Neuronal pathways for spinal reflexes activated by group I and group II muscle afferents in the spinal segment (Co1) innervating the tail in the low spinalized cat

 We studied neuronal pathways for spinal reflexes activated by group-I and group-II muscle afferents in the spinal segments innervating the tail in unanesthetized and spinalized (L1) cats. Experiments were performed on 25 adult cats of both sexes. The effects of stimulating nerves innervating six tail muscles on both sides were recorded from tail motoneurons in the first coccygeal spinal segment (Co1) using glass microelectrodes. Stable recordings were obtained from 150 tail motoneurons. Stimulation of group-I muscle afferents (stimulus intensity <1.8 T) often produced EPSPs (82/150) after stimulating nerves innervating neighboring tail muscles. Motoneurons innervating the long-tendoned muscles, M. extensor caudae lateralis and M. flexor caudae longus (ECL and FCL), received heteronymous monosynaptic connections from group-I muscle afferents innervating the ipsilateral tail muscles. The motoneurons innervating segmental muscles, M. extensor caudae medialis and M. flexor caudae brevis (ECM and FCB), received heteronymous monosynaptic connections from group-I muscle afferents innervating tail muscles on both sides. The motoneurons innervating tail muscles originated from the Ossa coxae, M. abductor caudae externus and M. abductor caudae internus (ACE and ACI), received monosynaptic connection from group-I muscle afferents innervating most of the tail muscles on both sides. Crossed disynaptic inhibitory pathways activated by primary muscle afferent inputs were observed in ECM, ACE, FCL, and FCB motoneurons. The effects of group-II afferent inputs were not dependent on the kind of motoneuron, and alternative excitatory and inhibitory pathways were not clearly observed in the tail motoneuron pool. It is suggested that variability of the neuronal pathways from group-I and -II muscle afferents to tail motoneurons corresponds to functional relationships among tail muscles, depending on the tail movements. Received: 9 April 1997 / Accepted: 8 August 1998     

Monosynaptic and dorsal root reflexes during locomotion in normal and thalamic cats

1. In normal and thalamic walking cats electrical stimulation of muscle nerves via chronically implanted electrodes produced electromyographic (EMG) and neurographic responses that were modulated in amplitude depending on the phase of the step cycle. These responses were examined for possible indications of effects of primary afferent depolarization (PAD) during stepping. 2. Monosynaptic reflexes (MSRs) produced by stimulating the lateral gastrocnemius (LG) and medial gastrocnemius (MG) nerves were recorded as EMGs in MG or LG muscles during treadmill locomotion in normal cats. These heteronymous MSR responses were greatest during the stance (extensor) phase. 3. In the same animals, after decerebration, similar modulation of the heteronymous ankle extensor MSRs occurred during spontaneous locomotion with the use of the same stimulus and recording sites. 4. In both normal and thalamic cats the amplitude of neurogram responses recorded from LG or MG nerve after stimulation of the other muscle nerve varied with phase of stepping but did not parallel the variations of the MSR measured as EMG amplitude in the same muscle. The nerve responses were largest during the flexion phase of the step cycle and had a calculated central latency of 0.6-1.0 ms. These are interpreted as arising from antidromic activity in large-caliber afferent nerve fibers (i.e., dorsal root reflexes). 5. Spontaneous antidromic activity in severed L7 dorsal rootlet fibers to triceps surae was observed in the thalamic cats during episodes of locomotion and was closely correlated with flexion phase EMG activity in semitendinosus, a bifunctional muscle. 6. In decerebrate cats, dorsal root reflexes (DRRs) in severed filaments of L4-L7 dorsal roots were produced by stimulation of saphenous and posterior tibial nerves. These DRRs were always smaller during locomotion than during rest and were smallest during the flexion phase. 7. The short-latency antidromic activity produced in muscle nerves by stimulating heteronymous muscle nerves thus appears to be a DRR produced in Group I terminal arborizations that are depolarized close to threshold during the flexion phase. Such PAD could account for changes in the MSR that do not always parallel the levels of recruitment of the motor pools as manifest by background EMG amplitude.     

