Tendon Organ Afferents Research Articles

Ensemble coding of muscle stretches in afferent populations containing different types of muscle afferents

Ensemble coding of simple mechanical stimuli (small sinusoidal stretches) was studied in simultaneously recorded mixed ensembles of primary- and secondary muscle spindle afferents (MSAs), and Golgi tendon organ (GTO) afferents recorded from L7-S1 dorsal root filaments. The experiments were made on 48 recorded afferents (29 primary MSAs, 6 secondary MSAs and 13 GTO afferents) in chloralose anaesthetised cats. For the analyses, we used a combination of principal component analysis and algorithms for quantification of stimulus discrimination. Mixed ensembles of primary- and secondary MSAs, and GTO afferents, discriminated significantly better between different muscle stretches than ensembles of only one or two types of these afferents. All kinds of ensembles showed a successive increase in discriminative ability with increased ensemble size, and this ability seemed to level at larger populations. However, the increase in discriminative ability was significantly greater for the mixed ensembles. It is hypothesised that the main reason for the greater discriminative ability achieved by mixed ensembles, might be that the variation in response profiles (sensitivity tuning) among the individual afferents of the mixed ensemble will be larger than that for ensembles of only one type of afferent. Finally, the results in the present study give experimental support to some of the teleological arguments in favour of the ensemble coding theory.

Ensemble coding of muscle stretches in afferent populations containing different types of muscle afferents

Ensemble coding of simple mechanical stimuli (small sinusoidal stretches) was studied in simultaneously recorded mixed ensembles of primary- and secondary muscle spindle afferents (MSAs), and Golgi tendon organ (GTO) afferents recorded from L7-S1 dorsal root filaments. The experiments were made on 48 recorded afferents (29 primary MSAs, 6 secondary MSAs and 13 GTO afferents) in chloralose anaesthetised cats. For the analyses, we used a combination of principal component analysis and algorithms for quantification of stimulus discrimination. Mixed ensembles of primary- and secondary MSAs, and GTO afferents, discriminated significantly better between different muscle stretches than ensembles of only one or two types of these afferents. All kinds of ensembles showed a successive increase in discriminative ability with increased ensemble size, and this ability seemed to level at larger populations. However, the increase in discriminative ability was significantly greater for the mixed ensembles. It is hypothesised that the main reason for the greater discriminative ability achieved by mixed ensembles, might be that the variation in response profiles (sensitivity tuning) among the individual afferents of the mixed ensemble will be larger than that for ensembles of only one type of afferent. Finally, the results in the present study give experimental support to some of the teleological arguments in favour of the ensemble coding theory.

Loss of tendon organ inhibition in Parkinson's disease.

Electrical stimulation via skin electrodes placed over human tendons results in a reflex inhibition of voluntary activity in the stimulated muscle, probably due to activation of Golgi tendon organ afferents. The characteristics of this response in the extensor digitorum communis (EDC) muscles of subjects with Parkinson's disease were compared with those in age-matched controls. The threshold of the inhibitory response was significantly increased in the patient population compared with controls (159 +/- 34 V for tremulous patients; 134 +/- 10 V for rigid patients, 90 +/- 5.5 V for age-matched controls and 70 +/- 16 V for all normals). The latency of the inhibitory wave was increased (onset latency was 68.01 +/- 5.5 ms in patients was 51.5 +/- 4.9 ms in controls). The duration of I was also increased in patients (60 +/- 20.8 ms) relative to controls (46 +/- 11.8 ms). This was associated with slow development of the inhibition with the result that maximal inhibition was delayed by approximately 20 ms. Other features of the patient response were its oscillatory character whereby the initial inhibitory and excitatory components were followed by further prominent peaks and troughs which gave the appearance of continuing response cycles. Such behaviour was not seen in normal records. Also electrical stimulation of the extensor muscle tendon produced concurrent records in the forearm flexor muscle, which resembled those from the stimulated muscle. This was in contrast to normal records which showed no response in the flexor. The possible contribution of a disorder of tendon organ reflexes to the rigidity and tremor of Parkinson's disease is discussed.

Reflex inhibition following electrical stimulation over muscle tendons in man.

