Stimulation Of Superficial Radial Nerve Research Articles

Changes in intra- and interlimb reflexes from forelimb cutaneous afferents after staggered thoracic lateral hemisections during locomotion in cats.

In quadrupeds, such as cats, cutaneous afferents from the forepaw dorsum signal external perturbations and send inputs to spinal circuits to co-ordinate the activity in muscles of all four limbs. How these cutaneous reflex pathways from forelimb afferents are reorganized after an incomplete spinal cord injury is not clear. Using a staggered thoracic lateral hemisections paradigm, we investigated changes in intralimb and interlimb reflex pathways by electrically stimulating the left and right superficial radial nerves in seven adult cats and recording reflex responses in five forelimb and ten hindlimb muscles. After the first (right T5-T6) and second (left T10-T11) hemisections, forelimb-hindlimb co-ordination was altered and weakened. After the second hemisection, cats required balance assistance to perform quadrupedal locomotion. Short-, mid- and long-latency homonymous and crossed reflex responses in forelimb muscles and their phase modulation remained largely unaffected after staggered hemisections. The occurrence of homolateral and diagonal mid- and long-latency responses in hindlimb muscles evoked with left and right superficial radial nerve stimulation was significantly reduced at the first time point after the first hemisection, but partially recovered at the second time point with left superficial radial nerve stimulation. These responses were lost or reduced after the second hemisection. When present, all reflex responses, including homolateral and diagonal, maintained their phase-dependent modulation. Therefore, our results show a considerable loss in cutaneous reflex transmission from cervical to lumbar levels after incomplete spinal cord injury, albeit with preservation of phase modulation, probably affecting functional responses to external perturbations. KEY POINTS: Cutaneous afferent inputs co-ordinate muscle activity in the four limbs during locomotion when the forepaw dorsum contacts an obstacle. Thoracic spinal cord injury disrupts communication between spinal locomotor centres located at cervical and lumbar levels, impairing balance and limb co-ordination. We investigated cutaneous reflexes from forelimb afferents during quadrupedal locomotion by electrically stimulating the superficial radial nerve bilaterally, before and after staggered lateral thoracic hemisections in cats. We showed a loss/reduction of mid- and long-latency homolateral and diagonal reflex responses in hindlimb muscles early after the first hemisection that partially recovered with left superficial radial nerve stimulation, before being reduced after the second hemisection. Targeting cutaneous reflex pathways from forelimb afferents projecting to the four limbs could help develop therapeutic approaches aimed at restoring transmission in ascending and descending spinal pathways.

Enhanced somatosensory feedback modulates cutaneous reflexes in arm muscles during self-triggered or prolonged stimulation.

Somatosensory feedback plays important roles in regulating all animal movement. The effects of sensory feedback on spinally mediated neural excitability are widely studied using cutaneous electrical stimulation paradigms. Cutaneous reflex amplitudes are reduced when stimulation is self-triggered instead ofexternally triggered. Altered spinal excitability and motor output are also observed following sustained stimulation with various parameters. Our purpose was to probe for interactions between mode and duration by investigating muscle responses following enhanced cutaneous stimulation. Fifteen neurologically intact participants were recruited. Cutaneous reflexes in the extensor carpi radialis (ECR) were evoked with brief (15ms, 300Hz) or sustained (300ms, 50Hz) stimulation trains. Stimulation was applied to the superficial radial or median nerve at the wrist and triggered by: (1) a computer program (random-triggered); (2) muscle contraction (EMG-triggered); (3) the participant pressing a button themselves (button-triggered). During each condition, isometric contractions were performed with ECR muscle activity maintained at 10, 25, 35, and 50% of maximal voluntary contraction. Stronger inhibitory reflexes were found following brief superficial radial nerve stimulation was EMG-triggered suggesting that modulation of cutaneous reflex excitability is specific to the timing when sensory 'cues' are applied during muscle contraction. No difference was observed following sustained stimulation applied to the superficial radial nerve meaning that brief and sustained stimulation affect the cutaneous pathways differentially. Nerve-specific responses were found between superficial radial and median nerve stimulation, such that greater inhibition was induced by EMG-triggered sustained stimulation to the median nerve. These observations are critical in moving beyond pathway phenomenology toward targeted sensory enhancement and amplified motor output in rehabilitation and training.