A comparison of homonymous and heteronymous connectivity in the spinal monosynaptic reflex arc of the cat

Summary Multi-unit spike triggered averaging was used to determine functional connectivity between spindle afferent fibers from the medial gastrocnemius muscle and the motoneurons innervating the medial (homonymous connections) and the lateral gastrocnemius-soleus muscle (heteronymous connections). As many as 288 possible connections between 24 motoneurons and 12 afferent fibers were studied in single, acute experiments. The influences of morphological and topographical factors, as well as of motoneuron species on functional connectivity were analysed. The probability that a motoneuron would receive functional connections from a given population of afferent fibers was related to its size and its proximity to the spinal entry level of the afferent fibers. The faster the axonal conduction velocity of the motoneuron (i.e. the larger the motoneuron) and the closer its location to the entry zone of the afferent fibers, the higher was its probability of receiving functional connections. The greater the conduction velocity (i.e. diameter) of a stretch receptor afferent fiber, the higher was its probability of making functional connections with motoneurons. These relationships were qualitatively similar for homonymous and heteronymous connections. 58% (233/399) of the Ia and group II afferents (combined) had functional connections with homonymous motoneurons, 32% (75/234) with heteronymous motoneurons. However, homonymous and heteronymous motoneurons of similar sizes were equally likely to receive functional connections when located at the same craniocaudal level. Differences in the locations and mean sizes of homonymous and heteronymous motoneurons however, cannot account completely for the observed overall differences in homonymous and heteronymous connectivity.     

Analysis of synaptic efficacy in spinal motoneurones from `quantum' aspects

1. Synaptic responses of triceps surae motoneurones of the cat following stimulation of single afferent fibres were examined by intracellular recording techniques.

2. The mean quantum content (m) of monosynaptic excitatory postsynaptic potentials (EPSPs) was independent of the type of motoneurone recorded and of the afferent fibre stimulated. There was no significant difference in m value between homonymous and heteronymous synapses.

3. A positive correlation was found between the amplitude of unit EPSPs and the input resistance of motoneurones. The difference in the amplitude of unit EPSPs appears to be responsible for the higher synaptic efficacy in slow-conducting motoneurones than in fast-conducting motoneurones.

4. There was no significant difference in the time course of monosynaptic EPSPs evoked by impulses from homonymous and heteronymous afferent fibres.

5. The ratio of monosynaptic connexions from a given afferent fibre was significantly greater on to homonymous than to heteronymous motoneurones. It is concluded that the difference in efficacy between homonymous and heteronymous synaptic transmission is due to the difference in the number of afferent fibres converging upon these motoneurones.

     

Influence of load type on presynaptic modulation of Ia afferent input onto two synergist muscles

The present work was designed to investigate presynaptic modulation of Ia afferents in the extensor (ECR) and flexor carpi radialis (FCR) when the two muscles acted as synergists during radial deviation to either support an inertial load (position task) or exert an equivalent constant torque against a rigid restraint (force task). H reflexes were evoked in the ECR and FCR by stimulating at the elbow level (1-ms duration) the radial and median nerves, respectively. Conditioning stimulation was applied to the median and radial nerves at the elbow level to assess presynaptic inhibition of homonymous Ia afferent input (D1 inhibition) from the ECR and FCR, respectively. The ongoing presynaptic inhibition of heteronymous Ia afferents that converges onto ECR and FCR motor neuron pools (heteronymous Ia facilitation) was assessed by stimulating the median nerve at the wrist level (palmar branch) prior to the stimulus applied over the radial or median nerve. The heteronymous monosynaptic Ia facilitation was greater (P < 0.05) during the position task (ECR 121%; FCR 147%) compared with the force task (ECR 115%; FCR 132%), and was paralleled by the depression of D1 inhibition (P < 0.05) during the position task (ECR 75.4%; FCR 79.0%) compared with force task (ECR 58.7%; FCR 58.8%). These data indicate that Ia presynaptic inhibition is reduced during the position task relative to the force task. Such differential modulation of Ia afferent input onto the motor neuron pool likely reflects the requirement to heighten reflex responsiveness during the unstable task of maintaining limb position.     

