C3 Zone Research Articles

Inhibition of classically conditioned eyeblink responses by stimulation of the cerebellar cortex in the decerebrate cat.

The purpose of the present study was to test the hypothesis that neurones in the anterior interpositus nucleus, under the control of Purkinje cells in the c1 and c3 zones of the cerebellar cortex, exert some control over classically conditioned responses. In particular, the experiments were designed to determine whether the cerebellar control of conditioned and unconditioned responses is different. The experiments were performed on cats decerebrated rostral to the red nucleus under halothane anaesthesia. The cats were conditioned using either a 1000 Hz tone or trains of stimuli through the skin of the proximal forelimb as the conditioned stimulus, and periorbital electrical stimulation as the unconditioned stimulus. A large proportion of the animals acquired conditioned responses at normal rates. It could be shown that these were true conditioned responses and did not result from sensitization or pseudoconditioning. For instance, unpaired presentations of conditioned and unconditioned stimuli caused rapid extinction. Cerebellar areas controlling eyeblink were identified by recording climbing fibre responses in the cerebellar cortex and recording EMG activity in the eyelid evoked by stimulation of the cerebellar cortex. When single shocks of 40-70 microA were applied to these areas during the emission of conditioned eyeblink responses, the latter were strongly inhibited. The inhibition had a latency of about 10 ms and a duration of 25-75 ms. It was shown that this inhibition of the conditioned responses was topographically specific and could only be evoked from cortical sites identified as controlling eyeblink. Stimulation of the periphery of an eyeblink area caused little or no inhibition. The effect of cortical stimulation on unconditioned reflex responses in the orbicularis oculi muscle was also tested. Some inhibition of unconditioned responses was observed, but quantitative analysis showed that this inhibition was considerably weaker than the corresponding inhibition of conditioned responses. The magnitude of the inhibition was determined for unconditioned responses of different sizes including responses which were weaker than the conditioned responses. It is concluded that conditioned eyeblink responses are under strong cerebellar control from areas in the c1 and c3 zones receiving climbing fibre input from the periorbital area. This effect is not likely to be due to a reduction in the background facilitation of facial motoneurones. In contrast, the weak inhibition of the unconditioned response was probably due to this mechanism. The results, therefore, suggest that the conditioned responses are dependent on the cerebellum in a way that is not true of unconditioned responses.

Topographical organization of the cerebellar cortical projection to nucleus interpositus anterior in the cat.

1. A new methodological approach for detailed study of the organization of the cerebellar corticonuclear projection was evaluated in barbiturate-anaesthetized cats. Extracellular field potentials were simultaneously recorded in nucleus interpositus anterior and in the forelimb area of the C3 zone, at the cerebellar surface. On electrical and natural stimulation of the forelimb skin, the evoked positive field potentials in the nucleus and the climbing fibre field potentials in the cerebellar cortex had similar characteristics, indicating that the nuclear potentials were related to climbing fibre activity. 2. Application of a local anaesthetic to the cerebellar surface reversibly diminished the positive field potentials in the nucleus, demonstrating that the potentials were dependent on cerebellar cortical activity. It was thus concluded that the positive field potentials were mainly generated by climbing fibre-activated Purkinje cells and reflected synaptic inhibitory potentials in nuclear neurones. Accordingly, the positive field potentials in the nucleus could be used to reveal the termination area of Purkinje cells activated by a specific climbing fibre input evoked on peripheral stimulation. 3. The topographical organization of the cerebellar cortical projection to the forelimb part of nucleus interpositus anterior was then investigated by systematically mapping the cutaneous tactile and nociceptive 'receptive fields' of the positive field potentials at different sites in the nucleus. Five groups of receptive fields were distinguished and tentatively divided into a total of nineteen subgroups. 4. Each group of receptive fields corresponded to one or two of the previously described receptive field classes of climbing fibres to the C1, C3 and Y zones and was represented in a single area of the nucleus. Within each area there was an orderly representation of different receptive fields. The results suggest that microzones in the C1, C3 and Y zones with similar climbing fibre input project to a common set of neurones in nucleus interpositus anterior. 5. We propose a modular organization for the cerebellar control of forelimb movements through the rubrospinal tract. Each module would consist of a set of neurones in nucleus interpositus anterior and their afferent microzones in the C1, C3 and Y zones. A module would control a specific group of muscles and receive a homogeneous climbing fibre input related to the movement controlled.

