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.     

Organization of neurones in the cat cerebral cortex that are influenced from Group I muscle afferents

1. Neurones in the Group I projection area of the first sensori-motor cortex were investigated with extra- and intracellular technique.2. The majority of the neurones influenced by volleys in Group I afferents of contralateral forelimb nerves received exclusively excitation, but some received exclusively inhibition, and some mixed excitation and inhibition.3. The Group I influenced cells were usually found 500-1500 mu beneath the cortical surface.4. The EPSPs and IPSPs evoked from Group I afferents had a steep rising phase and a slow, approximately exponential decay. The duration was sometimes more than 50 msec. The EPSP evoked by a maximal Group I volley was often formed by a small number of large unitary EPSPs.5. Latency measurements indicate that the majority of the Group I activated neurones were monosynaptically excited from the thalamic fibres, and hence constitute the fourth-order neurones in the Group I projection system. The latencies of the IPSPs suggest a disynaptic linkage with thalamic fibres. Hence, the exclusively inhibited cells would constitute fifth-order neurones. It is suggested that most or all of the fourth-order neurones are inhibitory.6. The convergence of excitation and/or inhibition to individual cells was usually extensive and included effects not only from muscle groups working at different joints but also effects from antagonistic groups at the same joint. In addition cutaneous afferents contributed synaptic actions which had a longer latency than the synaptic actions from Group I afferents.7. The neurones influenced from Group I afferents were not antidromically activated on stimulation of the pyramidal tract.     

Projection from area 3a to the motor cortex by neurons activated from group I muscle afferents

Two receiving areas in the pericruciate cortex are known for inputs from group I muscle afferents of forelimb nerves. One focus is near the postcruciate dimple of area 3a, and the other in the lateral sigmoid gyrus of the motor cortex (area 4gamma). The cortico-cortical projection of area 3a to 4gamma, and the relay by this projection of group I muscle afferent input to the motor cortex were investigated in cats. The following results were obtained. 1. Seventy-four neurons within area 3a were antidromically activated by intracortical microstimulation of the motor cortex. 2. Although excitation evoked by stimulation of group I muscle afferents could be demonstrated for only a few (8 of 48) cortico-cortical neurons in extracellular recordings, due to the methodological limitations discussed, this input evoked EPSPs in 8 of 9 cortico-cortical neurons recorded intracellularly. Therefore, it is likely that the majority of neurons projecting from area 3a to the motor cortex have an excitatory synaptic input from group I afferents. 3. Neurons projecting from area 3a to the motor cortex were most commonly found in cortical layer III, although some were found in layer V. 4. Five of nine pyramidal tract neurons of area 3a had a strong excitatory synaptic input from group I muscle afferents. 5. A new type of pyramidal tract neuron was found which has cortico-cortical axon collaterals connecting the two cytoarchitectonic regions. These various neurons may be part of a feedback system from muscle afferents to the motor cortex.     

Integration in descending motor pathways controlling the forelimb in the cat

Summary A previously described disynaptic pathway from cortex to forelimb motoneurones whose intercalated neurones were excited both from other descending pathways and from forelimb afferents (Illert et al., 1976a, b) has been further analysed, mainly with respect to the location of the relay cells and their axons.Disynaptic EPSPs evoked in forelimb motoneurones by stimulation of the pyramid remained after complete transection of the corticospinal tract in C5 rostral to the forelimb segments but were abolished after a more rostral transection of the tract in the C2 segment. Corresponding findings were made with disynaptic rubral EPSPs after transection of the rubrospinal tract in these segments. It is concluded that disynaptic cortico-motoneuronal and rubro-motoneuronal excitation is relayed by propriospinal neurones originating in the C3–C4 segments. Other lesion experiments revealed that the axons of these propriospinal neurones descend to forelimb motoneurones in the ventrolateral part of the lateral funicle.Spatial facilitation of transmission from the corticospinal and rubrospinal tracts after transection of them in C5 occurred with a time course showing monosynaptic convergence from these pathways on common propriospinal neurones.Facilitation of disynaptic pyramidal EPSPs from the dorsal tegmentum remained after transection of the corticospinal tract at C5 but was abolished after a transection at C2. It is postulated that corticospinal and presumed tectospinal fibres converge onto common neurones in the propriospinal relay but evidence is also given for a more rostral relay (probably bulbar) with a similar convergence.The oligo- (probably mono-)synaptic facilitation of the disynaptic pyramidal EPSP evoked by volleys in cutaneous and group I muscle afferents from the forelimb likewise remained after a C5 transection of the corticospinal tract but was abolished after an additional C5 lesion in the dorsal column. It is concluded that propriospinal relay cells receive excitatory action from forelimb afferents ascending in the dorsal column. Spatial facilitation experiments using three tests revealed that propriospinal neurones monosynaptically excited from both corticospinal and rubrospinal fibres also receive excitation from cutaneous forelimb afferents.It is postulated that the propriospinal relay provides an important route for fast activation of forelimb motoneurones from the brain. The convergent monosynaptic excitation from several important motor centres in the brain is considered in relation to the general problem of the functional relationship between higher motor centres. The convergent action from forelimb afferents is taken to suggest that a descending command for a forelimb movement can be modified from the forelimb while on its way to the motoneurones.Supported by the Deutsche Forschungsgemeinschaft     

