Ascending tract neurones processing information from group II muscle afferents in sacral segments of the feline spinal cord.

1. Ascending tract neurones located in the dorsal horn of sacral segments of the spinal cord have been investigated by extracellular and intracellular recording in the anaesthetized cat. The aim was to determine whether information from group II afferents that terminate within the sacral segments is conveyed to supraspinal structures and which types of neurones are involved. 2. A considerable proportion of ascending tract neurones found in the dorsal horn in the same segments as the pudendal (Onuf's) motor nucleus were excited by group II muscle afferents. The great majority (93%) of these neurones had axons ascending in ipsilateral funiculi. Spinocervical tract neurones constituted the largest proportion (82%) of such neurones, while very few spinocerebellar tract and propriospinal neurones and no postsynaptic dorsal column neurones were found among them. 3. In addition to activation by group II muscle afferents all of the neurones were strongly excited by cutaneous afferents. The most potent excitation was evoked by afferents of the posterior biceps-semitendinosus and gastrocnemius muscle nerves and by afferents of the cutaneous femoris, sural and pudendal nerves. The latencies of intracellularly recorded excitatory potentials were indicative of a high incidence of monosynaptic coupling between the afferents and ascending tract neurones. 4. The highly effective monosynaptic excitation of spinocervical tract neurones in the sacral segments by group II afferents is in contrast to the weak disynaptically mediated actions of group II afferents on such neurones in the L6-L7 segments but comparable to the actions of group II afferents on ascending tract neurones in the midlumbar segments. 5. Both the patterns of peripheral input and the latencies of synaptic actions in ascending tract neurones were similar to those in interneurones at the same locations (accompanying report). Similar information is therefore likely to be processed by both categories of neurones. 6. The role of sacral spinocervical tract neurones as a system for transmitting information from group II muscle afferents to supraspinal centres and the potential contribution of this system to the perception of limb position are discussed.     

Recurrent Control from Motor Axon Collaterals of la Inhibitory Pathways to Ventral Spinocerebellar Tract Neurones

The effects of impulses in recurrent motor axon collaterals on transmission in different inhibitory pathways to ventral spinocerebellar tract (VSCT) neurones were investigated in the cat by conditioning of IPSPs evoked in intracellularly recorded VSCT cells. Disynaptic IPSPs from large muscle spindle (la) afferenls were depressed in many but not all VSCT cells following an antidromic stimulation of ventral roots. The effect was found in VSCT neurones which themselves did not receive recurrent inhibition from motor axon collaterals. In cells with affected la IPSPs also some polysynaptic IPSPs evoked from ipsi- and contralateral group II muscle afferents and low threshold cutaneous afferents were depressed by a ventral root volley as well as disynaptic IPSPs from fibres descending on the ipsilateral side of the spinal cord. In unanesthetized preparations recurrent facilitatory potentials similar to those in motoneurones were evoked in VSCT neurones with la IPSPs. The findings indicate that some VSCT neurones receive collateral connexions from the interneurones which mediate la recipiocal inhibition to motoneurones and support the hypothesis that the VSCT conveys information about transmission in inhibitory reflex pathways to motoneurones (Lundberg 1971).     

Cortically evoked pre-and postsynaptic inhibition of impulse transmission to the dorsal spinocerebellar tract

Summary Synaptic actions evoked from primary afferents and the sensorimotor cortex in neurones of the dorsal spinocerebellar tract were investigated: 1. Stimulation of the anterior lobe of the cerebellum produced a small IPSP in only one but not in the other six neurones examined. 2. IPSPs were induced not only from group I fibres (in 41% of group I neurones) but also from cutaneous and/or high threshold muscle afferents (in 37%). 3. Stimulation of the contralateral sensorimotor cortex evoked IPSPs in 80% of group I neurones. The IPSP had a latency of 10–15 msec and lasted for 40–100 msec. EPSPs were evoked from the cortex in a small number of neurones. 4. Effects from the cortex were compared with those from primary afferents in individual neurones. The cortical IPSPs were induced independently of whether the neurone received monosynaptic EPSP from extensor or flexor group I fibres. The cortical IPSPs (or EPSPs) occurred more frequently in neurones which exhibited polysynaptic IPSPs (or EPSPs) from primary afferents. 5. The few FRA neurones encountered were all excited from the cortex.Excitability measurements of primary afferent terminals in or near Clarke's column showed that a terminal depolarization is evoked from the cortex in group Ib but not in Ia afferents.The relative importance of post-and presynaptic inhibition of transmission to the DSCT is discussed.     

