Tonic descending inhibition of the spinal cardio-sympathetic reflex in the cat

Electrical stimulation of the left inferior cardiac nerve elicited a two-component reflex potential (spinal and supraspinal reflexes) in the ipsilateral white ramus T3 from which recordings were made in chloralose-anaesthetised cats. Reversible interruption of all spinal pathways achieved by cooling the spinal cord at C2/C3 produced an enhancement of the spinal reflex and abolished the supraspinal reflex, the latter usually being the more prominent reflex potential prior to spinal cord block. The spinal cord block-induced increase in the amplitude of the spinal reflex was, however, less than the increase observed during stimulation of the somatic intercostal nerve T4. Recordings of the afferent volley following cardiac nerve stimulation and analysis of the stimulus-reflex response relationship in neuraxis-blocked cats indicated that the spinal reflex as determined here was activated by A delta afferent fibres. However, if stimulus strength was raised above C-fibre threshold, spinal cord block revealed in addition a late spinal reflex response. In some cases, the appearance of this late potential was accompanied by a secondary decline of the earlier spinal reflex potential, possibly indicating C-fibre-mediated afferent inhibition. Neither baroreceptor activation nor denervation had any effect on spinal reflex amplitudes. Pharmacologically, clonidine given i.v. to cats with a blocked neuraxis reduced the spinal reflex amplitudes to pre-block values, an action which could be antagonised by the subsequent administration of the alpha 2-adrenoceptor antagonist rauwolscine. When given to non-pretreated cats with intact neuraxis, however, neither rauwolscine nor its analog yohimbine were capable of inducing a persistent release from tonic inhibition. The results suggest that both purely visceral and somato-visceral reflexes are subject to tonic descending inhibition, but they do not support the hypothesis that a catecholamine is the responsible transmitter mediating this inhibition.     

Plasticity of urinary bladder reflexes evoked by stimulation of pudendal afferent nerves after chronic spinal cord injury in cats

Bladder reflexes evoked by stimulation of pudendal afferent nerves (PudA-to-Bladder reflex) were studied in normal and chronic spinal cord injured (SCI) adult cats to examine the reflex plasticity. Physiological activation of pudendal afferent nerves by tactile stimulation of the perigenital skin elicits an inhibitory PudA-to-Bladder reflex in normal cats, but activates an excitatory reflex in chronic SCI cats. However, in both normal and chronic SCI cats electrical stimulation applied to the perigenital skin or directly to the pudendal nerve induces either inhibitory or excitatory PudA-to-Bladder reflexes depending on stimulation frequency. An inhibitory response occurs at 3–10 Hz stimulation, but becomes excitatory at 20–30 Hz. The inhibitory reflex activated by electrical stimulation significantly (P < 0.05) increases the bladder capacity to about 180% of control capacity in normal and chronic SCI cats. The excitatory reflex significantly (P < 0.05) reduces bladder capacity to about 40% of control capacity in chronic SCI cats, but does not change bladder capacity in normal cats. Electrical stimulation of pudendal afferent nerves during slow bladder filling elicits a large amplitude bladder contraction comparable to the contraction induced by distension alone. A bladder volume about 60% of bladder capacity was required to elicit this excitatory reflex in normal cats; however, in chronic SCI cats a volume less than 20% of bladder capacity was sufficient to unmask an excitatory response. This study revealed the co-existence of both inhibitory and excitatory PudA-to-Bladder reflex pathways in cats before and after chronic SCI. However our data combined with published electrophysiological data strongly indicates that the spinal circuitry for both the excitatory and inhibitory PudA-to-Bladder reflexes undergoes a marked reorganization after SCI.     