The impact of evoked cutaneous afferents on voluntary reaching movement in patients with Parkinson’s disease

Objective. Resting tremor may compound the effects of bradykinesia to further prolong the initiation of voluntary movement in patients with Parkinson’s disease (PD). However, the interaction between resting tremor and voluntary movements in these PD patients has not been well understood. Recently, we demonstrated that cutaneous afferents evoked by surface stimulation of superficial radial nerve can inhibit resting tremor effectively. The inhibition appears to take effect via spinal interneuronal pathways. This study evaluates how evoked cutaneous afferents would impact the performance of voluntary movements in PD subjects when tremor is inhibited. Approach. Ten PD patients with tremor and eight age-matched control subjects were recruited to participate in this study. Both groups of subjects performed fast reaching movements, while cutaneous stimulation was delivered during reaching tasks on or off randomly. Kinematic performance, such as reaction time (RT), movement time (MT), and movement variability, as well as muscle synergy of tasks were evaluated and compared to assess the impact of evoked cutaneous afferents on movement performances. Main results. Results indicated that the cutaneous stimulation significantly reduced RT in PD patients by 17.7%; but had an insignificant effect on RT in control subjects. Cutaneous stimulation, however, caused a significantly longer MT both in control subjects (8.6%) and in PD subjects (15.7%). Movement variability was not significantly altered in both groups of subjects by the cutaneous stimulation. Muscle synergy analysis revealed that cutaneous stimulation affected the power spectral densities (PSD) of time profiles of muscle synergies more significantly than the vector patterns of synergies in both control subjects and PD subjects. Significance. These findings provide evidence that tremor increases the RT of voluntary motor control in PD patients, and demonstrate that cutaneous stimulation reduces the RT of voluntary movements significantly, in addition to suppressing tremor, yet without interrupting voluntary control of movements.

Double-Shock Stimulation of the Superficial Radial Nerve Compared With Standard Medial Plantar Nerve Conduction in the Early Detection of Asymptomatic Diabetic Neuropathy: A Pilot Study.

Diabetes mellitus is a major public health problem. Diabetic polyneuropathy (DP) is one of the most common complications of diabetes mellitus. The early detection of DP is very important for timely treatment of symptoms and preventative foot care. Participants were sorted into 3 age- and sex-matched groups: 20 "healthy" individuals; 21 diabetic patients without DP symptoms, "asymptomatic"; and 24 diabetic patients suffering from symptoms consistent with DP, "symptomatic." All study participants had normal results on conventional nerve conduction studies. All groups underwent both medial plantar mixed nerve conduction (as a single-shock stimulation technique) and superficial radial nerve conduction (double-shock stimulation) measurements. Interstimulus intervals of 2 to 8 ms were used to record sensory nerve action potentials (SNAP) 1 and SNAP 2 for both stimuli. We found statistically significant decreases in medial plantar NAPs' amplitude and conduction velocity, and SNAP1/SNAP2 ratios between the three groups, especially at smaller interstimulus intervals. Both medial plantar mixed nerve conduction and double-shock superficial radial nerve stimulation are reliable methods for the early detection of asymptomatic DP. However, the medial plantar mixed nerve technique is easier and less time-consuming.

Neurophysiological study of the radial nerve variant in the innervation of the dorsomedial surface of the hand.