Afferent connexions to Group I activated cells in the main cuneate nucleus of the cat

1. Extracellular recordings were made from a total of 240 group I activated cells in the main cuneate nucleus. Cuneothalamic relay neurones (128) were identified by antidromic stimulation of the medial lemniscus in the ventrobasal thalamic complex.2. A majority of the relay neurones were activated by afferents in only one of six dissected forelimb nerves innervating muscle groups at various joints. Even among afferents from adjacent synergistic muscles, convergence to individual neurones was infrequent.3. Some of the relay neurones received excitation from group II muscle afferents in the same nerve that provided group I excitation. Excitation from group II muscle afferents in other nerves was uncommon. Some neurones were weakly excited by cutaneous volleys.4. Inhibition of group I relay cells was produced from cutaneous afferents and group II muscle afferents. Weak inhibition was sometimes observed from group I afferents. The relay cells were also inhibited by stimulation of the cerebral cortex with a focus around the lateral end of the cruciate sulcus. A good correspondence was found between the inhibition and the depolarization of group I afferent terminals in the cuneate nucleus.5. A majority of the group I activated cells not antidromically activated from the ventrobasal complex (;non-relay cells') were excited by cortical stimulation. Excitation from cutaneous afferents and group II muscle afferents was frequently found among these cells.6. The group I activated cells were found almost exclusively in the ventral part of the nucleus.7. The pattern of convergence found in eleven group I activated cells in the dorsal horn of the spinal cord from C 2 to C 4 is described.     

Neck muscle afferent input to spinocerebellar tract cells of the central cervical nucleus in the cat

Summary Extracellular and intracellular recordings were made from spinocerebellar tract neurones of the central cervical nucleus (CCN) in C1–C3 segments of the anaesthetized cat. These neurones were identified by antidromic activation from the cerebellar peduncle. Stimulation of the ipsilateral dorsal root elicited extracellular spikes or EPSPs with a monosynaptic latency in almost all CCN neurones in the same segment (segmental input). Late excitatory effects were also observed in about one third of CCN neurones. The monosynaptic EPSP was occasionally followed by an IPSP. The excitatory input from the dorsal root to CCN neurones was extended over several segments for some CCN neurons (extrasegmental input). Monosynaptic excitation was evoked in CCN neurones after stimulation of dorsal neck muscle nerves as well; i.e. splenius (SPL), biventer cervicis and complexus (BCC), rectus capitus dorsalis, and obliquus capitus caudalis. Thresholds for this excitation were near the threshold of the nerve, suggesting that it originated from group I fibres. The component of excitation added after strong stimulation of neck muscle nerves would be attributed to group II fibres. When a CCN neurone received excitatory input from the nerve of one muscle, it was generally not affected by stimulation of other nerves in the same segment. Such muscle specificity of segmental input was the principal pattern of connexion of neck muscle afferents with CCN neurones. In some cases, however, excitatory convergence from SPL and BCC nerves onto single CCN neurones or excitation from the SPL nerve and inhibition from the BCC nerve were also observed. Nearly half of the CCN neurones received input from one muscle nerve of the same segment and not from the afferent of the same muscle of different segments, indicating a segment specificity of input. In the remaining CCN neurones, weaker excitatory effects were induced from afferents of different segments as well. In such extrasegmental effects, inputs to CCN neurones from caudal segments predominated in frequency over those from rostral segments. The origin of extrasegmental input was generally confined to the same muscle. Low threshold muscle afferents from the SPL and BCC were intraaxonally stained with HRP. The collaterals of the stained fibre distributed branchlets and terminals to the CCN, laminae VII, VIII, and motor nuclei. Two fibres responding to local muscle prodding or stretch showed a similar morphology. The findings indicated that muscle spindle afferents from primary endings projected to the CCN.Supported by a Grant-in-Aid for Scientific Research from the Ministry of Education, Science and Culture of Japan     

Axon-collateral activation by dorsal spinocerebellar tract fibres of group I relay cells of nucleus Z in the cat medulla oblongata.

