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Paired, simultaneous recordings were made in anaesthetized cats from the peripheral and central axons of individual tactile and kinaesthetic sensory fibres. The aim was to determine whether failure of spike propagation occurred at any of the three major axonal branch points in the path to their cuneate target neurones, and whether propagation failure may contribute, along with synaptic transmission failures, to limitations in transmission security observed for the cuneate synaptic relay. No evidence for propagation failure was found at the two major axonal branch points prior to the cuneate nucleus, namely, the T-junction at the dorsal root ganglion, and the major branch point near the cord entry point, even for the highest impulse rates (approximately 400 impulses s(-1)) at which these fibres could be driven. However, at the highest impulse rates there was evidence at the central, intra-cuneate recording site of switching between two states in the terminal axonal spike configuration. This appears to reflect a sporadic propagation failure into one of the terminal branches of the sensory axon. In conclusion, it appears that central impulse propagation over group II sensory axons occurs with complete security through branch points within the dorsal root ganglion and at the spinal cord entry zone. However, at high rates of afferent drive, terminal axonal propagation failure may contribute to the observed decline in transmission security within the cuneate synaptic relay.

1. The present experiments examined the capacity of external cuneate nucleus (ECN) neurones in the anaesthetized cat to respond to static and vibrotactile stretch of forearm extensor muscles. The aim was to compare their signalling capacities with the known properties of main cuneate neurones in order to determine whether there is differential processing of muscle spindle inputs at these parallel relay sites. 2. Static stretch (<= 2 mm in amplitude) and sinusoidal vibration were applied longitudinally to individual muscle tendons and responses recorded from single ECN neurones. The muscle-related ECN neurones that were sampled displayed a high sensitivity to both static and dynamic components of stretch, including muscle vibration at frequencies of 50-800 Hz, consistent with their dominant input being derived from primary spindle afferent fibres. 3. In response to ramp-and-hold muscle stretch, ECN neurones resembled their main cuneate counterparts in the pattern of their responses and in quantitative response measures. Their coefficients of variation in interspike intervals during steady stretch ranged from approximately 0.3 to 0.7, as they do in main cuneate responses, and their stimulus-response relations were graded as a function of stretch magnitude with low variability in responses at a fixed stretch amplitude. 4. In response to muscle vibration, ECN activity was tightly phase locked to the vibration waveform, in particular at frequencies of <= 150 Hz, where vector strength measures (R) were high (R >= 0.8) before declining as a function of frequency, with R values of approximately 0.6 at 300 Hz and <= 0.4 at 800 Hz. Both the qualitative and quantitative aspects of ECN responsiveness to the vibro-stretch disturbances were indistinguishable from those of the main cuneate neurones. 5. The results demonstrate a high transmission fidelity for muscle signals across the ECN and no evidence for differential synaptic transmission across the parallel main and external cuneate nuclei. Earlier limitations observed in the capacity of cerebellar Purkinje cells to respond to primary spindle inputs must therefore be imposed at synapses within the cerebellum.

1. The capacity of cuneate neurones to signal information derived from muscle spindle afferent fibres about static stretch or vibration of forearm extensor muscles was examined electrophysiologically in anaesthetized cats. 2. Static stretch (>= 2 mm in amplitude) and sinusoidal vibration (at frequencies of 50-800 Hz) were applied longitudinally to individual muscle tendons by means of a feedback controlled mechanical stimulator, and responses were recorded from individual cuneate neurones and from individual spindle afferent fibres. 3. Cuneate neurones sampled were located caudal to the obex and displayed a sensitivity to both vibration and static stretch of forearm muscles that was consistent with their input arising from primary spindle endings. In response to static muscle stretch, they displayed graded and approximately linear stimulus-response relations, and a stability of response level at fixed lengths that was consistent with these neurones contributing discriminative information about static muscle stretch. 4. In response to sinusoidal muscle vibration the cuneate neurones also showed graded stimulus-response relations (in contrast to spindle afferents which at low vibration amplitudes attain a plateau response level corresponding to a discharge of 1 impulse on each vibration cycle). Lowest thresholds were at 100-300 Hz and bandwidths of vibration sensitivity extended up to approximately 800 Hz. 5. Temporal precision in cuneate responses to muscle vibration was assessed by constructing phase scatter and cycle histograms from which measures of vector strength could be calculated. Cuneate responses displayed somewhat poorer phase locking (and lower vector strengths) than spindle afferent responses to vibration (a reflection of uncertainties associated with synaptic transmission). Nevertheless, the remarkable feature of cuneate responses to muscle vibration is the preservation of tight phase locking at frequencies up to 400-500 Hz, which presumably enables these central neurones to contribute accurate temporal information for the kinaesthetic sense in a variety of circumstances involving dynamic perturbations to skeletal muscle.