Electrical stimulation over selected muscle tendons in alert human subjects produced, in each muscle, a reflex inhibition of muscle activity. This inhibition, when maximal, was seen in the surface EMG as an interval of complete electrical silence during a sustained voluntary contraction. The inhibition was clearly visible in single sweeps and in averaged records. Its onset latency and duration were respectively, 56 +/- 4.9 and 46 +/- 11.8 ms in extensor digitorum communis, 71 +/- 6.1 and 46 +/- 10.5 ms in extensor pollicis brevis, 77 +/- 11.2 and 47 +/- 10.5 ms in extensor pollicis longus, 72 +/- 7.3 and 43 +/- 8.6 ms in abductor digiti minimi, and 97 +/- 3.5 and 43 +/- 2.8 ms in tibialis anterior. The inhibitory response was produced at low stimulus intensities (< 10 mA) without electrical (M wave) or mechanical (muscle twitch) signs of direct muscle stimulation. It therefore did not arise from stimulation of la afferents (muscle spindles). The response arose from tendons since it occurred at lowest threshold when stimulation was applied directly over the tendons of the five different muscles studied. At low stimulus intensities, the response declined sharply when the stimulating electrodes were moved to the skin immediately adjacent to the tendons. The response did not arise from skin afferents since it was also presented when stimuli were delivered to the tendon by subcutaneous needle electrodes and it was not reproduced by stimulation of cutaneous nerves in the region of the tendon. In another series of experiments on extensor pollicis brevis, five skin locations were stimulated while overlying the tendon and again while the skin was stretched so that they were lying 0.6-0.8 cm dorsal to the tendon. In these experiments the response was again greatly attenuated when the stimulation was not directly over the tendon, although the same cutaneous sites were stimulated. The inhibition was followed by a pronounced excitatory component (E1) of peak latency 120-140 ms. The results of the study provide evidence for a powerful autogenic inhibitory reflex in man. The evidence is consistent with the possibility that the response arises from Golgi tendon organ afferents.

Changes in PAD patterns of group I muscle afferents after a peripheral nerve crush.

In the anesthetized cat we have analyzed the changes in primary afferent depolarization (PAD) evoked in single muscle spindle and tendon organ afferents at different times after their axons were crushed in the periphery and allowed to regenerate. Medial gastrocnemius (MG) afferents were depolarized by stimulation of group I fibers in the posterior biceps and semitendinosus nerve (PBSt), as soon as 2 weeks after crushing their axons in the periphery, in some cases before they could be activated by physiological stimulation of muscle receptors. Two to twelve weeks after crushing the MG nerve, stimulation of the PBSt produced PAD in all MG fibers reconnected with presumed muscle spindles and tendon organs. The mean amplitude of the PAD elicited in afferent fibers reconnected with muscle spindles was increased relative to values obtained from Ia fibers in intact (control) preparations, but remained essentially the same in fibers reconnected with tendon organs. Quite unexpectedly, we found that, between 2 and 12 weeks after crushing the MG nerve, stimulation of the bulbar reticular formation (RF) produced PAD in most afferent fibers reconnected with muscle spindle afferents. The mean amplitude of the PAD elicited in these fibers was significantly increased relative to the PAD elicited in muscle spindle afferents from intact preparations (from 0.08 +/- 0.4 to 0.47 +/- 0.34 mV). A substantial recovery was observed between 6 months and 2.5 years after the peripheral nerve injury. Stimulation of the sural (SU) nerve produced practically no PAD in muscle spindles from intact preparations, and this remained so in those afferents reconnected with muscle spindles impaled 2-12 weeks after the nerve crush. The mean amplitude of the PAD produced in afferent fibers reconnected with tendon organs by stimulation of the PBSt nerve and of the bulbar RF remained essentially the same as the PAD elicited in intact afferents. However, SU nerve stimulation produced a larger PAD in afferents reconnected with tendon organs 2-12 weeks after the nerve crush (mean PAD changed from 0.05 +/- 0.04 to 0.32 +/- 0.17 mV). The results obtained indicate that the PAD patterns of the afferent fibers reconnected with muscle spindle and tendon organ afferents are changed after crushing their axons in the periphery: stimulation of the bulbar RF appears to produce larger PAD in fibers reconnected with muscle spindles, and stimulation of cutaneous afferents produces larger PAD in fibers reconnected with tendon organs. It is suggested that these alterations in the patterns of PAD of muscle afferents result from central changes in the balance of excitatory and inhibitory influences acting on the segmental pathways mediating the PAD. Although the functional role of these changes has not been established, they may reflect compensatory changes aimed to adjust information arising from damaged afferents.