Suppression of cerebellar Purkinje cells during conditioned responses in ferrets.

Decerebrate ferrets were conditioned, using electrical stimulation of the forelimb as conditioned stimulus and periorbital stimulation as unconditioned stimulus, until they produced conditioned eyeblink responses. The latency of these was 125-250 ms. Microelectrode recordings were made from single Purkinje cells in an eyeblink controlling area in the c3 zone of the cerebellar cortex. Whereas Purkinje cells in animals, which had only received unpaired stimulus presentations, responded weakly or not at all to the conditioned stimulus, some cells in conditioned animals responded with a powerful suppression of simple spike firing. The latency of this suppression was 50-200 ms. The results support the hypothesis that classical conditioning involves plastic changes in cerebellar Purkinje cells.

Zonal Organization Within the Projection from the Inferior Olive to the Rostral Paramedian Lobule of the Cat Cerebellum

The projection from the inferior olivary nucleus to the forelimb-related regions of the c1, c2 and c3 zones within rostral folia of the paramedian lobule of the cat cerebellum was studied using a combined electrophysiological and neuroanatomical tracing technique. In each experiment, the cerebellar cortical zones were identified by their receipt of spino-olivocerebellar input evoked by percutaneous electrical stimulation of the limbs. A small (15-30 nl) injection of wheatgerm agglutinin-horseradish peroxidase (WGA-HRP) was then made into the centre of one of the zones. The olivary regions projecting to each zone were compared with the regions within the olive that have previously been shown to project to the corresponding cortical zones within lobule V of the anterior lobe. The results show that some (but not all) of the paravermal zones in the anterior lobe are also represented in the paramedian lobule (PML). The most medial (c1) zone in rostral PML receives climbing fibre input from an area in the rostral dorsal accessory olive which overlaps partially with the area that projects to the medial half of the c1 zone in lobule V. A zone equivalent to the lateral half of the c1 zone (the cx zone) in lobule V does not appear to be present within rostral PML. A middle (c2) zone within rostral PML receives olivary input from a region within the rostral medial accessory olive that overlaps partially with the area that projects to the c2 zone in lobule V. The presence of a c3 zone within rostral PML was found to be variable between animals and it could be identified electrophysiologically in only two out of a possible nine cases. In summary, the results demonstrate that, although a number of similarities exist between the olivocerebellar projections to corresponding cortical zones in the two forelimb-receiving regions of the cat paravermal cortex, the differences in olivocerebellar connectivity between the two regions suggest that functional differences between them may also exist.

Somatotopic termination of the spino-olivary fibers in the cat, studied with the wheat germ agglutinin-horseradish peroxidase technique.

Terminal sites of the spino-olivary fibers (SOFs) were examined by the anterograde transport of wheat germ agglutinin-horseradish peroxidase in the cat. The tracer was injected at various spinal cord levels from the first cervical to the caudal segments. The SOFs derived from the C1-T1 segments terminated medially in the caudal half (levels II-VIII of Brodal) of the medial accessory olive (MAO), which projects to the A zone of the cerebellar cortex, whereas the SOFs derived from the L6-S1 segments terminated laterally in the caudal half (levels I-VIII) of the MAO. No projections were found from the T2-L5 segments to the MAO. In the dorsal accessory olive (DAO), the SOFs terminated at levels III-XIV; the DAO projects to the B zone and the C1 and C3 zones of the cerebellar cortex. The SOFs derived from the C1-C4 segments terminated in the most medial part of the DAO (levels III-XIV), followed laterally by those from the C5-T1 segments. Further laterally, the SOFs derived from the T2-L5 and the L6-S1 segments terminated in the mediolateral order at levels V-XIV. The SOFs from the L6-S1 segments occupied the most lateral part of the DAO. The present study demonstrates that there is a distinct somatotopic termination of the SOFs in the mediolateral order in the caudal MAO and the DAO.

Distribution of Cutaneous Nociceptive and Tactile Climbing Fibre Input to Sagittal Zones in Cat Cerebellar Anterior Lobe.

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.

Organization of inferior olivary projections to the flocculus and ventral paraflocculus of the rat cerebellum.