1. Peripheral nerves arising in joint capsules are known to contain a 'contaminating' contribution from muscle afferent fibres. In the present report we provide the first electrophysiological evidence that some joint afferent fibres may take an 'ectopic' path to the central nervous system via a nearby muscle nerve. 2. Experiments were conducted in anaesthetized cats in which a distal extension of the indicis proprius nerve was observed to project beyond its own muscle to the dorsal surface of the wrist joint capsule which is also supplied by the 'classic' wrist joint nerve, a branch of the dorsal interosseous nerve. Both the proximal and distal segments of the indicis proprius nerve were exposed for recording, by means of silver hook electrodes, while each segment remained in continuity. 3. Individual wrist joint afferent fibres with receptive fields on the dorsal surface of the joint capsule could be identified electrophysiologically within the distal segment of the indicis proprius nerve. In each of these cases the same fibre could also be identified at the proximal recording site. The identity of each of these simultaneously recorded units was established (1) by the short fixed interval between their times of spike occurrence, (2) from the exact correspondence of the capsular receptive field for the simultaneously recorded spikes, and (3) by the unfailing correlation in the presence, or absence, of the distally and proximally recorded spikes in association with either manual or controlled stimulation of the wrist joint capsule. Most joint afferent fibres identified with this projection path were in the group II range of conduction velocities and had conventional properties but group III fibres also appeared to be represented. 4. The present demonstration that some joint afferent fibres may be located within 'muscle' nerves emphasizes the importance of activating deep inputs, of joint or muscle origin, by adequate stimulation of the peripheral receptors in order to examine selectively the central actions of either source of input. Electrical stimulation of the peripheral nerves may lead to interpretative ambiguities.

1. The capacity of cuneate neurones to attain normal functional properties following neonatal median nerve injury was investigated with single neurone recording in anaesthetized cats, 12-24 months subsequent to a controlled crush injury. Effectiveness of the peripheral nerve injury was confirmed by the abolition of the median nerve compound action potential following the crush. 2. Cuneate recording was carried out after denervation of the forearm, apart from the median nerve, to ensure that neurones studied had receptive fields within the distribution zone of the regenerated median nerve. Controlled and reproducible tactile stimuli were used to evaluate the functional capacities of neurones to determine whether they were consistent with those reported earlier for cuneate neurones in cats that had normal peripheral nerve development. 3. Twenty-two cuneate neurones with well-defined tactile receptive fields within the distribution zone of the regenerated median nerve were classified according to their adaptation characteristics and functional properties. Slowly adapting neurones responded throughout static skin indentations and had graded and approximately linear stimulus-response relations over indentation ranges up to 1.5 mm. Rapidly adapting neurones responded to the dynamic phases of skin indentations and could be divided into two broad classes, one most sensitive to vibrotactile stimuli at 200-400 Hz which appeared to receive a predominant input from Pacinian corpuscle receptors, and a non-Pacinian group that included neurones most sensitive to skin vibration at 5-50 Hz which appeared to receive glabrous skin input from the rapidly adapting class of afferent fibres. 4. Based on the stimulus-response relations and on measures of phase locking in the responses to vibrotactile stimuli, it appears that the functional properties of cuneate neurones activated from the field of a regenerated median nerve subsequent to a neonatal nerve crush injury were consistent with those reported previously for 'control' cuneate neurones. The results indicate that cuneate neurones can acquire normal tactile coding capacities despite the disruption caused by prior crush injury to their peripheral nerve source.

