Reflex pathways from group II muscle afferents

Summary A hypothesis is forwarded regarding the role of secondary spindle afferents and the FRA (flexor reflex afferents) in motor control. The hypothesis is based on evidence (cf. Lundberg et al. 1987a, b) summarized in 9 introductory paragraphs. Group II excitation. It is postulated that subsets of excitatory group II interneurones (transmitting disynaptic group II excitation to motoneurones) may be used by the brain to mediate motor commands. It is assumed that the brain selects subsets of interneurones with convergence of secondary afferents from muscles whose activity is required for the movement. During movements depending on coactivation of static -motoneurones impulses in secondary afferents may servo-control transmission to -motoneurones at an interneuronal level. The large group II unitary EPSPs in interneurones are taken to indicate that, given an adequate interneuronal excitability, impulses in single secondary afferents may fire the interneurone and produce EPSPs in motoneurones; interneuronal transmission would then be equivalent to that in a monosynaptic pathway but with impulses from different muscles combining into one line. It is postulated that impulses in the FRA are evoked by the active movements and that the role of the multisensory convergence from the FRA onto the group II interneurones is to provide the high background excitability which allows the secondary spindle afferents to operate as outlined above. The working hypothesis is put forward that a movement governed by the excitatory group II interneurones is initiated by descending activation of these interneurones, but is maintained in a later phase by the combined effect of FRA activity evoked by the movement and by spindle secondaries activated by descending activation of static -motoneurones. As in the original follow up length servo hypothesis (Rossi 1927; Merton 1953), we assume that a movement at least in a certain phase can be governed from the brain solely or mainly via static -motoneurones. However, our hypothesis implies that the excitatory group II reflex connexions have a strength which does not allow transmission to motoneurones at rest and that the increase in the gain of transmission during an active movement is supplied by the movement itself. Group II inhibition. It is suggested that the inhibitory reflex pathways like the excitatory ones have subsets of interneurones with limited group II convergence. When higher centres utilize a subset of excitatory group II interneurones to evoke a given movement, they may mobilize inhibitory subsets to inhibit muscles not required in the movement. Inhibition may be reciprocal of extensors during flexor activation (the spinal pattern), of flexors during extensor activation or of flexors and extensors in more complex movements involving cocontraction of other flexors and extensors. It is postulated that group II inhibition depends on conjoint activation from spindle afferents and other sources (descending and/or the FRA) so that inhibition may be coupled to group II excitation of other motoneurones. Such a coupling would correspond to the --linkage in reciprocal Ia inhibition (Lundberg 1970) and is denoted --linkage in lateral group II inhibition. FRA and other reflex pathways. Results are summarized showing that the FRA evoke convergent excitation in interneurones not only in group II reflex pathways but also in other reflex pathways like the reciprocal Ia inhibitory, the nonreciprocal group I inhibitory and probably also in specialized reflex pathways from cutaneous afferents. It is inferred that facilitation of reflex transmission by impulses in the FRA evoked by the active movement may be a general principle. In this way reflex transmission to -motoneurones may be weak at rest and not disturb passive movements but have a high gain when the reflexes are required to regulate active movement.This work was supported by the Swedish Medical Research Council (project no. 94)     

Regularity in the generation of discharge patterns by primary and secondary muscle spindle afferents,as recorded under a ramp-and-hold stretch

The discharge frequency of primary (Ia) and secondary (II) muscle spindle afferents from the tibial anterior muscle of the cat were recorded under a rampand-hold stretch of the host muscle. The rate of ramp stretch and the prestretch of the muscle were varied systematically. The degree of stretch was kept constant. For a discharge pattern recorded at a ramp rate of 10 mm/s, the peak dynamic discharge, the maximum static value and the final static value were determined. These three discharge rate values were plotted against the maximum static value. In the resulting charts the II afferents presented themselves as a homogeneous group of spindle afferents, whereas the Ia fibers separated into three subgroups. The existence of three subpopulations of Ia fibers was verified by the method of Hald. Furthermore, it is shown that each subpopulation generated its discharge patterns in its own regularly systematic manner. It was concluded that, as one of the three Ia subpopulations exhibits much the same dynamic and static stretch properties as the II fibers, the encoder of this subpopulation must receive its receptor current from the sensory terminals of passive intrafusal chain fibers. The encoder of a second Ia subpopulation indicates its action potentials using the receptor current stemming from the bag₁ sensory terminals, these Ia fibers eliciting a slow adaptation component of a high magnitude which is assumed to be the consequence of a high level of creep in the passive intrafusal bag₁ fiber. The third Ia subpopulation initiates its action potential sequences by means of the receptor current stemming from the passive bag₂ fiber, producing behavior patterns that lie between those of the other two Ia subpopulations.     

