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Notes: Climbing fibres projecting to the cerebellar C3 zone (and the related C1 and Y zones) receive spatially well organized tactile and nociceptive inputs from the skin. In the present study, cutaneous tactile and nociceptive input to climbing fibres projecting to the X, B, C2 and D1 zones in lobule V were investigated in pentobarbitone-anaesthetized cats. From the present results and previous studies, it is concluded that the X, C1, CX, C3 and Y zones receive cutaneous nociceptive climbing fibre input. By contrast, climbing fibres to the B, C2 and D1 zones lack cutaneous nociceptive input. Tactile input was found in all zones. The spatial organization of receptive fields of climbing fibres projecting to the X and D1 zones was similar to that in the C3 zone. They were located on the ipsilateral forelimb, mainly its lateral and distal parts, and their proximal borders were located close to joints. In the B zone, more than half of the receptive fields of climbing fibres were confined to the ipsilateral hind- or forelimb. However, frequently more than one limb and parts of the trunk were included. In the C2 zone, the majority of climbing fibres had distal ipsi- or bilateral receptive fields on the forelimbs, often also including the head/face. Some of the bilateral forelimb receptive fields additionally included the hindlimbs ipsi- or bilaterally. The results indicate that each zone has a characteristic set of climbing fibre receptive fields, which is probably related to its efferent control functions.

Notes: Inferior olivary cells projecting as climbing fibres to the forelimb area of the cerebellar C3 zone were investigated with respect to their cutaneous and muscle afferent input in barbiturate-anaesthetized cats. Climbing fibre responses were recorded from single cerebellar cortical Purkinje cells on natural stimulation of the skin and on electrical stimulation of nerves to m. biceps brachii, m. triceps brachii and to nine muscles acting as dorsal or palmar flexors of the paw (and, in some cases, the digits). The analysis was focused on the functional organization of convergence between cutaneous and muscle afferents onto single olivary neurons. Cutaneous receptive fields on the dorsal side of the paw and on the digits were generally associated with moderate to strong input from dorsal flexors, but little or no input from palmar flexors or proximal muscles. Receptive fields on the ventral side of the paw and forearm were associated with relatively strong input from biceps and palmar flexors. Climbing fibres with cutaneous receptive fields extending on the ulnar side of the paw and forearm usually received strong input from the triceps and moderate to strong input from dorsal flexors, whereas input from the palmar flexors was weak or lacking. In conclusion, the results indicate that the cutaneous receptive fields in many cases are associated with input from muscles the action of which would tend to move the receptive field towards a stimulus applied to the skin.

Notes: Summary The spino-olivocerebellar paths ascending through the dorsal funiculi (DF-SOCPs) were studied by recording climbing fiber field potentials in the cerebellar cortex. Several DF-SOCPs were identified on the basis of their response latencies, peripheral inputs, and projection areas. The projection areas consist of eight sagittal zones on each side of the anterior lobe denoted a, x, b, C1, c2, c3, d1, and d2. The a and b zones, which are activated exclusively from hindlimb nerves (Oscarsson, 1969a) were not studied. The shortest response latencies in the x, c1, c3 and d2 zones indicate that these zones are activated by direct paths between the dorsal funiculus nuclei and the inferior olive. These zones also have long latency responses evoked through indirect paths. The direct paths are activated from the flexor reflex afferents and the indirect paths from distal cutaneous afferents. The x zone is activated from forelimb nerves only. The c1 and c3 zones and presumably also the d2 zone are activated from hindlimb and forelimb nerves and have a detailed somatotopical organization. The c2 and d1 zones have long latency responses evoked through indirect paths. Both zones are activated from distal cutaneous afferents. The c2 zone has no distinct somatotopical organization, whereas the d1 zone has largely separate forelimb and hindlimb areas. In contrast to all other zones, the c2 zone is activated not only from ipsilateral but also from contralateral nerves.

Notes: Summary The somatotopical organization of the projection through the dorsal spino-olivocerebellar path (DF-SOCP) to the c3 zone in the cerebellar anterior lobe was studied by recording climbing fiber field potentials in the cerebellar cortex evoked on stimulation of peripheral nerves. The c3 zone was shown to contain a detailed and systematic representation of the ipsilateral body half with the following characteristics: 1. Single nerves project to one or two sagittal strips of cortex which extend across several folia. The width of the strips varies between 0.2 and 1 mm and the length may be as long as 15–20 mm along the unfolded cortex. 2. The strips activated from different nerves occur in an order which usually follows the segmentai innervation, the hindlimb being represented rostrally and the forelimb caudally in the zone. 3. The double representation of some nerves makes it possible to distinguish one medial and one lateral part of the c3 zone with the projection areas organized approximately as mirror images. Additional observations on other zones activated by the DF-SOCP system (Ekerot and Larson, 1979a) indicate that the c1 zone has a somatotopical organization which is reminiscent of that of the c3 zone, whereas the x and d1 zones have different topographical organizations.