Sensory innervation of the dorsomedial surface of the hand usually depends on the dorsal ulnar nerve (DUN). Innervation in this area by the superficial radial nerve (SRN) has been described as a normal variant. We studied 358 patients using nerve conduction of the DUN and SRN with dorsomedial recording. Each hand was classified into usual innervation (only DUN response), mixed variant (response of both nerves), or complete variant (only SRN response). Mixed innervation was found in 14.2% of hands and complete innervation was found in 6.8% of hands, mostly unilaterally. No statistically significant differences were observed in age, sex, or clinical suspicion between usual and variant innervation. The potential amplitude after SRN stimulation was greater in the complete variant. It is important to know the characteristics of this variant in order to avoid diagnostic errors and to prevent iatrogenic lesions in surgery performed on the dorsum of the wrist. Muscle Nerve 58: 732-735, 2018.

Effects of wrist position on reciprocal inhibition and cutaneous reflex amplitudes in forearm muscles

In the leg, amplitudes of cutaneous reflexes and reciprocal inhibition are significantly affected by joint and limb position. Comparatively little is known about such modulation in the arm. In this study, amplitudes of reciprocal inhibition (from median nerve stimulation near elbow) and cutaneous reflexes (from median or superficial radial nerve stimulation at the wrist) were measured in forearm muscle extensor carpi radialis with the hand pronated or neutral during graded voluntary activation. Significant correlations with muscle activation were found for reciprocal inhibition and cutaneous reflex amplitudes at both positions. Only cutaneous reflexes from superficial radial nerve were modulated by wrist position. This study reveals that effect of limb position is nerve-specific in cutaneous reflexes and not significant on reciprocal inhibition in the arm. This has implications for measurement and study design in those who have mobility and motor activation challenges (e.g. neurotrauma) that affect hand function.

A common neural element receiving rhythmic arm and leg activity as assessed by reflex modulation in arm muscles.

Neural interactions between regulatory systems for rhythmic arm and leg movements are an intriguing issue in locomotor neuroscience. Amplitudes of early latency cutaneous reflexes (ELCRs) in stationary arm muscles are modulated during rhythmic leg or arm cycling but not during limb positioning or voluntary contraction. This suggests that interneurons mediating ELCRs to arm muscles integrate outputs from neural systems controlling rhythmic limb movements. Alternatively, outputs could be integrated at the motoneuron and/or supraspinal levels. We examined whether a separate effect on the ELCR pathways and cortico-motoneuronal excitability during arm and leg cycling is integrated by neural elements common to the lumbo-sacral and cervical spinal cord. The subjects performed bilateral leg cycling (LEG), contralateral arm cycling (ARM), and simultaneous contralateral arm and bilateral leg cycling (A&L), while ELCRs in the wrist flexor and shoulder flexor muscles were evoked by superficial radial (SR) nerve stimulation. ELCR amplitudes were facilitated by cycling tasks and were larger during A&L than during ARM and LEG. A low stimulus intensity during ARM or LEG generated a larger ELCR during A&L than the sum of ELCRs during ARM and LEG. We confirmed this nonlinear increase in single motor unit firing probability following SR nerve stimulation during A&L. Furthermore, motor-evoked potentials following transcranial magnetic and electrical stimulation did not show nonlinear potentiation during A&L. These findings suggest the existence of a common neural element of the ELCR reflex pathway that is active only during rhythmic arm and leg movement and receives convergent input from contralateral arms and legs.

Medial and lateral branches of the superficial radial nerve: Cadaver and nerve conduction studies

To establish a reliable technique for nerve conduction study (NCS) of medial and lateral branches of the superficial radial nerve (SRN). The distribution of SRN and its branches were examined in 10 cadavers. Based on the cadaveric study, anti-dromic nerve conduction studies were performed on 31 healthy subjects. The SRN became superficial approximately 8.4 cm from the radial styloid process (RS) and divided into medial and lateral branches 4.9 cm proximal to RS. The medial branch divided into two branches: proximal to extensor pollicis longus (EPL) tendon in 16 hands. The lateral branch ran parallel and 0.2 cm lateral to abductor pollicis longus (APL) tendon. Sensory nerve action potential (SNAP) was significantly shorter in latencies when recording over snuff box, compared with that from the EPL and APL tendons. SNAP obtained from APL tendon was significantly smaller in amplitude, compared with those from snuff box and EPL tendon. Anti-dromic stimulation of SRN should be performed 10 cm proximal to the recording electrode. The optimal recording position for each branch of SRN is snuff box for medial branch, and lateral to APL tendon for lateral branch. The SRN can be evaluated appropriately by NCS of medial and lateral branches of SRN.