1. The problem whether the group I hind limb cerebral tract and the dorsal spinocerebellar tract (DSCT) have common spinal axons has been investigated in the present study. 2. Thirty-two cells located in the nucleus Z of the cat medulla oblongata and activated by spinal fibres in the dorsolateral fascicle were selected for the study. 3. Extracellular recording from these neurones demonstrated that most of them were monosynaptically linked to spinal fibres excited by ipsilateral hind limb group I muscle afferents. The cells exhibited a restricted spatial convergence and had a limited excitatory convergence from group II muscle and from skin afferents. 4. Antidromic activation from the contralateral thalamus showed that they were bulbothalamic relay cells. 5. Cerebellar surface or depth stimulation activated 88% of the twenty-six cells tested at a short latency. With a collision technique it was demonstrated that twelve out of twenty-three (52%) of these group I relay neurones were activated by axon-collaterals of the DSCT. 6. 43% of the cells activated from the cerebellum, but not proven to be linked to the DSCT, could, nevertheless, have been excited by DSCT axon-collaterals, if it is assumed that different fibres converging with excitation on the group I relay cells were activated in the collision test.     

Do noradrenergic descending tract fibres contribute to the depression of transmission from group II muscle afferents following brainstem stimulation in the cat?

Two alpha 2 noradrenaline antagonists, idazoxan and yohimbine, were injected in midlumbar segments of the spinal cord to test whether they counteract depression of field potentials evoked by group II muscle afferents by conditioning stimuli applied in the brainstem. The tested field potentials were those evoked monosynaptically in the intermediate zone of midlumbar segments. Their depression reflected thus the depression of transmission between group II fibres and their first relay neurones. The conditioning stimuli were applied either within the ipsilateral locus coeruleus/subcoeruleus or outside these nuclei (in the raphe magnus, raphe obscurus, or cuneiform nuclei). The brainstem evoked depression of the tested field potentials (n = 12) was reduced following injection of idazoxan or yohimbine to about two thirds of that which was evoked originally but in three cases to about one half. The study leads thus to the conclusion that noradrenergic descending tract neurones contribute to the depression of transmission from group II afferents to spinal interneurones and that such noradrenergic neurones are activated by stimuli applied within as well as outside their nuclei. However, the relative contribution of monoaminergic and non-monoaminergic descending tract neurones to the control of transmission from group II afferents to these neurones remains to be established.     

Functional differentiation and organization of feline midlumbar commissural interneurones

Interneurones interconnecting the two sides of the spinal cord (commissural interneurones) are critically important for interlimb coordination, but little is known about their organization. We have examined the inputs to commissural interneurones located in the midlumbar segments with projections to contralateral motor nuclei, aiming to determine whether they form distinct subpopulations. Based on intracellular records from 78 interneurones, two major non-overlapping subpopulations were identified: one monosynaptically excited by group II muscle afferents ( n = 10), the other monosynaptically excited by reticulospinal neurones ( n = 52). Monosynaptic input from group I muscle afferents and/or from vestibulospinal tract neurones was found in those with monosynaptic reticulospinal, but not group II input, and in a few other neurones ( n = 6). Only disynaptic input from these sources was found in the remaining 10 interneurones. Disynaptic excitatory input from ipsilateral and contralateral muscle afferents and from descending tracts was distributed less selectively and might mediate coexcitation of interneurones with monosynaptic afferent or descending input. The dominant disynaptic and polysynaptic input was, however, inhibitory. IPSPs were evoked from the descending tracts in a high proportion of the commissural interneurones that were monosynaptically excited by group II afferents (55%) and from group II afferents in a high proportion of the commissural interneurones that were monosynaptically excited by reticulospinal fibres (78%). This distribution suggests that the two subpopulations are activated differentially, rather than being coactivated, in either centrally initiated movements or reflex adjustments. This would be consistent with the previous demonstration that noradrenaline differentially affects commissural neurones of the two subpopulations.     