Influences of morphology and topography of motoneurons and muscle spindle afferents on amplitude of single fiber excitatory postsynaptic potentials in cat

Summary Excitatory postsynaptic potentials (e.p.s. p.s) elicited by impulses in single muscle spindle afferent fibers from the medial gastrocnemius (m.g.) muscle were recorded intracellularly from homonymous and heteronymous motoneurons in order to study factors that influence the amplitudes of such responses. Impulses in large afferent fibers elicited larger single-fiber e.p.s.p.s than those in smaller afferents. Mean e.p.s.p. amplitudes were related exponentially to afferent conduction velocities of both Ia and spindle group II fibers. The closer a motoneuron was to the spinal entry point of an afferent fiber, the larger was the mean e.p.s.p. amplitude evoked in it. Impulses in the same afferent fiber elicited larger e.p.s.p.s in small than large motoneurons when the two cells were located at the same craniocaudal levels. Other factors being equal, the single-fiber e.p.s.p.s evoked in homonymous and heteronymous motoneurons were approximately equal in amplitude. Relatively simple morphological and topographical explanations for the findings were advanced and their functional significance for orderly recruitment in partitioned and unpartitioned reflexes was described.     

Effects on the fusimotor-muscle spindle system induced by intramuscular injections of hypertonic saline

Intramuscular injection of hypertonic saline (HS) is a procedure widely adopted to experimentally induce deep muscle pain in humans. This study was undertaken to test whether intramuscular injections of HS (5%) influence the activity of primary and secondary muscle spindle afferents (MSAs) from homonymous as well as heteronymous muscles. The experiments were performed on six cats anaesthetised with alpha-chloralose. Usually responses of two to nine MSAs from gastrocnemius medialis (GM) and/or gastrocnemius lateralis (GL) muscles were recorded simultaneously, while HS was injected either into the receptor-bearing muscle (homonymous responses) or into a close (GM/GL) or remote synergistic muscle (posterior biceps, PB, heteronymous responses). The mean rate of discharge and the depth of modulation of the MSA responses to sinusoidal stretching of the receptor-bearing muscle were calculated. Out of the 42 afferents tested (7 from GM and 35 from GL), 38 (90%) exhibited statistically significant responses to injections of HS into homonymous and/or heteronymous muscles. With injections into the homonymous muscle, the average maximal increase in mean rate of discharge was 74% and the average decrease in depth of modulation was --18%. The mean duration of the effects was 2.1 min. The corresponding values for heteronymous injections into a close synergist were 87%, -17% and 2.1 min (GM or GL), and for injections into PB 52%, -11%, and 1.8 min. The majority of the responses (72%) were compatible with reflex action on static fusimotor neurones, whereas 20% of the responses could be attributed to mixed static and dynamic fusimotor action. The remaining 8% of the responses were attributed to inhibition of fusimotor activity. There were no statistically significant differences between the responses following injections into homonymous or heteronymous muscles. Injections of Tyrode's solution did not induce any significant alterations in MSA responses, implying that they were not induced by direct and/or injury effects of the injections. HS-related changes in MSA activity were completely abolished after the nerves to corresponding muscles were cut, confirming the reflex nature of the effects. Thus, intramuscular injections of HS induce reflex changes in MSA activity from both homonymous and heteronymous muscles, most likely via fusimotor reflexes. Predominantly static fusimotor neurones were activated. The possible role of the fusimotor-muscle spindle system in altered motor control during experimentally induced muscle pain is discussed.     

Pattern of monosynaptic heteronymous Ia connections in the human lower limb

Changes in the firing probability of single motor units in response to electrical stimulation of muscle nerves were used to derive the projections of muscle spindle Ia afferents to the motoneurones of various leg and thigh muscles. Discharges of units in soleus, gastrocnemius medialis, peroneus brevis, tibialis anterior, quadriceps, biceps femoris and semitendinosus were investigated after stimulation of inferior soleus, gastrocnemius medialis, superficial peroneal, deep peroneal and femoral nerves. Homonymous facilitation, occurring at the same latency as the H reflex and therefore attributed to monosynaptic Ia EPSPs, was found in virtually all the sampled units. In many motor nuclei an early facilitation was also evoked by heteronymous low-threshold afferents. The heteronymous facilitation was considered to be mediated through a monosynaptic pathway when the difference between the central latencies of heteronymous and homonymous peaks was not more than 0.2 ms. The heteronymous Ia connections were widely distributed. In particular, monosynaptic coupling between muscles operating at different joints appears to be the rule in humans, though it is rare between ankle and knee muscles in the cat and the baboon.     