Organization of group I activated cells in the main and external cuneate nuclei of the cat: Identification of muscle receptors

Summary Extracellular recording was made from 77 primary afferent fibres, 106 cells in the external cuneate nucleus, and 60 cells in the main cuneate nucleus, all activated by slowly adapting muscle stretch receptors. The nature of the muscle receptors responsible for the activation was determined by various types of receptor stimulation.Primary group I afferents from muscle spindles and tendon organs in distal forelimb muscles showed complete overlap of conduction velocities and thresholds to electrical stimulation. Both types of group I afferents as well as group II muscle spindle afferents were shown to ascend through the dorsal funiculus to the level of the cuneate nuclei.Three groups of cells were identified in the external cuneate nucleus, activated by group I muscle spindle afferents, tendon organ afferents and group II muscle spindle afferents, respectively.Almost all group I activated cells in the main cuneate nucleus, including all 34 cells identified as cuneo-thalamic relay cells, received their afferent input from muscle spindle afferents. Three cells were activated by tendon organ afferents.     

The afferents and projections of the ventroposterolateral thalamus in the monkey

Summary In seven monkeys (M. fascicularis), recordings were made from neurones in subnuclei VPL_o and VPL_c of the thalamus. The peripheral inputs to these cells from the forelimb were established by stimulating muscle, skin and joint nerves, and cutaneous receptive fields were examined by natural stimulation of the skin. The projection of the same cells to motor and sensory cortex was examined by stimulation of the cortex to demonstrate antidromic responses and by retrograde axonal transport of horseradish peroxidase injected into these regions of cortex. Of the 139 neurones which were driven by stimulation of peripheral nerves, 73 were located in VPL_o. This suggests that there is a significant projection of forelimb afferents to VPL_o in the monkey. Of the 73 neurones located in VPL_o, 60 responded to stimulation of muscle or joint nerves. The projection to VPL_o is therefore predominantly from deep receptors. Twenty-one of these 73 neurones had convergent inputs. The latencies of activation of units in VPL_o were short (4–8 ms) and consistent with a lemniscal pathway via the dorsal column nuclei. Injection of horseradish peroxidase into the arm area of the motor cortex labelled cells in VPL_o and not in VPL_c. The neurones from which recordings were made in VPL_o were located within the population of cells labelled with horseradish peroxidase after injections into the primary motor area. This suggests that this subnucleus is the thalamic relay for the sensory input from peripheral receptors to cells of the motor cortex.     

Thalamic projection of muscle nerve afferents in the cat

1. Evoked responses on stimulation of hind- and forelimb muscle nerves have been recorded in the ventro-postero-lateral (VPL) and centre median (CM) nuclei of the thalamus of the cat.2. The stimulation of hind-limb muscle afferents with increasing strength usually did not evoke a thalamic response, either in the CM or in the VPL, until the threshold for Group III muscle afferents was reached.3. Short latency potentials were evoked in the VPL on stimulation of low threshold Group I muscle afferents from forelimbs. The projection zone occupies the dorso-medio-rostral part of the nucleus.4. An attempt was made to locate the medullary relay for forelimb Group I afferents either in the lateral part of the cuneate or in external cuneate.5. Low threshold afferents from forelimb muscle nerves do not project to the CM. CM responses began to appear only when the threshold for Group II muscle nerve fibres was reached.6. Group II afferents from hind- or forelimb muscle nerves projecting to the CM are probably not connected to spindle secondary endings.7. The role played by muscle afferents in the somesthetic mechanisms is briefly discussed.     

