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Monotremes, perhaps more than any other order of mammals, display an enormous behavioural reliance upon the tactile senses. In the platypus, Ornithorhynchus anatinus, this is manifest most strikingly in the special importance of the bill as a peripheral sensory organ, an importance confirmed by electrophysiological mapping that reveals a vast area of the cerebral cortex allocated to the processing of tactile inputs from the bill. Although behavioural evidence in the echidna, Tachyglossus aculeatus, suggests a similar prominence for tactile inputs from the snout, there is also a great reliance upon the distal limbs for digging and burrowing activity, pointing to the importance of tactile information from these regions for the echidna. In recent studies, we have investigated the peripheral tactile neural mechanisms in the forepaw of the echidna to establish the extent of correspondence or divergence that has emerged over the widely different evolutionary paths taken by monotreme and placental mammals. Electrophysiological recordings were made from single tactile sensory nerve fibres isolated in fine strands of the median or ulnar nerves of the forearm. Controlled tactile stimuli applied to the forepaw glabrous skin permitted an initial classification of tactile sensory fibres into two broad divisions, according to their responses to static skin displacement. One displayed slowly adapting (SA) response properties, while the other showed a selective sensitivity to the dynamic components of the skin displacement. These purely dynamically-sensitive tactile fibres could be subdivided according to vibrotactile sensitivity and receptive field characteristics into a rapidly adapting (RA) class, sensitive to low frequency (

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.

The functional organization of the primate somatosensory system at thalamocortical levels has been a matter of controversy, in particular, over the extent to which the primary and secondary somatosensory cortical areas, SI and SII, are organized in parallel or serial neural networks for the processing of tactile information. This issue was investigated for the marmoset monkey by recording from 55 single tactile-sensitive neurons in the lateral division of the ventral posterior nucleus of the thalamus (VPL) with a projection to either SI or SII, identified with the use of the antidromic collision technique. Neurons activated from the hand and distal forearm were classified according to their peripheral source of input and characterized in terms of their functional capacities to determine whether the direct thalamic input can account for tactile processing in both SI and SII. Both the SI- and SII-projecting samples contained a slowly adapting (SA) class of neurons, sensitive to static skin displacement, and purely dynamically sensitive tactile neurons that could be subdivided into two classes. One was most sensitive to high-frequency (> or =100 Hz) cutaneous vibration whose input appeared to be derived from Pacinian sources, while the other was sensitive to lower frequency vibration (< or =100 Hz) or trains of rectangular mechanical pulse stimuli, that appeared to receive its input from rapidly adapting (RA) afferent fibers presumed to be associated with intradermal tactile receptors. There appeared to be no systematic differences in functional capacities between SI- and SII-projecting neurons of each of these three classes, based on receptive field characteristics, on the form of stimulus-response relations, and on measures derived from these relations. These measures included threshold and responsiveness values, bandwidths of vibrational sensitivity, and the capacity for responding to cutaneous vibrotactile stimuli with phase-locked, temporally patterned impulse activity. The analysis indicates that low-threshold, high-acuity tactile information is conveyed directly to both SI and SII from overlapping regions within the thalamic VP nucleus. This direct confirmation of a parallel functional projection to both SI and SII in the marmoset is consistent with our separate studies at the cortical level that demonstrate first, that tactile responsiveness in SII largely survives the SI inactivation and second, that SI responsiveness is largely independent of SII. It therefore reinforces the evidence that SI and SII occupy a hierarchically equivalent network for tactile processing.

