Stimulation Of Muscle Afferents Research Articles

Regulation of afferent connectivity in the adult spinal cord by nerve growth factor.

During development, nerve growth factor (NGF) regulates the density and character of peripheral target innervation (Barde, Neuron, 2, 1525 - 1534, 1989; Ritter et al., Soc. Neurosci. Abstr., 17, 546.2, 1991); its role in adult animals is less well defined. Here we have asked if the availability of growth factors such as NGF in peripheral tissues can influence the pattern of primary afferent connections in the CNS. Using osmotic minipumps, we raised the levels of NGF in rat skeletal muscle in vivo, a tissue where the levels of this factor are normally very low (Korsching and Thoenen, Proc. Natl. Acad. Sci. USA, 80, 3513 - 3516, 1983; Shelton and Reichardt, Proc. Natl. Acad. Sci. USA, 81, 7951 - 7955, 1984; Goedert et al., Mol. Brain Res., 1, 85 - 92, 1986). After 2 weeks of treatment we asked if the sensory neurons innervating this tissue showed an altered strength and distribution of connections with dorsal horn neurons. The contralateral (vehicle-treated) muscle, and totally untreated animals, served as controls. In normal and vehicle-treated animals, electrical stimulation of muscle afferents excited relatively few neurons in the dorsal horn, and these generally showed only weak responses. In contrast, on the NGF-treated side many more dorsal horn neurons in the lumbar enlargement of the spinal cord were excited by muscle afferents. The increased responsiveness could not be explained by a generalized increase in dorsal horn excitability, since spontaneous activity was not enhanced, nor by a change in A-fibre-mediated inhibitions from the treated afferents. Thus, these afferents appeared to establish new synaptic connections or strengthened previously weak ones as a result of increased neurotrophic factor availability. The data suggest that, in the adult rat, the levels of growth factors in peripheral targets may be used to regulate an appropriate degree of afferent connectivity within the central nervous system.

Stimulation of craniofacial muscle afferents induces prolonged facilitatory effects in trigeminal nociceptive brain-stem neurones

Stimulation of small-diameter afferents supplying deep tissues has been shown to increase the excitability of spinal cord neurones responding to cutaneous afferent inputs. This facilitation has been implicated as integral central mechanisms of deep pain that may contribute to the tenderness and spread and/or referral of pain following injury of deep tissues. In view of the recent documentation of deep craniofacial afferent inputs, as well as cutaneous afferent inputs to the trigeminal (V) spinal tract nucleus, we wished to determine the effects of deep inputs excited by the small-fibre irritant mustard oil on trigeminal nociceptive neurones. The extracellular activity of single brain-stem neurones was recorded in subnuclei caudalis and oralis of the V spinal tract nucleus of anaesthetized rats. The neurones were classified as low-threshold mechanosensitive (LTM), wide dynamic range (WDR) and nociceptive specific (NS) on the basis of their cutaneous mechanoreceptive field properties and their responses evoked by electrical stimulation of their cutaneous afferent inputs. Injection of 5% mustard oil (2–5, μl) into the deep masseter muscle produced a facilitatory effect in 12 of 27 nociceptive neurones tested in caudalis and in 5 of 12 nociceptive neurones in oralis. This effect was reflected in an expansion of the cutaneous mechanoreceptive field, an increase in spontaneous activity or an increase in responsivity to electrical stimulation of cutaneous afferent inputs to the neurones. The facilitation was reversible and typically became apparent within 3–5 min of the injection, reached its peak at 5–10 min, and lasted for 20–30 min. These findings document for the first time the sensitivity of the cutaneous receptive field properties of V brain-stem nociceptive neurones to facilitatory influences from deep craniofacial afferents, and point to possible mechanisms that may contribute to the tenderness, spread and referral of pain that may result from trauma to deep craniofacial tissues.

Reciprocal Ia inhibition between elbow flexors and extensors in the human.