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.     

Classification and response characteristics of muscle spindle afferents in the primate.

A study was made of the response characteristics of spindle afferents in the baboon soleus muscle. Afferents were isolated from the dorsal roots, their conduction velocities were determined, and their responses were recorded to muscle stretch at rates of 2.5-45 mm/s and amplitudes of 2-10 mm. Spindle afferents could be classified as primary or secondary on the basis of two criteria. The first criterion was conduction velocity. The conduction velocity histogram was bimodal, with peaks at about 45 and 80 m/s and an intermediate region from 55 to 70 m/s. The second criterion was the pattern of adaptation following the peak of ramp stretch. This latter criterion has the advantage of allowing units with intermediate conduction velocities also to be confidently classified as primary or secondary. The velocity and position sensitivities of primate spindle afferents were determined. The mean dynamic index and mean dynamic sensitivity of secondary afferents were about 45% of the corresponding values for primary afferents. On the other hand, the position sensitivities of primary and secondary spindle afferents in the baboon were not significantly different.     

Event-related cross-correlations between spike trains illustrated on interactions between motor units and muscle spindle afferents

A method is presented for computing correlation coefficients of two (or more) output spike trains in temporal relation to one (or more) input event trains. These event-related correlation functions are computed by convolving the output spike trains, represented as point processes, with rectangular pulses of selectable width, and by then calculating linear correlation coefficients for the pairs of amplitude values obtained from the two convolved processes in temporal relation to the input events. The merits of this technique are illustrated on stimulus trains delivered to motor units (MUs) and output spike trains recorded from muscle spindle afferents of the same cat hindlimb muscle. The correlation functions obtained show the temporal course of the correlated firings of the two afferents (mostly Ia afferents from primary muscle spindle endings) as a function of time from MU activation; they are compared with the conventional cross-correlation histograms (CCHs) between afferents and with peri-stimulus time histograms (PSTHs) between stimulus and afferent firing patterns. Stimulus-related cross-correlation functions as displayed here can be calculated for any three spike trains. Possible extensions of the method to larger numbers of input and output channels are also discussed.     

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.     

Segmental and supraspinal control of synaptic effectiveness of functionally identified muscle afferents in the cat

The present investigation documents the patterns of primary afferent depolarization (PAD) of single, functionally identified muscle afferents from the medial gastrocnemius nerve in the intact, anesthetized cat. Classification of the impaled muscle afferents as from muscle spindles or from tendon organs was made according to several criteria, which comprised measurement of conduction velocity and electrical threshold of the peripheral axons, and the maximal frequency followed by the afferent fibers during vibration, as well as the changes in discharge frequency during longitudinal stretch, the projection of the afferent fiber to the motor pool, and, in unparalyzed preparations, the changes in afferent activity during a muscle twitch. In confirmation of a previous study, we found that most muscle spindle afferents (46.1–66.6%, depending on the combination of criteria utilized for receptor classification) had a type A PAD pattern. That is, they were depolarized by stimulation of group I fibers of the posterior biceps and semitendinosus (PBSt) nerve, but not by stimulation of cutaneous nerves (sural and superficial peroneus) or the bulbar reticular formation (RF), which in many cases inhibited the PBSt-induced PAD. In addition, we found a significant fraction of muscle spindle primaries that were depolarized by stimulation of group I PBSt fibers and also by stimulation of the bulbar RF. Stimulation of cutaneous nerves produced PAD in 9.1–31.2% of these fibers (type B PAD pattern) and no PAD in 8.2–15.4% (type C PAD pattern). In contrast to muscle spindle afferents, only the 7.7–15.4% of fibers from tendon organs had a type A PAD pattern, 23–46.1% had a type B and 50–61.5% a type C PAD pattern. These observations suggest that the neuronal circuitry involved in the control of the synaptic effectiveness of muscle spindles and tendon organs is subjected to excitatory as well as to inhibitory influences from cutaneous and reticulospinal fibers. As shown in the accompanying paper, the balance between excitation and inhibition is not fixed, but can be changed by crushing the afferent axons in the peripheral nerve and allowing subsequent reconnection of these afferent fibers with muscle receptors.     