Neural organization of the viscero-cardiac reflexes

The reflex responses evoked in the postganglionic nerves to the heart were tested in chloralose-anaesthetized cats. Electrical stimulation of the A delta afferent fibres from the left inferior cardiac nerve evoked spinal and supraspinal reflex responses with the onset latencies of 36 ms and 77 ms respectively. The most effective stimulus was a train of 3-4 electrical pulses with the intratrain frequency of 200-300 Hz. Electrical stimulation of the high threshold afferent fibres (C-fibres) from the left inferior cardiac nerve evoked the reflex response with the onset latency of 200 ms. The C-reflex was present in intact animals and disappeared after spinalization. The most effective stimulus to evoke this reflex was a train of electrical pulses delivered at a frequency of 1-2 Hz with an intratrain frequency of 20-30 Hz. The most prominent property of the C-reflex was its marked increase after prolonged repeated electrical stimulation. We conclude that: (1) viscero-cardiac sympathetic reflexes may be organized at the spinal and supraspinal level; (2) viscero-cardiac sympathetic reflexes evoked by stimulation of the A delta and C afferent fibres from the left inferior cardiac nerve have different central organization.     

Functional organization of long ascending propriospinal pathways linking lumbo-sacral and cervical segments in the cat

In high spinal cats propriospinal pathways ascending from lumbo-sacral levels of the spinal cord can mediate strong excitatory and inhibitory changes in reflexes to different groups of motoneurones supplying muscles of the forelimb. Discharges evoked by electrical stimulation of hindlimb nerves could be evoked in 41% of experiments in the motoneurones of pectoralis major and minor. The latency of the discharge (8–18 msec) could be shortened by increasing the repetition frequency of the stimulus, the greatest reduction occurring in the range 1–4 Hz. Contralateral hindlimb nerves were less effective and the discharge generally occurred at a latency 1–2 msec longer than for ipsilateral nerves.Monosynaptic reflexes to pectoralis major and deep radial motoneurones supplying the physiological flexor muscles were strongly facilitated by hindlimb nerve stimulation, ipsilateral nerves being more effective than contralateral. Monosynaptic reflexes to latissimus dorsi showed a reciprocal pattern of conditioning, being depressed by ipsilateral and facilitated by contralateral hindlimb extensor nerves, the flexor nerves giving the reverse pattern. Monosynaptic reflexes to median and ulnar nerves supplying physiological extensor muscles were not significantly affected by hindlimb nerve conditioning.Polysynaptic reflexes to pectoralis major and deep radial motoneurones received initial strong facilitation followed by prolonged depression, ipsilateral hindlimb nerves being more effective than contralateral. In latissimus dorsi a reciprocal pattern similar to that for monosynaptic reflex testing was found. Polysynaptic reflexes to median and ulnar motoneurones received only prolonged depression.The hindlimb afferent nerves responsible for the discharge in forelimb motoneurones and for the facilitation and depression of forelimb reflexes include groups II and II muscle afferents and group II skin afferents, especially from quadriceps and sartorius muscles, and sural and superficial peroneal nerves, respectively.The ascending long propriospinal pathways are influenced bilaterally from hindlimb nerves and are located in the lower thoracic segments in the ventrolateral funiculus. The pathways mediate effects on ipsilateral and contralateral forelimb reflex systems, the ipsilateral projections being dominant. Part of the long ascending projection terminates monosynaptically on the motoneurones of pectoralis major. It is likely that group II afferents from ipsilateral quadriceps muscle activate the ascending tract monosynaptically and those from contralateral quadriceps disynaptically.The hypothesis is suggested that long propriospinal paths primarily represent intrinsic links between hindlimb and forelimb ‘motor centres’. The pattern of long ascending influences to groups of forelimb motoneurones corresponds closely to the sequences of hindlimb and forelimb stepping observed in normal cats. A functional role in stepping is therefore proposed for long ascending propriospinal pathways.     

Somatosplanchnic reflex discharges in rats

Splanchnic efferent reflex discharges caused by electrical stimulation of limb afferent nerves or intercostal afferent nerves were studied in chloralose-urethane anesthetized rats. Stimulation of the limb afferent nerve produced late supraspinal reflex discharges via group II and III afferent excitation. Stimulation of the intercostal afferent nerve produced early spinal reflex discharges via group II and III afferent excitation and also late spinal reflex discharges via group IV afferent excitation. Intercostal afferent nerve stimulation seemed to strongly depress the splanchnic late supraspinal reflex discharges.     