1. Reciprocal inhibition between elbow flexor and extensor muscles (biceps and triceps brachii) has been investigated in nine healthy subjects. Two techniques were used to assess changes in motoneurone excitability after stimulation of antagonist muscle afferents: (1) monosynaptic reflexes elicited by a mechanical stimulation of the distal muscle tendon (tendon tap); (2) post-stimulus time histograms (PSTH) of voluntarily activated motor units. 2. Electrical stimulation of the antagonist muscle nerve produced a short-latency and short-lasting inhibition of the flexor and extensor motoneurones. The amount of this inhibition was found to be similar in both motor nuclei. 3. The inhibition could be evoked with conditioning electrical stimuli as low as 0.7 x motor threshold (MT) or by very weak tendon taps applied to the antagonist tendon. In the former case the threshold of this inhibition was found to be consistently increased after raising the threshold of Ia afferent fibres by a long-lasting muscle vibration. Since a contribution from cutaneous afferent fibres was ruled out, it is concluded that this inhibition was Ia in origin. 4. Post-stimulus time histograms of voluntarily activated triceps and biceps motor units were made following electrical stimulation of homonymous and antagonist muscle afferents. This enabled an estimate of the central synaptic delay of the inhibitory process. An average central delay of 0.94 ms in excess of that of monosynaptic facilitation was found, thus suggesting that the inhibitory process could be mediated by only one interneurone. 5. A conditioning reflex discharge elicited in the antagonist muscle by a tendon tap depressed or suppressed this inhibition. This depression was maximal when the reflex discharge was elicited 10-20 ms before the conditioning stimulus for the inhibition and never lasted more than 30 ms. It is argued that the only mechanism compatible with such a depression is the inhibitory activity of Renshaw cells acting on the pathway mediating reciprocal inhibition. 6. We conclude that group Ia afferent fibres from elbow extensor and flexor muscles project monosynaptically onto Ia inhibitory interneurones to mediate disynaptic reciprocal inhibition of antagonist motoneurones.

Respiratory muscle activation by limb muscle afferent stimulation in anesthetized dogs

In 10 chloralose anaesthetized and spontaneously breathing dogs, we assessed the effect of limb muscle afferents on the peak integrated EMG activities of the genioglossus, alae nasi, costal diaphragm, parasternal intercostal, triangularis sterni, and transverse abdominis muscles. The influence of vagal and baroreceptor afferents were eliminated by vagotomy and perfusion of carotid sinuses at a constant pressure. Muscle afferents were activated by stimulating the central end of the gastrocnemius nerve for 1 min at 40 Hz and at different voltages. Stimulation at voltages equal to 5, 10 and 20 times twich-threshold increased minute ventilation to 165, 216 and 250% of pre-stimulation values, respectively, which was achieved by increasing breathing frequency (shortening of the inspiratory and expiratory times) and tidal volume. The activity of the parasternal intercostal and alae nasi muscles increased by a similar degree to that of the diaphragm while the activities of the genioglossus and transverse abdominis were augmented to a greater degree than that of the diaphragm. On the other hand, the motor drive to triangularis sterni increased significantly only at 20 times twitch-threshold and to a lesser degree than that to the diaphragm. These results suggest that upper airway, inspiratory and expiratory rib cage and abdominal muscles may be independently regulated. Differences in the sensitivity of these muscles to the activation of limb muscle afferents can be explained by a complex pattern of central projections of these afferents on the central respiratory controllers or by intrinsic properties of the motor output of these controllers.

Intrathecal galanin blocks the prolonged increase in spinal cord flexor reflex excitability induced by conditioning stimulation of unmyelinated muscle afferents in the rat

The effect of intrathecal (i.t.) galanin (GAL) on the prolonged increase of spinal reflex excitability produced by conditioning stimulation (CS) of unmyelinated muscle afferents was studied in decerebrate, spinalized, unanesthetized rats. A CS train (1 Hz, 20 s) applied to the unmyelinated fibers in the gastrocnemius muscle nerve facilitated the ipsilateral flexor reflex for about 1 h. Pretreatment with GAL (0.1–10 μg) decreased reflex facilitation induced by the gastrocnemius nerve CS. The present results indicate that GAL is capable of blocking the prolonged increase in spinal cord excitability after stimulation of unmyelinated muscle afferents, possibly by antagonizing the facilitatory effect of tachykinins and calcitonin gene-related peptide released at the intraspinal terminals of these fibers.