The climbing fiber projection to the rat flocculus and adjacent ventral paraflocculus was investigated by using Phaseolus vulgaris-leucoagglutinin as an anterograde and horseradish peroxidase as a retrograde tracer. Large injections of horseradish peroxidase in the flocculus and ventral paraflocculus indicated that the climbing fibers to this region are derived exclusively from any of the following contralateral olivary regions: the dorsal cap of Kooy, the ventrolateral outgrowth, the caudal half of the ventral leaf of the principal olive near its lateral bend, and the rostral pole of the medial accessory olive. Subsequent anterograde and retrograde studies with small injections demonstrated that the latter area projects to the C2 zone, which runs caudally in the ventral paraflocculus and enters the caudal most aspect of the flocculus. The ventral leaf of the principal olive is connected to a D zone in the cerebellar hemisphere and paraflocculus, which, upon entering the ventral paraflocculus, divides into a caudal and rostral strip, termed FD and FD', respectively. The dorsal cap and the ventrolateral outgrowth each project to two distinct zones in the flocculus and part of the ventral paraflocculus. Two floccular zones, which are continuous with the parafloccular FD and FD' zones, receive their climbing fibers from the ventrolateral outgrowth. Two other zones, (FE and FE') receive their climbing fibers from the dorsal cap. The FE' zone is found at the rostral pole of the flocculus and is followed caudalwards by the FD', FE, FD, and C2 zones, respectively. The rostromedial part of the dorsal cap is connected to the continuation of the FE zone into the ventral paraflocculus. The observation that the dorsal cap and the ventrolateral outgrowth are both connected to a set of two alternating zones of floccular/ventral parafloccular Purkinje cells is in agreement with recent studies in the rabbit, and suggests that these zones reflect functionally distinct and discrete units related to specific aspects of visuomotor control.

Changes in light scatter and width measurements from the human lens cortex with age.

Light scatter and width measurements of the anterior cortical layers of the human lens were made in 50 eyes of 50 subjects using computerised linear scanning densitometry of Scheimpflug images. It was demonstrated that the amount of light scatter increased with age in all of the three major zones and that zone C3 showed the most marked increase. Most lens growth occurs in zone C2 with C3 showing little increase in width once it has become established. Zone C1 showed a tendency to decrease in width with age. In addition it was shown that the C3 zone, which is not present at birth and during early childhood, appears as a distinct layer during the second decade of life. Its scattering properties continue to increase throughout life, exceeding all other zones after approximately 30 years of age, in the absence of cataract. Possible explanations for the lens zone pattern are discussed.

Spread of synaptic activity along parallel fibres in cat cerebellar anterior lobe.

1. Mossy fibre evoked activity in the cerebellar cortex elicited by peripheral electrical stimulation was studied in chloralose anesthetized cats. The distribution of intracortical field potentials in the C3 and D zones was mapped in order to determine if there is a spread of synaptic activity outside the mossy fibre termination area. This area was identified by the presence of short latency synaptic field potentials in the granular layer. 2. Molecular layer field potentials were recorded up to 1.5 mm outside the mossy fibre termination area. The latencies of these potentials increased with increasing distance from the mossy fibre termination area, corresponding to a conduction velocity of about 0.4 m/s. 3. Recordings from Purkinje cells, within and outside the mossy fibre termination area, revealed an increase of simple spike activity at latencies corresponding to those of the field potentials in the same location. 4. From the spatial and temporal characteristics of the evoked activity, it is concluded that a mossy fibre input results in spread of synaptic activity along the parallel fibres. 5. The findings are discussed in relation to the recently discovered microzonal organization of the C3 zone. It is proposed that the organization of this zone offers a possibility for the control of muscle synergies, each synergy being represented by a mossy fibre input and the specific set of microzones activated by this input via the parallel fibres.