Efferent activity in the phrenic nerve during the startle reflex in chloralose-anesthetized cats

Reflex activity in the phrenic nerve was studied in chloralose anesthetized cats during development of somatic startle reflexes in limb and lower intercostal nerves. It was shown that the main component of this activity during low-threshold reflexes evoked by acoustic, tactile and low-threshold somatic afferent stimulation was depression of phrenic inspiratory activity. The following reflex discharges were prevalent components of phrenic responses to high-threshold afferent stimulation: early, propriospinal (intercostal-to-phrenic reflex) and late, suprasegmental ones. The latter were of two types: inspiratory (observed mainly during inspiration in about 75% of experiments) and expiratory (observed during expiration in 25% of experiments) which could be classified as "phrenic startle reflexes". Modulation of all responses during the respiratory cycle was described. Structural characteristics of reflex responses evoked in the phrenic nerve by stimulation of various respiratory and nonrespiratory bulbar sites as well as their respiratory modulation have been analyzed. Organization of possible neurophysiological mechanisms of phrenic responses during startle reflexes is discussed.     

Electrophysiological analysis of the ascending and descending components of the micturition reflex pathway in the rat

Electrophysiological techniques were used to examine the organization of the spinobulbospinal micturition reflex pathway in the rat. Electrical stimulation of afferent axons in the pelvic nerve evoked a long latency (136 +/- 41 ms) response on bladder postganglionic nerves, whereas stimulation in the dorsal pontine tegmentum elicited shorter latency firing (72 +/- 25 ms) on these nerves. Transection of the pelvic nerve eliminated these responses. Firing on the bladder postganglionic nerves was evoked by stimulation in a relatively limited area of the pons within and close to the laterodorsal tegmental nucleus (LDT) and adjacent ventral periaqueductal gray. Stimulation at sites ventral to this excitatory area inhibited at latencies of 107 +/- 11 ms the asynchronous firing on the bladder postganglionic nerves elicited by bladder distension. Electrical stimulation of afferents in the pelvic nerve evoked short latency (13 +/- 3 ms) negative field potentials in the dorsal part of the periaqueductal gray as well as long latency (42 +/- 7 ms) field potentials in and adjacent to the LDT. The responses were not altered by neuromuscular blockade. Similar responses were elicited by stimulation of afferent axons in the bladder nerves. The sum of the latencies of the ascending and descending pathways between the LDT and the pelvic nerve (i.e. 72 ms plus 42 ms = 114 ms) is comparable although somewhat shorter (22 ms) than the latency of the entire micturition reflex. These results provide further evidence that the micturition reflex in the rat is mediated by a spinobulbospinal pathway which passes through the dorsal pontine tegmentum, and that neurons in the periaqueductal gray as well as the LDT may play as important role in the regulation of the micturition.     

The possibility of different effects of primary afferent depolarization on spinal reflexes]

Intensive depolarization of central primary afferent terminals evoked by strong stimulation of afferent nerves or dorsal root produces recurrent discharges which may be recorded as antidromic dorsal root reflexes. It is shown that the discharges are simultaneously propagating in the dorso-ventral direction and thus produce facilitation of spinal reflexes. The obtained results allow suggesting the existence of two types of influences of the primary afferent depolarization on the reflex transmission to the spinal cord.     

Somato-vesical reflexes in chronic spinal cats

The effects of afferent volleys in hindlimb cutaneous and muscle nerves on vesical tone and contractility and on the discharges in pelvic nerves to the bladder were measured in anesthetized CNS-intact and 2-19 months chronic spinal cats. In chronic spinal cats volleys in group III and IV fibers increased the tone of the quiet, empty bladder (excitatory somato-vesical reflex). The same volleys inhibited the slow, large, rhythmic micturition contractions of the expanded bladder (inhibitory somato-vesical reflex). In CNS intact cats single or short tetanic volleys induced a reflex discharge in pelvic vesical nerve branches with 3 distinct components. These reflexes could be observed during micturition contractions, not markedly between the contractions or when the bladder was empty and quiet. The latencies of the 3 components were 90, 320 and 770 ms, respectively. The two early components (AI- and A2-reflex) were evoked by volleys in group II and III hindlimb afferents. The late component (C-reflex) was induced by group IV volleys. In chronic spinal cats a group II and III-induced A-reflex (latency 90 ms) and a group IV-induced C-reflex (latency 340 ms) were observed. The central pathways and the physiological significance of the various somato-vesical reflexes are discussed.     