Increased resting discharge of human spindle afferents following voluntary contractions.

1. The aim of this study was to assess the incidence of lasting alterations in discharge rate of muscle spindle afferents innervating human ankle and toe dorsiflexor muscles following isometric contractions. 2. The subjects performed controlled isometric ankle dorsiflexions maintained for approximately 5 s. During the contraction the discharge of all but one spindle afferent increased above the precontraction level. After complete relaxation, there was prolonged enhancement of the discharge rate of nineteen of fifty-five muscle spindle afferents and none of three Golgi tendon organ afferents. Ten of the nineteen spindle afferents had been silent prior to the contraction. For the population of fifty-five spindle afferents, the mean 'postcontraction' discharge rate was 65% higher than the mean precontraction discharge rate, with the mean rate increasing from 2.3 to 3.9 Hz (P < 0.001). The mean duration of the enhanced postcontraction discharge was 52 s (range, 8-240 s). 3. Stretch applied to the tendon of the receptor-bearing muscle in twelve of fourteen spindle afferents with an enhanced postcontraction discharge rate eliminated or reduced the enhanced discharge rate. 4. The high incidence of an enhanced spindle discharge after voluntary contraction (35% of spindle afferents) suggests that muscle 'history' should be taken into account when interpreting changes in spindle discharge rates. The enhanced discharge rates following contraction probably reflect a long-lasting effect of the contraction-associated increase in fusimotor drive on intrafusal stiffness, rather than the persistence of fusimotor drive following relaxation.

Reflex effects from Golgi tendon organ (Ib) afferents are unchanged after spinal cord lesions in humans.

The reflex actions of medial gastrocnemius (MG) group I afferents on soleus motoneurons were examined in 20 normal subjects and 13 patients with well-defined chronic spinal cord lesions. In normal subjects, stimulation of the MG nerve with near-nerve electrodes at or below the threshold of the alpha motoneuron axons resulted in inhibition of the soleus H reflex at a condition-test interval of 6 msec. The central delay was estimated to be 2.5 msec. This inhibition was attributed to Ib afferents. We found that this inhibition was present in patients with spinal cord lesions. This contrasts with previous reports that stimulation of MG afferents facilitates the soleus H reflex in hemiplegic limbs, and it may represent a physiologic difference between cerebral and spinal spasticity.

Ankle extensor group I afferents excite extensors throughout the hindlimb during fictive locomotion in the cat.

1. The effects of stimulating hindlimb extensor nerves (100-200 ms trains, 100 Hz, < or = 2 times threshold) during the flexor and extensor phases of the locomotor step cycle were analysed in the decerebrate, paralysed cat during fictive locomotion evoked by stimulation of the mesencephalic locomotor region. 2. Stimulation during extension of either the medial gastrocnemius (MG), lateral gastrocnemius-soleus (LGS) or plantaris (Pl) nerves was equally effective in increasing the duration and amplitude of electroneurogram (ENG) activity recorded in ipsilateral ankle, knee and hip extensor nerves. Enhancement of extensor ENG activity could be evoked with near threshold stimulation intensity and appeared within 10-40 ms of the onset of ankle extensor nerve stimulation. Stimulation of anterior biceps during extension occasionally evoked a modest increase in the duration of activity of hip, knee and ankle extensors. Stimulation of quadriceps during extension enhanced the activity of proximal extensors and soleus, but inhibited other ankle extensors. 3. Selective activation of ankle extensor Ia spindle afferents by muscle stretch also enhanced ipsilateral extension. It is argued that both muscle spindle and tendon organ afferents can contribute to the increase in extensor nerve activity evoked by group I stimulation intensity during fictive locomotion. 4. During flexion, stimulation of either the MG, Pl or LGS nerves at group I strength terminated on-going activity in ipsilateral flexors and initiated a burst of activity in ipsilateral hip, knee and ankle extensors, i.e. reset the step cycle to extension. 5. Low strength stimulation of the mixed muscle and cutaneous nerve innervating the plantar aspect of the foot produced extension enhancement and resetting similar to that evoked by group I muscle afferent stimulation. Stimulation of the cutaneous nerve supplying the dorsal aspect of the foot during extension enhanced extensor activity, and during flexion, enhanced the activity of flexors. 6. The effects reported here during fictive locomotion may also occur during overground locomotion with natural activation of group I muscle spindle and tendon organ afferents. Extensor spindle and tendon organ afferents may thus serve as an excitatory reflex system helping to shape the amplitude, duration and timing of ipsilateral extensor activity. Increased or unexpected activation of group I ankle extensor afferents or plantar foot afferents during locomotion could also compensate for increased loading of the limb.