1. Partial deafferentation, based on peripheral nerve section or local anaesthetic blockade, has been reported to induce both immediate loss of responsiveness and/or immediate reorganization in receptive fields of neurones in the somatosensory system. In the present study, in anaesthetized cats, we have used a rapid, reversible deafferentation procedure based on cold block of the median nerve in order to evaluate quantitatively the response characteristics of cuneate neurones (n = 39) before, during and after partial deafferentation. 2. The first hypothesis tested was that cuneate neurones with input from ulnar or superficial radial nerve fields in the vicinity of the median nerve field should undergo, in association with median nerve blockade, an increased level of responsiveness to tactile stimuli within the ulnar or radial nerve zone, and an expansion of their cutaneous receptive fields. However, among eighteen cuneate neurones of this type, there was no evidence for any systematic enhancement of responsiveness nor, in at least sixteen of the eighteen neurones, any evidence for receptive field expansion. 3. The second hypothesis tested was that cuneate neurones whose input came from both the median nerve and another peripheral nerve source should undergo, in association with median nerve blockade, an increase in responsiveness to the remaining input and an expansion of the receptive field into the field of that remaining nerve source. However, in none of thirteen neurones of this type tested was there evidence of such a change. 4. The third hypothesis was that cuneate neurones whose control' receptive fields were within the median nerve zone of deafferentation should show an emergence of novel receptive fields and responsiveness from areas around the field of innervation of the median nerve. However, in none of eight neurones of this type was there evidence for such changes in adjacent skin areas. 5. In conclusion, with the use of cold block of the median nerve for partial deafferentation, the present study has confirmed previous findings of denervation-related loss of responsiveness in dorsal column nuclei neurones. The conflicting findings in studies of central nervous system plasticity indicate the need to understand better factors that do and do not lead to acute central changes.

1. Studies in monkeys indicate that proprioceptive and tactile inputs are conveyed from the thalamus to the primary somatosensory cortex (SI) and thence to the secondary somatosensory area (SII) in a serial scheme. In contrast, in the cat, tactile information is conveyed in parallel from the thalamus to SI and SII. The present study, in the cat, employed reversible inactivation of SI to determine whether proprioceptive inputs to SII from joint receptors depend on an indirect serial path via SI or are conveyed over a direct path from the thalamus. 2. SI and SII foci for knee joint inputs were determined with evoked potential mapping. Reversible inactivation of the SI focus by cooling had no effect on the amplitude, latency or time course of SII potentials evoked by joint inputs. There was also no consistent effect on the response levels of individual SII neurones examined during SI inactivation. Furthermore, there was no attenuation of the later components of the responses, and therefore no evidence that these depended on an indirect path to SII via SI. 3. Results demonstrate that proprioceptive inputs project directly from thalamus to SII over a pathway organized in parallel with that to SI, in contrast to the serial scheme reported for proprioceptive processing in primates.

1. The synaptic linkage between single, identified slowly adapting type I (SAI) fibres and their central target neurones of the cuneate nucleus was examined in pentobarbitone-anaesthetized cats. Simultaneous extracellular recordings were made from individual cuneate neurones and from fine, intact fascicles of the lateral branch of the superficial radial nerve in which it was possible to identify and monitor the activity of each group II fibre. Individual SAI fibres were activated by static displacement and by vibration delivered with a fine probe (0.25-2 mm diameter) to their associated touch domes in the hairy skin of the forelimb. 2. Transmission properties across the synapse were analysed for nine SAI-cuneate pairs in which the single SAI fibre of each pair provided a suprathreshold input to the cuneate neurone. Neither spatial nor temporal summation was required for effective impulse transmission, and often more than 80% of SAI impulses led to a response in the cuneate neurone. Responses of the cuneate neurones to single SAI impulses occurred at a short, fixed latency (S.D. often < 0.1 ms), and frequently consisted of a burst of two or three impulses, at low SAI input rates in particular. 3. The tight phase-locking in the responses to vibration of single SAI fibres was preserved in the cuneate responses for frequencies up to approximately 400 Hz. However, as the impulse rates of the cuneate neurones were less than 150 impulses s-1, their impulse patterns could not directly signal the vibration periodicity at frequencies > 100-150 Hz despite 1:1 responses in their single SAI input fibres up to approximately 500 Hz. 4. The reliable transmission of touch dome-associated SAI input across the cuneate nucleus indicates that transmission failure at this first relay is unlikely to be responsible for the reported failure of touch dome-SAI inputs to contribute to tactile perception. 5. The transmission characteristics for the SAI fibres were very similar to those demonstrated previously for fibres associated with Pacinian corpuscles, which argues against any marked differential specialization in transmission characteristics for dorsal column nuclei neurones that receive input from different tactile fibre classes.