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. 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. Responsiveness within the hand region of the second somatosensory area of cortex (SII) was investigated in the marmoset monkey (Callithrix jacchus) in association with cooling-induced, reversible inactivation of the primary somatosensory area, SI. The aims were to determine whether thalamocortical systems in this primate species are organized according to a serial scheme in which tactile information is conveyed from the thalamus to SI and thence to SII as the next hierarchical level of processing and to establish whether primates are fundamentally different, in this respect, from mammals in which tactile information is conveyed in parallel from the thalamus to both SI and SII. 2. Inactivation of the SI had area was achieved when the temperature at the face of the silver cooling block over this SI region was lowered to < or = 13 degrees C. Inactivation was confirmed by abolition of the SI surface potential evoked by a brief tap stimulus to the hand and by the abolition of responsiveness in single SI neurons located beneath and around the edge of the block. 3. The effect of SI inactivation on SII-evoked potentials was investigated in 20 experiments by simultaneous recording of the SI- and SII-evoked potentials. The SII response was never abolished and was unchanged in the majority (12/20) of experiments. In the remainder, the SII-evoked potentials underwent a reduction in amplitude that was usually < 30% but never > 50%. 4. Tactile responsiveness was examined quantitatively in 47 individual SII neurons of different functional classes before, during, and after the inactivation of SI. Controlled tactile stimuli consisted of trains of sinusoidal vibration or rectangular pulses delivered to the glabrous or hairy skin of the hand. 5. Thirteen of the 47 SII neurons (28%) were unaffected in their response levels in association with SI inactivation. The remaining 34 SII neurons underwent some reduction in responsiveness, but in only 6% (3/47) was responsiveness abolished by SI inactivation. As the same range of functional classes of tactile neurons were represented among the affected and unaffected SII neurons, there was no evidence for a differential susceptibility among SII tactile neurons to the effect of SI inactivation. 6. Where reductions in amplitude of the SII-evoked potential or in response levels of SII neurons were observed, the effects were not attributable to direct spread of cooling from SI to the SII hand area as there was no cooling-induced prolongation of either the evoked potential or spike waveform in SII, an effect that is known to precede cooling-induced reductions in responsiveness. 7. These lines of evidence indicate that reductions in SII responsiveness in association with SI inactivation may be attributable to a loss of a background facilitatory influence rather than to a blockage of a component of peripheral input that comes over a putative serial path to SII via SI. First, as SI was cooled, there was a progressive increase in latency and time course of the SI responses before their disappearance, but no comparable delay in the SII responses as might be expected if SI were placed earlier than SII in a strict hierarchical scheme of thalamocortical processing. Second, SI inactivation failed to bring about a tightening in the phase-locking of SII responses to vibrotactile stimuli as might have been expected if the inputs to the SII neurons come via both a direct path from the thalamus and an indirect intracortical path via SI. Blockage of the indirect intracortical pathway through SI might be expected to reduce temporal dispersion in the input to SII neurons and result in an improvement in phase-locking in the SII responses to skin vibration. Third, the background activity of some SII neurons was reduced during SI inactivation along with the reduction in their responses to tactile stimulation.

Transmission from single, identified, slowly adapting type II (SAII) tactile fibers to their target neurons in the cuneate nucleus was examined in anesthetized cats. Simultaneous recordings were made from cuneate neurons and from fine, intact fascicles of the superficial radial nerve in which it was possible to identify and monitor the activity of each group II fiber. Selective activation of individual SAII fibers was achieved by means of skin stimulation with fine probes, in conjunction with extensive forelimb denervation. Responses were studied for seven SAII-driven cuneate neurons. For three there was unequivocal monitoring of the identified SAII input fiber. However, in six of the seven there was evidence that just one SAII fiber provided suprathreshold input to the cuneate neuron, and neither temporal nor spatial summation was required for reliable transmission. Cuneate impulse rates, in response to SAII inputs lasting 1 s, were less than 250 impulses per second, even though the SAII impulse rates could be 500 s-1. Responses to individual SAII impulses consisted of a burst of 2-3 impulses at low SAII input rates, but burst responses disappeared at high SAII rates. In all three SAII-cuneate pairs studied, the transmission security (the percentage of SAII impulses that evoked cuneate spike output) exceeded 80% in response to static skin displacement and in response to certain frequencies of skin vibration, in particular, at 100-200 Hz, exceeded 98% when the SAII fiber responded near the 1:1 level (one impulse per vibration cycle).(ABSTRACT TRUNCATED AT 250 WORDS)

1. Previous observations on the effect of ablation or inactivation of the primary somatosensory cortex (SI) on the responses of neurons within the second somatosensory area (SII) to tactile stimuli point to profound differences between monkeys and certain other mammals in the organization of thalamocortical systems. In the cat, for example, tactile information appears to be conveyed in parallel from the thalamus to both SI and SII, whereas, in macaque and marmoset monkeys, it is conveyed in a serial (or hierarchical) scheme from the thalamus to SI and thence to SII. The present study examined the responses of individual SII neurons during reversible, cooling-induced inactivation of SI in another nonprimate placental mammal, the rabbit, to obtain further evidence on whether the above differences might reflect a fundamental distinction between simian primates and other mammalian species. 2. When the temperature at the face of a silver cooling block over the forepaw and hindpaw regions of SI was lowered to 5-13 degrees C, the SI surface potentials evoked by brief tactile stimuli were abolished (indicative of SI inactivation), whereas SII potentials remained intact. 3. The responses of 25 SII neurons to controlled tactile stimuli (consisting of 1- to 1.5-s trains of vibration or rectangular mechanical pulses) were studied before, during, and after inactivation of SI. The effects on the spontaneous activity of a further three SII neurons that lacked identified receptive fields were also studied. 4. The response or activity levels of 26 of the 28 SII neurons examined (93%) were unaffected by SI inactivation.(ABSTRACT TRUNCATED AT 250 WORDS)