Reduced motor unit activation of muscle spindles and tendon organs in the immobilized cat hindlimb

Six weeks of limb immobilization of a healthy muscle (cat tibialis posterior) at a short length resulted in a significant reduction of mean fiber area for all fiber types (I, 71% of control; IIa, 77% of control; IIb, 79% of control), whereas fiber type proportions were unchanged. For motor units, there was a reduction in peak tetanic force (type slow > fast fatigue resistant > fast fatigable); an increase in the twitch-to-tetanus ratio for fast fatigue-resistant and slow units; and no effect on the twitch force, twitch time course, or fatigability. The reduction in peak force was greater than expected because of fiber atrophy in slow units. Immobilization had a minimal effect on muscle spindle afferent (Ia and spindle group II) responses to a ramp-and-hold stretch of the passive muscle. Tendon organ (Ib) afferents had an increased responsiveness to stretch after immobilization but only when the muscle was stretched from a short resting length. However, immobilization reduced the modulation of muscle afferent discharge in response to tetanic contractions of single motor units. The decline in responsiveness of spindles was a result of the reduced tetanic force of motor units. In contrast, tendon organs in immobilized muscle were twice as likely to convey no information on the contraction of a single motor unit and were more likely to be unloaded, suggesting that immobilization caused the functional denervation of some muscle fibers. Thus the responses of muscle spindles and tendon organs in immobilized muscle reflected atrophic changes in extrafusal fibers but did not provide evidence for substantial disturbance of receptor function.

Contraction-induced excitation in cat peroneal motoneurons.

1. Motoneurons innervating peroneal muscles were recorded intracellularly in anesthetized cats during sustained submaximal isometric contractions of peroneus brevis produced by repetitive electrical stimulation of motor axons in the distal portion of cut ventral root filaments. 2. In contrast with the inhibition previously observed during contractions of gastrocnemius medialis muscle in triceps surae motoneurons, the afferent input generated by peroneus brevis contraction elicited excitatory potentials in nearly all motoneurons supplying peroneus brevis, peroneus tertius, or peroneus longus muscles. 3. We ascribed the contraction-induced excitation of peroneal motoneurons to spindle afferents for two reasons. First, the amplitude of contraction-induced excitatory potentials increased when the ventral root stimulation strength was increased to recruit gamma-axons. Second, with stimulation strengths under gamma-threshold, peroneus brevis contraction still excited peroneal motoneurons, and we obtained evidence that activation of spindles by skeletofusimotor beta-axons could account at least partly for this excitation. 4. The lack of contraction-induced inhibition in peroneal motoneurons and the prevalence of contraction-induced excitation raised the possibility that, in contrast to the usual effects of tendon organ afferents, Ib afferents from peroneus brevis might exert an excitatory influence on homonymous motoneurons. The fact that electrical stimulation of group I afferents in the nerve to peroneus brevis only exceptionally evoked inhibition in peroneal motoneurons would appear compatible with this hypothesis. Furthermore, stimulation of cutaneous afferents, known to facilitate transmission in Ib pathways, only exceptionally revealed a weak contraction-induced inhibition.(ABSTRACT TRUNCATED AT 250 WORDS)

Static sensitivity of tendon organs to tetanic contraction of in‐series motor units in feline peroneus tertius muscle.

1. The results of several studies have indicated an absence of any consistent relationship between the discharge of Ib Golgi tendon organ afferents and the steady-state tetanic tension generated by activating motor units. This question has been re-examined by recording the responses of individual tendon organs to tetanic, isometric contractions of one or more motor units from the peroneus tertius muscle of anaesthetized cats. 2. In three experiments, seventy-three individual tendon organ-motor unit interactions were recorded. The motor units were stimulated at 30-150 s-1 and for each tendon organ-motor unit pair a linear relationship was found between steady-state tetanic tension and Ib afferent discharge. The slopes of these relationships (the static sensitivities) were steepest for the weakest units. 3. When motor units were stimulated in combination, the relationship between discharge frequency and plateau tension was again linear but the static sensitivities were generally much lower than for single units and approached 1 impulse s-1 g-1. 4. Expression of these relationships in terms of the relative tensions generated revealed that the tendon organs were activated most strongly by the IIb muscle fibres, the static sensitivities being reduced by unloading effects. 5. The linear relationships observed during stimulation of single, and groups of, motor units suggest that the patterns of discharge from the tendon organs can mirror the steady-state contractile tensions within the muscle.