Activity in receptor afferents from the anterior cruciate ligament evokes reflex effects on fusimotor neurones

Responses from 2-4 muscle spindle afferents from triceps surae and/or posterior biceps and semitendinosus muscles were simultaneously recorded in cats anaesthetized with alpha-chloralose. It was demonstrated that stretch of the anterior cruciate ligament (ACL) of the ipsilateral knee causes changes in dynamic and/or static sensitivity of primary and secondary spindle afferents to sinusoidal stretching. The changes were due to reflex actions of stretch/tension-sensitive receptor afferents from ACL on dynamic and static fusimotor neurones. The findings support the hypothesis that ACL-afferents contribute to the regulation of muscular stiffness around the knee, and thereby also to dynamic knee joint stability.     

Projection of la and lb afferents from forelimb muscles to the C3-C4 segments of the cat spinal cord

Group la muscle spindle afferents were activated separately by small stretches applied to the tendons of antibrachial muscles in the forelimb in the cat. Group lb tendon organ afferents were stimulated electrically after a selective increase of the threshold of the la afferents. Recordings of focal synaptic potentials were made in the C₃-C₄ segments in the medial part of the base of the dorsal horn. It has been found that both la and lb afferents have monosynaptic connections with neurones in this medial region. Quantitatively, these two groups of afferents produced focal synaptic potentials of approximately the same size. The connections may be to inhibitory interneurones projecting to the C₃-C₄ propriospinal neurones, which are known to receive disynaptic IPSPs from group I muscle afferents.     

Cold-sensitive afferents from the abdomen

In cats afferent impulses from splanchnic and vagus nerve preparations were studied. In each splanchnic nerve serving the stomach or the adjacent part of the duodenum, cold-sensitive afferents could be recorded. There were also numerous mechanosensitive fibers originating from stomach, intestine, mesentery and the region of blood vessels. No thermal afferents were found in the vagus nerve. The cardiovascular and respiratory myelinated afferents were insensitive to thermal stimuli.     

Linear and nonlinear effects in the interactions of motor units and muscle spindle afferents

Summary Successive motor unit (MU) twitches often do not sum linearly. Also, muscle spindle (MS) afferents may react nonlinearly to MU contractions occurring at short intervals. Little data is presently available on the interactions between two (or more) MUs regarding their effects on tension output and MS responses. We have studied these effects in cat Mm. gastrocnemius medialis (MG), soleus and semitendinosus. In adult anaesthetized cats, MUs of the muscle under study were electrically stimulated via their ventral root axons with random sequences of brief pulses having mean rates between 6 and 12 pulses per second. Isometric tension fluctuations were recorded from the muscle under study, and discharge patterns of MS afferents (Ia and group II) were recorded from dorsal root filaments. A crosscorrelation analysis was performed to display linear and nonlinear effects evoked by selected time constellations of MU activations. 1) 18 (67%) of 27 MG MUs showed marked potentiation of the second of two twitches in response to pairs of stimuli separated by 5 to about 25 ms. The remainder of these and 16 of the soleus MUs did not exhibit conspicuous nonlinearities. — 2) MS responses to such pairs of MU activations usually showed a prolonged spindle pause. — 3) About 28% of 36pairs of MG MUs produced twitch tension less than expected for linear summation if activated nearly simultaneously. — 4) If two MUs both produced a spindle pause and possibly a relaxation discharge in an MS afferent, the near-synchronous activation of the units produced respective discharge variations that were less than expected for linear summation. If one MU produced an early discharge, contraction of another MU would often prevent it. — These results are discussed in regard to mechanisms of tremor suppression.     