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

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

Post-tetanic influences on primary afferent depolarization in the cat spinal cord

Summary In the spinal cord of pentobarbitone anaesthetised cats, increases in the electrical threshold of the terminations of extensor muscle group Ia afferent fibres, produced by tetanic stimulation of either the appropriate peripheral nerve or the central termination, were associated with parallel changes in the bicuculline-sensitive reduction in electrical threshold of the termination produced synaptically by impulses in flexor muscle low threshold afferent fibres (primary afferent depolarization, PAD) or by microelectrophoretic piperidine-4-sulphonic acid (P4S), an analogue of GABA. Since this post-tetanic hyperpolarization (PTH) could be produced by tetanic stimulation of a single termination centrally, and not by peripheral stimulation of heteronymous nerves, it presumably resulted from changes intrinsic to the tetanized termination. Increases in PAD and the effectiveness of P4S were probably associated with post-tetanic activation of an electrogenic Na⁺/K⁺ pump as the predominant cause of PTH, whereas decreases may have been largely the consequence of post-tetanic increases in intracellular Ca²⁺ levels. These results provide further evidence that GABA is the depolarizing transmitter at axo-axonic synapses upon primary afferent terminals, and that the underlying membrane conductance increase has a reversal potential at a more depolarized level than the resting potential.     

Intramuscular localization of monosynaptic Ia reflex effects in the cat biceps femoris muscle

Intracellular recordings from biceps femoris (BF) motoneurons were made in anesthetized low spinal cats during periods of electrical stimulation of the nerve branches supplying the anterior, middle and posterior portions of the BF muscle and the nerves to semimembranosus and semitendinosus. Measurements were made of each cell's composite intrahomonymous and heteronymous monosynaptic Ia-EPSP responses to stimulation of the test nerves (branches). We have found evidence for an intramuscular localization of these monosynaptic Ia reflex effects not only when comparing responses between the two functional components of the BF muscle as is well established [6] but, in addition, when comparing responses between different parts of each functional (hip extensor and knee flexor) component as well. It is argued that both somatotopic and neuronal recognition factors may contribute to the localization of these monosynaptic reflex effects.     

Individual EPSPs produced by single triceps surae Ia afferent fibers in homonymous and heteronymous motoneurons.

1. The individual EPSPs evoked by the action of single Ia fibers from cat triceps surae (MG, LG, SOL) were recorded in homonymous and heteronymous motoneurons innervating these same three muscles. 2. In general, Ia fibers projected to a greater percentage of homonymous than heteronymous motoneurons. One class of Ia afferent evoked EPSPs in virtually all homonymous motoneurons; the other had a substantially lower projection frequency. Possible difficulties introduced by the limited resolution of the averaging technique are discussed. 3. Individual EPSPs were larger on the average if evoked a) in SOL rather than in MG or LG motoneurons, b) by LG rather than by MG or SOL afferent fibers, or c) in homonymous rather than in heteronymous motoneurons. The mean EPSP was larger in homonymous than in heteronymous motoneurons because the largest EPSPs (greater than 150 muV) were found mainly in homonymous motoneurons. 4. Rise times of EPSPs were only slightly shorter in homonymous than in heteronymous motoneurons, suggesting that other factors besides relative location of Ia terminals account for the observed EPSP amplitude differences. Rise times in SOL motoneurons were longer than those in MG or LG. 5. LG afferent fibers tended to produce larger EPSPs in rostral than in caudal LG motoneurons, and MG afferents produced larger EPSPs in caudal than in rostral MG motoneurons. These spatial effects were in accord with the more rostral entry of LG than MG Ia afferents into the spinal cord. The differential projection of SOL afferents to MG and SOL motoneurons which overlap spatially in the spinal cord suggests a species specificity in addition to a location specificity.     