Mutual inhibition between olivary cell groups projecting to different cerebellar microzones in the cat

Summary Intracellular recording was made in the C3-C4 segments from cell bodies of a previously described system of propriospinal neurones (PNs), which receive convergent monosynaptic excitation from different higher motor centres and mediate disynaptic excitation and inhibition from them to forelimb motoneurones. Inhibitory effects in these PNs have now been investigated with electrical stimulation of higher motor centres and forelimb nerves. Short-latency IPSPs were evoked by volleys in the cortico-, rubro- and tectospinal tracts and from the reticular formation. Latency measurements showed that those IPSPs which required temporal summation were disynaptically mediated. After transection of the corticospinal tract in C2, only small and infrequent disynaptic IPSPs were evoked from the pyramid. It is postulated that disynaptic pyramidal IPSPs only to a small extent are evoked by monosynaptic excitation of reticulospinal inhibitory neurones known to project directly to the PNs, and that they are mainly mediated by inhibitory interneurones in the C3-C4 segments. Tests with spatial facilitation revealed monosynaptic excitatory convergence from tecto-, rubro- and probably also from reticulospinal fibres on inhibitory interneurones monosynaptically excited from corticospinal fibres (interneuronal system I). Disynaptic IPSPs were also evoked in the great majority of the PNs by volleys in forelimb muscle and skin nerves. A short train of volleys was usually required to evoke these IPSPs from group I muscle afferents. In the case of cutaneous nerves and mixed nerves single volleys were often effective, and the lack of temporal facilitation of IPSPs produced by a train of volleys showed strong linkage from these nerves. The results obtained after transection of the dorsal column at different levels show that the relay is almost entirely rostral to the forelimb segments. Test with spatial facilitation revealed that interneurones monosynaptically activated from forelimb afferents receive convergent excitation from corticospinal but not or only weakly so from tecto- or rubrospinal fibres. There was also convergence from group I muscle afferents and low threshold cutaneous afferents on common interneurones. It is postulated that the disynaptic IPSPs from forelimb afferents are mediated by inhibitory interneurones (interneuronal system II) other than those receiving convergent descending excitation. Volleys in corticospinal fibres, in addition to the disynaptic IPSPs, evoke late IPSPs in the PNs. Similar late IPSPs were evoked from the ipsilateral forelimb by stimulation of the FRA. Monosynaptic IPSPs were evoked in the majority of the PNs on weak stimulation of the lateral reticular nucleus (LRN) and from regions dorsal to it. Results from threshold mapping suggest that these IPSPs are due to antidromic stimulation of ascending inhibitory neurones which also project to the C3-C4 PNs, and that the ascending collaterals terminate in the LRN or/and the base of the cuneate nuclei. Activity in the ascending collaterals may give higher centres information regarding inhibitory control of the PNs. It is postulated that interneuronal system I subserves descending feed-forward inhibition and interneuronal system II feed-back inhibition from the forelimb of transmission through the C3-C4 PNs to motoneurones.This work was supported by the Swedish Medical Research Council (project no. 94)     

REORGANIZATION OF BARREL CIRCUITS LEADS TO THALAMICALLY-EVOKED CORTICAL EPILEPTIFORM ACTIVITY

We studied circuit activities in layer IV of rat somatosensory barrel cortex containing microgyri induced by neonatal freeze lesions. Structural abnormalities in GABAergic interneurons are present in the epileptogenic paramicrogyral area (PMG) and we therefore tested the hypothesis that decreased postsynaptic inhibition within barrel microcircuits occurs in the PMG and contributes to epileptogenesis when thalamocortical afferents are activated. In thalamocortical (TC) slices from na?ve animals, single electrical stimuli within the thalamic ventrobasal (VB) nucleus evoked transient cortical multi-unit activity lasting 65±42 ms. Similar stimuli in TC slices from lesioned barrel cortex elicited prolonged 850 ±100 ms paroxysmal discharges that originated in the PMG and propagated laterally over several mm. Paroxysmal discharges were shortened in duration by ~70 % when APV was applied, and were totally abolished by CNQX. The cortical paroxysmal discharges did not evoke thalamic oscillations. Whole cell patch clamp recordings showed that there was a shift in the balance of TC evoked responses in the PMG that favored excitation over inhibition. Dual whole-cell recordings in layer IV of the PMG indicated that there was selective loss of inhibition from fast-spiking interneurons to spiny neurons in the barrel circuits that likely contributed to unconstrained cortical recurrent excitation with generation and spread of paroxysmal discharges.     