Short-Term Plasticity of Spinal Reflex Excitability Induced by Rhythmic Arm Movement

Rhythmic arm movement reduces Hoffmann (H)-reflex amplitudes in leg muscles by modulation of presynaptic inhibition in group Ia transmission. To date only the acute effect occurring during arm movement has been studied. We hypothesized that the excitability of soleus H-reflexes would remain suppressed beyond a period of arm cycling conditioning. Subjects used a customized arm ergometer to perform rhythmic 1-Hz arm cycling for 30 min. H-reflexes were evoked before, during, and after arm cycling via stimulation of the tibial nerve in the popliteal fossa. The most important finding was that the H-reflex amplitudes were significantly suppressed during and <or=20 min after arm cycling had been terminated. Thus remote arm cycling can induce adaptive plasticity in the soleus H-reflex pathway that persists beyond the period of conditioning. In an additional experiment, the prolonged effect of arm cycling combined with cutaneous superficial radial (SR) nerve stimulation was investigated. Cutaneous stimulation cancelled the prolonged suppression of H-reflex amplitude induced by arm cycling. Because SR nerve stimulation facilitates soleus H-reflex via reducing the level of Ia presynaptic inhibition, persistence in presynaptic inhibitory pathways is suggested as the underlying neural mechanism. The simplest explanation of this observation is plateau potential-like behavior of interneurons mediating presynaptic inhibition of Ia afferent transmission.

A propriospinal-like contribution to electromyographic responses evoked in wrist extensor muscles by transcranial stimulation of the motor cortex in man.

We tested the hypothesis that some of the electromyographic (EMG) responses elicited in preactivated forearm muscles by transcranial stimulation of the human motor cortex are produced by activity in a disynaptic corticospinal linkage involving propriospinal-like interneurones with cell bodies in the spinal C3-4 segments. The experimental design incorporated a previous observation that stimulation of afferents in the superficial radial nerve inhibits propriospinal-like neurones projecting to the extensor carpi radialis (ECR) muscle. Surface EMG responses were recorded from the active ECR muscle after transcranial electrical or magnetic stimulation over the motor cortex. In random trials, single conditioning stimuli at twice perceptual threshold were given to the superficial radial nerve at the wrist at different times before a cortical shock. When the cortex was stimulated electrically, the conditioning stimulus suppressed the EMG responses when the interval between the shocks was 11 ms or more. This was about 3.5 ms longer than the minimum time calculated for a possible direct cutaneous effect on spinal motoneurones. The time course of suppression began earlier and was more complex during magnetic stimulation of the cortex. It is argued that this difference is due to the repetitive I waves generated by the magnetic shock. Whether electrical or magnetic stimulation was used, the first 1-3 ms of the EMG response was relatively unaffected by superficial radial nerve stimulation at any interstimulus interval, whereas clear suppression was seen in the later portion of the response. In contrast, if the EMG response in ECR was suppressed by a conditioning stimulus to the median nerve at the elbow, then all portions of the EMG response were inhibited including the first 1-3 ms. The median nerve effect is thought to be due to direct reciprocal inhibition of the extensor motoneurones. Thus sparing of the initial part of the cortically evoked response with superficial radial stimulation suggests that the latter type of inhibition occurs at a premotoneuronal level. The timing of the effect is compatible with the explanation that corticospinal excitation is produced in ECR motoneurones through both monosynaptic and disynaptic (including propriospinal premotoneuronal) pathways, with superficial radial nerve inhibition being exerted at the propriospinal level.