Premotor interneurones contributing to actions of feline pyramidal tract neurones on ipsilateral hindlimb motoneurones

The aim of the study was to analyse the potential contribution of excitatory and inhibitory premotor interneurones in reflex pathways from muscle afferents to actions of pyramidal tract (PT) neurones on ipsilateral hindlimb motoneurones. Disynaptic EPSPs and IPSPs evoked in motoneurones in deeply anaesthetized cats by group Ia, Ib and II muscle afferents were found to be facilitated by stimulation of the ipsilateral, as well as of contralateral, PT. The ipsilateral actions were evoked by either uncrossed or double-crossed pathways. The results show that interneurones mediating reflex actions of muscle afferents may be activated strongly enough by PT stimulation to contribute to movements initiated by ipsilateral PT neurones and that PT actions relayed by them might be enhanced by muscle stretches and/or contractions. However, in some motoneurones disynaptic IPSPs and EPSPs evoked from group Ib or II afferents were depressed by PT stimulation. In order to analyse the basis of this depression, the transmitter content in terminals of 11 intracellularly labelled interneurones excited by PT stimulation was defined immunohistochemically and their axonal projections were reconstructed. The interneurones included 9 glycinergic and 2 glutamatergic neurones. All but one of these neurones were mono- or disynaptically excited by group I and/or II afferents. Several projected to motor nuclei and formed contacts with motoneurones. However, all had terminal projections to areas outside the motor nuclei. Therefore both inhibitory and excitatory interneurones could modulate responses of other premotor interneurones in parallel with direct actions on motoneurones.     

Cerebellar Golgi cells in the rat receive multimodal convergent peripheral inputs via the lateral funiculus of the spinal cord

We recently showed that the activity of cerebellar Golgi cells can be powerfully modulated by stimulation of peripheral afferents, in a pattern different to local Purkinje cells. Here we have examined the pathways underlying these responses. Graded electrical stimulation of muscle and cutaneous nerves revealed that long-lasting depressions and short-lasting excitations of Golgi cells were evoked by stimulation of cutaneous nerves at stimulus intensities that activated large mechanoreceptive afferents, and grew as additional afferents were recruited. In contrast, none of the neurones responded to stimulation of muscle nerves at intensities that activated group I afferents, although about half responded with long-lasting depressions, but not excitations, to stimuli that recruited group II and III afferents. Selective lesions of the spinal dorsal columns did not affect either of these types of response. After lesions of one lateral funiculus in the lumbar cord the responses evoked by stimulation of the hindlimb contralateral to the lesion were reduced or abolished, leaving responses evoked by ipsilateral hindlimb afferents unaltered. Since both ipsi- and contralateral afferents generate responses in Golgi cells, the convergence from the two sides must occur supraspinally. It is difficult to reconcile these properties with any of the direct spinocerebellar pathways or spinoreticulocerebellar pathways that have been described. Instead, it is likely that the responses are evoked via the multimodal 'wide dynamic range' neurones of the anterolateral system. Golgi cell activity may thus be powerfully enhanced or depressed during arousal via the anterolateral system.     

Rubrospinal effects on ventral spinocerebellar tract neurones.

Stimulation of the contralateral red nucleus evoked monosynaptic EPSPs in 14 of 82 ventral spinocerebellar tract neurones. In some of these cells the monosynaptic EPSP was followed by a disynaptic IPSP. The remaining cell population received di- or polysynaptic PSPs from the rubrospinal tract, either EPSPs or IPSPs or both. Convergence of the rubrospinal tract onto interneurones of the segmental pathways projecting to VSCT cells was demonstrated. Rubrospinal volleys facilitated disynaptic Ia IPSPs evoked in VSCT neurones from both flexors and extensors, as well as disynaptic Ib IPSPs. Facilitation of the Ia interneurones was disynaptic whereas facilitation of Ib interneurones was monosynaptic. Disynaptic rubrospinal EPSPs and IPSPs were facilitated by volleys in ipsi- as well as in contralateral cutaneous and high threshold muscle afferents. The complex pattern of projections from the rubrospinal tract onto VSCT neurones and the related reflex pathways gives further support to the hypothesis that these tract cells convey information on transmission through interneurones of the spinal segmental mechanisms.     