Projection of Ia and Ib afferents from forelimb muscles to the C3-C4 segments of the cat spinal cord

Group Ia muscle spindle afferents were activated separately by small stretches applied to the tendons of antibrachial muscles in the forelimb in the cat. Group Ib tendon organ afferents were stimulated electrically after a selective increase of the threshold of the Ia afferents. Recordings of focal synaptic potentials were made in the C3-C4 segments in the medial part of the base of the dorsal horn. It has been found that both Ia and Ib afferents have monosynaptic connections with neurones in this medial region. Quantitatively, these two groups of afferents produced focal synaptic potentials of approximately the same size. The connections may be to inhibitory interneurones projecting to the C3-C4 propriospinal neurones, which are known to receive disynaptic IPSPs from group I muscle afferents.     

Columnar distribution of U-fibres from the postcruciate cerebral projection area of the cat's group I muscle afferents

Needle stitch lesions were made in the maximal point of the cerebral projection area of the low threshold muscle afferents near the postcruciate dimple of the cat's posterior sigmoid gyrus. The lesions did not exceed 500 mum in diameter and were restricted to the cortical grey matter. Degenerating nerve fibres and terminals were investigated with Fink-Heimer technique in four cats (survival times: 26, 48 and 96 hours). The cytoarchitectonic areas of the sensori-motor cortex were determined in cresyl violet and van Gieson sections. All lesions were made in area 3a. Degenerating U-fibres originating from the lesion travelled in the white matter to the cortex of area 4 gamma, 3b and 2. They reentered the cortex and branched in layer III. Terminal degeneration was found in layer I. The degeneration was distributed to distinct columns with a diameter of about 1mm. Such columns were observed laterally and medially in area 4 gamma, in area 3b near the caudal end of the coronal sulcus, in area 2 near the lateral ansate sulcus and in the forelimb region of SII. The distribution of the cortico-cortical connections from the cerebral projection area of the forelimb group I muscle afferents was discussed in relation to the known cerebral projections of group I muscle afferents, low threshold joint afferents, pacinian afferents and low threshold skin afferents.     

Integration in descending motor pathways controlling the forelimb in the cat

Summary With intracellular recording from forelimb motoneurones the spatial facilitation technique has been used to investigate interaction between descending pathways and forelimb afferents.As previously shown for the hindlimb, pyramidal volleys effectively facilitate interneuronal transmission in reflex pathways from different primary afferents. Evidence is presented suggesting disynaptic excitation from corticospinal fibres of interneurones in the reciprocal Ia inhibitory pathway. Interneurones of other reflex pathways from group I muscle afferents receive monosynaptic pyramidal excitation. During pyramidal facilitation volleys in cutaneous afferents may evoke PSPs in motoneurones after a central delay of 1.3 ms suggesting that the minimal linkage is disynaptic.Information regarding convergence on the neurones intercalated in the disynaptic cortico-motoneuronal pathway was obtained by investigating the effect from primary afferents and from other descending pathways on the disynaptic pyramidal EPSPs. Volleys in cutaneous and group I muscle afferents facilitate transmission in the disynaptic cortico-motoneuronal pathway with a time course showing oligosynaptic (probably monosynaptic) action on the intercalated neurone. Rubrospinal volleys likewise effectively facilitate disynaptic cortico-motoneuronal transmission with a time course showing monosynaptic action on the intercalated neurone. Spatial facilitation experiments involving three tests revealed that those intercalated neurones which receive convergent monosynaptic excitation from corticospinal and rubrospinal fibres are excited also from cutaneous forelimb afferents.Disynaptic cortico-motoneuronal transmission was also monosynaptically facilitated by stimuli in the dorsal mesencephalic tegmentum probably activating tectospinal fibres. Disynaptic, presumed tectospinal EPSPs were facilitated from cutaneous forelimb afferents.The convergence onto the neurones intercalated in the disynaptic excitatory cortico-motoneuronal pathway suggests that these neurones integrate the activity in different descending pathways and primary forelimb afferents.Supported by the Deutsche ForschungsgemeinschaftIBRO/UNESCO Fellow     

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.     

Projection of la and lb afferents from forelimb muscles to the C3-C4 segments of the cat spinal cord

Group la muscle spindle afferents were activated separately by small stretches applied to the tendons of antibrachial muscles in the forelimb in the cat. Group lb tendon organ afferents were stimulated electrically after a selective increase of the threshold of the la afferents. Recordings of focal synaptic potentials were made in the C₃-C₄ segments in the medial part of the base of the dorsal horn. It has been found that both la and lb afferents have monosynaptic connections with neurones in this medial region. Quantitatively, these two groups of afferents produced focal synaptic potentials of approximately the same size. The connections may be to inhibitory interneurones projecting to the C₃-C₄ propriospinal neurones, which are known to receive disynaptic IPSPs from group I muscle afferents.     