The mode of synaptic activation of pyramidal neurons in the cat primary somatosensory cortex: an intracellular HRP study

A total of 141 pyramidal neurons in the cat primary somatosensory cortex (SI) were recorded intracellularly under Nembutal anesthesia (7 in layer II, 43 in layer III, 8 in layer IV, 58 in layer V and 25 in layer VI). Most neurons were identified by intracellular staining with HRP, though some layer V pyramidal neurons were identified only electrophysiologically with antidromic activation of medullary pyramid (PT) or pontine nuclear (PN) stimulation. Excitatory synaptic potentials (EPSPs) were analyzed with stimulation of the superficial radial nerve (SR), the ventral posterolateral nucleus (VPL) in the thalamus and the thalamic radiation (WM). The pyramidal neurons in layers III and IV received EPSPs at the shortest latency: 9.1 +/- 2.1 ms (Mean +/- S.D.) for SR and 1.6 +/- 0.7 ms for VPL stimulation. Layer II pyramidal neurons also responded at a short latency to VPL stimulation (1.7 +/- 0.5 ms), though their mean latencies for SR-induced EPSPs were relatively longer (10.6 +/- 1.9 ms). The mean latencies were much longer in layers V and VI pyramidal neurons (10.2 +/- 2.4 ms and 2.9 +/- 1.5 ms in layer V pyramidal neurons and 9.9 +/- 2.5 ms and 2.8 +/- 1.6 ms in layer VI pyramidal ones, respectively for SR and VPL stimulation). The comparison of the latencies between VPL and WM stimulation indicates that most layer III-IV pyramidal neurons and some pyramidal cells in layers II, V and VI received monosynaptic inputs from VPL. These findings are consistent with morphological data on the laminar distribution of thalamocortical fibers, i.e., thalamocortical fibers terminate mainly in the deeper part of layers III and IV with some collaterals in layers V, VI and II-I. The time-sequences of the latencies of VPL-EPSPs indicate that corticocortical and/or transcallosal neurons (pyramidal neurons in layers II and III) fire first and are followed by firing of the output neurons projecting to the subcortical structures (pyramidal neurons in layers V and VI).

Effect of midbrain raphe nucleus stimulation on somatosensory evoked potential in cat

Experiments were carried out to clarify the effect of serotonin [5-hydroxytryptamine (5-HT)] upon the somatosensory evoked potential (SEP) elicited by superficial radial nerve stimulation in cat, and the following results were obtained: (i) in untreated (control) animals, a significant reduction occurred in amplitude of the SEP in response to conditioning stimulation of the dorsal raphe nucleus in the midbrain, within 100 ms of the interval between conditioning and test stimuli; (ii) in 5-HT depleted animals prepared by intraperitoneal injection of p-chlorophenylalanine (PCPA), the amplitude of SEP significantly increased in comparison with that of nontreated animals. Conditioning stimulation of the raphe nucleus in 5-HT depleted animals failed to cause any remarkable change in the SEP; (iii) the diminuation of the SEP by raphe nucleus conditioning reappeared following intraventricular administration of a large dose of 5-HT (200 micrograms) in the PCPA-treated animals. These results have demonstrated that the serotonergic system with its cell bodies located in the raphe nucleus of the midbrain has the function of reducing the amplitude of the SEP, thus suggesting that it acts as a suppressive control on the excitability of the cortex to sensory input.

Contribution of cutaneous and muscle afferent fibres to cortical SEPs following median and radial nerve stimulation in man

Somatosensory evoked potentials were recorded after stimulation of motor and cutaneous nerves in the upper limb. Stimulation of the thenar motor branch of the median nerve and the deep motor branch of the radial nerve produced only broad, ill-defined and small-amplitude scalp-recorded responses. In contrast, stimulation of purely cutaneous nerves (digital and the superficial radial) gave responses of large amplitude. The cortical responses following combined deep and superficial radial nerve stimulation were of smaller amplitude than the two individual responses combined. These findings suggest that, contrary to an earlier report (Gandevia et al. 1984), muscle afferents do not make a major positive contribution to the scalp-recorded cortical responses produced by electrical stimulation of mixed nerves in the upper limb.

