Lateral Gastrocnemius Nerve Research Articles

Task Specific Expression of Sensory Loss Following Gastrocnemius Self-reinnervation in the Cat

PURPOSE: In this study we investigated the effects of self-reinnervation of feline medial gastrocnemius (MG) and lateral gastrocnemius (LG) muscles on joint kinematics of the whole hindlimb during overground walking on surfaces of varying slope (−50%, 0%, +50%). METHODS: Hindlimb kinematics during stance were assessed (1) with little or no activity in MG and LG (short-term effects of self-reinnervation), and (2) after motor function of these muscles was presumably recovered but their proprioceptive feedback permanently disrupted (long-term effects of self-reinnervation). Measurements were performed prior to and at consecutive time points after transecting and immediately suturing MG and LG nerves. RESULTS: Substantial short-term effects were observed in the ankle joint (e.g., increased flexion in early stance) as well as in metatarsophalangeal and knee joints, leading to altered interjoint coordination. Hindlimb kinematics in level and upslope walking progressed back towards baseline within 14-19 weeks, despite the absence of length feedback from gastrocnemius, suggesting that the proprioceptive loss is compensated by other sensory sources or altered central drive. In contrast, ankle joint kinematics as well as interjoint coordination in downslope walking gradually progressed towards, but never reached their baseline patterns (>32 weeks post self-reinnervation). CONCLUSIONS: The muscle contractile conditions and the corresponding potential importance of length and force feedback for the different slope conditions are discussed. It is concluded that proprioceptive feedback from MG and LG muscles is important for regulating locomotor activity of ankle extensors. Supported by NIH Grants HD032571 and NS048844.

The effects of self-reinnervation of cat medial and lateral gastrocnemius muscles on hindlimb kinematics in slope walking

The aim of this study was to investigate the effects of self-reinnervation of the medial (MG) and lateral gastrocnemius (LG) muscles on joint kinematics of the whole hindlimb during overground walking on surfaces of varying slope in the cat. Hindlimb kinematics were assessed (1) with little or no activity in MG and LG (short-term effects of self-reinnervation), and (2) after motor function of these muscles was presumably recovered but their proprioceptive feedback permanently disrupted (long-term effects of self-reinnervation). The stance phase was examined in three walking conditions: downslope (-50%, i.e. -26.6 degrees ), level (0%) and upslope (+50%, +26.6 degrees ). Measurements were performed prior to and at consecutive time points (between 1 and 57 weeks) after transecting and immediately suturing MG and LG nerves. It was found that MG-LG self-reinnervation did not significantly change hip height and hindlimb orientation in any of the three walking conditions. Substantial short-term effects were observed in the ankle joint (e.g., increased flexion in early stance) as well as in metatarsophalangeal and knee joints, leading to altered interjoint coordination. Hindlimb kinematics in level and upslope walking progressed back towards baseline within 14-19 weeks. Thus in these two conditions the cats were walking without any detectable kinematic deficits, despite the absence of length feedback from two major ankle extensors. This was verified in a decerebrate preparation for four of the five cats. In contrast, ankle joint kinematics as well as interjoint coordination in downslope walking gradually progressed towards, but never reached their baseline patterns. The short-term effects can be explained by both mechanical and neural factors that are affected by the functional elimination of MG and LG. Permanent changes in kinematics during downslope walking indicate the importance of proprioceptive feedback from the MG and LG muscles in regulating locomotor activity of ankle extensors. Full recovery of hindlimb kinematics during level and upslope walking suggests that the proprioceptive loss is compensated by other sensory sources (e.g. cutaneous receptors) or altered central drive.

Electrode Array for Reversing the Recruitment Order of Peripheral Nerve Stimulation: Experimental Studies