Effect of ammonium acetate on depolarization processes in the central terminals of primary afferents]

Experiments on cats determined that ammonium acetate injected intravenously (2-4 mM/kg) supressed the processes of primary afferent depolarization (PAD) which are thought to be responsible for the presynaptic inhibition of spinal reflexes. The supression was transient and proceeded in paralle to depression of postsynaptic inhibition of monosynaptic reflexes. Ammonium acetate slightly decreased the amplitude of the negative postsynaptic potentials recorded form the dorsal surface of lumbar cord in response to stimulation of hind limb afferent nerves and increased polysynaptic reflex discharges in appropriate ventral roots. These findings make it unlikely that the ammonium depression of PAD is a result of impairment of interneuronal activity. A suggestion is made that ammonium depression of PAD results from diminition of the EMF for synaptic currents producing PAD.     

Colonosphincteric electromyographic responses to sacral root stimulation: evidence for a somatosympathetic reflex

Abstract  The aim of the present study was to determine the effects of selectively stimulating the afferent fibres running in the dorsal sacral roots (S1, S2, S3) and the somatic (radial and sciatic) nerves on colonic and internal anal sphincter (IAS) electromyographic (EMG) activity in anaesthetized cats to try to understand how sacral nerve stimulation can improve fecal continence in human. Electrically stimulating the afferent fibres present in the sacral dorsal roots and somatic nerves inhibited the colonic spike potential frequency ( n  = 97) and increased the slow variations in the sphincteric membrane potential ( n  = 76). These effects were found to have disappeared after administering an α-noradrenergic receptor blocker ( n  = 64) or sectioning the sympathetic efferent fibres innervating these organs ( n  = 69) suggesting the involvement of the sympathetic system in the effects observed. Moreover, no significant differences were observed between the effects of sacral dorsal root vs somatic nerve stimulation on colonic and sphincteric EMG activity. In conclusion, the data obtained here show that neurostimulation applied to the sacral spinal roots may improve fecal continence by inhibiting colonic activity and enhancing IAS activity via a somatosympathetic reflex.     

Spinal cholinergic modulation of cardiovascular tone and a somatosympathetic reflex response

The purpose of this study was to examine the relationship between spinal cholinergic pressor neurons and a somatosympathetic reflex response in rats. Intrathecal (IT) injection of the cholinesterase inhibitor, neostigmine (NEO), produced marked pressor and tachycardic responses without any changes in respiratory parameters. On the other hand, stimulation of the sciatic nerve produced increases in both cardiovascular and respiratory (tidal volume, minute volume, respiratory rate) responses. These cardiorespiratory responses to nerve stimulation were inhibited by IT NEO. A pressor response could also be induced by topical application of NEO to the surface of lower spinal cord, which was not altered by prior dorsal rhizotomy. These results indicate that two independent cholinergic systems exist in the spinal cord, one of which participates in the inhibitory modulation of the somatosympathetic reflex, and the other which mediates a sympathoexcitatory response. It is unlikely that the pressor response to spinal administration of NEO is mediated through this somatosympathetic response.     

Responses of sympathetic postganglionic neurons to peripheral nerve stimulation in pigeon (Columba livia).

Reflex changes in heart rate and arterial blood pressure can be elicited in pigeons with high cervical transection by stimulation of brachial or lumbosacral peripheral and spinal nerves. This extends the phenomenon of spinally mediated, somatosympathetic reflexes to another vertebrate class. In a preliminary attempt to explore the spinal circuitry mediating these reflexes, the responses of single sympathetic postganglionic neurons were studied during spinal and peripheral nerve stimulation. With stimulation and recording at the same spinal segment, calculation of the central delay suggests the segmental reflex circuitry may be relatively simple, possibly trisynaptic. As the distance between stimulating and recording sites increases, postganglionic neuronal responsiveness decreases and becomes more variable. However, there is clear evidence that lumbosacral afferents can activate postganglionic neurons at brachial levels, indicating an effective propriospinal circuitry for somatosympathetic reflexes. Experiments on birds with intact spinal cords demonstrate that these spino-spinal pathways are also functional in the intact animal. While the segmental reflex is not different in the intact bird, the propriospinal pathways do behave somewhat differently, possible suggesting tonic central control.     