Investigations on integration of mossy fiber inputs to Purkyně cells in the anterior lobe

Summary The 275 Purkyn cells identified by the criteria of the previous paper have been investigated with respect to their role as units integrating the input to the anterior lobe from various limb nerves. The discharges from single Purkyn cells have been studied in lightly anesthetized (pentothal) or in decerebrate unanesthetized cats, there being averaging usually of 128 responses in the form of post-stimulus time histograms and cumulative frequency distributions.Single Purkyn cells exhibited a wide variation in their responses to the diverse inputs from the various afferent nerves. Attention was focussed on excitatory and inhibitory responses evoked by mossy fibers with a short latency, usually 10–15 msec for hindlimb afferents. With most Purkyn cells these responses were predominantly evoked from cutaneous nerves, low threshold fibers being particularly effective. A few Purkyn cells were preponderantly excited by afferent volleys from muscle nerves, but there was a large group with a mixed input from cutaneous and muscle nerves. Graded strengths of stimulation of muscle nerves showed that sometimes group I volleys were prepotent, but other Purkyn cells were selectively excited by group II volleys. Though sometimes the afferent volleys from antagonistic muscles had a reciprocal action on a Purkyn cell, as on a motoneurone, it was more common to find similar actions. Also convergence of inputs from forelimb and hindlirnb nerves, both cutaneous and muscular, was not uncommon, particularly in marginal areas between hindlimb and forelimb zones. A special design feature is the convergence onto a Purkyn cell of mossy fiber and climbing fiber inputs evoked by the same afferent volley. This convergence was of particular interest along the parasagittal strip of hindlimb climbing fiber distribution in lobule V.It was not possible to translate the observations into some map of the cerebellar cortex on which are marked the territorial distributions from the various limb afferent nerves. Rather, there was an ill-defined patchy character, closely adjacent Purkyn cells often receiving very different subsets of the total input from the various limb nerves. The unitary integrations accomplished by the individual Purkyn cells are further integrated when their axons converge onto and inhibit the neurones of the cerebellar nuclei, and this integration by convergence would occur in each successive relay on the output pathways from the cerebellum.It is pointed out that the experimental findings on the integrative action of the individual Purkyn cells provide basic information for attempts to construct models simulating cerebellar performance and control.Post-Doctoral Fellow NINDS (1F2NB40, 545101 NSRB).Post-Doctoral Fellow UHF Grant No. FTF-3-UB-70.     

H-reflexes in masseter and temporalis muscles in man

In contrast with limb muscles, studies on H-reflexes in the trigeminal system are scarce. The present report aimed at reevaluating the responses obtained in the masseter and temporalis muscles after electrical stimulation of their nerves. Twenty-four subjects participated in the experiments. The reflexes were elicited in the masseter and temporal muscles by monopolar stimulation and recorded using surface electrodes. Stimulation of the masseteric nerve evoked an M-response in the masseter and an H-reflex in both the masseter and the temporal muscles. In contrast with the masseter muscle, where the homonymous H-reflex disappeared at higher stimulation intensities, the heteronymous temporal H-reflex remained and reached a plateau. Simultaneous stimulation of the masseteric and deep temporal nerves resulted in an M-response and an H-reflex in both the masseter and temporal muscles. Increasing stimulus intensitites led to disappearance of the H-reflex in both muscles. The results were compared with those obtained by others on limb muscles. As in these muscles, the presence of heteronymous H-reflexes in the jaw muscles can be used in future studies of motoneuronal excitability.     

Joint receptors modulate short and long latency muscle responses in the awake cat