Lateral and medial sub-divisions within the olivocerebellar zones of the paravermal cortex in lobule Vb/c of the cat anterior lobe

The olivocerebellar projection to the c1, c2 and c3 zones in the paravermal cortex of lobule Vb/c has been investigated in the cat using a combined electrophysiological/neuroanatomical tracing technique. The zonal boundaries in the paravermal cortex were located by recording, on the cerebellar surface, climbing fibre field potentials evoked in response to percutaneous stimulation of one or more paws. A small (10-30 nl) injection of WGA-HRP was then made either into the centre or into the medial or lateral geographical half of a chosen zone and the resultant distribution of retrogradely labelled cells within the contralateral inferior olive was plotted. The c1 and c3 zones were each found to consist of two mediolaterally oriented 'sub-zones' which could be distinguished by their olivocerebellar input. The medial part of the c1 zone received climbing fibre input from the rostromedial part of the dorsal accessory olive (DAO) while the lateral part of the c1 zone received climbing fibre input from middle/rostral regions of the medial accessory olive (MAO). Both medial and lateral 'sub-zones' within the c3 zone were found to receive climbing fibre input from the rostral pole of DAO but, whereas there was heavy overlap between the olivary territories projecting to the medial c1 and medial c3 subzones, olivary cells projecting to the lateral part of c3 were located more rostrally within DAO. The c2 zone was found not to be divisible into mediolaterally oriented subzones and to receive climbing fibre input from a region of MAO located rostral and somewhat lateral to the region projecting to the lateral part of the c1 zone. The sub-zonal organisation of the olivocerebellar projection to the c1, c2 and c3 zones is discussed in relation to the functional properties of the different zones.

A study of branching in the projection from the inferior olive to the x and lateral c1 zones of the cat cerebellum using a combined electrophysiological and retrograde fluorescent double-labelling technique

The pattern of transverse branching in the olivocerebellar projection to the x zone in the vermis and the lateral c1 zone in the paravermis of the cat anterior lobe was studied using a combined electrophysiological and retrograde double-labelling tracer technique. Fluorochrome-tagged latex microspheres were well suited for this purpose. The results show that the region of olive that supplies climbing fibres to the two zones forms a continuous, rostrocaudally directed column about 2.25 mm in length, in a caudo-lateral to rostromedial part of the medial accessory olive (MAO), on average between A-P levels 12.50-10.50. This column may be divided into caudal and rostral halves that project respectively to the x and lateral c1 zones in the apical folia of lobules V/VIa. Partial overlap between these two territories occurs in an intermediate region (A-P levels 12.00-11.00) in middle MAO where olive cells that supply climbing fibres to either x or lateral c1 are intermingled with a smaller population of cells whose axons branch to provide climbing fibres to both zones. Quantitative analysis showed that, when different tracers were injected into each zone in the same animal, double-labeled cells represented only 5-7% of either single-labelled cell population within this area of overlap. It is concluded that, although some transverse branching is present within the olivocerebellar projection to the x and lateral c1 zones in the apical folia of lobule V, such branching is not extensive.

Topography and nociceptive receptive fields of climbing fibres projecting to the cerebellar anterior lobe in the cat.

1. The cutaneous receptive fields of 225 climbing fibres projecting to the forelimb area of the C3 zone in the cerebellar anterior lobe were mapped in the pentobarbitone-anaesthetized cat. Responses in climbing fibres were recorded as complex spikes in Purkinje cells. 2. A detailed topographical organization of the nociceptive climbing fibre input to the C3 zone was found. In the medial C3 zone climbing fibres with receptive fields covering proximal and/or lateral parts of the forelimb projected most medially. Climbing fibres with receptive fields located more medially on the forelimb projected successively more laterally. The sequence of receptive fields found in the lateral C3 zone was roughly the reverse of that in the medial C3 zone. Climbing fibres with receptive fields restricted to the digits projected preferentially to the caudal part of the forelimb area, whereas those with receptive fields covering both proximal and ventral areas of the forearm projected to more rostral parts. 3. The representation of the forelimb was uneven. Receptive fields with a focus on the digits or along the lateral side of the forearm dominated. 4. The proximal borders of the receptive fields were located close to joints. The area from which maximal responses were evoked was usually located eccentrically within the receptive field. Based on spatial characteristics the receptive fields could be divided into eight classes, which in turn were tentatively divided into subclasses. Similar subclasses of receptive fields were found in different cats. This classification was further supported by the results of a quantitative analysis of eighty-nine climbing fibres. The receptive fields of these climbing fibres were mapped with standardized noxious stimulation. 5. Climbing fibres terminating within sagittal strips (width, 100-300 microns; length, greater than 1 mm) had receptive fields which belonged to the same subclass. There were commonly abrupt changes in receptive fields between such microzones. Most classes of receptive fields were found in both the medial and the lateral parts of the C3 zone. However, receptive fields with a focus on the ventral side of either the metacarpals, the wrist or the forearm were found only in the medial part of the C3 zone. Furthermore, the class of receptive fields restricted to the lateral side of the upper arm and shoulder was only found in the lateral part of the C3 zone. 6. In the discussion, it is proposed that climbing fibres projecting to each microzone carry information from spinal multireceptive reflex arcs acting on a single muscle or a group of synergistic muscles.(ABSTRACT TRUNCATED AT 400 WORDS)