Rubrospinal Effects on Ventral Spinocerebellar Tract Neurones

Stimulation of the contralateral red nucleus evoked monosynaptic EPSPs in 14 of 82 ventral spinocerebellar tract neurones. In some of these cells the monosynaptic EPSP was followed by a disynaptic IPSP. The remaining cell population received di- or polysynaptic PSPs from the rubrospinal tract, either EPSPs or IPSPs or both. Convergence of the rubrospinal tract onto interneurones of the segmental pathways projecting to VSCT cells was demonstrated. Rubrospinal volleys facilitated disynaptic Ia IPSPs evoked in VSCT neurones from both flexors and extensors, as well as disynaptic Ib IPSPs. Facilitation of the Ia interneurones was disynaptic whereas facilitation of Ib interneurones was monosynaptic. Disynaptic rubrospinal EPSPs and IPSPs were facilitated by volleys in ipsi- as well as in contralateral cutaneous and high threshold muscle afferents. The complex pattern of projections from the rubrospinal tract onto VSCT neurones and the related reflex pathways gives further support to the hypothesis that these tract cells convey information on transmission through interneurones of the spinal segmental mechanisms.     

Interneurones in pathways from group II muscle afferents in sacral segments of the feline spinal cord.

1. Properties of dorsal horn interneurones that process information from group II muscle afferents in the sacral segments of the spinal cord have been investigated in the cat using both intracellular and extracellular recording. 2. The interneurones were excited by group II muscle afferents and cutaneous afferents but not by group I muscle afferents. They were most effectively excited by group II afferents of the posterior biceps, semitendinosus, triceps surae and quadriceps muscle nerves and by cutaneous afferents running in the cutaneous femoris, pudendal and sural nerves. The earliest synaptic actions were evoked monosynaptically and were very tightly locked to the stimuli. 3. EPSPs evoked monosynaptically by group II muscle afferents and cutaneous afferents of the most effective nerves were often cut short by disynaptic IPSPs. As a consequence of this negative feedback the EPSPs gave rise to single or double spike potentials and only a minority of interneurones responded with repetitive discharges. However, the neurones that did respond repetitively did so at a very high frequency of discharges (0.8-1.2 ms intervals between the first 2-3 spikes). 4. Sacral dorsal horn group II interneurones do not appear to act directly upon motoneurones because: (i) these interneurones are located outside the area within which last order interneurones have previously been found and (ii) the latencies of PSPs evoked in motoneurones by stimulation of the posterior biceps and semitendinosus, cutaneous femoris and pudendal nerves (i.e. the main nerves providing input to sacral interneurones) are compatible with a tri- but not with a disynaptic coupling. Spatial facilitation of EPSPs and IPSPs following synchronous stimulation of group II and cutaneous afferents of these nerves shows, however, that sacral interneurones may induce excitation or inhibition of motoneurones via other interneurones. 5. Comparison of the properties of group II interneurones in the sacral segments with those of previously studied group II interneurones in the midlumbar segments leads to the conclusion that these two populations of neurones are specialized for the processing of information from different muscles and skin areas. In addition, equivalents of only one of the two subpopulations of midlumbar interneurones have been found at the level of the pudendal nucleus: neurones with input from group II but not from group I muscle afferents. Neurones integrating information from group I and II muscle afferents and in direct contact with motoneurones thus seem to be scarce in the sacral segments.(ABSTRACT TRUNCATED AT 400 WORDS)     

Dopaminergic control of transmission from group II muscle afferents to spinal neurones in the cat and guinea-pig

The effects of dopamine and its agonists on transmission from muscle afferents to spinal neurones were investigated in the cat and guinea-pig spinal cord, by measuring the drug effects on the amplitude of monosynaptic field potentials evoked by electrical stimulation of group I and group II muscle afferents. Local iontophoretic application of dopamine, the dopamine D1/D5 agonist SKF-38393 and the D2/D3/D4 agonist quinpirole all depressed the group II field potentials evoked at the base of the dorsal horn. Group II field potentials in the intermediate zone were depressed by dopamine to a similar degree as the dorsal horn field potentials, whereas the dopamine agonists were without effect upon them. The intermediate zone field potentials evoked by group I muscle afferents were not depressed by any of the drugs. The dopamine-evoked depression of the group II-evoked field potentials in the dorsal horn in the guinea-pig spinal cord was reduced by the simultaneous application of haloperidol. The results demonstrate that dopamine receptors mediate the depression of transmission from group II muscle afferents to interneurones in the dorsal horn, but not to neurones in the intermediate zone of the spinal cord.     