Synaptic and mechanical coupling between type-identified motor units and individual spindle afferents of medial gastrocnemius muscle of the cat

Experiments were performed to test the possibility that motor unit-muscle spindle pairs that are coupled especially strongly mechanically will also be coupled especially strongly synaptically ("weighted ensemble input": Ref. 4). Synaptic and mechanical coupling between one or two individual muscle spindle afferents and individual motor units of the medial gastrocnemius (MG) muscle were measured in barbiturate-anesthetized cats. Synaptic coupling was assessed by measuring the amplitude of single-fiber monosynaptic excitatory postsynaptic potentials (EPSPs) generated in motoneurons by individual spindle afferents. Mechanical coupling was assessed by measuring the alteration in discharge rate of these spindle afferents caused by tetanic activation of the same motor units. Afferents were classified as primary or secondary on the basis of conduction velocity and response to muscle stretch and contraction. Motor units were classified as slow twitch (S); fast twitch, fatigue resistant (FR); fast twitch, intermediate fatigue resistance (FI); and fast twitch, fatigue sensitive (FF) on the basis of twitch contraction time and resistance to fatigue. In 85% of 138 motor unit-primary afferent interactions tested, tetanic activation of the single motor unit unloaded (i.e., decreased the discharge rate of) the primary afferent. A very weak though significant correlation was found between tetanic contraction strength and primary afferent unloading. In 66% of 155 motor unit-secondary afferent interactions tested, tetanic activation of the single motor unit unloaded the secondary afferent. Again, afferent unloading was but weakly related to tetanic contraction strength. Single-fiber EPSPs generated by primary or secondary muscle spindle afferents were recorded in type-identified motor units. EPSPs generated by primary afferents were significantly larger in oxidative (S + FR) than in glycolytic (FF) motor units. No such differences were seen for EPSPs generated by secondary afferents. The magnitude of the EPSP generated in a motoneuron by a spindle afferent was compared to the magnitude of the unloading of that afferent by tetanic activation of the corresponding motor unit. Overall, no relationship was found between these measures.(ABSTRACT TRUNCATED AT 400 WORDS)     

Effects of fusimotor stimulation on dynamic and position sensitivities of spindle afferents in the primate.

The effects of stimulation of single static and dynamic fusimotor fibers on the dynamic sensitivity and position sensitivity of primary and secondary spindle afferents have been studied in the soleus muscle of the baboon. Static fusimotor fibers decreased the mean dynamic sensitivity of primary afferents at all rates of stretch and stimulation. The magnitude of the decrease in dynamic sensitivity increased as the rate of fusimotor stimulation was increased. Qualitatively similar effects were observed in secondary afferents. Static fusimotor stimulation had a strong excitatory effect on spindle afferent resting discharge and greatly increased the mean position sensitivity of both primary and secondary afferents. Dynamic fusimotor fibers increased the mean dynamic index of primary afferents at all rates of stretch and stimulation. The effect of dynamic fusimotor fibers on the mean dynamic sensitivity, however, was dependent on the rate of muscle stretch; at rates below 15 mm/s the dynamic sensitivity was substantially increased, whereas at rates greater than 15 mm/s it was either unchanged or decreased. Dynamic fusimotor fibers slightly decreased the mean position sensitivity of primary afferents.     

Classification of human muscle stretch receptor afferents: a Bayesian approach

1. A sample of 124 human muscle afferents originating from the finger extensor muscles were recorded from the radial nerve in the upper arm. A method is described to formalize the classification of units in muscle spindle primary and secondary afferents and Golgi tendon organ afferents on the basis of a few, nonrigorous assumptions. The classification was based on experimental data that largely have been described in a series of previous papers, although some additional data were collected in the present study. 2. The units were subjected to five tests providing identification data: twitch contraction test, ramp-and-hold stretch, small-amplitude sinusoidal stretches superimposed on ramp stretch, stretch sensitization, and isometric contraction/relaxation. From these five tests the following eight response features were extracted: response to maximal isometric twitch contractions, type of stretch sensitization, correlation between discharge rate and contractile force, response to sudden isometric relaxation, presence or absence of an initial burst, deceleration response, prompt silencing at slow muscle shortening, and driving by small-amplitude sinusoidal stretches. 3. A Bayesian decision procedure was adopted to classify the units on the basis of the eight discriminators. As a first step, units were provisionally classified into muscle spindle primary and secondary afferents, and Golgi tendon organ afferents, by intuitively weighting their responses to the identification tests. Prior probabilities were estimated on the basis of the provisional classification. The eight response features were analyzed and tabulated for all afferents, and the likelihood functions of the tests were directly calculated on the basis of these data.(ABSTRACT TRUNCATED AT 250 WORDS)     