The postsynaptic dorsal column pathway mediates cutaneous nociceptive information to cerebellar climbing fibres in the cat.

1. The location in the spinal cord of the pathway mediating cutaneous nociceptive C fibre input to climbing fibres projecting to the forelimb area of the C3 zone in the cerebellar anterior lobe was investigated in pentobarbitone-anaesthetized cats. Lesions of the spinal cord at the segmental level of C3 sparing the dorsal funiculi (DF preparation) or lesions of the ipsilateral and part of the contralateral dorsal funiculi were made. 2. In the DF preparation, the cutaneous input to climbing fibres projecting to the C3 zone was the same as in cats with an intact spinal cord. Also, the topography of tactile and nociceptive receptive fields and the distribution of A- and C fibre-evoked climbing fibre field potentials was similar to that in cats with an intact spinal cord. 3. In cats with an initially intact spinal cord the cutaneous nociceptive C fibre input and the topographically well organized tactile input to the C3 climbing fibres disappeared following a lesion of the ipsilateral and part of the contralateral dorsal funiculi. Following this lesion the receptive fields of the climbing fibres became indistinct and only irregular responses were evoked on skin stimulation. 4. It is concluded that the cutaneous nociceptive C fibre input from the forelimb to climbing fibres projecting to the C3 zone is mediated by the ipsilateral dorsal funiculus. Since cutaneous C fibres terminate exclusively in the spinal cord close to their entrance zone the postsynaptic dorsal column pathway must be part of this spino-olivocerebellar pathway.

Evidence of an x zone in lobule V of the squirrel monkey (Saimiri sciureus) cerebellum: the distribution of corticonuclear fibers.

The distribution of corticonuclear fibers to medial-most parts of the posterior interposed nucleus (NIP) from lateral areas of the vermis was studied in the squirrel monkey (Saimiri sciureus), using a silver impregnation method. The origin and course of degenerated fibers were studied in serial sections. The distribution pattern of corticonuclear fibers from a series of small well localized lesions placed in the vermis and paravermal cortex of lobule V is compatible with the interpretation that an x zone is present in Saimiri. A comparison of the positions of lesions and the trajectory of fibers arising therein suggests that corticonuclear input to medial-most parts of the NIP originated from a narrow cortical area (about 0.5-0.7 mm wide) located between a cortical area projecting into the medial cerebellar nucleus (the A zone) and a laterally adjacent area (the B zone) which related to the lateral vestibular nucleus. This NIP-projecting cortical area, located about 1.7 mm to 2.5 mm off the midline in lobule V, is interpreted as the x zone in this primate; it extends from lobule IV into lobule VI in squirrel monkey. Corticonuclear fibers of zone x in this primate form a comparatively small terminal field in the medial-most portions of NIP. This contrasts with the distribution of corticonuclear fibers of the C2 zone which consistently distribute to terminal fields that are shifted into more central areas of NIP. There appears to be no overlap of the corticonuclear terminal fields in the NIP for zone x versus the C2 zone. These results were correlated with data from the literature on the distribution of olivocerebellar fibers to the x zone and the C2 zone and the arrangement of cerebellar nucleoolivary projections into the inferior olive from the NIP. The x zone and the C2 zone both receive input from the contralateral medial accessory olive (MAO), both zones project into the NIP, and the NIP projects into those regions of the MAO which, in turn, project to these respective cortical zones and into the NIP. This suggest that the x zone is a component of the NIP-MAO circuit. Furthermore the proposed function of the x zone would support the view that this sagittal strip may have a more extensive rostrocaudal distribution in primates as compared to the cat.