Characteristics of sympathetic reflexes evoked by electrical stimulation of phrenic nerve afferents.

In chloralose-anaesthetized cats, sympathetic reflex responses were recorded in left cardiac and renal nerve during stimulation of afferent fibres in the ipsilateral phrenic nerve. In cardiac nerve, a late reflex potential with a mean onset latency of 75.6 +/- 13.8 ms was regularly recorded which, in 20% of the experiments, was preceded by an early, very small reflex component (latency between 35 and 52 ms). In contrast, in renal nerve only a single reflex component after a mean latency of 122.1 +/- 13.1 ms was observed. Bilateral microinjections of the GABA-agonist muscimol into the rostral ventrolateral medulla oblongata resulted in a nearly complete abolition of sympathetic background activity and in an 88% reduction of the late reflex amplitude with only small effects on the latency of the evoked potentials. Under this condition, an early reflex component was never observed to appear. After subsequent high cervical spinalization, the residual small potentials which persisted after bilateral muscimol injections were completely abolished and in cardiac nerve an early reflex potential with a mean latency of 45 +/- 10 ms was observed in all but one experiment. The early reflex was therefore referred to as a spinal reflex component which, however, is suppressed in most animals with an intact neuraxis. In the renal nerve a spinal response was only observed in one experiment after spinalization. The results suggest that sympathetic reflexes evoked by stimulation of phrenic nerve afferent fibres possess similar spinal and supraspinal pathways as previously described for somato-sympathetic and viscero-sympathetic reflexes.(ABSTRACT TRUNCATED AT 250 WORDS)     

Longitudinal distribution of negative cord dorsum potentials following stimulation of afferent fibres in the left inferior cardiac nerve

Cord dorsum potentials were recorded along the spinal cord following electrical stimulation of afferent fibres of the left inferior cardiac nerve in chloralose anaesthetized cats. The potentials were more pronounced in spinal than in intact cats. Afferent fibres which generated cord dorsum potentials in the cervical spinal cord were localized mainly in T2 and T3 and to a smaller extent in C8 and T1 dorsal roots. The responses consisted of two waves: with short (7.0 ms; N3 wave) and long (56 ms; N4 wave) latency to the onset of potentials. N3 and N4 waves were generated by group III and group IV afferent fibres, respectively. The N3 wave was maximal at C8 and T1 spinal cord level and could be detected at least 5-6 segments rostrally from the level of afferent input responsible for its generation. The N4 wave could be detected at least 4 segments rostrally from its afferent fibre input. We conclude that afferent fibres from the left inferior cardiac nerve activate neurones in the cervical spinal cord. The implications of such finding are discussed.     

Effects of morphine on somatocardiac sympathetic reflexes in spinalized cats

The effects of morphine on sympathetic reflexes, recorded in the inferior cardiac nerve, to myelinated A and unmyelinated C afferent stimulation were tested in 17 acutely spinalized cats. Stable sympathetic A and C reflexes of short latency (approximately 30 ms and 140 ms in the case of the ulnar nerve, respectively) could be recorded in the inferior cardiac sympathetic nerve to stimulation of somatic A and C afferents in the ulnar and upper thoracic intercostal nerves, ipsilaterally. Spinal sympathetic A reflexes, which were primarily evoked from stimulation of A delta afferent fibers, could be elicited from more segmental levels than could sympathetic C reflexes. Additionally, smaller reflexes, only from A afferent fiber activation, were identified from stimulations on the contralateral side of the body. Small doses of morphine (0.02 mg kg-1, i.v.) proved to be ineffective at altering sympathetic A and C reflexes, while somewhat larger doses (0.2 mg kg-1, i.v.) produced a clear 62% decrease in C reflexes and a 33% decrease in A reflexes, Dosages of 1 and 2 mg kg-1 severely depressed both A and C reflexes. All of the above effects of morphine administration were completely and immediately reversible by naloxone (i.v.). The results are discussed with regard to the effects of morphine on sympathetic A and C reflexes in CNS intact, anesthetized cats.     