Influence of the ipsilateral hemisphere upon somatosensory evoked potential in cats.

Somatosensory evoked potentials (SEPs) were recorded on the skull corresponding to the (left) SI area by (right) superficial radial nerve stimulation. Amongst the various components of the SEP, special attention was directed to the negative component (N15) with a latency of approximately 15 ms. Changes in this potential followed by conditioning stimulation of the ipsilateral (right) hemisphere were observed and the following results were obtained: (i) when conditioning stimuli were applied to the contralateral (left) superficial radial nerve, the ipsilateral (right) thalamic VPL nucleus and the ipsilateral (right) sensory cortex, the amplitude of N15 decreased to 65-80% of the control level at C-T intervals less than 100 ms and (ii) following functional elimination of the unilateral sensory cortex by KCl application, the amplitude of N15 recorded at the opposite side significantly increased. In this condition, the inhibitory effects of the ipsilateral thalamus and contralateral peripheral nerve disappeared. From these observations, ipsilateral homologous cortex may well have an inhibitory influence upon the near field potential (N15) of the SEP.

Origin of the initial negative potential (N15) recorded on the skull by superficial radial nerve stimulation in the cat.

The origin of the initial prominent negative potential with a latency of about 15 msec (N15), recorded on the skull by superficial radial nerve stimulation was studied in the cat, and the following results were obtained. In the direct recording from the cortex, SEPs were elicited from the SI and SII areas, as well as the lateral gyrus and anterior suprasylvian gyrus. Among various wave forms recognized in the SI area, diphasic positive-negative (P-N) potential obtained from the postero-lateral part was most distinct, representing the primary evoked potential. Responses recorded on the dura and those directly on the cortex showed similar wave patterns over many sites. Although different forms of evoked potentials were recorded extensively over the skull, the most prominent negative potential was elicited at the site corresponding to the postero-lateral part of the SI area. The latency of this potential was approximately in agreement with that of the negative component of the P-N potential recorded on the cortex. Based on the intracortical laminar analysis of P-N potential, the positive component of this potential was assumed to reflect the activity of cells in the deeper layer of the cortex. The negative component, on the other hand, might represent activities of apical dendrites of the cortex. From the result of functional elimination of the cortex, however, this positive component was thought to contain potentials from far field neural structures. N15 recorded on the skull completely disappeared during cortical spreading depression.(ABSTRACT TRUNCATED AT 250 WORDS)

Changes of somatosensory evoked potential accompanying ischaemia and hypoxia in cats.

Changes of evoked potential accompanying haemorrhagic hypotension and hypoxia were investigated on cats to evaluate the usefulness of SEP as a monitor in an intensive care unit (ICU), and the following results were obtained. Positive-negative diphasic potential was elicited at posterior sigmoid gyrus(PSG) by contralateral superficial radial nerve stimulation. This potential was recorded at the restricted area of the posterior border of PSG and regarded as primary somatosensory evoked potential. In the initial stage of haemorrhagic hypotension, both positive and negative components of SEP occasionally increased in amplitude. In profound hypotension in which CBF fell to less than the critical level of 30 ml 100 g-1 min-1, the latency was retarded and the amplitude was decreased. At CBE less than 10 ml 100 g-1 min-1, SEP disappeared. Within the range of CBF between 10 and 30 ml 100 g-1 min, a close correlation was noted between CBF and SEP amplitude. Transient increase of SEP amplitude was also observed during hypoxia induced by inhalation of nitrogen gas. (3) In normal state SEP was decreased in amplitude by conditioning stimulation of the nucleus lateralis posterior (LP nucleus) of the thalamus. This might be explained by the fact that intracortical inhibitory interneurons were activated by stimulation of LP nucleus. After haemorrhagic hypotension and hypoxia, however, the inhibitory effect on SEP elicited by LP nucleus stimulation attenuated or disappeared. Because of the initial impairment of the inhibitory interneurons by ischaemia and hypoxia, the amplitude of SEP might increase transiently. In conclusion, the authors thought that SEP might be less useful than EEG in ICU, because of its insensible change to hypoxia and ischaemia.