Sensory integration in the spino-olivocerebellar pathways of the anaesthetized cat.

1. The responses evoked by peripheral nerve stimulation in the c1 and c3 zones of the cerebellar cortex have been examined in barbiturate-anaesthetized cats. The responses evoked via the spino-olivocerebellar pathways (SOCPs), which terminate in the cortex as climbing fibres, were recorded as positive multiunit field potentials from the cerebellar surface of lobule V. 2. Low-strength conditioning stimulation of the superficial radial, ulnar or median nerve frequently modified the climbing fibre-mediated responses evoked by a subsequent test stimulus to one of the other nerves. In most cases this modification involved a depression of the evoked response. The depression was not dependent on the conditioning stimulus evoking climbing fibre-mediated responses in the cortex. 3. The depression of the evoked responses increased as the conditioning stimulus intensity was raised within the range of 1.1-1.5 x threshold (1.1-1.5T). 4. Topical application of bicuculline to the surface of the dorsal column nuclei reduced the depression evoked by conditioning stimulation and it is therefore concluded that GABAergic inhibition in the cuneate nucleus contributes to the depression. 5. The inhibition is discussed in relation to its possible contribution to movement-related regulation of the excitability of SOCPs which occurs during locomotion in awake cats.

Topographic relationship between sagittal Purkinje cell bands revealed by a monoclonal antibody to zebrin I and spinocerebellar projections arising from the central cervical nucleus in the rat

We have examined the topographic relationship between the sagittal bands of zebrin I immunoreactive Purkinje cells revealed by a monoclonal antibody, mabQ113, and the distribution of spinocerebellar fibers originating from the central cervical nucleus in the rat. The mossy fiber terminals were anterogradely labeled following injections of cholera toxin subunit B into the C1-C3 segments and visualized immunohistochemically. Zebrin I positive Purkinje cells appeared in seven sagittal bands (P1+ to P7+ bands). In lobules I-V of the anterior lobe, labeled mossy fiber terminals were distributed in the midline region, subjacent to the P1+ bands and at around 0.5 mm from the midline region, subjacent to the P2+ band in the lateral A1 to the medial A2 zones of Voogd et al. (1985). Labeled terminals were seen in the entire B zone and those distributed in its medial part were related to the P3+ band. In lobule VIII, labeled terminals were seen subjacent to the P1+, P2+ and P3+ bands, which were located in the lateral A1-A3 (or B) zones. In the copula pyramidis, labeled terminals appeared subjacent to the P4+, P5+ and the P6+ bands in the C1 and C2 zones (or the C1-C3 zones). Although the labeled terminals were seen beneath the zebrin I positive bands, the borders of terminal distribution were not well-delineated, and did not respect the borders of zebrin I positive bands.

Gating in the spino‐olivocerebellar pathways to the c1 zone of the cerebellar cortex during locomotion in the cat.

1. The field potentials evoked in the cerebellar cortical c1 zone by single-pulse, non-noxious stimulation of the superficial radial nerve have been recorded with tungsten-in-glass microelectrodes in awake cats. Responses that were due to transmission in the spino-olivocerebellar pathways (SOCPs), which terminate in the cortex as climbing fibres, were identified and studied while the cat walked on a moving belt. 2. The size of the climbing fibre-evoked potentials varied systematically during the step cycle. They were invariably largest in mid- to late swing of the ipsilateral forelimb and, at most recording sites (5/6), they were smallest during the first half of stance. 3. With low stimulus strength, the probability of evoking a measurable response also varied. The probability was greatest in mid- to late swing and least in early stance. 4. Similar variations were shown to occur when the analysis was restricted to responses evoked by a single functionally homogenous SOCP, the dorsal funiculus SOCP. 5. It is proposed that these variations reflect the operation of a gating mechanism which modulates the excitability of the SOCPs and prevents them transmitting self-generated tactile inputs to the cerebellum while facilitating transmission when unexpected inputs are most likely to arise. 6. The present data are compared with those from a similar study of the c2 zone SOCPs (Apps, Lidierth & Armstrong, 1990) and are discussed in relation to a study of the effects of unexpected mechanical perturbations of stepping (Andersson & Armstrong, 1987).