1. Slowly adapting type II (SAII) afferent fibres that supply the forelimb were isolated from the medial cutaneous nerve of anaesthetized cats and examined for their capacity to signal information about vibrotactile events in the hairy skin. 2. The SAII fibres had a single spot-like receptive field focus where they were highly sensitive to steady indentation and vibration applied with probes normal to the skin surface. However, their sensitivity was affected profoundly by the size of the stimulus probe, its position in relation to the receptive field focus and, to a lesser extent, the magnitude of any pre-indentation on which vibration was superimposed. Small stimulus probes (e.g. 250 microns diameter) were much more effective than larger (> or = 1-2 mm) ones, and small shifts in the position of the perpendicularly applied probe away from the receptive field focus led to a marked decline in responsiveness. 3. With appropriate choice of stimulus parameters for vibratory stimuli applied at the receptive field focus, the SAII fibres could respond at low threshold (< 100 microns), with a tightly phase-locked, regular 1:1 impulse pattern (one impulse per vibration cycle) that accurately signalled the vibration frequency over a bandwidth that extended to 600 Hz. Furthermore, their responses remained phase-locked up to 1000 Hz. Phase-locking in SAII fibres was marginally tighter than that in SAI fibres and comparable to that of Pacinian corpuscle fibres. 4. The sensitivity of forelimb SAII fibres to tangential skin stretch was directionally selective; stretch across the forelimb was much more effective than along its long axis. Vibration associated with tangential skin stretch led to a marked spatial expansion of the field of vibration sensitivity. SAII fibres could therefore signal information about natural stimuli that contain elements of skin stretch and vibration, as may be encountered when the forelimb brushes against textured surfaces. Should the SAII fibres fail to contribute to the sensory experience of vibrotactile stimuli, the explanation may be related to limitations imposed centrally on the processing of their signals. Nevertheless, the present results demonstrate that, with appropriate stimulus conditions, the SAII afferent fibres have much greater vibrotactile sensitivity than has been suggested by past studies.

1. Recordings were made from single slowly adapting type I (SAI) afferent fibres associated with touch domes in the cat hairy skin. Controlled vibratory stimuli were used first, to characterize the precision with which these SAI afferents reflect the temporal aspects of vibrotactile stimuli, and second, to determine whether earlier disparate reports of SAI responsiveness to vibration may be attributable to highly specific stimulus requirements. 2. Eighteen SAI fibres from femoral cutaneous nerve branches were examined; each was associated with one to three touch domes. SAI responses to both steps and sinusoidal vibration (1-1.5 s in duration) were affected profoundly by both probe size and position. Punctate stimulus probes (250 microns) produced much higher response levels and steeper stimulus-response relations than those elicited with large (2 mm) probes, probably on account of focal distortion created within the dome by the smaller probes. SAI sensitivity to vibration was also affected markedly by the amplitude of any pre-indentation on which the vibration was superimposed; sensitivity was much lower when the pre-indentation exceeded 100 microns, in particular with larger stimulus probes. 3. Measures of both vibration sensitivity and the precision of impulse patterning demonstrated that, if appropriate stimulus parameters are chosen, the SAI fibres can respond to 1 s trains of vibration (amplitude < or = 100 microns) in a tightly phase-locked, 1:1 manner for frequencies up to 500 Hz. At frequencies from approximately 100-500 Hz the SAI fibres displayed broad 1:1 plateaus, where their response rate remained constant over a range of amplitudes, and phased-locking was tightest. Responses remained phase-locked up to 1000 Hz, but could not follow the vibration with a 1:1 pattern above 500 Hz. 4. The results demonstrate that with appropriate stimulus parameters, touch dome-associated SAI fibres are capable of signalling vibrotactile information over a similar bandwidth of frequencies as do Pacinian sensory fibres. The variability in past reports of SAI vibration sensitivity may relate principally to differences in stimulus conditions. However, in view of the SAI capacity for responding to vibration with temporally precise, patterned activity, it appears that their reported failure to contribute to vibrotactile sensibility must be attributed to limitations imposed in the central processing of SAI signals.