Effects on the Ventral Spinocerebellar Tract Neurones from Deiters' Nucleus and the Medial Longitudinal Fascicle in the Cat

Effects from the vestibulospinal tract (VST) and from fibres descending in the medial longitudinal fascicle (MLF) on the cells of origin of the ventral spinocerebellar tract (VSCT) have been studied with intracellular recording. Out of 110 VSCT neurones, the VST evoked monosynaptic EPSPs in 27, di- or polysynaptic EPSPs in 56 and disynaptic lPSPs in 26. In 93 tested VSCT cells, MLF stimulation evoked monosynaptic EPSPs in 26, monosynaptic IPSPs in 2, di- or polysynaptic EPSPs in 25 and disynaptic IPSPs in 21, Convergence of monosynaptic EPSPs from VST and MLF was found in a small proportion of cells whereas the two descending pathways evoked reciprocal effects in another small group of neurones. Convergence of monosynaptic EPSPs from VST or MLF and from group 1 afferents was also modest. In 9 VSCT neurones there was convergence of monosynaptic excitation and disynaptic inhibition from the vestibulospinal tract and the same pattern from MLF was recorded in 9 neurones. The results are discussed in view of the hypothesis that VSCT neurones carry information on the interneuronal transmission in the spinal cord.     

Input from muscle and cutaneous nerves of the hand and forearm to neurones of the precentral gyrus of baboons and monkeys

1. The precentral bank of the Rolandic fissure of the cortical arm area has been explored with extracellular micro-electrodes in primates (baboons and monkeys) under nitrous oxide and oxygen anaesthesia, supplemented by small doses of Parkesernyl(R) and chloralose. The results in baboons and monkeys were the same.2. Single units were classified as pyramidal tract neurones or non-pyramidal tract neurones according to their antidromic responsiveness to stimuli applied in the dorsolateral funiculus at C1-2.3. Responses to electrical stimulation of the deep (motor) radial nerve, the deep palmar (motor) branch of the ulnar nerve, and the superficial (cutaneous) radial nerve could be recorded in the majority of neurones of the motor cortex provided that short trains of strong stimuli were used. Minimal responses to muscle nerve stimulation were observed in a few neurones at 1.4 x group I threshold, but most units reacted only with higher stimulus intensities (2-3 x group I threshold).4. The latencies to peripheral nerve stimulation were measured from the first peak of the incoming volley recorded at the root entry zone. The mean response latencies of pyramidal tract cells were between 20 and 25 msec; non-pyramidal tract cells were activated at slightly shorter mean latencies, the difference being significant for superficial radial nerve stimulation only (4 msec). These latencies are more than twice as long as those recorded in the postcentral gyrus, and the probability of discharge is lower than for postcentral neurones.5. A further difference between neurones of the postcentral and precentral gyrus is the pronounced convergence from different nerves and also from different modalities (cutaneous and muscle afferents) in units of the precentral cortex in contrast to units of the postcentral cortex.6. The high thresholds, necessary to activate precentral neurones by muscle nerve stimulation, make it unlikely that group I muscle afferents are involved. This is, furthermore, indicated by the lack of responsiveness to intravenous injection of succinylcholine which was, however, effective for driving neurones of the specific projection area for group I afferents, area 3a. The present experiments are consistent with the view that sensitivity of precentral neurones to muscle stretch (described in previous studies) is due to activation of secondary muscle spindle endings and their ascending pathways.7. The original hypothesis of a load compensating ;pyramidal reflex' with an oligosynaptic afferent contribution from the spindle primaries can be discarded. The present findings indicate that there is a feed-back from secondary muscle spindle afferents which, by way of a more complex pathway, can modulate the firing frequency of neurones in the motor cortex.     

Effects on the ventral spinocerebellar tract neurones from deiters' nucleus and the medial longitudinal fascicle in the cat.

Effects from the vestibulospinal tract (VST) and from fibres descending in the medial longitudinal fascicle (MLF) on the cells of origin of the ventral spinocerebellar tract (VSCT) have been studied with intracellular recording. Out of 110 VSCT neurones, the VST evoked monosynaptic EPSPs in 27, di- or polysynaptic EPSPs in 56 and disynaptic IPSPs in 26. In 93 tested VSCT cells, MLF stimulation evoked monosynaptic EPSPs in 26, monosynaptic IPSPs in 2, di- or polysynaptic EPSPs in 25 and disynaptic IPSPs in 21. Convergence of monosynaptic EPSPs from VST and MLF was found in a small proportion of cells whereas the two descending pathways evoked reciprocal effects in another small group of neurones. Convergence of monosynaptic EPSPs from VST or MLF and from group I afferents was also modest. In 9 VSCT neurones there was convergence of monosynaptic excitation and disynaptic inhibition from the vestibulospinal tract and the same pattern from MLF was recorded in 9 neurones. The results are discussed in view of the hypothesis that VSCT neurones carry information on the interneuronal ttransmission in the spinal cord.     