Thalamocortical projections activated by phrenic nerve afferents in the cat

This study identified thalamocortical projections activated by respiratory afferents. Cortical evoked potentials were recorded in the right primary somatosensory cortex of the cat following electrical stimulation of the left C5 root of the phrenic nerve. The majority of primary sites were located in the vicinity of the postcruciate dimple, in area 3a near the 3a/3b border, corresponding to the trunk region of the cortical body map. Retrograde fluorescent tracers injected at the sites of primary activation produced labeled cells in the oralis nucleus of the ventroposterior complex [4]. Control injections made in adjacent cortical areas not activated by phrenic stimulation resulted in labeling in the ventroposterior complex which did not overlap that seen with injections of primary activation sites. We conclude that respiratory muscle afferents in the phrenic nerve elicit activity in the trunk region of primary somatosensory cortex via specific thalamocortical projections originating in the oralis portion of the thalamic ventroposterior complex.     

Integration in descending motor pathways controlling the forelimb in the cat

Summary Recording was made in the C3-C4 segments from cell bodies of propriospinal neurones identified by their antidromic activation from more caudal segments. Monosynaptic excitatory effects from descending motor pathways and primary afferents were investigated by electrical stimulation of higher motor centres and peripheral nerves in the forelimb and neck.The cell bodies were located mainly laterally in Rexed's layer VII. Threshold mapping for single axons showed that they descend in the lateroventral part of the lateral funicle. Antidromic stimulation at different spinal cord levels showed that some neurones terminated in the forelimb segments, others in the thoracic cord or in the lumbar segments. Terminal slowing of the conduction velocity suggested axonal branching over some segments.Monosynaptic EPSPs were evoked in the neurones by stimulation of the contralateral pyramid, red nucleus and dorsal tegmentum-superior colliculus. It is concluded that corticospinal, rubrospinal and tectospinal fibres project directly to both short and long propriospinal neurones. There was marked frequency potentiation in tectospinal synapses. Convergence from two descending tracts was common and in half of the tested cells all three tracts contributed monosynaptic excitation. Experiments with collision of descending volleys and antidromic volleys from the brachial segments demonstrated that the corticospinal and rubrospinal monosynaptic projection to the propriospinal neurones is by collaterals from fibres continuing to the forelimb segments.Stimulation of cervical primary afferents in the dorsal column gave monosynaptic EPSPs in somewhat less than half of the tested propriospinal neurones. The further analysis with stimulation of forelimb nerves and C2-C3 dorsal rami showed that monosynaptic EPSPs may be evoked from low threshold cutaneous and group I muscle afferents in the forelimb and from C2-C3 neck afferents entering close to the spinal ganglia, possibly from joint receptors. Convergence from cervical afferents and at least two of the above descending tracts was common.It is postulated that the propriospinal neurones previously indirectly defined by their action on motoneurones as relaying disynaptic excitation from higher motor centres to forelimb motoneurones (Illert et al., 1977) belong to those neurones of the C3-C4 propriospinal systems which terminate in the cervical enlargement. The function of the neurones projecting beyond the upper thoracic segments is discussed.Supported by the Deutsche ForschungsgemeinschaftIBRO/UNESCO Fellow     

Pyramidal actions in identified radial motornuclei of the cat

This study aimed to establish the projection from the corticospinal tract (CST) to the motoneurones innervating the deep radial (DR) forelimb muscles. In the anaesthetized cat stimulation of the contralateral pyramid and intracellular recording from identified forelimb motoneurones was used. A train of pyramidal stimuli evoked disynaptic EPSPs in DR motoneurones. The effects were very similar in the different nuclei. Pyramidal IPSPs had a slightly longer latency and occurred in most cases together with disynaptic EPSPs. It is suggested that the inhibitory actions to the distal forelimb are predominantly relayed in a trisynaptic pathway, but that a disynaptic linkage seems possible as well. The disynaptic pyramidal EPSPs remained after CST transection in C5. They were abolished after CST transections in C2. It is concluded that disynaptic corticospinal excitation of distal DR motornuclei is relayed in a short midcervical propriospinal system. Transection experiments at different cervical levels suggest that the majority of the propriospinal neurones is located in C3-C4. The CST facilitated a variety of reflex pathways to motoneurones innervating distal forelimb muscles. Disynaptic excitatory and inhibitory effects from cutaneous and low threshold group I muscle afferents were common. They were present in all investigated nuclei and powerfully facilitated from the CST. It is suggested that this allows the brain to adapt the reflex mechanisms of the distal forelimb to the synergistic-antagonistic relations between the muscles, which are changing according to the performed movement.     

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.     

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.     

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.