1. The effect of changes in amplitude on the perceived pitch of cutaneous vibratory stimuli was studied in psychophysical experiments designed to test whether the coding of information about the frequency of the vibration might be based on the ratio of recruitment of the PC (Pacinian corpuscle-associated) and RA (rapidly adapting) classes of tactile sensory fibres. The study was based on previous data which show that at certain vibration frequencies (e.g. 150 Hz) the ratio of recruitment of the PC and RA classes should vary as a function of vibration amplitude. 2. Sinusoidal vibration at either 30 Hz or 150 Hz, and at an amplitude 10 dB above subjective detection thresholds was delivered in a 1 s train to the distal phalangeal pad of the index finger in eight human subjects. This standard vibration was followed after 0.5 s by a 1 s comparison train of vibration which (unknown to the subject) was at the same frequency as the standard but at a range of amplitudes from 2 to 50 dB above the detection threshold. A two-alternative forced-choice procedure was used in which the subject had to indicate whether the comparison stimulus was higher or lower in pitch (frequency) than the standard. 3. Marked differences were seen from subject to subject in the effect of amplitude on perceived pitch at both 30 Hz and 150 Hz. At 150 Hz, five out of the eight subjects reported an increase in pitch as the amplitude of the comparison vibration increased, one experienced no change, and only two experienced the fall in perceived pitch that is predicted if the proposed ratio code contributes to vibrotactile pitch judgements. At 30 Hz similar intersubject variability was seen in the pitch-amplitude functions. 4. The results do not support the hypothesis that a ratio code contributes to vibrotactile pitch perception. We conclude that temporal patterning of impulse activity remains the major candidate code for pitch perception, at least over a substantial part of the vibrotactile frequency bandwidth.

1. Recordings were made in decerebrate cats from gracile and cuneate neurones responding to vibration-induced inputs from Pacinian corpuscle (P.c.) receptors of the hind-limb and forelimb footpads. The two groups of neurones were compared, in particular for their capacities for responding to cutaneous vibration with phase-locked impulse patterns. 2. In both nuclei the P.c. neurones were most sensitive to vibration in the range 80 to greater than 600 Hz. Stimulus-response relations were similar for the two groups, as were measures derived from these relations such as response levels, absolute thresholds and the dynamic range (defined as the vibration amplitude range over which responses were graded). 3. At frequencies up to 300-400 Hz, responses for some neurones in both nuclei remained well phase locked to the vibration; however, quantitative analysis using a factorial analysis of variance indicated that the phase locking was poorer in gracile than cuneate neurones. 4. In both nuclei there was marked variability from neurone to neurone in measures of phase locking which may reflect variations in the extent of convergence of P.c. fibres upon different target neurones. For neurones in either nucleus that had comparatively tight phase locking of responses to vibration it is proposed that their output is functionally dominated by one or a few of their convergent P.c. input fibres.

The receptive fields of sixteen spinocervical tract (s.c.t.) cells whose responses were recorded extracellularly were mapped using discrete and uniform jets of air given at equally spaced locations on the clipped fur of cats anaesthetized with chloralose. All the cells whose receptive fields were on the thigh or upper hind limb showed approximately unimodal gradients of sensitivity to stimulation within their excitatory receptive fields. The response magnitudes declined steadily as the stimuli were moved sequentially from the centres to the peripheries of the fields and abrupt edges were not found. Spatial summation from within the excitatory receptive field was studied in twelve s.c.t. cells. These cells showed a poor ability to summate the responses to two spatially separated air jets when these stimuli were applied simultaneously within their receptive fields. No significant summation was found in twenty-five out of thirty-one trials and in six of these trials (four cells) the responses were significantly reduced. Summation was found in six trials (four cells). Lack of summation or response reduction was more prevalent when the individual response levels were low (less than impulses stimulus-1). These results are discussed in relation to similar findings for cells of somatosensory relay nuclei and cortex.