Notes: Summary The termination patterns in the cerebellar anterior lobe of one mossy fiber path, the exteroceptive component of the cuneocerebellar tract (E-CCT), and one climbing fiber system, the dorsal spino-olivocerebellar paths (DF-SOCPs), which both relay in the main cuneate nucleus, were compared in the cat. The E-CCT terminates in the ipsilateral half of the anterior lobe in five sagittal zones which overlap five of the eight zones activated from the DFSOCPs. In at least one zone the E-CCT projection has a somatotopical organization which is similar to and overlaps that of the DF-SOCP.

Notes: Summary Intracellular recordings were obtained from 204 neurones in the lateral reticular nucleus (LRN). LRN neurones contacted by the bVFRT were identified by the responses evoked on stimulation of descending fibres in the contralateral ventral quadrant of the spinal cord (cVQ) at cervical (C5cVQ) and lumbar (L2cVQ) levels. Stimulation of the cVQ evoked excitatory or inhibitory responses in 124 of the 204 LRN neurones. EPSPs were evoked in 45, IPSPs in 52 and both EPSPs and IPSPs in 27 LRN neurones. The shortest latencies of the responses evoked from the cVQ indicated that both EPSPs and IPSPs were disynaptic. This finding was confirmed by direct stimulation of the ascending fibres in the ipsilateral ventrolateral funiculus at C3 (C3iVLF) or LI (LliVLF). In most LRN neurones activated or inhibited from the cVQ, stimulation of the iVLF evoked similar responses at a monosynaptic latency. These results indicate that the bVFRT consists of roughly equally large groups of excitatory and inhibitory neurones monosynaptically connected with the LRN. Excitatory and inhibitory bVFRT neurones had similar peripheral receptive fields and termination areas in the LRN. LRN neurones were divided into those contacted by cervical bVFRT neurones and lumbar bVFRT neurones. The former group consisted of LRN neurones responding to C5cVQ stimulation at latencies below 5 ms, whereas the latter group contained LRN neurones responding to stimulation of the L2cVQ. Cervical bVFRT neurones projected to most parts of the LRN whereas the projection of lumbar bVFRT neurones were confined to the ventrolateral part of the nucleus. Excitatory and inhibitory bVFRT neurones of each group had similar termination areas. About half of the LRN neurones contacted by cervical bVFRT neurones did not respond to stimulation of the contralateral forelimb (cF) nerve. These bVFRT neurones formed a separate group which terminated in the dorsomedial part of the LRN. Cervical bVFRT neurones activated by the cF terminated in the ventrolateral part of the nucleus. The conduction velocity between the L1 and C3 segments was determined for axons of lumbar bVFRT neurones. The velocities ranged between 55–137 m/s (mean 90.6 m/s) for excitatory neurones and between 35–120 (mean 87.5 m/s) for inhibitory neurones. Monosynaptic responses, particularly EPSPs, were frequently evoked from the L1iVLF in LRN neurones with a bVFRT input from cervical segments only. The results suggest that many excitatory cervical bVFRT neurones have fast conducting descending axon branches projecting to the lumber cord. Long descending axon branches seemed to be less common among inhibitory cervical bVFRT neurones.

Notes: Summary The afferent paths from the spinal cord and from trigeminal afferents to the lateral reticular nucleus (LRN) were investigated by intracellular recording from 204 LRN neurones in preparations with a spinal cord lesion at C3 that spared only the ipsilateral ventral quadrant. Stimulation of nerves in the limbs evoked EPSPs and JPSPs in 201 of 204 tested LRN neurones. The strongest input was from the ipsilateral forelimb (iF) which evoked EPSPs in 49% and IPSPs in 73% of the LRN neurones. Each of the other limbs evoked EPSPs in approximately 20% and IPSPs in approximately 25% of the neurones. Stimulation of the ipsilateral trigeminal nerve (iTrig) evoked EPSPs in 32% and IPSPs in 46% of the neurones. The shortest latencies of the EPSPs and IPSPs indicated a disynaptic connection between primary afferents in the iF and iTrig and the LRN. The most direct pathways for excitatory and inhibitory responses from the other limbs were trisynaptic. Stimulation of the ventral part of the ipsilateral funiculus (iVLF) at C3 (C3iVLF) evoked monosynaptic responses in 189 of 201 tested LRN neurones. Monosynaptic EPSPs were recorded in 104 neurones and monosynaptic IPSPs in 126 neurones. Monosynaptic EPSPs and IPSPs were encountered in all parts of the LRN. Stimulation of the iVLF at L1 (LliVLF) evoked monosynaptic EPSPs and IPSPs in the ventrolateral part of the LRN. The termination areas of excitatory and inhibitory fibres appeared to be the same. LRN neurones without monosynaptic EPSPs or IPSPs from the LliVLF were located mainly in the dorsal part of the magnocellular division. Stimulation of the dorsal funiculi (DF) at C2 and the ipsilateral trigeminal nerve (iTrig) evoked excitatory and inhibitory responses in the LRN. The shortest latencies of EPSPs and IPSPs indicated disynaptic connections.