One of the most challenging problems in peripheral nerve stimulation is the ability to activate selectively small axons without large ones. Electrical stimulation of peripheral nerve activates large diameter fibers before small ones. Currently available techniques for selective activation of small axons without large ones require long-duration stimulation pulses (>500 micros) and large stimulation amplitude, which shorten battery life of the implanted stimulator and could lead to electrode corrosion. In the current study, the hypothesis that small axons can be recruited before large ones with narrow pulse width (50 micros) using an electrode array was tested in both simulations simulation and experiments in the cat lateral gastrocnemius (LG) model. The LG nerve innervates both LG and soleus muscle groups with axons within 10-13 and 8-12 microm diameter ranges, respectively. A finite element model of LG nerve was constructed and simulations showed that, when activating 40% of LG, a conventional tripolar electrode activated only 9% of soleus whereas the electrode arrays of 5, 7, and 11 contacts activated 39, 46, and 60% of soleus respectively, suggesting that the arrays could activate small axons before fully recruiting large axons. In animal experiments, peak twitch force of LG and soleus were plotted as a function of stimulation amplitude to indicate the recruitment curve. At 40% activation of LG, a conventional tripolar electrode activated only 7% of soleus whereas the electrode arrays of 5, 7, and 11 contacts activated 43, 48, and 72% of soleus respectively. The electrode arrays also decreased significantly the recruitment curve slopes to only 10-20% of the value obtained for the tripolar electrode in both computer simulations and experiments. In conclusion, the 5-, 7-, and 11-contact arrays can be used to reverse the recruitment order of peripheral nerve stimulation with a narrow pulse.

Neurohistochemical analysis of regeneration in rat peripheral nerve after end-to-side neurorrhaphy

We investigated the regenerative capacity of motor nerves repaired by end-to-side or end-to-end neurorrhaphy, using choline-acetyltransferase (ChAT) activity measurement or histological analysis. The right medial gastrocnemius nerves (MGNs) of 62 male Fisher strain rats were transected and divided into three groups. In group 1, the distal ends of the MGN were coapted to the side of the lateral gastrocnemius nerve, using a Y-shaped silicone tube in end-to-side neurorrhaphy. In group 2, the nerve ends were reconnected by the traditional end-to-end technique. In group 3, the nerve ends were separated and remained unrepaired. The MGNs were sampled 1, 2, and 3 months postoperatively for histological examinations and ChAT activity measurement. The medial gastrocnemius muscle (MGM) was also sampled for histological evaluations. Axonal regeneration of MGN and the recovery of MGM to nearly normal histology and weight were observed in groups 1 and 2 3 months postoperatively. Although there were no significant differences in ChAT values between groups 1 and 2, the values were significantly larger than that of group 3 3 months postoperatively. These findings suggested that end-to-side neurorrhaphy would be an alternative treatment for peripheral nerve injury in certain clinical situations.

Collateral sprouting mechanism after end-to-side nerve repair in the rat.

The collateral sprouting mechanisms of axons from an uninjured donor nerve after end-to-side nerve repair was investigated in motor nerves of rats, with special reference to the neurotrophins related to nerve regeneration. In addition, growth cone formation at the tip of the regenerating nerve was examined. A transected medial gastrocnemius nerve (MGN) was transferred to the side of an intact lateral gastrocnemius nerve (LGN) using a Y-shaped silicone tube. At 3, 7, or 14 days later, the MGN with the LGN was transected and was stained immunohistologically. Expression of neurotrophin-3 (NT-3) and Trk C (receptor of NT-3) was most significantly observed 3 days postoperatively around the site of coaptation. Brain-derived neurotrophic factor (BDNF) and Trk B (receptor of BDNF) was weakly detected at the coaptation site 3 days after-operation. Growth-associated protein 43 (GAP-43), which is a marker of growth cone formation, was observed at the site of coaptation in the LGN 7 days postoperatively and in the MGN at the site of coaptation at 14 days. We concluded that motor nerve regeneration due to collateral sprouting of axons after end-to-side nerve repair is possible. We thus demonstrated the involvement of at least one neurotrophin, NT-3, in the process of collateral sprouting of motor nerves.

Effects of PAD on conduction of action potentials within segmental and ascending branches of single muscle afferents in the cat spinal cord.