1. Responses were recorded from individual tactile afferent fibres isolated by microdissection from the median nerve of pentobarbitone-anaesthetized neonatal kittens (1-5 days post-natal age). Experiments were also conducted on adult cats to permit precise comparisons between neonatal and adult fibres.2. Neonatal fibres with receptive fields on the glabrous skin of the foot pads were classified into two broad groups, a slowly adapting class (40%) which responded throughout a 1 sec period of steady indentation and a rapidly adapting or dynamically sensitive class comprising 60% of units. Fibres in these two groups had overlapping conduction velocities in the range 4.3 to 7.5 m/sec and were believed to be the developing Group II afferents of the adult.3. Neonatal slowly adapting fibres qualitatively resembled their adult counter-parts. They displayed graded stimulus-response relations which, over the steepest segment of the curves, had mean slopes of 15.7 impulses/100 mum of indentation. Plateau levels of response were often reached at amplitudes of skin indentation of < 0.5-0.7 mm.4. Dynamically sensitive fibres with receptive fields on the glabrous skin were studied using sinusoidal cutaneous vibration which in the adult enables them to be divided into two distinct classes. However, in the neonate, they formed a continuum whether criteria of sensitivity or responsiveness were used.5. In response to vibration neonatal fibres differed from adult ones according to the following quantitative indices: (i) sensitivity as measured by both absolute thresholds and thresholds for a 1: 1 pattern of response, both of which were higher in the neonate than in the adult at all frequencies > 50 Hz and differed by an order of magnitude at frequencies >/= 200 Hz; (ii) responsiveness based on the mean impulse rate evoked at a fixed amplitude of cutaneous vibration; (iii) band width of vibratory sensitivity which in the neonate was confined to approximately 5-300 Hz whereas in the two classes of adult units it covered the range 5-800 Hz; (iv) capacity for coding information about vibration frequency. Impulse activity of neonatal fibres was less tightly phase-locked to the vibratory stimulus and showed a poorer reflection of the periodic nature of the vibratory stimulus than impulse patterns of adult units.6. The results reveal that tactile receptors and afferent fibres in the neonate are functionally immature. Their restricted coding capacities suggest that peripheral tactile sensory mechanisms impose limits on the ability of the new-born animal to derive information about its tactile environment.

1. The synaptic linkage between single, identified sensory fibres associated with Pacinian corpuscle (P.c.) receptors and central neurones of the dorsal column nuclei was examined in decerebrate or anaesthetized cats. Paired recordings were made from individual neurones in the gracile division of the dorsal column nuclei and from the hind-limb interosseous nerve in which it is possible to identify and monitor the activity of each P.c. fibre activated when recording from the intact nerve with a platinum hook electrode. Individual P.c. fibres were activated by vibration delivered with an 0.2 mm diameter probe to the interosseous P.c. receptors. 2. Thirty-five P.c. fibre-gracile neurone pairs were isolated in which activity in the single, identified P.c. fibre evoked suprathreshold responses (mean latency +/- S.D., 10.3 +/- 1.5 ms) in the gracile neurone. A single impulse arriving over one P.c. fibre could generate pairs or triplets of output spikes from several target neurones thus revealing a potent synaptic organization within the dorsal column nuclei for the transmission and amplification of weak sensory signals. 3. The potency of the linkage for some pairs resulted in post-synaptic response levels of up to 400 impulses s-1 when a single input fibre was discharging one impulse on each vibration cycle at 200-400 Hz. 4. Gracile neurones driven by single P.c. fibres had phase-locked responses to vibration at frequencies of up to 400-500 Hz. However, the responses displayed much greater phase dispersion than those of P.c. fibres, indicating that a major component of phase dispersion in the vibration-induced responses of dorsal column nuclei neurones is attributable to the properties of the synaptic linkage between an individual fibre and the target neurone. 5. The potent actions of single, identified P.c. fibres on their target neurones are consistent with the hypothesis that phase-locked responses in dorsal column nuclei neurones to vibration at 100-400 Hz may reflect the functional domination of the target neurone's output by one or a few of its converging fibres.