Pudendal-to-bladder reflex in chronic spinal-cord-injured cats

The effects of pudendal nerve stimulation on reflex bladder activity were investigated in cats with chronic spinal cord injury (6-12 months) under alpha-chloralose anesthesia. Electrical stimulation of the pudendal nerve on one side at different frequencies and intensities induced either inhibitory or excitatory effects on bladder activity. The inhibitory effect peaked at a stimulation frequency of 3 Hz and gradually decreased at lower or higher frequencies. The inhibitory effect could occur at stimulation intensities between 0.3 and 1 V (pulse width 0.1 ms) and increased at intensities up to 10 V. Stimulation of the central end of transected pudendal nerve also inhibited bladder activity, indicating that afferent axons in pudendal nerve are involved. Nerve transections also showed that both hypogastric and pelvic nerves might be involved in the inhibitory pudendal-to-bladder spinal reflex. Pudendal nerve stimulation at 20 Hz and at the same intensities (1-10 V) elicited a bladder excitatory response. Although this excitatory effect could not sustain a long lasting bladder contraction at small bladder volumes, it did induce continuous rhythmic bladder contractions at large bladder volumes. This study indicated the possibility of developing a neuroprosthetic device based on pudendal nerve electrical stimulation to restore micturition function after spinal cord injury.     

An intracellular study of the synaptic input to sympathetic preganglionic neurones of the third thoracic segment of the cat

In chloralose anaesthetized, paralyzed and artificially ventilated cats intracellular recordings were obtained from sympathetic preganglionic neurones (SPN) of the third thoracic segment of the spinal cord identified by antidromic stimulation of the white ramus T3. The synaptic input to SPNs was assessed, in cats with intact neuraxis or spinalized at C3, by electrical stimulation of segmental afferent fibres in intercostal nerves and white rami of adjacent thoracic segments and by stimulation of the ipsi- and contralateral dorsolateral funiculus and of the dorsal root entry zone of the cervical spinal cord. In both preparations SPNs showed on-going synaptic activity which predominantly consisted of excitatory post-synaptic potentials (EPSPs). Inhibitory post-synaptic potentials (IPSPs) were rarely observed. EPSPs were single step (5 mV) or, less frequently, large (up to 20 mV) summation EPSPs. The proportion of SPNs showing very low levels of on-going activity was markedly higher in spinal than in intact cats. Stimulation of somatic and sympathetic afferent fibres evoked early EPSPs (amplitude 3 mV, latency 5-22.3 ms), and late, summation EPSPs (amplitude up to 20 mV, latency 27-55 ms). Early and late EPSPs were evoked in nearly all SPNs in which this synaptic input was tested in the intact preparation (from 79-93% of the SPNs). In spinal cats, early EPSPs were evoked in 88% of the SPNs, whereas late EPSPs were recorded only in half of the neurones. No evidence for a monosynaptic pathway from these segmental afferent fibres to SPNs was obtained. In both intact and spinal cats, stimulation of the dorsolateral funiculus evoked early and late EPSPs in SPNs. Late EPSPs were recorded in 70% and 37% of the SPNs in intact and spinal cats, respectively. Early EPSPs, however, were evoked in all neurones. The early EPSPs evoked by stimulation of the dorsolateral funiculus had several components which are suggested to arise from stimulation of descending excitatory pathways with different conduction velocities. The following conduction velocities were calculated in intact (spinal) cats: 9.5-25 m/s (7.8-13.2 m/s), 5.7-9.5 m/s (5.5-7.8 m/s), 3.8-5.7 m/s (3.2-5.5 m/s), and 2.6-3.8 m/s (2.1-3.2 m/s). EPSPs of these various groups were elicited in a varying percentage in SPNs. EPSPs of the most rapidly conducting pathway were subthreshold for the generation of action potentials; some EPSPs of this group had a constant latency suggesting a monosynaptic pathway to SPNs. Stimulation of the dorsal root entry zone at the cervical level yielded essentially the same results as stimulation of the dorsolateral funiculus.(ABSTRACT TRUNCATED AT 400 WORDS)     