1. Localized cortical cooling was employed in anesthetized cats for the rapid reversible inactivation of the distal forelimb region within the primary somatosensory cortex (SI). The aim was to examine the responsiveness of individual neurons in the second somatosensory area (SII) in association with SI inactivation to evaluate the relative importance for tactile processing of the direct thalamocortical projection to SII and the indirect projection from the thalamus to SII via an intracortical path through SI. 2. Response features were examined quantitatively before, during, and after SI inactivation for 29 SII neurons, the tactile receptive fields of which were on the glabrous or hairy skin of the distal forelimb. Controlled mechanical stimuli that consisted of l-s trains of either sinusoidal vibration or rectangular pulses were delivered to the skin by means of small circular probes (4- to 8-mm diam). 3. Twenty-three of the 29 SII neurons (80%) showed no change in response level (in impulses per second) as a result of SI inactivation. These included seven neurons activated exclusively or predominantly by Pacinian corpuscle (PC) receptors, six that received hair follicle input, four activated by convergent input from hairy and glabrous skin, and six driven by dynamically sensitive but non-PC inputs from the glabrous skin. 4. Six SII neurons (20%), also made up of different functional classes, displayed a reduction in response to cutaneous stimuli when SI was inactivated. 5. Stimulus-response relations, constructed by plotting response level in impulses per second against the amplitude of the mechanical stimulus, showed that the effect of SI inactivation on individual neurons was consistent over the whole response range. 6. The reduced response level seen in 20% of SII neurons in association with SI inactivation cannot be attributed to direct spread of cooling from SI to the forelimb area of SII, as there was no evidence for a cooling-induced prolongation in SII spike waveforms, an effect that is known to precede any cooling-induced reduction in responsiveness. 7. As SI inactivation produced a fall in spontaneous activity in the affected SII neurons, we suggest that the inactivation removes a source of background facilitatory influence that arises in SI and affects a small proportion of SII neurons. 8. Phase-locking and therefore the precision of impulse patterning were unchanged in the responses of SII neurons to vibration during SI inactivation. This was the case whether response levels of neurons were reduced or unchanged by SI inactivation.(ABSTRACT TRUNCATED AT 400 WORDS)

Responses of neurons in the ventroposterolateral nucleus of the thalamus to vibration applied to the forelimb footpads were analyzed in anesthetized cats in order to describe the signalling properties of thalamic neurons that received input from the different classes of tactile afferents innervating the glabrous skin of the distal forelimb. Seventy-six thalamic neurons, the majority of which (60 of 76) were positively identified as thalamocortical projection neurons, were classified into two broad groups according to their responses to 1-s step indentations of the skin. A minority (24%) comprised neurons that had slowly adapting (SA) responses, whereas the remainder (76%), the dynamically sensitive neurons, had transient responses to the onset and offset phase of the step and were further classified according to their sensitivity to cutaneous vibrotactile stimuli into those activated by low-frequency vibration (rapidly adapting, RA, neurons) and those activated by high frequencies (Pacinian afferent, PC, neurons). Thalamic RA neurons displayed phaselocked responses to vibration at frequencies up to approximately 100 Hz, while PC neurons displayed phaselocked responses to vibration up to 400-500 Hz. Thalamic SA neurons varied in their responses to vibrotactile stimuli; half were most sensitive to vibration frequencies of 50 Hz or less, while the others responded over a broader range of frequencies. Although three major classes of footpad-related thalamic neurons were identified, there was evidence of convergent input to a small proportion of them. The study demonstrates that thalamic neurons have the capacity for responding to cutaneous vibration with phaselocked, patterned impulse trains, which would enable them to encode information about vibrotactile frequencies up to approximately 300 Hz.

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.

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.

Comparison of the information transmitted about the intensity of a steady "tactile" stimulus applied to facial skin by single trigemino-thalamic neurons in nucleus oralis and nucleus caudalis indicates that little information loss occurs at the medial lemniscal synaptic relay (nucleus oralis), but that it is gross within the nucleus caudalis.

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.