Excitatory responses of the dorsal root discharge to stimulation of cutaneous and muscle afferents in the cat

Excitatory responses of the dorsal root discharge (DRD) consisting in transient increase in its frequency have been studied in non-anaesthetized low spinal cats. They were evoked by stimulation of cutaneous nerves (superficial peroneal and posterior tibial) and muscle nerves (gastrocnemius-soleus and posterior biceps-semitendinosus), recorded from the central cut ends of single fibres of L7 dorsal roots. The excitatory responses were elicited by single volleys in low threshold cutaneous and group II muscle afferents. Group III afferents of gastrocnemius-soleus nerve were also effective. Increase in strength of stimulation of posterior biceps-semitendinosus nerve from 4T to 40T which activated group III muscle afferents significantly decreased the excitatory effects of the DRD. Incidence of excitatory responses of the DRD to volleys both in low and high threshold cutaneous afferents, was 100%. Frequency of occurrence of excitatory effects to volleys in group II muscle afferents ranged from 23% to 43% (for responses to posterior biceps-semitendinosus and gastrocnemius-soleus volleys, respectively). It was increased to 47% and 72% when excitatory effects were elicited by group III afferent volleys. These findings indicate that only some types of afferent fibres evoke excitatory responses of the DRD. Variable incidence of the excitatory responses to stimulation of cutaneous and muscle afferents suggests important difference in effectiveness of connections between both types of fibres and interneurones generating the DRD.

Projection of thenar muscle afferents to frontal and parietal cortex of human subjects

There is some controversy about the projection of muscle afferents from the human upper limb to cerebral cortex and about their contribution to somatosensory evoked potentials. In 8 normal volunteers, the somatosensory projections of muscle and cutaneous afferents from the hand were recorded at 21 scalp sites, using a non-cephalic reference. Low-threshold thenar muscle afferents were selectively activated by intramuscular microstimulation. In addition, the averaged data for the projections were mapped for each individual. In each subject a focal parietal negativity was detected over the contralateral parietal cortex at a mean latency of 20.8 msec (S.D. 1.15 msec) following stimulation of thenar muscle afferents. The amplitude of the parietal ‘N20-P25’ was relatively small (mean 0.49 μV, range 0.18–1.56 μV). A small focal positivity was detected, maximal over contralateral frontal cortex at 22.8 msec (S.D. 2.05 msec) but recorded bilaterally. In all subjects subcortical positive waves (P9 and P14) were defined for the muscle afferent volley. This pattern of cortical activity was similar to that for the projection from the digital nerves of the index finger. For the cutaneous input the latency of the parietal ‘N20’ was 21.7 msec (S.D. 1.17 msec) and of the frontal ‘P22’ was 24.2 msec (S.D. 3.09 msec). The amplitude of the parietal ‘N20-P25’ was larger for the cutaneous projection (mean 1.59 μV; range 0.65–4.28 μV).

Responses of inspiratory neurons of the dorsal respiratory group to stimulation of expiratory muscle and vagal afferents

In decerebrate, paralyzed and ventilated cats, we monitored the intracellular responses of 30 inspiratory neurons of the dorsal respiratory group (DRG) to stimulation of vagal and expiratory muscle (internal intercostal and abdominal) afferents. We hypothesized that the inhibitory effects of stimulation of expiratory muscle afferents, previously reported, would block the excitatory responses of inspiratory neurons of the DRG to vagal stimulation. Although prolonged stimulus trains to expiratory muscle afferents caused respiratory phase-switching, single shocks or short trains elicited no responses in 17 bulbospinal neurons, excitatory responses in 6, and inhibitory responses in 2. Of the 4 propriobulbar neurons tested, 2 had inhibitory responses and 2 did not respond. In only 2 neurons, both bulbospinal, did conditioning stimuli to expiratory muscle afferents block or reduce the excitatory effects of vagal stimulation. These results suggest that interaction of vagal and expiratory muscle afferents, which might account for the absence of a change in inspiratory duration despite increased vagal afferent feedback at elevated end-expiratory lung volumes, does not occur within the DRG.