Notes: Summary Intracellular recording from neurones in the lateral reticular nucleus (LRN) demonstrated that, in addition to the previously identified excitatory ipsilateral forelimb tract (iF tract) (Clendenin et al. 1974c) there is an inhibitory tract mediating information from the ipsilateral forelimb to the LRN. The excitatory and inhibitory tracts were similarly organized. The tract neurones were monosynaptically activated by affcrents in the ipsilateral forelimb and projected to the same area of the LRN. They will be considered as excitatory and inhibitory components of the iF tract and denoted the excitatory and inhibitory iF tract (EiF and IiF tracts). Stimulation of the descending ipsilateral dorsolateral funiculus (iDLF) in the C3 segment evoked disynaptic EPSPs and IPSPs in LRN neurones contacted by the EiF and IiF tracts. The responses in individual LRN neurones evoked from the iDLF were similar to the responses evoked from the forelimb nerves suggesting that the EiF and IiF tracts are monosynaptically activated by fibres in the iDLF. The dorsal portion of the magnocellular part of the LRN constituted the main termination area of both the EiF and IiF tracts. Neurones in this area have previously been shown to project ipsilaterally to lobule V in the pars intermedia of the cerebellar anterior lobe and to the paramedian lobule (Clendenin et al. 1974a). IPSPs evoked from the IiF tract in LRN neurones outside the main termination area had smaller amplitudes and longer latencies. This finding suggests that these responses were generated by thin axon collaterals given off from dorsally located stem axons.

Notes: Summary 1. The areas of the cerebellar cortex receiving fibres from the mLRN (major portion of the lateral reticular nucleus comprising its parvi- and magnocellular parts) were determined by identifying the low-threshold spots on the cerebellar surface from which 286 mLRN neurones could be antidromically activated. 2. The mLRN fibres terminate bilaterally in the anterior lobe and pyramis, and ipsilaterally in the paramedian lobule. Some fibres terminate also in the rostral part of lobule VI but hardly any in other parts of the cortex. It follows that the mLRN projects almost exclusively to the classical spinal receiving areas. 3. Neurones in the parvi- and magnocellular parts project to similar cortical areas. 4. Two regions in the mLRN were distinguished on the basis of their different projection areas. Region A, a dorsolateral portion of the magnocellular part, projects to the ipsilateral parts intermedia of the anterior lobe and the ipsilateral paramedian lobule. Region B, a ventral portion of the parvi- and magnocellular parts, projects bilaterally to the pars intermedia and vermis of the anterior lobe and sparsely to the pyramis. 5. Neurones throughout the parvi- and magnocellular parts receive monosynaptic excitation from fibres ascending in the ventral part of the ipsilateral lateral funiculus.

Notes: Abstract The functional relation between receptive fields of climbing fibres projecting to the C1, C3 and Y zones and forelimb movements controlled by nucleus interpositus anterior via the rubrospinal tract were studied in cats decerebrated at the pre-collicular level. Microelectrode tracks were made through the caudal half of nucleus interpositus anterior. This part of the nucleus receives its cerebellar cortical projection from the forelimb areas of these three sagittal zones. The C3 zone has been demonstrated to consist of smaller functional units called microzones. Natural stimulation of the forelimb skin evoked positive field potentials in the nucleus. These potentials have previously been shown to be generated by climbing fibre-activated Purkinje cells and were mapped at each nuclear site, to establish the climbing fibre receptive fields of the afferent microzones. The forelimb movement evoked by microstimulation at the same site was then studied. The movements usually involved more than one limb segment. Shoulder retraction and elbow flexion were frequently evoked, whereas elbow extension was rare and shoulder protraction never observed. In total, movements at the shoulder and/or elbow occurred for 96% of the interpositus sites. At the wrist, flexion and extension movements caused by muscles with radial, central or ulnar insertions on the paw were all relatively common. Pure supination and pronation movements were also observed. Movements of the digits consisted mainly of dorsal flexion of central or ulnar digits. A comparison of climbing fibre receptive fields and associated movements for a total of 110 nuclear sites indicated a general specificity of the input-output relationship of this cerebellar control system. Several findings suggested that the movement evoked from a particular site would act to withdraw the area of the skin corresponding to the climbing fibre receptive field of the afferent microzones. For example, sites with receptive fields on the dorsum of the paw were frequently associated with palmar flexion at the wrist, whereas sites with receptive fields on the ventral side of the paw and forearm were associated with dorsiflexion at the wrist. Correspondingly, receptive fields on the lateral side of the forearm and paw were often associated with flexion at the elbow, whereas sites with receptive fields on the radial side of the forearm were associated with elbow extension. The proximal movements that were frequently observed also for distal receptive fields may serve to produce a general shortening of the limb to enhance efficiency of the withdrawal. It has previously been suggested that the cerebellar control of forelimb movements via the rubrospinal tract has a modular organisation. Each module would consist of a cell group in the nucleus interpositus anterior and its afferent microzones in the C1, C3 and Y zones, characterised by a homogenous set of climbing fibre receptive fields. The results of the present study support this organisational principle, and suggest that the efferent action of a module is to withdraw the receptive field from an external stimulus. Possible functional interpretations of the action of this system during explorative and reaching movements are discussed.