Presynaptic modulation of spinal reflexes

Recent evidence suggests that independent sets of interneurons mediate presynaptic inhibition of primary and secondary muscle spindles and of tendon organ afferents. There is also evidence that the information which flows through different intraspinal collaterals of a single muscle spindle or tendon organ afferent fiber is selectively affected by electrical stimulation of the motor cortex. These studies suggest that presynaptic inhibition plays an important role in the selection of the sensory signals required for the execution of a specific motor task.

Reticulospinal actions on primary afferent depolarization of cutaneous and muscle afferents in the isolated frog neuraxis.

The effects of the brainstem reticular formation on the intraspinal excitability of low threshold cutaneous and muscle afferents were studied in the frog neuraxis isolated together with the right hindlimb nerves. Stimulation of low threshold fibers (less than two times threshold) in cutaneous nerves produced short latency, negative field potentials in the ipsilateral dorsal neuropil (200-400 microns depth) that reversed to positivity at deeper regions (500-700 microns). Stimulation of low threshold fibers (less than two times threshold) in muscle nerves produced, instead, negative response that acquired their maximum amplitude in the ventral neuropil (700-900 microns depth). These electrophysiological findings suggest, in agreement with observations in the cat, that low threshold cutaneous and muscle afferents end at different sites in the spinal cord. Intraspinal microstimulation applied within the dorsal neuropil produced antidromic responses in low threshold cutaneous afferents that were increased in size following stimulation of the dorsal or ventral roots, as well as of the brainstem reticular formation. This increase in excitability is interpreted as being due to primary afferent depolarization (PAD) of the intraspinal terminals of cutaneous fibers. Antidromic responses recorded in muscle nerves following intraspinal stimulation within the ventral neuropil were also increased following conditioning stimulation of adjacent dorsal or ventral roots. However, stimulation of the bulbar reticular formation produced practically no changes in the antidromic responses, but was able to inhibit the PAD of low threshold muscle afferents elicited by stimulation of the dorsal or ventral roots. It is suggested that the PAD of low threshold cutaneous and muscle afferents is mediated by independent sets of interneurons. Reticulospinal fibers would have excitatory connections with the interneurons mediating the PAD of cutaneous fibers and inhibitory connections with the interneurons mediating the PAD of muscle afferents. Although our results provide no direct information on whether the reticulospinal depression of the PAD elicited in low threshold muscle afferents is due to inhibition along the pathways producing PAD of muscle spindle or of tendon organ afferents, it seems likely-by analogy with what has been seen in the cat spinal cord-that these inhibitory actions are mostly restricted to the pathways producing PAD in the terminal arborizations of muscle spindle afferents. These results emphasize the specificity of the descending control of the synaptic efficacy of low threshold cutaneous and muscle afferents which could be of importance for motor performance.

Functional properties of spinal interneurons activated by muscular free nerve endings and their potential contributions to the clasp-knife reflex.

1. The goal of this study was to characterize the functional properties of spinal interneurons that are excited by muscular free nerve endings and to assess their contributions to the clasp-knife reflex. 2. The patterns of activity of 82 spinal interneurons that were excited by squeezing the Achilles tendon or manipulation of the muscle surfaces, preferential stimuli for muscular free nerve endings, were extracellularly recorded in lamina V-VII of the L5-S1 spinal cord in decerebrated and spinalized cats. 3. Interneurons were uniformly excited by increases in muscular length and force. Responses to muscle stretch exhibited gradual decay during maintained stretch, afterdischarge after stretch release, and adaptation to repeated stretch. Responses to isometric contraction induced by electrical stimulation of motor axons was also prolonged after contraction, but did not decay during maintained contraction. For similar increases in force, stretch evoked greater excitation than contraction, indicating that both stretch and contraction contributed to interneuronal activity. Overall, the time course and magnitude of the interneuronal responses to stretch and contraction paralleled the time course and magnitude of the clasp-knife reflex. 4. Interneurons were powerfully excited by muscular free nerve endings, which mediate the clasp-knife reflex, and by cutaneous receptors. Only occasionally were they excited by primary spindle or Golgi tendon organ afferents, which suggests that activation of muscular free nerve endings mediated the interneuronal responses to stretch and contraction. 5. Simultaneous recordings of interneuronal activity and the clasp-knife reflex revealed a broad correlation between interneuronal activity and clasp-knife inhibition. 6. Because the patterns of activity of free nerve ending-responsive interneurons during stretch and contraction were similar to the clasp-knife reflex, were closely correlated with clasp-knife inhibition during simultaneous interneuronal and reflex recordings, and were powerfully excited by muscular free nerve endings, it is likely that the interneurons described above contributed to the clasp-knife reflex. 7. In contrast, a small number (n = 16) of interneurons were recorded that were only weakly excited by muscular free nerve endings but strongly excited by group I afferents, exhibited less spontaneous and evoked activity, and had significantly different responses to stretch and contraction. These interneurons are less likely to have contributed to the clasp-knife reflex.