The cortical distribution of muscle and cutaneous afferent projections from the human foot

Somatosensory evoked potentials to electrical stimulation of muscle and cutaneous afferents from the foot were recorded in normal human subjects using multiple channels centered on the vertex and referenced to the contralateral earlobe. Low-threshold muscle afferents were selectively activated by an insulated microelectrode inserted percutaneously at the motor point of abductor hallucis. Low-threshold cutaneous and joint afferents of the hallux or second toe were stimulated with ring electrodes. The posterior tibial and sural nerves were stimulated at the ankle through surface electrodes. The cerebral distribution of the initial cortical responses (N33-P40) to stimulation of muscle afferents largely paralleled that to stimulation of its parent nerve, the posterior tibial nerve (which contains afferents of muscle, cutaneous and joint origin). They were maximal slightly posterior and ipsilateral to the vertex. The cutaneous-joint afferent projection from the hallux paralleled that from the sural nerve and both were less lateralized than the tibial and abductor hallucis projections.

Modulation of reflex responses during fictive locomotion in the forelimb of the cat

During cat forelimb fictive locomotion, short-latency reflex pathways were examined by recording nerve discharges and intracellularly from motoneurones. Stimulation of cutaneous afferents, superficial radial nerves, evoked trisynaptic excitation of the elbow flexors, biceps brachii and brachialis, and stimulation of muscle afferents, deep radial nerves, evoked oligosynaptic, i.e. monosynaptic and disynaptic excitation of the flexors. The short-latency excitatory postsynaptic potentials (EPSPs) evoked from both nerves were rhythmically modulated; they were facilitated during the flexion phase and suppressed during the extension phase. Stimulation of high threshold muscle afferents evoked EPSPs with a central delay of ca. 4.2 ms, which were depressed throughout episodes of fictive locomotion. Since the short-latency EPSPs and longer-latency EPSPs in the same motoneurone were differently influenced during fictive locomotion, the effects observed could not be explained by changes occurring at only the motoneuronal level but they probably occurred at the premotoneuronal level. In addition, short-latency cutaneous excitation of the distal muscles, innervated by the median and ulnar nerves, was little modulated during fictive locomotion.

Do muscle afferents contribute to the cervical response evoked by electrical stimulation of the median nerve in man?

The possible contribution of low threshold muscle afferents to the postsynaptic component (N13) of the cervical response evoked by electrical stimulation of the median nerve (MN) was investigated in normal subjects. Electroneurographic (ENG) and electromyographic (EMG) correlates of the reflex motoneuronal discharge (RMND) were recorded simultaneously. A. No reflex activity could be elicited by stimulation of the MN at the wrist, at least in the resting subjects, while well developed ENG (P2 efferent volley) and EMG (H reflex) monosynaptic responses occurred following stimulation of the MN at the elbow at suitable strengths. In neither case could a surface correlate of interneuronal activity evoked by muscle afferents be demonstrated. B. Recruitment curves showed that at stimulus intensities above maximal for the H reflex both P2 and H responses started to decrease until they completely disappeared, while N13 showed further enhancement. C. Subthreshold conditioning stimulation of the MN enhanced both P2 and H responses, while vibratory muscle stimuli provoked a clearcut suppression of these two responses. In contrast, N13 was completely unaffected by either manoeuvre. D. No cervical evoked activity could be detected following tendon tapping of the anterior forearm muscles in spite of the appearance of well developed cortical responses and the ENG and EMG correlates of the T reflex. E. Conditioning volleys elicited by tendon taps of the anterior forearm muscles suppressed both P2 and H responses following stimulation of the MN at the elbow without affecting the related N13 component. Conditioning supramaximal stimulation of the MN at the wrist suppressed the N13 component of the cervical response evoked by stimulation of the MN at the elbow without affecting the related reflex responses. No component chronologically related to the RMND could be recorded at the posterior neck region during suppression of N13, thus ruling out the possibility that failure to detect the RMND (as well as its interneuronal concomitants) with cervical electrodes is due to a masking effect of the N13 component. G. Conditioning tendon taps of anterior forearm muscles provoked a clearcut reduction of the primary cortical response to finger stimulation without affecting the postsynaptic component of the related cervical response. It is concluded that neither segmental (motoneuronal or interneuronal in origin) nor ascending postsynaptic impulses generated in the spinal cord by stimulation of low threshold muscle afferents contribute to N13, the latter being probably due to activation of both short and long axoned spinal neurons by cutaneous afferents.