Differential Effects of a Distant Noxious Stimulus on Hindlimb Nociceptive Withdrawal Reflexes in the Rat

Recent studies indicate that the nociceptive withdrawal reflexes to individual muscles are evoked by separate reflex pathways. The present study examines whether nociceptive withdrawal reflexes to different muscles are subject to differential supraspinal control in rats. A distant noxious stimulus was used to activate a bulbospinal system which selectively inhibits 'multireceptive' neurons (i.e. neurons receiving excitatory tactile and nociceptive inputs) in the dorsal horn of the spinal cord. Withdrawal reflexes, recorded with electromyographic techniques in single hindlimb muscles, were evoked by standardized noxious pinch. Thirty-seven rats, anaesthetized with halothane and nitrous oxide, were used. Whereas withdrawal reflexes to the extensor digitorum longus and brevis, tibialis anterior and biceps posterior muscles were strongly inhibited, reflexes to interossei muscles were potentiated during noxious pinch of the nose. Reflexes to peronei muscles were not significantly changed. The effects on the reflexes usually had an onset latency of <0.5 s and outlasted the conditioning stimulation by up to 2 s. The monosynaptic la reflex to the deep peroneal nerve, innervating dorsiflexors of the digits and ankle, was not significantly changed during noxious pinch of the nose. Hence, the inhibitory effects on the hindlimb withdrawal reflexes induced by the conditioning stimulation were presumably exerted on reflex interneurons. It is concluded that nociceptive withdrawal reflexes to different hindlimb muscles are differentially controlled by descending pathways activated by a distant noxious stimulus. The results support our previous conclusion that there are separate nociceptive withdrawal reflex pathways to different hindlimb muscles.     

The cutaneous contribution to the hamstring flexor reflex in the rat: an electrophysiological and anatomical study

The location and properties of the cutaneous receptive fields responsible for detecting the flexor withdrawal reflex in the posterior head of biceps femoris (pBF) and semitendinosus (ST) components of the hamstring muscle have been examined in unanaesthetized decerebrate rats, spinalized at T10-T11. Single alpha-motoneurone efferents were recorded from the nerve to pBF and the principal head of ST and their responses to ipsi- and contralateral hindlimb skin stimulation investigated. The efferents to both muscles characteristically had a low or absent background discharge and they all had mechanoreceptive fields on the ipsilateral foot. The mechanical threshold of these fields was high with no response to light touch or brush. Fifty-four percent of these units also had a smaller and weaker contralateral mechanoreceptive field. The only apparent difference between ST and pBF efferents was that more ST efferents had contralateral fields than pBF units. Noxious, hot and cold thermal stimuli applied to the ipsilateral foot activated 56% of the efferents. Mustard oil, a chemical irritant, produced a long-lasting flexor response when applied to the ipsilateral foot. The responses of these efferents to stimulation of A beta, A delta and C cutaneous afferents in the sural nerve were also studied. Short latency reflexes were elicited in all efferents by A beta inputs, longer latency reflexes were elicited in 64% by A delta inputs and very long latency responses with long afterdischarges were found in 73% of the units to C inputs. Retrograde labelling of the hamstring motoneurones with WGA-HRP indicated that they lay in ventrolateral lamina IX extending from the caudal portion of the third lumbar segment to the junction of the 5th and 6th lumbar segments. Transganglionic labelling of small diameter primary afferent terminals in the dorsal horn of cutaneous nerves innervating the foot revealed that the longitudinal distribution corresponded closely with that of the hamstring motor nucleus. The flex-or reflex in the spinal rat provides a useful model therefore, for studying how the input in nociceptive afferents is processed and transformed within the spinal cord, to produce appropriate outputs.