Thalamic projection of muscle nerve afferents in the cat

1. Evoked responses on stimulation of hind- and forelimb muscle nerves have been recorded in the ventro-postero-lateral (VPL) and centre median (CM) nuclei of the thalamus of the cat.2. The stimulation of hind-limb muscle afferents with increasing strength usually did not evoke a thalamic response, either in the CM or in the VPL, until the threshold for Group III muscle afferents was reached.3. Short latency potentials were evoked in the VPL on stimulation of low threshold Group I muscle afferents from forelimbs. The projection zone occupies the dorso-medio-rostral part of the nucleus.4. An attempt was made to locate the medullary relay for forelimb Group I afferents either in the lateral part of the cuneate or in external cuneate.5. Low threshold afferents from forelimb muscle nerves do not project to the CM. CM responses began to appear only when the threshold for Group II muscle nerve fibres was reached.6. Group II afferents from hind- or forelimb muscle nerves projecting to the CM are probably not connected to spindle secondary endings.7. The role played by muscle afferents in the somesthetic mechanisms is briefly discussed.     

The preferred sensory direction of muscle spindle primary endings influences the velocity coding of two-dimensional limb movements in humans

The present study compares how accurately two different but close velocities of movement are discriminated by populations of muscle spindle primary afferents whether or not one takes into account the direction of the movement and the preferred sensory directions of the units (i.e., the direction of movement to which the afferents are the most sensitive). The activities of 26 muscle spindle primary endings originating from the tibialis anterior, the extensor digitorum longus, the extensor hallucis longus, and the peroneus lateralis muscles were recorded in the lateral peroneal nerve. Their responses to movements imposed at two velocities (12.5 and 18 mm/s) were analyzed. These movements were straight-line movements imposed in eight directions and circular movements in both clockwise and anticlockwise directions. The encoding of the movement velocity was analyzed in two ways. First, the mean frequencies of discharge of the muscle spindle afferents were compared for the two velocities. Second, the data were analyzed using a "neuronal population vector model." This model is based on the idea that such neuronal coding can be analyzed in terms of a series of population vectors (i.e., mean contribution of all the muscle spindle afferents within one directionally tuned muscle) and by finally calculating a sum vector. The results showed no clear and consistent difference in the response frequency of the muscle spindle afferents for the two velocities of movement compared. Rather, the most consistently significant differences between the two velocities were in the lengths of the sum vectors. It is concluded that the encoding of two-dimensional movement velocity relies on populations of muscle spindle afferents coming from the whole set of muscles surrounding a particular joint, each muscle making an instantaneous, oriented, and weighted contribution to the sensory coding of the kinematics parameters.     

Static stretch sensitivity of Ia and II afferents in the cat's gastrocnemius

The static discharge characteristics of deefferented Ia and II spindle afferents in the cat medical gastrocnemius muscle have been compared with respect to incidence and firing rate of spontaneously firing units, length threshold of non spontaneous units, linearity of the response to stretch, and position sensitivity. Measurements of the stretch response were taken 20 s after each step increase in length so as to obtain the nearly fully adapted rate. Under this condition the frequency-length relationship was quite linear, especially for group II units. The mean position sensitivity of group II fibers exceeded that of I a fibers in the ratio of 1.51.0. The incidence of spontaneously discharging units was about 20% for both types of unit.The relationship between some of these measures and axonal conduction rate were examined. No correlation of position sensitivity with axonal conduction velocity was detected for either afferent type, but the length thresholds of the group II afferents showed a significant trend for slower conducting, that is, smaller fibers to be less sensitive. There was some indication of a direct relationship between conduction velocity and length threshold for I a fibers. The contradiction to the size principle and mechanisms contributing to the static discharge behavior of the spindle are discussed.Supported by U. S. Public Health Service Grants NS 11950 and HL 07249     

Naloxone-reversible respiratory inhibition induced by muscular thin-fiber afferents in decerebrated cats

Stimulation of muscular thin-fiber afferents of cats causes two types of respiratory suppression: one is stimulus-locked suppression which is not affected by naloxone, and the other is naloxone-reversible respiratory suppression after cessation of the stimulation. Both types of respiratory suppression could still be evoked after decerebration of cats at the midcollicular level. The present experiment revealed that muscular thin-fiber afferents, presumably polymodal receptor afferents, caused respiratory inhibition mediated through an opiate system in the brain structure below the caudal part of the brain stem.     