In anesthetized and paralyzed cats under artificial respiration, we examined the extent to which primary afferent depolarization (PAD) might affect invasion of action potentials in intraspinal axonal and/or terminal branches of single muscle afferents. To this end, one stimulating micropipette was placed at the L6 spinal level within the intermediate or motor nucleus, and another one at the L3 level, in or close to Clarke's column. Antidromically conducted responses produced in single muscle afferents by stimulation at these two spinal levels were recorded from fine lateral gastrocnemius nerve filaments. In all fibers examined, stimulation of one branch, with strengths producing action potentials, increased the intraspinal threshold of the other branch when applied at short conditioning testing stimulus intervals (<1.5-2.0 ms), because of the refractoriness produced by the action potentials invading the tested branch. Similar increases in the intraspinal threshold were found in branches showing tonic PAD and also during the PAD evoked by stimulation of group I afferent fibers in muscle nerves. It is concluded that during tonic or evoked PAD, axonal branches in the dorsal columns and myelinated terminals of muscle afferents ending deep in the L6 and L3 segmental levels continue to be invaded by action potentials. These findings strengthen the view that presynaptic inhibition of muscle afferents produced by activation of GABAergic mechanisms is more likely to result from changes in the synaptic effectiveness of the afferent terminals than from conduction failure because of PAD.

Neuromuscular compartments of cat lateral gastrocnemius produce different torques about the ankle joint.

The primary purpose of this study was to establish whether the neuromuscular compartments of cat lateral gastrocnemius produce different mechanical actions on the skeletal system, by determining the contributions made by these compartments to the torques produced about the ankle joint. It was postulated that neuromuscular compartments might represent output elements of the spinal circuits. If so, they should produce unique mechanical actions. Isometric torques about the center of the ankle joint produced by the neuromuscular compartments of the cat lateral gastrocnemius were measured with a multiaxis force-moment sensor connected to the plantar surface of the foot. Individual compartment torques were elicited by activation of the primary compartment branches of the lateral gastrocnemius nerve. The magnitude of the individual torque components, and thus of the resultant torque, was significantly different between compartments. In three of the four lateral gastrocnemius compartments, significantly different torque trajectories were noted. The results, together with those from previous studies demonstrating that compartments can be activated in a task-dependent manner, suggest that neuromuscular compartments represent anatomical substrates that can be used by the nervous system for regulating movement.

Central projections and entries of capsaicin-sensitive muscle afferents

The entry pathway and central distribution of Aδ and C muscle afferents within the central nervous system (CNS[ were investigated by combining electron microscopy and electrophysiological analysis after intramuscular injection of capsaicin. The drug was injected into the rat lateral gastrocnemius (LG) and extraocular (EO) muscles. The compound action potentials of LG nerve and the evoked field potentials recorded in semilunar ganglion showed an immediate and permanent reduction in Aδ and C components. The morphological data revealed degenerating unmyelinated axons and terminals in the inner sublamina 11 and in the border of laminae 1–11 of the dorsal horn at L4-L5 and CI-C2 (subnucleus caudalis trigemini[ spinal cord segments. Most degenerating terminals were the central bouton (C) of type I and 11 synaptic glomeruli. Furthermore, degenerating peripheral axonal endings (V 2) presynaptic to normal C were found. Since V 2 were previously found degenerated after cutting the oculomotor nerve (ON) or L4 ventral root, we conclude that some Aδ and C afferents from LG and EO muscles entering the CNS by ON or ventral roots make axoaxonic synapses on other primary afferents to promote an afferent control of sensory input.

Effects of Capsaicin in the Motor Nerve

The injection of capsaicin into the lateral gastrocnemius (LG) muscle of the rat induced an immediate and sustained reduction in the Aδ and C components of the compound action potential (CAP) of the LG motor nerve. Conversely, the drug did not immediately affect the CAP wave belonging to fast-conducting fibers or the motor responses to LG nerve stimulation. It seems that capsaicin only affects the group III and IV afferents of LG nerve. However, a week after the injection the capsaicin also altered the motor responses, as shown by the threshold enhancement and amplitude reduction of the muscle twitch and by the decrease of the Aα-β CAP components. This late motor impairment was attributed to a central depression following a reduction of capsaicin-sensitive neuron input into the CNS. However, this motor effect was transient since the LG nerve regained the preinjection excitability level in a week and the muscle twitch amplitude reached the control value in a month.

Interactions between Respiratory, Cardiac and Stepping Rhythms in Decerebrated Cats: Functional Hierarchical Structures of Biological Oscillators

Interactions are described of central origin between respiratory, cardiac and stepping rhythms during fictive locomotion in paralyzed, vagotomized, and decerebrated cats. Fictive locomotion was induced by tonic electrical stimulation of the mesencephalic locomotor region (MLR). The coherence between heart beat fluctuation, the efferent discharges of the phrenic, and the lateral gastrocnemius nerves was used to evaluate the strength of the coupling between those three rhythms. The heart beat rhythm was modulated by the centrally generated respiratory and stepping rhythms. The central respiratory rhythm was modulated by the centrally generated stepping rhythm. Based on the present findings, we have proposed a new model concerning the functional hierarchical structures of the three biological oscillators.