Regional differences in evoked response latencies in the cat motor cortex (area 4 gamma).

The latencies of responses evoked in the cat motor cortex (area 4 gamma) on contralateral forelimb nerve stimulation were measured. In the caudolateral part, the latencies were 4-5 ms on ulnar and superficial radial nerve stimulation. Further, rostromedially the response latencies were longer, 7-9 ms. The latency-to-peak of the evoked responses was also measured and showed a parallel increase. In many cases, inflexions could be observed on the rising phase of the evoked potential, indicating that it consisted of several components. In most experiments, four components (I-IV) were observed. Their mean latencies were 4.3, 6.0, 7.2 and 8.5 ms, respectively. The early components were largest in the caudolateral part of area 4 gamma and decreased when recording further rostromedially such that, in the anterior sigmoid gyrus, usually only component IV could be recorded. The findings suggest that the cat motor cortex is functionally divided into regions which can be identified by their peripheral input latencies.

A non-linear analysis of afferent modulatory activity in the cat somatosensory system

The effects of prior input on afferent activity in the somatosensory system of the cat have been studied using both twin-pulse experiments and random stimulus trains for which evoked activity is characterized by a set of mathematical functions known as kernels. The kernels quantitatively characterize transformational processes occurring in the system. The first-order kernel is analogous to a classical averaged evoked potential, but to temporally interacting stimuli; the second-order kernel may be likened to a generalized recovery function. Responses to superficial radial nerve stimulation were obtained at sensory (S1) cortex, thalamic n. ventralis posterolateralis (VPL), and the nucleus cuneatus of the dorsal column nuclei (DCN). Comparison of first-order kernels with averaged evoked potentials showed good agreement in chronic awake, chronic anesthetized, and acute anesthetized animals. Second-order kernels indicated complex interactions with an overall depression of subsequent response amplitudes. Comparison between second-order kernels and the results of the two-pulse experiments at both S1 cortex and VPL showed good agreement for early latency response components, but significant deviations for later components suggest higher than second-order non-linearities for this system.

Labelling of cerebellar mossy fiber afferents with intra-axonal horseradish peroxidase

Several unidentified units recorded in the intermediate zone of lobules IV and V of the cerebellar cortex responded to superficial radial nerve stimulation with a distinctive short train of 5 to 11 low amplitude spikes (less than 5 mV) at a frequency of 800 to 1200 Hz. Responses of this type have in the past been attributed to mossy fiber afferents. Following the electrophysiological characterization horseradish peroxidase was successfully injected by iontophoresis into four axon terminals. The labelled axons presented the typical structure of mossy fibers with terminal rosettes and corresponded to the morphology of these axons described in Golgi material.

Influence of drugs on evoked potentials in the cat cerebellum: III. Effects of morphine hydrochloride (2).

In intact cats, morphine hydrochloride (2.0 mg/kg, i.v.) depressed spontaneous discharge of the Purkinje cell in 8 of the 15 neurons tested. These effects of morphine were antagonized by naloxone hydrochloride (0.4 mg/kg, i.v.). This opiate significantly decreased the evoked potentials produced by superficial radial nerve (SR) stimulation in the molecular layer of the cerebellar cortex. Although morphine decreased the potentials evoked by SR or tooth pulp stimulation in the nucleus fastigii and nucleus interpositus, this drug hardly influenced them in the nucleus dentatus. In the decerebrated cats, morphine significantly increased the potentials evoked in the cerebellar cortices by SR stimulation. In the somatosensory area I (SSA-I)- or motor area (MA)-lesioned cats, morphine decreased the cerebellar potentials evoked by SR stimulation, but increased them in the SSA-I- and MA-lesioned cats. Naloxone antagonized these effects of morphine. From our results, it appears that the decreasing effects of morphine on the cerebellar potentials evoked by SR stimulation may be affected by the functions of the SSA-I and MA of the cerebral cortex and that morphine may influence the cerebellum.