Neurons in the dorsal column nuclei of the rat respond to stimulation of neck mechanoreceptors and project to the thalamus

Electrophysiological recordings were made from neurons in the dorsal column nuclei which were activated by stimulation of muscle and cutaneous receptors in the neck of the rat. 222 units were studied, 158 (71%) of which responded to activation of cutaneous mechanoreceptors while 64 (29%) were activated by muscle receptors. The response patterns of 12 neurons with input from receptors in neck muscles were tested more fully. Their response patterns strongly suggested that 6 were activated by muscle spindle afferents while the other 6 were activated by Golgi tendon organ afferents. 18 (8%) of the neck-responsive neurons in the medulla were shown to project rostrally to the thalamus.

Effect of abdominal compression on diaphragmatic tendon organ activity.

Previous studies suggest that afferents in the diaphragm participate in the reflex reduction in phrenic nerve efferent activation when the length of the diaphragm is increased by abdominal compression. The present study determined the response of tendon organ afferents in the diaphragm to increases in abdominal pressure. Five cats were anesthetized with thiopental sodium (60 mg/kg ip to induce, supplemented intravenously). Extracellular recordings from nine individual tendon organ afferents were made from right cervical dorsal root ganglia 5 and 6. Right crural electromyographic activity was recorded. The right extrathoracic phrenic nerve was isolated and stimulated to identify tendon organs on the basis of conduction velocity and response to twitch. The response to ramp-and-hold stretch of the diaphragm was used as an additional test to differentiate tendon organs from muscle spindles. The mean level of activity of the tendon organs during the 1st s of the inspiratory phase was 47 +/- 10 (SD) Hz. Abdominal compression was associated with a significant increase in the activity of these afferents to 61 +/- 11 Hz. Results indicate that increases in the activity of diaphragmatic tendon organs are associated with moderate increases in abdominal pressure and are likely the result of elevations in the active tension developed by the diaphragm. Combined with results from previous studies, it is possible that diaphragmatic tendon organs may play a role in the attenuation of respiratory muscle activation when abdominal pressure is increased.

Presynaptic inhibition of muscle spindle and tendon organ afferents in the mammalian spinal cord

More than 30 years ago, Frank and Fuortes proposed that the synaptic effectiveness of muscle spindle afferents associated with spinal motoneurones could be diminished by the activation of nerves from flexor muscles. Since that time, research has focused on disclosing the mode of operation and the spinal pathways involved in this presynaptic inhibitory control. Initially, it was assumed that the same last-order interneurones mediated presynaptic inhibition of both muscle spindle and tendon organ afferent fibres. More recent evidence indicates that the synaptic effectiveness of these two groups of afferents is controlled by separate sets of GABAergic interneurones synapsing directly with the intraspinal terminals of the afferent fibres. This unique arrangement allows for selective control of the information on muscle length or muscle tension, despite the convergence of muscle spindle and tendon organ afferents on second-order interneurones.