The Microtubular Pattern Changes at the Spinal Cord-Root Junction and Reverts at the Root-Peripheral Nerve Junction in Sensory and Motor Fibres of the Rat.

In the rat, we studied the microtubular content of central nervous system (CNS) axons (pyramidal tract, dorsal funiculus, and intracord domain of motor axons), of radicular axons (ventral and dorsal roots), and of peripheral axons (sural and lateral gastrocnemius nerves). The microtubular density had an inverse relationship with the size of the axon. Within the CNS, values ranged from over 120 microtubules/microm2 for axons smaller than 0.1 microm2 of the pyramidal tract and dorsal funiculus to 24 for 3-microm motor axons (area, 7 microm2) in their spinal cord domain. Peripheral nerve and CNS axons of the same size had comparable microtubular densities. In contrast, the microtubular density of dorsal and ventral root axons was one half that of CNS or peripheral nerve axons of equal calibre. Considered along the axon, the microtubular density of motor and sensory fibres is high in the CNS domain, low in the root, and high again in the peripheral nerve domain. These observations are inconsistent with the notion that the cytoskeleton moves coherently away from the perikaryon. We conclude that the axonal microtubular content accords with the calibre of the fibre and with the anatomical region where it courses. We propose that axonal microtubules are regulated by local cues.

Excitability changes of ankle extensor group Ia and Ib fibers during fictive locomotion in the cat.

The present study examines the modulation of gastrocnemius-soleus (GS) monosynaptic reflexes as well as the intraspinal threshold changes of GS group I primary afferent terminals ending in the intermediate and motor nuclei during fictive locomotion in high decerebrate cats. The amplitude of the monosynaptic reflexes (MSR's) evoked in the medial gastrocnemius by stimulation of the lateral gastrocnemius nerve was increased during the extensor (E) phase, decreased during the flexion (F) phase of the step cycle and remained transiently increased after spontaneous episodes of fictive stepping. The intraspinal threshold of populations and of single group Ia GS afferent fibers ending in the motor pool, as well as of single Ia and Ib fibers ending in the intermediate nucleus, showed a sustained reduction during the episodes of fictive locomotion with superimposed cyclic changes in phase with the step cycle. During fictive walking and trotting the reduction of the intraspinal threshold of both Ia and Ib fiber terminals was maximal during the middle or late portion of the F-phase. During fictive gallop elicited by stimulation of the superficial peroneus nerve, the decrease in the intraspinal threshold of the Ia afferent fibers occurred however in phase with the activity of the GS motoneurons. During episodes of fictive locomotion slow, sustained negative DC potential shifts lasting tents of seconds, reflecting an increase in the extracellular potassium concentration were recorded at the base of the dorsal horn and in the intermediate nucleus. The present findings support the existence of tonic and phasic depolarization of the intraspinal terminals of GS group Ia and Ib primary afferents during spontaneous fictive locomotion. It is suggested that accumulation of potassium ions in the extracellular space contributes mainly to the sustained depolarization of group I fibers. The phasic depolarization would be mostly due to the activation of specific sets of interneurons and may, in the case of Ia fibers, contribute to the cyclic modulation of the MRS elicited during fictive locomotion.

Spinal cord projections from hindlimb muscle nerves in the rat studied by transganglionic transport of horseradish peroxidase, wheat germ agglutinin conjugated horseradish peroxidase, or horseradish peroxidase with dimethylsulfoxide.