Summary Nerve cuff electrodes were chronically implanted around multiple peripheral nerves in adult cats, including the medial and posterior articular nerves (MAN and PAN) to the knee while EMG electrodes were implanted into seven hindlimb muscles. Randomized load perturbations producing mid-range knee flexions at varying angular velocities were subsequently applied to awake cats. Recordings were initially obtained with knee joint innervation intact and then after local anaesthetic or saline control solution was injected into the knee. Averaged neurogram and EMG responses to the imposed movements were utilized to assess the contribution of joint mechanoreceptor activity to the evoked muscle responses. Additionally, spike-triggered averaging techniques and peri-stimulus time histograms of single joint afferent units isolated from the articular nerve cuffs were utilized to characterize unitary joint receptor responses. The averaged whole nerve response to knee joint perturbations on each of the cuffed articular nerves revealed phasic increases in activity relative to constant background levels. The earliest phasic responses on the articular nerves were initiated at latencies that were too short to be voluntary, occurring in the short latency (reflex) period. Detectable joint receptors were not recruited until after the earliest excitatory responses of agonist/antagonist muscle pairs acting across the knee had occurred, presumably resulting in mechanical loading of the knee joint capsule and subsequent activation of articular mechanoreceptors. Introduction of local anaesthetic into the knee was accompanied by marked diminution in joint afferent activity. Perturbation-evoked muscle responses were characterized by increased activity above background levels in all seven muscles studied, including antagonist muscle pairs. Local anaesthetic-mediated loss of knee joint mechanoreceptor input altered the latency, amplitude and duration of EMG responses in each muscle. The effect of joint anaesthesia in the short latency period was a generalized decrease in all muscle responses relative to normal and saline controls. The loss of afferent input after joint anaesthesia was also associated with altered muscle responses during the long latency period, when both reflex and voluntary mechanisms could potentially contribute to the generation of EMG activity. Interestingly, long latency activity after joint anaesthesia was characterized by unbalancing in the EMG responses of some antagonist muscle pairs. This alteration of normal antagonist pair co-contraction patterns served to increase the magnitude of the imposed perturbations, rather than to bring the movements under control. Analysis of single joint afferents isolated from whole joint nerve recordings demonstrated that some joint afferent units were tonically active at quiescent, mid-range knee positions. Additionally, isolated afferents demonstrated different time courses of response to imposed perturbations. Some afferents responded with decreased or absent firing activity (TONIC units) while other joint afferents responded with phasic bursts of activity which were greatly increased above their relatively low levels of tonic, background activity (TONIC-PHASIC units). In addition to TONIC and TONIC-PHASIC units, other joint afferents were identified which were only active after imposition of passive knee movements (PHASIC units). In conclusion, data obtained from whole joint nerve recordings as well as data from isolated, single joint afferents demonstrate that joint receptors can modulate short and long latency muscle responses to passively-imposed knee movements in the awake cat.Supported by Medical Research Council of Canada (MRC) grant MT5218. KWM is an MRC fellow     

Haemodynamic responses and renin release during stimulation of afferent renal nerves in the cat.

1. Experiments were done on anaesthetized cats to study the effect of electrical stimulation of afferent renal nerves on the circulatory system and on the release of renin from the kidney. 2. Stimulation of afferent renal nerves over a wide range of parameters consistently elicited an increase in arterial pressure and heart rate. This response was still present in paralysed animals and was not accompanied by changes in respiration or in sympathetic autonomic activity usually associated with painful stimulation. Mesenteric and iliac vasoconstriction was observed concomitantly with the increase in arterial pressure. 3. Release of renin from the contralateral innervated kidney was not significantly changed by stimulation of afferent renal nerves. 4. The existence of renal vascular mechanoreceptors was investigated by altering renal circulation. Stenosis of the renal artery or a marked reduction in renal perfusion pressure elicited an increase in arterial pressure while stenosis of the renal vein elicited a decrease in arterial pressure. These responses, however, were not affected by denervation of the kidney and were therefore interpreted as not being due to neural mechanisms. 5. The precise nature, location and physiological role of renal receptors involved in the cardiovascular responses observed during electrical stimulation of afferent renal nerves remain to be determined.     

Distribution of heteronymous Ia facilitation and recurrent inhibition in the human deltoid motor nucleus

Summary Distribution of heteronymous Ia facilitation and of heteronymous recurrent inhibition in motoneurones innervating the anterior part of the deltoid muscle were investigated in normal human subjects following electrical stimulation of the nerves innervating the main muscles of the upper limb. Activation of group I afferents originating from deltoid, biceps, triceps and extensor carpi radialis (ECR) muscles resulted in an early increase in firing probability of voluntarily activated motor units belonging to the anterior part of the deltoid muscle whereas activation of motor axons supplying deltoid, triceps, ECR and flexor carpi radialis (FCR) muscles resulted in an early and long-lasting decrease in firing probability. No effect was seen following activation of group I afferents and motor axons contained in the ulnar nerve. The characteristics of the early facilitation suggest that it is at least partly due to heteronymous Ia monosynaptic connections while these of the long-lasting inhibition suggest that it is at least partly due to heteronymous recurrent inhibition. Their patterns of distribution are discussed with regards to the functional role of the human deltoid.     