Locomotion‐related variations in excitability of spino‐olivocerebellar paths to cat cerebellar cortical c2 zone.

1. Cutaneous nerve stimulation was used to study the excitability of the spino-olivocerebellar pathways (SOCPs) to the c2 zone of the paravermal cerebellar cortex in the cat. Non-noxious single-shock stimulation of the right and left superficial radial (SR) nerves via implanted cuff electrodes was used to evoke field potentials in the cerebellar cortex via the SOCPs. 2. The evoked potentials were recorded extracellularly either in lobule V of the anterior lobe (three cats) or within the paramedian lobule of the posterior lobe (one cat) with glass-coated tungsten microelectrodes. Measurement of the amplitudes of the responses was used to monitor transmission in the SOCPs in cats at rest and during walking. 3. A total of eleven c2 recording sites were investigated in detail. At seven of these sites, responses were recorded both during locomotion and at rest. For all seven sites responses during locomotion were smaller, more variable in amplitude and less securely evoked (average reduction 59%). 4. At five out of the eleven recording sites (45%) the mean amplitude of responses elicited during different tenths of the step cycle fluctuated sufficiently that the largest response was more than twice the smallest. In the majority of these cases (4/5) the responses were largest in either mid-stance or late swing. These fluctuations in response size occurred without parallel fluctuation in the amplitude of the peripheral nerve volley. At the remaining sites fluctuation of the cerebellar field size was less and in some cases practically absent. 5. At six recording sites it was possible to record the climbing fibre potentials evoked by stimulation of both the ipsilateral and contralateral superficial radial nerves. In all six cases the fluctuations in size of the response during locomotion occurred in phase, despite the fact that the two limbs move out of phase. 6. The probability that an individual stimulus would evoke any cerebellar response also varied between the different tenths of the step cycle and such variations occurred in parallel with the fluctuations in response size. This shows that the SOCP regulatory mechanism(s) must, at least in part, operate at a precerebellar level.

Topographical organisation within the cerebellar nucleocortical projection to the paravermal cortex of lobule Vb/c in the cat

The projection from the intracerebellar nuclei to the paravermal (intermediate) cerebellar cortex of lobule Vb/c has been investigated in the cat using a combined electrophysiological and neuroanatomical technique. A small (10-30 nl) injection of WGA-HRP was made into one of the three paravermal zones (c1, c2 or c3) after the mediolateral boundaries of the zones had been delimited on the cerebellar surface by recording climbing fibre field potentials evoked in response to percutaneous stimulation of one or more paws. The distribution of retrogradely labelled cell bodies within the intracerebellar nuclei was compared with the distribution of terminal labelling arising from anterograde transport by cerebellar Purkinje cells. The three paravermal zones displayed marked heterogeneity in their receipt of a projection from the intracerebellar nuclei. The c1 and c3 zones received virtually no such input, although injections in either zone resulted in significant terminal labeling (which was largely restricted to nucleus interpositus anterior). By contrast, the intervening c2 zone received a much heavier nucleocortical input which arose almost exclusively from nucleus interpositus posterior (to which the zone also projected). A sparse contralateral nucleocortical input to the c2 zone was also demonstrated. This arose primarily from nucleus fastigius. It is concluded that the nucleocortical projection to the paravermal cortex of lobule Vb/c displays marked topographical specificity and some functional implications of this are discussed.

The olivocerebellar projection to lobules I and II.

The olivocerebellar projection to lobules I and II was studied by means of retrograde transport from implants of the crystalline WGA-HRP complex. Retrogradely labelled neurons were found in the medial and dorsal accessory olives. Judged from the distribution of labelled cells, we conclude that parasagittally the olivocerebellar terminal zones A and B (i.e., the cerebellar cortical strips receiving axons from the olivary A and B regions) extend anteriorly into lobule Ia, whereas the fused olivocerebellar terminal C1/C3 zones reach lobule IIa. The olivocerebellar terminal C2 zone extends into lobule IIIa but not into lobule IIa. In lobule I the medial border of the B zone lies about 1 mm from the midline, in lobule II the B zone extends somewhat more medially. The lateral border of this zone is 1.9-2 mm from the midline. Compared to previous results, it appears that most of the Purkinje cells in lobule I projecting to the vestibular nuclei lie medially to the olivocerebellar terminal B zone.