according to the size principle.     

Motor unit recruitment for dynamic tasks: current understanding and future directions

Skeletal muscle contains many muscle fibres that are functionally grouped into motor units. For any motor task there are many possible combinations of motor units that could be recruited and it has been proposed that a simple rule, the ‘size principle’, governs the selection of motor units recruited for different contractions. Motor units can be characterised by their different contractile, energetic and fatigue properties and it is important that the selection of motor units recruited for given movements allows units with the appropriate properties to be activated. Here we review what is currently understood about motor unit recruitment patterns, and assess how different recruitment patterns are more or less appropriate for different movement tasks. During natural movements the motor unit recruitment patterns vary (not always holding to the size principle) and it is proposed that motor unit recruitment is likely related to the mechanical function of the muscles. Many factors such as mechanics, sensory feedback, and central control influence recruitment patterns and consequently an integrative approach (rather than reductionist) is required to understand how recruitment is controlled during different movement tasks. Currently, the best way to achieve this is through in vivo studies that relate recruitment to mechanics and behaviour. Various methods for determining motor unit recruitment patterns are discussed, in particular the recent wavelet-analysis approaches that have allowed motor unit recruitment to be assessed during natural movements. Directions for future studies into motor recruitment within and between functional task groups and muscle compartments are suggested.     

Fusimotor reflex profiles of individual triceps surae primary muscle spindle afferents assessed with multi-afferent recording technique.

The experiments were performed on 21 cats anaesthetized with alpha-chloralose. The aim of the study was to investigate sets of simultaneously recorded spindle afferents (2-4 in each set) from the triceps surae muscle (GS) with respect to the pattern of fusimotor reflex effects evoked by different types of ipsi- and contralateral reflex stimulation. The afferents' responses to sinusoidal stretching of the GS muscle were determined and the fusimotor reflex effects were assessed by comparing the afferent responses (i.e. the mean rate of firing and the depth of modulation) elicited during reflex stimulation with those evoked in absence of any reflex stimulus. Natural of electrical activations of ipsi- and contralateral muscle, skin and joint receptor afferents were used as reflex stimuli. The spindle afferents were influenced by several modalities and from wide areas, with a majority responding to both ipsi- and contralateral stimuli. A particular reflex stimulus often caused different effects on different afferents, and the various reflex stimuli seldom gave similar effects on a particular afferent. Multivariate analysis revealed that the variation in response profiles among simultaneously recorded afferents were as great as between afferents recorded on different occasions. This suggests that the individualized response prifiles, observed in earlier investigations, represent a very diversified reflex control of the spindle primary afferents, and are not a reflection of changes in the setting of the spinal interneuronal network, occurring during the time interval between the recordings of different units. Also, there was no relation between the conduction velocity of the afferents and the reflex profiles of the afferents, but non-linear relations were found between effects elicited by different types of stimuli. Indications were also found that it may be possible to separate the population of GS muscle spindles into subgroups, according to the fusimotor effects exhibited by activation of various categories of ipsi- and contralateral receptor afferents. It is concluded that one possible way of making the very complex reflex system controlling the muscle spindles intelligible may be a combination of multiple simultaneous recordings of spindle afferents and multivariate analysis.     

Motor unit behavior during clonus.

Medial gastrocnemius surface electromyographic activity and intramuscular electromyographic activity were recorded from six individuals with chronic cervical spinal cord injury to document the recruitment order of motor units during clonus. Four subjects induced clonus that lasted up to 30 s while two subjects induced clonus that they actively stopped after 1 min. Mean clonus frequency in different subjects ranged from 4.7 to 7.0 Hz. Most of the 166 motor units recorded during clonus (98%) fired once during each contraction but at slightly different times during each cycle. Other motor units fired during some clonus cycles (1%) or in bursts (1%). When 59 pairs of units were monitored over consecutive clonus cycles (n = 5-89 cycles), only 8 pairs of units altered their recruitment order in some cycles. Recruitment reversals only occurred in units that fired close together in the clonus cycle. These data demonstrate that orderly motor unit recruitment occurs during involuntary contractions of muscles paralyzed chronically by cervical spinal cord injury, providing further support for the importance of spinal mechanisms in the control of human motor unit behavior.     

Modulation of spinal inhibitory reflexes depends on the frequency of transcutaneous electrical nerve stimulation in spastic stroke survivors

Neurophysiological studies in healthy subjects suggest that increased spinal inhibitory reflexes from the tibialis anterior (TA) muscle to the soleus (SOL) muscle might contribute to decreased spasticity. While 50?Hz is an effective frequency for transcutaneous electrical nerve stimulation (TENS) in healthy subjects, in stroke survivors, the effects of TENS on spinal reflex circuits and its appropriate frequency are not well known. We examined the effects of different frequencies of TENS on spinal inhibitory reflexes from the TA to SOL muscle in stroke survivors. Twenty chronic stroke survivors with ankle plantar flexor spasticity received 50-, 100-, or 200-Hz TENS over the deep peroneal nerve (DPN) of the affected lower limb for 30?min. Before and immediately after TENS, reciprocal Ia inhibition (RI) and presynaptic inhibition of the SOL alpha motor neuron (D1 inhibition) were assessed by adjusting the unconditioned H-reflex amplitude. Furthermore, during TENS, the time courses of spinal excitability and spinal inhibitory reflexes were assessed via the H-reflex, RI, and D1 inhibition. None of the TENS protocols affected mean RI, whereas D1 inhibition improved significantly following 200-Hz TENS. In a time-series comparison during TENS, repeated stimulation did not produce significant changes in the H-reflex, RI, or D1 inhibition regardless of frequency. These results suggest that the frequency-dependent effect of TENS on spinal reflexes only becomes apparent when RI and D1 inhibition are measured by adjusting the amplitude of the unconditioned H-reflex. However, 200-Hz TENS led to plasticity of synaptic transmission from the antagonist to spastic muscles in stroke survivors.     

The absence of an effect of L-tryptophan on the C-fiber reflex in the chronic spinal cat

L-tryptophan (150 mg/kg) had no effect on the C-fiber reflex in the acute decerebrate chronic spinal cat. L-tryptophan had been previously shown to facilitate this reflex in the acute decerebrate acute spinal cat. This finding supports the hypothesis that tryptaminergic neurons in the brain have axons which descend the spinal cord to produce modulation of some spinal reflexes.