Reflex responses evoked in the adrenal sympathetic nerve to electrical stimulation of somatic afferent nerves in the rat

The present study was initiated to determine the role of somatic A (myelinated) and C (unmyelinated) afferent fibers in both responses of increases and decreases in adrenal sympathetic nerve activities during repetitive mechanical pinching and brushing stimulations of the skin in anesthetized rats with central nervous system (CNS) intact. Accordingly, changes in adrenal sympathetic nerve activity resulting from repetitive and single shock electrical stimulation of various spinal afferent nerves, especially the 13th thoracic (Th13) spinal nerve and the sural nerve, were examined in urethane/chloralose-anesthetized rats. Repetitive electrical stimulation of A afferent fibers in Th13 spinal or sural nerve decreased the adrenal nerve activity similarly as brushing stimulation of skin of the lower chest or hindlimb did, while repetitive stimulation of A plus C afferent fibers of those nerves increased the adrenal nerve activity as pinching stimulation of those skins did. Single shock stimulation of spinal afferent nerves evoked various reflex components in the adrenal nerve: an initial depression of spontaneous activity (the early depression); the following reflex discharge due to activation of A afferent fibers (the A-reflex); a subsequent reflex discharge due to activation of C afferent fibers (the C-reflex); and following post-excitatory depressions. These reflexes seem to be mediated mainly via supraspinal pathways since they were abolished by spinal transection at the C1-2 level. Although the supraspinal A- and C-reflexes could be elicited from stimulation of a wide variety of spinal segmental afferent levels, the early depression was more prominent when afferents at spinal segments closer to the level of adrenal nerve outflow were excited. It is suggested that the decreased responses of the adrenal nerve during repetitive electrical stimulation of A afferent nerve fibers are attributable to summation of both the early depression and post-excitatory depression evoked by single shock stimulation, while the increased responses during repetitive stimulation of A plus C afferent fibers are attributable to summation of the C-reflex after single shock stimulation. In spinalized rats, repetitive stimulation of Th13 always increased the adrenal nerve activities regardless of whether A fibers alone or A plus C fibers were stimulated, just as brushing and pinching of the lower chest skin always increased them. The increased responses in spinal animals seem to be related to the fact that single electrical stimuli of Th13 produced A- and C-reflexes of spinal origin without clear depressions.     

Nervous control of bronchial circulation in pigs: application to the airway stimulation

Vascular responses in airways were studied in an anesthetized pig model. Nervous control of bronchial mucosa blood flow was found to involve mainly the non-adrenergic, non-cholinergic (NANC) system (activation of afferent C-fibers) and the sympathetic system. Nervous control of tracheal and laryngeal vascularization involved the cholinergic and non-cholinergic parasympathetic system and the sympathetic system. Exposure of airways to irritants was followed by vasodilatation in the tracheobronchial mucosa, partly as a result of activation of afferent C-fibers. In sensitized animals, respiratory challenge with the specific allergen produced activation of the NANC system with antidromic vasodilatation in the mucosa. This response can be likened to the "axon reflex" seen in skin. Activation of airway autonomic nerves thus was found to be a central step in the genesis of inflammatory reactions in the lungs.     

Expansion of afferent vestibular signals after the section of one of the vestibular nerve branches

The anterior branch of N. VIII was sectioned in adult frogs. Two months later the brain was isolated to record in vitro responses in the vestibular nuclei and from the abducens nerves following electric stimulation of the anterior branch of N. VIII or of the posterior canal nerve. Extra- and intracellularly recorded responses from the intact and operated side were compared with responses from controls. Major changes were detected on the operated side: the amplitudes of posterior canal nerve evoked field potentials were enlarged, the number of vestibular neurons with a monosynaptic input from the posterior canal nerve had increased, and posterior canal nerve stimulation recruited stronger abducens nerve responses on the intact side than vice versa. Changes in the convergence pattern of vestibular nerve afferent inputs on the operated side strongly suggest the expansion of posterior canal-related afferent inputs onto part of those vestibular neurons that were deprived of their afferent vestibular input. As a mechanism we suggest reactive synaptogenesis between intact posterior canal afferent fibers and vestibularly deprived second-order vestibular neurons.