The spinal cord projections of four different groups of hindlimb muscle nerve branches--the medial and lateral gastrocnemius nerves, muscle branches of the deep peroneal nerve, muscle branches of the femoral nerve, and a nerve to the hamstring muscles--were studied with transganglionic transport of horseradish peroxidase (HRP) in the rat. The influence of varying the postoperative survival (3, 6, and 10 days) and of using wheat germ agglutinin-HRP conjugate (WGA-HRP), or HRP with dimethylsulfoxide (DMSO) instead of free HRP was studied for the gastrocnemius nerves. After 3 days' survival following application of HRP to the gastrocnemius nerves, fine granular labeling was found mainly in lamina V in L4-5, and coarse granular labeling was found in Clarke's column as far caudally as L2, and in laminae VI and VII predominantly in Th12-L2. After 6 or 10 days' survival, the fine labeling in lamina V was sparse or absent, whereas the coarse labeling appeared to remain or to be only slightly reduced in Clarke's column and in laminae VI and VII. No labeling suggestive of terminals was observed in laminae I-III from the gastrocnemius nerves. Except for sparse labeling in lamina I in some of the cases and some minor differences rostrocaudally, the spinal distribution of labeling was similar to that from the other nerves investigated. The distribution of labeling obtained after application of WGA-HRP or HRP with DMSO to the gastrocnemius nerves was very similar to that obtained with free HRP after 3 days' survival. The results indicate that the spinal cord projections of hindlimb muscle nerves in the rat distribute mainly in the deep part of the dorsal horn and in the intermediate zone. Furthermore, the lack of labeling suggestive of terminals in laminae I-III from the gastrocnemius nerves suggests, in conflict with earlier findings in the cat, that primary afferent fibers from muscles do not necessarily terminate in these laminae in the rat. The results suggest, furthermore, that fine granular labeling found in lamina V represents fine-calibered afferent fibers. Finally, the similar spinal projection patterns of the different muscle nerves investigated suggest either a less developed or an essentially different somatotopic organization for muscle afferents compared to cutaneous afferents, as revealed in earlier studies.

Inhibition of nociceptive responses of laminae V–VII dorsal horn neurones by stimulation of mixed and muscle nerves, in the cat

Low frequency (1–3 Hz) stimulation of muscle nerves (ipsilateral medial and lateral gastrocnemius nerves) had to be applied at intensities which would recruit C-fibre afferents, to produce inhibition of nociceptive responses of cat lumbar spinal cord laminae V–VII units. Inhibitory responses evoked by muscle nerve stimulation and also by mixed nerve stimulation could be blocked by microionophoretically applied bicuculline. No effects on these inhibitions were obtained with naloxone.

Motoneurons of the rat sciatic nerve

The sciatic nerve of the rat is a commonly used model for studies on nerve injury, regeneration, and recovery of function. To interpret the changes that occur in a neuron population subsequent to peripheral nerve injury, and to compare different repair procedures, it is essential to have a complete and accurate understanding of the population's normal cellular constituents and their locations. This study reports on the numbers, sizes, and topographic distributions of the motoneuron populations of individual branches of the rat sciatic nerve (peroneal, tibial, sural, and the medial and lateral gastrocnemius nerves), as determined by retrograde transport of HRP (or WGA-HRP) from cut proximal nerve ends isolated in wax to prevent spread of the tracer substance. Optimal labeling of motoneurons was evident between 42 and 73 h of survival. Reconstructions were made from 40-μm serial sections of spinal segments L6 through L2, usually in the coronal plane. Accurate motoneuron counts were obtained by detailed reconstructions in which an accounting of all “split cell” fragments was made to avoid double cell counts. The sciatic nerve of the albino rat contains a total population of about 2005 ± 89 motoneurons. The tibial nerve contained 982 ± 36 cells or 49% of the total. The common peroneal nerve contained 31% or 632 ± 27 motoneurons. The medial and lateral gastrocnemius nerve branches contained collectively 322 ± 16 (16%). The sural nerve accounted for only 68 ± 10 motoneurons (3%). The sciatic motoneurons form a continuous, compact cell column in the dorsolateral quadrant of the ventral horn extending from rostral L6 into the caudal third of L3 over a longitudinal distance of about 6.3 to 7.5 mm. This fusiform column shows its greatest width, 0.5 mm, in mid-L4. Within this compartment motoneurons of each branch of the sciatic occupy spatially distinct subcompartments. Their relative positions are described in detail.

Partitioning of monosynaptic Ia excitatory postsynaptic potentials in the motor nucleus of the cat lateral gastrocnemius muscle.