PS. Cerebral potential evoked by selective stimulation of muscle and skin afferents in man

PS. Cerebral potential evoked by selective stimulation of muscle and skin afferents in man

Peripheral specification of sensory neurons transplanted to novel locations along the neuraxis

Thoracic dorsal root ganglia in bullfrogs contain sensory neurons that innervate the skin of the trunk and have synaptic connections in the dorsal horn of the spinal cord. The ganglion that innervates the forelimb contains, in addition to cutaneous afferents, many muscle afferents that project more ventrally in the spinal cord and make monosynaptic connections with motoneurons. In the present study, we have transplanted thoracic sensory neurons to the brachial level in tadpoles to discover whether they can innervate forelimb muscles and, if so, whether they form central connections characteristic of forelimb muscle afferents. The ganglion that normally supplies the forelimb was removed from tadpoles and replaced with 2 thoracic ganglia. After the tadpoles completed metamorphosis, the peripheral and central connections of the transplanted thoracic sensory neurons were examined with anatomical and electrophysiological techniques. When the ganglia were transplanted at stage XIV or earlier, transplanted sensory neurons innervated the forelimb and projected into the brachial spinal cord. Electrical stimulation of forelimb muscle nerves evoked impulses in the dorsal root, indicating that some centrally projecting sensory neurons were muscle afferents. Furthermore, muscle afferents were also activated by stretching muscles which suggest that they terminated on spindles. HRP labeling of the central projections revealed that transplanted sensory neurons terminated at sites characteristic of both cutaneous and muscle afferents. The pattern of synaptic connections was assessed by recording intracellularly from motoneurons. Stimulation of muscle afferents produced monosynaptic EPSPs in motoneurons. As in normal frogs, triceps muscle afferents projected more strongly to triceps motoneurons than to subscapularis and pectoralis motoneurons, while subscapularis afferents projected to all 3 types of motoneurons. Thus, the transplanted sensory neurons formed central connections appropriate to their novel peripheral targets. These observations suggest that interactions between sensory neurons and their targets may be important in determining their central connections.

Presynaptic mechanisms of changes in segmental responses accompanying the formation of a spinal locomotor generator in the cat

It is shown on unanesthetized immobilized decorticated cats that spinalization of the animal leads to depolarization of the central afferent terminals, decrease of early polysynaptic responses in motoneurons and dorsal root potentials (DRP) evoked by stimulation of the low-threshold cutaneous and muscle afferents, increase of early polysynaptic responses and DRP evoked by stimulation of high-threshold muscle afferents, reduction in the activity of intermediate nucleus interneurons mono- and polysynaptically connected with primary afferents, rise of the activity of interneurons di- and oligosynaptically connected with primary afferents. Injection of DOPA into spinal animals leads to opposite changes. Dependence between changes in the state of segmental neuronal apparatus and the level of spinal locomotor generator activity are discussed on the basis of the data obtained.