1. In decerebrate or anaesthetized cats, the vibration-induced responses of dorsal column nuclei neurones were examined, first, when their input came from simultaneously recorded pairs or other combinations of identified Pacinian corpuscle (P.c.) afferent fibres of the interosseous nerve, and secondly, when different convergent sets of P.c. fibres were engaged by footpad vibration. 2. Suprathreshold actions were observed on individual dorsal column nuclei neurones from two or more identified P.c. fibres. Recruitment of these convergent fibres usually led to summation in the dorsal column nuclei neurone as reflected in higher response levels compared with those evoked by single-fibre inputs. 3. When the input was increased from one to two or more identified P.c. fibres the dorsal column nuclei neurones could retain a single, dominant phase of response to high-frequency (greater than 100 Hz) vibration even though these fibres, in isolation, evoked responses in the target neurone at substantially different latencies. However, on average, phase locking was significantly tighter in response to single-fibre input than to multiple P.c.-fibre input. 4. Dorsal column nuclei neurones were also able to retain phase-locked responses to high-frequency vibration when phase differences between different convergent inputs were systematically introduced to alter the degree of synchrony in the activity arriving over convergent, identified P.c. fibres. 5. When the input to dorsal column nuclei neurones came from the skin it was found that with the recruitment of two converging sets of P.c. fibres the dorsal column nuclei neurones were able to retain phase-locked responses to high-frequency vibration even when phase shifts were introduced between the two sets of P.c. inputs. 6. In conclusion, the observed integrative processing by dorsal column nuclei neurones of vibration-induced inputs arriving over identified, convergent P.c. fibres, or sets of P.c. fibres, is consistent with our hypothesis that the retention of phase-locked responses to vibration at frequencies greater than or equal to 100 Hz may reflect the functional domination of the target neurone by just one or a few of its convergent input fibres.

The sciatic and saphenous nerves of one hind limb were sectioned in young adult cats anaesthetized with halothane. Between 19 and 55 days later, under chloralose anaesthesia, dorsal horn neurones in the L6 and L7 segments were recorded and their receptive field properties examined. In seven animals recordings were made from identified spinocervical tract, post-synaptic dorsal column and dorsolateral funicular neurones as well as from neurones that did not project through these pathways. Thirty-one neurones were intracellularly stained with horseradish peroxidase, and fifty-three were recorded extracellularly and located by reference to stained cells. In two animals (both 31 days after nerve section) no attempt was made to identify axonal projections of the dorsal horn neurones in order to avoid any effects of cervical cord search stimuli on the cells' properties, but all isolated extracellularly recorded units were examined. On the side ipsilateral to the nerve sections 143 units were recorded. In all experiments, neurones in the medial three-quarters of the dorsal horn had no discernible cutaneous, mechanosensitive receptive fields between 19 and 55 days after nerve section. There were only two exceptions to this generalization, one neurone being one of the most rostral cells in the sample (in caudal L5) and the other being one of the most caudal cells (in caudal L7). We present evidence to show that neither of these two neurones had inappropriate receptive fields in terms of the somatotopic organization of the dorsal horn. All other neurones with receptive fields on the skin were appropriately located in the somatotopic map laid out in the dorsal horn. There was no evidence for gross anatomical changes in the dendritic trees of dorsal horn neurones following sciatic and saphenous nerve sections. We have been unable to confirm that, following loss of cutaneous receptive fields by peripheral nerve section, dorsal horn neurones in adult cats acquire 'inappropriate' receptive fields. Possible reasons for this are discussed.

1. In pentobarbitone-anaesthetized cats, responses were recorded in peripheral nerves or cervical dorsal columns from sensory fibres associated with Pacinian corpuscle (P.c.) receptors in the forelimb footpads. Factors affecting the phase of response to cutaneous vibration in individual P.c. fibres, and the extent of phase coherence in the responses of different P.c. fibres were examined when sinusoidal vibratory stimuli at 100-400 Hz were delivered using a 1 mm diameter probe. 2. Increases in vibration amplitude from the absolute to the 1:1 threshold for the P.c. fibre led to phase advances in the response, often of about 60 deg, in over 85% of fibres tested at 200 and 300 Hz, but further increases had little effect. 3. Variations in stimulus position within the receptive field led to unpredictable changes in the response phase that ranged from minimal change to shifts of 180 deg. As the response phase was unrelated to the distance from the point of peak sensitivity it is likely that at high vibration frequencies (greater than or equal to 100 Hz) the recruited population of P.c. fibres will respond over the whole range of phase angles. 4. The calculated phase of spike initiation in different pairs of P.c. fibres that shared coincident points of best sensitivity on the skin ranged from near synchrony to maximum asynchrony indicating that there is little phase coherence even in the subpopulation of somatotopically related P.c. fibres recruited by high-frequency cutaneous vibration. 5. Paired recordings from P.c. fibres within the cervical dorsal columns revealed a broad range of phase discrepancies in the responses of P.c. fibres to vibration at 200 and 300 Hz. 6. Several hypotheses are considered to explain the known presence of phase-locked responses to high-frequency (greater than or equal to 100 Hz) vibration in the central neurones of dorsal column nuclei.