Experiments were conducted to test the hypothesis that a partitioning of Ia monosynaptic excitatory postsynaptic potentials (Ia EPSPs) is present in motor nuclei supplying muscles with regions capable of different mechanical actions. Intracellular recordings of synaptic potentials were made in lateral gastrocnemius (LG) motoneurons in anesthetized low-spinal cats. The effects were tested of stimuli (group I range) to the four primary nerve branches of the LG nerve supplying muscle compartments LGm, LG1, LG2, and LG3 (terminology of English, Ref. 26) and the nerve to a heteronymous muscle, soleus. Stimulation of a given LG nerve branch produced monosynaptic Ia EPSPs of greater amplitude in "own-branch" motoneurons than "other-branch" cells. A significant partitioning of mean Ia EPSPs was found in three (LG1, LG2, LG3) out of the four homonymous pathways studied. An EPSP normalization (7) was performed to eliminate potential differences in cell type that might affect the amplitudes of the EPSPs between these four cell groups (e.g., differences in the number of cells supplying FF, FR, and S muscle units). This normalization confirmed that the partitioning of monosynaptic Ia inputs upon stimulation of LG1, LG2, and LG3 could not be attributed to differences in cell type. In addition, the effects of LGm stimulation were found to be significantly greater in the LGm motoneurons compared with the other cell groups. Heteronymous input (from soleus) to the LG motor nucleus showed some partitioned effects. Motoneurons innervating compartment LG2 received larger EPSPs from soleus than did the cells supplying compartments LG1, LG3, and LGm. The contributions of location specificity and species specificity (terminology of Scott and Mendell, Ref. 55) in the establishment of these Ia-afferent-motoneuronal connections were examined. Cell location sites within the spinal cord were consistent with location specificity making some contribution to the observed pattern of homonymous Ia connections. A more prominent role for species specificity was indicated by species-dependent differences in EPSP amplitude in pairs of LG motoneurons (e.g., LGm vs. LG2) at similar rostrocaudal locations upon stimulation of a given homonymous or heteronymous nerve/branch.

Differences in the responses of raphe nuclei to repetitive somatosensory stimulation

The responses of single units in raphe (R.) nuclei dorsalis, magnus, pallidus, and obscurus to repetitive stimulation of peripheral nerves were studied in the chloraloseanesthetized cat. Low-intensity electrical stimuli (1.5 T) which activated the largediameter fibers were applied to the common peroneal and lateral gastrocnemius nerves at 0.25, 0.5, 1 and 2 Hz. Responses evoked at each frequency were compared with the control response evoked at 0.1 Hz. All units isolated demonstrated response decrements during periods of stimulation ≥ 0.5 Hz. The long-term effects of repetitive stimulation, however, varied among the four nuclei. After 150 stimulus presentations at 0.5, 1, or 2 Hz, sensory responsiveness decreased in R. dorsalis but was enhanced in caudal R. obscurus units. Changes in the responsiveness of the other two nuclei were not significantly different from control. Responses to twin pulses indicated that R. neurons are not well suited to relay rapid, repetitive stimuli. The functional significance of these observations has implications for the role of the raphe in habituation and in the modulation of sensory traffic to higher centers. Raphe responses are also contrasted to the known responses of reticular formation neurons to repetitive stimulation.

The effect of lesions in the neural crest on the formation of synaptic connexions in the embryonic chick spinal cord.

1. The pattern of synaptic activity in lateral gastrocnemius (l.g.) motoneurones in the lumbar spinal cord of chick embryos (Stage 44-45, 19-21 d of incubation) has been examined using intracellular recording. In the motoneurones of normal chick embryos, stimulation of different peripheral, sciatic nerve branches gave rise to characteristic synaptic responses. Stimulation of the lateral gastrocnemius nerve caused a monosynaptic e.p.s.p. which was graded by the intensity of nerve stimulation. Stimulation of synergistic muscle afferents also caused a brief latency e.p.s.p., followed by longer latency excitatory and inhibitory synaptic potentials. Stimulation of antagonistic muscle afferents or cutaneous afferents gave rise to longer latency inhibitory and excitatory synaptic potentials respectively.2. The synaptic activity of l.g. motoneurones was also recorded in embryos in which short segments of the lumbar neural crest had been destroyed by microcautery at 3 d of incubation (Stage 18). The embryos developed without sensory ganglia and dorsal roots in the corresponding region.3. At 19-21 d of incubation, the amplitude of the l.g. e.p.s.p. of l.g. motoneurones in deafferented segments was on the average only a half to a third of the amplitude seen in motoneurones of intact spinal segments. However, both the l.g. and synergist e.p.s.p.s were larger than those seen in acutely deafferented segments of normal embryos.4. In spite of the weak monosynaptic input from l.g. and synergistic afferents, the pattern of synaptic activity evoked by antagonistic muscle afferent or cutaneous afferent stimulation was not different from normal. This was even the case for gastrocnemius motoneurones in which no early e.p.s.p. could be evoked by stimulating the l.g. or synergistic muscle nerves.5. No muscle spindles could be seen in sections of l.g. muscles from embryos with extensive lesions of the lumbosacral neural crest. Incomplete lesions of l.g. segments reduced the number of spindles in the muscle.6. These results suggest that when motoneurones are deprived of part of their normal synaptic input before the formation of peripheral connexions, the identity of the motoneurones (in terms of the origin of their synaptic input) is preserved. Missing synaptic inputs are either replaced by appropriate afferent fibres, if they are available, or not at all. The chick sensory ganglion cells with monosynaptic connexions to motoneurones appear to be unable to compensate significantly for peripheral or central defects in the innervation of the hind limb. They behave as if their developmental possibilities were quite rigidly determined at an early embryonic stage.