Bradykinin in reflex cardiovascular responses to static muscular contraction

We examined the contribution of bradykinin to the reflex hemodynamic response evoked by static contraction of the hindlimb of anesthetized cats. During electrical stimulation of ventral roots L7 and S1, we compared the cardiovascular responses to hindlimb contraction before and after the following interventions: inhibition of converting enzyme (kininase II) with captopril (3-4 mg/kg, n = 6); inhibition of kallikrein activity with aprotinin (Trasylol, 20,000-30,000 KIU/kg, n = 8); and injection of carboxypeptidase B (500-750 U/kg, n = 7). Treatment with captopril augmented the rise in mean arterial blood pressure and maximal time derivative of pressure (dP/dt) caused by static contraction from 21 +/- 3 to 39 +/- 7 mmHg and 1,405 +/- 362 to 2,285 +/- 564 mmHg/s, respectively. Aprotinin attenuated the contraction-induced rise in mean arterial blood pressure (28 +/- 4 to 9 +/- 2 mmHg) and maximal dP/dt (1,284 +/- 261 to 469 +/- 158 mmHg/s). Carboxypeptidase B reduced the cardiovascular response to static contraction. Thus the mean arterial blood pressure response was decreased from 36 +/- 12 to 24 +/- 11 mmHg, maximal dP/dt from 1,618 +/- 652 to 957 +/- 392 mmHg/s, and heart rate from 12 +/- 2 to 7 +/- 1 beats/min. These data suggest that stimulation of muscle afferents by bradykinin contributes to a portion of the reflex cardiovascular response to static contraction.

Responses of neurons in feline trigeminal subnucleus caudalis (medullary dorsal horn) to cutaneous, intraoral, and muscle afferent stimuli.

The extracellular activity of single neurons was recorded in subnucleus caudalis (medullary dorsal horn) of chloralose-anesthetized cats to test the effects of electrical and natural stimuli that activated afferents supplying the jaw and tongue muscles as well as the face, teeth, and intraoral mucosa. Many caudalis neurons that could be functionally classified on the basis of their cutaneous receptive-field properties as low-threshold mechanoreceptive (LTM), wide-dynamic-range (WDR), or nociceptive-specific (NS) neurons could be excited by muscle afferent stimuli. Only five neurons were encountered that received muscle afferent inputs and had no demonstrable cutaneous, dental, or mucosal input. The muscle afferent inputs were a particular feature of the cutaneous nociceptive (i.e., WDR and NS) neurons. Approximately two-thirds of this nociceptive neuronal population (n = 109) could be excited by jaw and/or tongue muscle stimulation, whereas only a small proportion of the LTM neuronal population (n = 247) was activated by muscle afferent stimulation. Neurons with a demonstrated direct axonal projection to the contralateral thalamus as well as nonprojection neurons received muscle afferent inputs. The caudalis nociceptive neurons receiving muscle as well as cutaneous afferent inputs had receptive-field properties comparable to those previously described for caudalis cutaneous nociceptive neurons; they were predominantly located in laminae I/II and V/VI, and many also received convergence of tooth pulp afferent inputs. These neurons generally had larger cutaneous receptive fields than neurons unresponsive to muscle afferent stimulation. The muscle afferent inputs were considered to be predominantly of a nociceptive character for several reasons. These included the long latency and high threshold of most neuronal responses evoked by electrical stimulation of the muscle afferents, the predominance of afferents of small diameter in some of the muscle nerves stimulated, the preferential responsiveness to the muscle afferent stimulation of neurons that were functionally identified as cutaneous nociceptive neurons, and the responsiveness of most of the neurons excited by electrical stimulation of the muscle nerves also to noxious mechanical or thermal stimulation of muscle and the injection of two or more algesic chemicals into small arteries supplying the jaw and tongue muscles. Of the algesic chemicals used in this study (7% NaCl, KCl, bradykinin, histamine, 5-HT), the first two were found to be the most effective and to cause the most rapidly induced excitation.(ABSTRACT TRUNCATED AT 400 WORDS)

Bradykinin-induced chemoreflexes from skeletal muscle: implications for the exercise reflex.