Phrenic afferents and ventilatory control

It has been understood since the late 1800s that the diaphragm has significant sensory innervation. The role of phrenic afferents in the control of breathing, however, has been obscure. The phrenic nerve has been shown to contain a full array of afferent fibers. However, proprioceptive (group 1 fibers) afferents are few compared to postural muscles or the intercostals. The diaphragm, unlike the inspiratory intercostal muscles, has a small complement of spindle afferents and not all of these spindles are supplied with fusorial fibers. Reduced spindle afferents under gamma control help to explain previous studies of the diaphragm that have failed to reveal autogenic facilitation, that is, a reflex-mediated increase in drive during inspiratory loading. Nevertheless, some clinical studies have revealed increased activation of the diaphragm when its length is reduced. Group 1 fibers, which are predominantly tendon organ afferents in the diaphragm, have been shown to have a phasic inhibitory function. A reduction in this inhibition brought about by a reduction in diaphragmatic length during lung inflation may explain the increased diaphragmatic activation reported in clinical studies. Phrenic afferents have been shown to have multiple spinal and supraspinal projections. Recent studies have explored the ventilatory effects of thin fiber afferents (group III and IV fibers) in the phrenic nerve. Stimulation of these afferents has been shown both to inhibit and excite ventilation. These afferents arise from polymodal receptors that respond to both mechanical and chemical stimulation. Activation of these receptors may occur in a variety of conditions and the ventilatory response may be determined by the specific receptor activated.

Reflex activation of muscle spindles in human pretibial muscles during standing

1. Experiments were performed in standing subjects to determine whether low-threshold cutaneous and muscle afferents from mechanoreceptors in the human foot reflexly influence fusimotor neurons innervating pretibial flexor muscles. Recordings were made from 30 identified muscle-spindle afferents, four tendon-organ afferents, and one alpha-motor axon innervating the pretibial flexor muscles. The subjects stood without support or vision on a force platform while trains of electrical stimuli (5 stimuli, 300 Hz) were delivered at nonpainful intensities to the sural nerve or to the posterior tibial nerve at the ankle. 2. Seventeen of the 30 spindle endings had no background discharge, and none was activated by the sural or posterior tibial stimuli. Five silent afferents were given a background discharge by sustained pressure on the relevant tendon, but with two the discharge was dominated by a tremor rhythm obscuring any reflex response to the stimuli. Based on peristimulus time histograms (PSTHs), the sural stimuli then produced increases in discharge of two of the remaining three endings at latencies of 84 and 90 ms. These effects could not be explained by muscle stretch and are presumed to have been fusimotor mediated. 3. When the subjects stood freely without support or vision, 13 muscle-spindle endings had a background discharge, but with three endings tremor developed at the ankle and dominated the spindle discharge. Sural stimuli affected the discharge of five of nine endings unaffected by tremor. With three of these endings, there were changes in discharge that could be explained by muscle stretch.(ABSTRACT TRUNCATED AT 250 WORDS)

Dynamic response of human muscle spindle afferents to stretch

1. One hundred and twenty-four muscle afferents from the finger extensor muscles were recorded from the radial nerve in human subjects. 2. The afferents were provisionally classified as muscle spindle primary (78/124) and secondary afferents (25/124), and Golgi tendon organ afferents (21/124), on the basis of their response to 1) maximal twitch contractions, 2) 20- and 50-Hz sinusoids superimposed on ramp-and-hold stretches, 3) stretch sensitization, and 4) isometric contractions and sudden relaxations. 3. Ramp-and-hold stretches at two velocities, 10 and 50 degrees/s, were applied to the appropriate metacarpophalangeal (MCP) joint while the parent muscle remained relaxed. For each unit three discrete parameters were assessed: the presence or absence of 1) an initial burst at the commencement of the ramp stretch, 2) a deceleration response at the beginning of the hold phase, and 3) a prompt silencing at muscle shortening. In addition, two kinds of dynamic indexes were calculated for 79 of the muscle spindle afferents. 4. Most spindle afferents responded readily to stretch, whereas the Golgi tendon organ afferents produced very poor stretch responses. All of them lacked a static response, whereas the dynamic response, when present at all, consisted of only a few impulses. 5. The dynamic index was higher for spindle primaries than for secondaries, and this difference was statistically significant although the distribution was unimodal for spindle afferents as a group. Hence, this parameter was a poor discriminator. 6. Initial bursts, deceleration responses, and silences during imposed shortening were more common in spindle primaries than in secondaries. The differences were significant in all these respects. 7. The three discrete parameters were statistically pairwise independent for the spindle afferents, justifying the combination of the three into a useful battery for discrimination between primary and secondary spindle afferents and the use of this battery as a partial data base for a probability approach towards a solid classification of human muscle afferents.