Nociceptor-driven dorsal horn neurones in the lumbar spinal cord of the cat

Single dorsal horn neurones have been recorded extracellularly in the lumbar spinal cord of cats anaesthetized with chloralose. Cold block at L 1 was used to provide reversible spinalization. The location of the units in the dorsal horn was marked by the electrophoretic deposition of pontamine sky blue from the recording microelectrode. There was a clear somatotopic representation of the ventrolateral surface of the foot in the L 6 segment. Thirty-five of the 46 units recorded in the marginal zone of the L 6 dorsal horn (lamina I) could only be excited by volleys in small afferent fibres and by noxious stimulation of the skin in the foot regions and were termed class 3 cells. The remaining 11 units could, in addition, be excited by sensitive cutaneous mechanosensitive afferent units — they were class 2 units. The ‘specific’ nociceptor-driven neurones could be divided into 2 subclasses on the basis of their excitability by afferent fibres. Class 3 (a) were excited only by Aδ cutaneous afferents and class 3 (b) by both Aδ and C cutaneous afferents. Some of the latter were also excited by Aδ and C afferent fibres in the lateral gastrocnemius nerve. When tested by natural stimulation all class 3 cells were excited by noxious mechanical stimuli, but only the 3 (b) units were effectively excited by heating the skin. This discharge in 3 (b) units could be suppressed by electrical stimulation of large (group II) cutaneous myelinated afferent fibres and a similar effect could be produced in responses evoked by Aδ and C afferent volleys. Additional inhibition was accomplished by electrical stimulation of the slower myelinated cutaneous (Aδ or group III) afferent fibres. The excitability of the class 3 cells was greater in spinal preparations but the tonic descending inhibition was weaker than the apparently similar descending tonic inhibition of class 2 cells. The results are discussed with reference to pain mechanisms.

Alpha motoneuron responses to natural stimuli in decerebellate cats

The responses of single alpha motoneurons to various ‘natural’ stimuli were recorded from dissected ventral root filaments in lightly anesthetized control and recently decerebellate cats. The identity of the motoneurons was established by recording the unitary responses to orthodromic stimulation of nerves innervating the triceps surae muscles in the hindlimb. We recorded only the activity of units which responded to electrical stimulation of the medial or lateral gastrocnemius nerves, though some units responded to stimulation of both these nerves, and some also responded to stimulation of the posterior tibial nerve. The distributions of unit latencies and amplitudes did not differ between the two groups of animals. Baseline firing rates were also similar. However, unit firing rates in response to neck extension, ipsilateral hindfoot dorsi- or ventral-flexion were significantly higher in decerebellate than in control cats. The discharge rates in response to other stimuli, including neck flexion, pinna stimulation, and noxious stimulation in the hindlimb, were not significantly different between the two groups. Since the alpha motoneurons showed an increase in responsiveness only to some stimuli, specific mechanisms probably explain this release. The most important mechanism seems to be a loss of inhibition normally exerted by the cerebellum on vestibular and joint proprioceptive reflexes. The pathways to the ventral horn cells are thought to involve the vestibulo-spinal and reticulo-spinal tracts.