Excitation of Group I activated thalamocortical relay neurones in the cat

1. Extracellular recordings have been made from 134 group I activated neurones in the ventrobasal thalamic complex. One hundred of the neurones were identified as thalamocortical relay neurones by antidromic activation from the projection areas for group I afferents.2. The discharge evoked by group I volleys showed characteristic fluctuations in latency and number of spikes. The mechanisms underlying these phenomena are discussed.3. Half of the relay neurones were activated from afferents in only one of six dissected forelimb nerves innervating muscle groups at the various forelimb joints. Two-thirds of the relay neurones were activated from at least two adjacent synergistic muscles.4. The pattern of group I convergence in the thalamic neurones is compared with that at the cuneate and cortical levels of the group I pathway to the cerebral cortex. It is suggested that integration of information from synergistic muscles occurs at the thalamic level and that integration of information from muscle groups of more unrelated function occurs at the cortical level.5. A few of the thalamocortical cells (15%) were activated by group II muscle afferents, often in the same nerve as that which provided group I excitation. Weakly linked excitation from cutaneous afferents was observed in 39% of the neurones.6. Stimulation of the group I projection area in the first sensorimotor cortex at moderately high stimulus frequencies produced a trans-synaptic excitation in most of the group I activated cells.     

Corticofugal influences on transmission to the dorsal spinocerebellar tract from hindlimb primary afferents

Summary Effects from the cerebral cortex on neurones of the dorsal spinocerebellar tract (DSCT) were examined: I. In group I units (units receiving monosynaptic excitation from group I fibres) repetitive stimulation of the contralateral sensorimotor cortex usually inhibited impulse transmission from the primary afferents. The inhibition had a latency of 10–20 msec and lasted for 82-100 msec or more. Discharges induced by muscle stretch were also inhibited by the cortical stimulation. DSCT units belonging to extensors and flexors were both inhibited from the cortex. In a small percentage of group I units the inhibition was preceded by a shorter-lasting excitation. 2. FRA units (units receiving excitation from cutaneous and/or high threshold muscle afferents) were typically excited by the cortical stimulation. The excitation was often followed by a period of depression of transmission from the periphery. 3. It is suggested from the effective cortical area and experiments with lesions in the medullary pyramid and in the spinal cord that the inhibition in group I units and the excitation of FRA units are both mediated by the corticospinal tract.Experiments were also made to determine the level where the cell body of a given DSCT unit is located, and the results from 56 units are presented.     

Input from ipsilateral proprio- and exteroceptive hind limb afferents to nucleus Z of the cat medulla oblongata.

1. Extracellular recordings have been made from proprio- and exteroceptive neurones in nucleus Z of the cat medulla oblongata. 2. Seventeen non-relay cells were excited by ipsilateral hind limb group I muscle afferents. Their functional were similar to those of the corresponding relay cells (Johansson & Silfvenius, 1977), i.e. activation by axon-collaterals of dorsolateral fascicle (DLF) fibres was observed, indicative of an input from the dorsal spinocerebellar tract (DSCT) to these neurones. The non-relay cells differed, however, from the relay cells in that a larger proportion of them were disynaptically connected to the spinal fibres. It is suggested that the non-relay cells by inhibition modulate the group I relay transmission in nucleus Z. 3. Fifteen cells were excited by ipsilateral hind limb low threshold skin afferents with their spinal fibres at the first cervical segment located either in the DLF or in the dorsal column (DC). Evidence for a cutaneous hind limb bulbothalamic path independent of that of the DC and of the spinocervicolemniscal path is presented. 4. Six cells were activated by ipsilateral hind limb joint afferents at low threshold. At the C1 level the spinal fibres were located either in the DLF or in the DC. No joint activated relay-cells were encountered. Axon-collateral activation of DLF-linked joint cells was observed during cerebellar stimulation, indicating that segmental DSCT cells mediated the joint input to nucleus Z. 5. The topographical organization within nucleus Z and its input relation to the DSCT is discussed.