Kinaesthetic role of muscle afferents in man,studied by tendon vibration and microneurography

Summary The characteristics of vibration-induced illusory joint movements were studied in healthy human subjects. Unseen by the subject, constant frequency vibration trains applied to the distal tendon of the Triceps or Biceps induced an almost constant velocity illusory movement of the elbow whose direction corresponded to that of a joint rotation stretching the vibrated muscle. Vibration trains of the same duration and frequency applied alternatively to the Biceps and Triceps evoked alternating flexion-extension illusory movements.During successive application of vibration trains at frequencies from 10 to 120 Hz, the perceived velocity of the illusory movements increased progressively from 10 to 70–80 Hz, then decreased from 80 to 120 Hz. The maximal perceived velocity was three times higher during alternating vibration of the Biceps and Triceps than during single muscle stimulation.Unit activity from 15 muscle spindle primary endings and five secondary endings located in Tibialis anterior and Extensor digitorum longus muscles were recorded using microneurography in order to study their responses to tendon vibration and passive and active movements of the ankle.Primary endings were all activated by low amplitude tendon vibration (0.2–0.5 mm) previously used to induce illusory movements of the elbow. The discharge of some was phase-locked with the vibration cycle up to 120 Hz, while others responded one-to-one to the vibration cycle up to 30–50 Hz, then fired in a sub-harmonic manner at higher frequencies. Secondary endings were much less sensitive to low amplitude tendon vibration.Primary and secondary ending responses to ramp and sinusoïdal movements of the ankle joint were compared. During the movement, the primary ending discharge frequency was almost constant, while the secondary ending activity progressively increased. During ankle movements the primary ending discharge appeared mainly related to velocity, while some secondary activities seemed related to both movement velocity and joint angle position.Muscle spindle sensory ending responses to active and passive ankle movements stretching the receptor-bearing muscle (plantar flexion) were qualitatively and quantitatively similar. During passive reverse movements (dorsiflexion) most of the sensory endings stopped firing when their muscle shortened. Active muscle shortening (isotonic contraction) modulated differently the muscle spindle sensory ending discharge, which could stop completely, decrease or some times increase during active ankle dorsiflexion. During isometric contraction most of the muscle spindle sensory endings were activated.The characteristics of the vibration-induced illusory movements and the muscle spindle responses to tendon vibration and to active and passive joint movements strengthened the possibility of the contribution of primary endings to kinaesthesia, as suggested by several previous works. Moreover, the present results led us to attribute to proprioception in the muscle stretched during joint movement a predominant, but not exclusive, role in this kind of perception.     

Dorsolateral spinal afferents to some medullary sensory nuclei

Summary The course and termination of afferents in the spinal dorsolateral fascicle to some medullary sensory nuclei were studied by tracing degeneration following lesions of spinal white matter. The main conclusions depend on successive degeneration experiments; other points were studied with single-stage lesions. The dorsal column nuclei were particu-larly studied; terminations in these nuclei following dorsolateral lesions followed a clear-cut pattern, with fibres arising from segments below T 6 terminating in the gracile nucleus and those with more rostral origin solely in the cuneate nucleus. In both nuclei, the major terminations were in their rostral third with most fibres traversing deep caudal regions where some termination also occurred. Some fibres ended contralaterally. These restricted regions of termination contrasted with the wide-spread terminations seen after lesions of the dorsal column. A region at the cuneate rostral pole, adjacent to but clearly separable from nucleus z, receives a dense projection from both caudal and rostral spinal levels, the former fibres terminating in the dorsal part of the region, the latter extending more ventrally. We treat this as a separate subnucleus. The afferents to the dorsal column nuclei (together with those terminating in the other nuclei studied) were confined to the extreme dorsolateral white matter. Our observations confirm the established view that only afferents arising from caudal segments (below at least T 4–5) terminate in nucleus z, and that afferents terminating in group x arise from all levels (at least between C 5 and L 5): also that neither receives any afferents through the dorsal columns. Dorsolateral fibres arising from segments above at least T 6 terminate in a clear-cut area at the lateral border of the external cuneate nucleus. Heavy terminal degeneration was also seen in the lateral cervical nucleus of afferents arising from both above and below T4–5Supported by the Swedish Medical Research Council, Project No. 553