We examined the cardiovascular response to bradykinin stimulation of skeletal muscle afferents and the effect of prostaglandins on this response. Intra-arterial injection of 1 microgram bradykinin into the gracilis muscle of cats reflexly increased mean arterial pressure by 16 +/- 2 mmHg, left ventricular end-diastolic pressure by 1.6 +/- 0.6 mmHg, maximal dP/dt by 785 +/- 136 mmHg/s, heart rate by 11 +/- 2 beats/min, and mean aortic flow by 22 +/- 3 ml/min. The hemodynamic responses were abolished following denervation of the gracilis muscle. The increases in mean arterial pressure and maximal dP/dt were reduced by 68 and 45%, respectively, following inhibition of prostaglandin synthesis with indomethacin (2-8 mg/kg iv). Treatment with prostaglandin E2 (PGE2, 15-25 micrograms ia) restored the initial increase in mean arterial pressure, but not dP/dt, caused by bradykinin stimulation. Injection of PGE2 (15-30 micrograms ia) into the gracilis, without prior treatment with indomethacin, augmented the bradykinin-induced increases in mean arterial pressure and dP/dt. We conclude that small doses of bradykinin injected into skeletal muscle are capable of reflexly activating the cardiovascular system and that prostaglandins are necessary for the full manifestation of the corresponding hemodynamic response. The pattern of hemodynamic adjustment following bradykinin injection into skeletal muscle is very similar to that induced by static exercise. Therefore, it is possible that intense exercise provides a stimulus for this bradykinin-induced reflex in vivo.

Activation of lumbar spinoreticular neurons by stimulation of muscle, cutaneous and sympathetic afferents

Forty-four spinal neurons located mainly in laminae IV–VII of the L7 and S1 segments projected primarily contralaterally and to the lateral reticular nucleus. Muscle and cutaneous input was via group II and III afferents, whereas sympathetic input was via group III or IV afferents. Close-arterial injection of bradykinin and/or caosaicin excited most of the neurons so tested. These responses make these neurons likely candidates for the ascending pathway of the exercise pressor reflex.

Crossed actions of group I muscle afferents in the cat.

Reflex actions evoked by electrical stimulation of contralateral quadriceps, hamstring and gastrocnemius-soleus muscle afferents were investigated using intracellular recording from motoneurones, in chloralose-anaesthetized, acute, low spinal cats. Contralateral group I afferents were found to evoke excitatory post-synaptic potentials only occasionally. Contralateral group II afferents evoked post-synaptic potentials (excitatory to extensor motoneurones and inhibitory to flexor motoneurones) from the quadriceps nerve, whereas contralateral group III afferents evoked post-synaptic potentials (with the same pattern as quadriceps group II) from all muscles tested. Contralateral group I afferents were found to facilitate Ia reciprocal inhibition, and both the excitatory and inhibitory reflex actions from Ib afferents. This is taken to indicate an action of these afferents on interneurones interposed in ipsilateral reflex pathways.

Segmental organization of rat lateral longissimus, a muscle involved in lordosis behavior: EMG and muscle nerve recordings

Following stimulation of lumbar cutaneous and high threshold muscle afferents in urethane-anesthetized ovariectomized rats, polysynaptic multi-unit responses were recorded from muscle nerves to lateral longissimus (LL) at a mean latency of 9.5 ms. These responses could be reciprocally evoked between segments with no apparent rostro-caudal asymmetry. Polysynaptic responses habituated to repeated stimulation within 0.5–1.5 min but could be evoked again after several minutes rest. In estradiol-treated rats, 60% of the 9.5 ms responses were followed by late discharges (> 40 ms) as compared to 24% in untreated control animals ( P = 0.05). Following stimulation to the L3 and L4 motor nerves to lateral longissimus, EMG responses were recorded from overlapping fields of innervation. The conduction velocity of the fast muscle fibers was determined to be 7 m/s. The short length of most of the LL muscle fibers and the intersegmental recruitment of LL motoneurons by cutaneous afferents is compatible with the integrated action of axial muscles in the performance of lordosis behavior.