1. In cats anaesthetized with Nembutal climbing fibre (CF) responses evoked in individual cerebellar Purkyne cells by mechanical stimulation of the skin were conditioned by preceding stimuli both to the periphery and to the precruciate area of the cerebral cortex.2. Cortical stimulation, generally subthreshold for evoking a CF response itself, induced an inhibition of cutaneous evoked CF responses in 65% of all Purkyne cells tested. The maximum inhibition ranged from 40 to 100% of the control responses at conditioning-testing intervals of 30-70 msec and the duration of the inhibition was usually 125 msec.3. In most cases the corticofugal inhibition of cutaneously evoked CF responses was mediated by inhibitory mechanisms outside the cerebellar cortex, probably at relays within the spino-olivocerebellar pathways. Purkyne cells undergoing corticofugal inhibition were distributed widely within both the vermis and the pars intermedia of the anterior lobe.4. In 40% of all Purkyne cells tested, there was evidence for afferent inhibition of their peripherally evoked CF responses as revealed by conditioning stimuli applied to the skin outside the receptive field. Again it was found that the inhibition was exerted at levels prior to the cerebellum.5. It is concluded that the afferent input transmitted to Purkyne cells via climbing fibres can be modified by corticofugal and peripheral influences exerted on the relays of the CF pathways outside the cerebellar cortex.

1. A technique of reversible block of synaptic transmission through the dorsal column nuclei and the trigeminal nucleus caudalis (n. caudalis) has been employed to assess the somatic afferent input to individual posterior thalamic neurones in the cat. The axon projection to the cerebral cortex of these neurones has been identified by antidromic activation following cortical stimulation.2. Unitary responses in the nucleus ventralis posterolateralis (VPL) evoked by cutaneous stimulation were abolished or depressed following block of transmission in the dorsal column nuclei. Block in n. caudalis, however, depressed unitary responses in nucleus ventralis posteromedialis (VPM) evoked by facial skin stimulation in less than 10% of cells.3. A more complex source of the somatic input to the posterior nuclear region of the thalamus (PO) was found. It was most commonly noted that PO unitary activity evoked by cutaneous stimulation of the face was unaffected by block of synaptic transmission in n. caudalis. No uniform effect was observed on unitary responses in PO evoked by limb stimulation when transmission in the dorsal column nuclei was blocked.4. Antidromic activation from the cerebral cortex was seen in 69% of ventrobasal neurones. Most cells (66%) had ;antidromic cortical fields' restricted to a region consisting of a third to a half of the specific somatic projection areas. In 10% of cells evidence was obtained for discontinuous cortical ;antidromic fields' suggesting subcortical bifurcation of the projecting axon.5. An axon projection to the cortex was found in 35% of PO cells, about half of which projected only to specific somatic projection areas. Evidence for subcortical branching of the axon was obtained for seven PO cells.

1. In cats anaesthetized with Nembutal, the cutaneous receptive fields of individual cerebellar climbing fibres were assessed by recording the climbing fibre responses of single Purkyne cells following controlled mechanical stimulation (air jets, vibration, taps, pressure) of the foot pads of all four limbs and of the hairy skin of the limbs and the body.2. Three major types of cutaneous receptive fields of individual climbing fibres were recognized: (a) restricted fields generally confined to the distal areas of one limb only; (b) circumscribed fields on the distal areas of two to four limbs (discontinuous fields); and (c) widespread continuous fields extending over all or almost all of the body surface. A fourth group appears to receive cutaneous inputs from one limb and sensory input from deeper structures in other limbs.3. Thresholds for tapping of the foot pads were often quite low (< 100 mum indentation), and there was no noticeable difference in the distribution of thresholds between the climbing fibres having restricted or more widespread cutaneous receptive fields. Similarly, the latencies of climbing fibre responses evoked by vibration and tapping were in the same ranges for climbing fibres with receptive fields restricted to one limb only and for those having more widespread fields.4. In regard to location in the anterior lobe of the cerebellum it was found that those climbing fibres receiving inputs only from the ipsilateral forelimb projected to Purkyne cells located almost entirely in lobule V, whereas all other climbing fibres with restricted or more widespread receptive fields projected to Purkyne cells distributed widely within lobules II to V.