The human capacity for vibrotactile frequency discrimination has been compared directly for glabrous and hairy skin regions by means of a two-alternative, forced-choice psychophysical procedure in five subjects. Sinusoidal vibratory stimuli, delivered by means of a 4-mm-diam probe, were first used to obtain detection threshold values for the two skin sites, the finger tip and the dorsal forearm, at four standard frequencies, 20, 50, 100, and 200 Hz. Values confirmed previous results showing detection thresholds were markedly higher on hairy skin than on glabrous skin. For the discrimination task, each standard frequency, at an amplitude four times detection threshold, was paired with a series of comparison frequencies, and discrimination capacity then was quantified by deriving from psychometric function curves, measures of the discriminable frequency increment (Deltaf) and the Weber Fraction (Deltaf/f), which, when plotted as a function of the four standard frequencies, revealed similar capacities for frequency discrimination at the two skin sites at the standard frequencies of 20, 100, and 200 Hz but an equivocal difference at 50 Hz. Cutaneous local anesthesia produced a marked impairment in vibrotactile detection and discrimination at the low standard frequencies of 20 and 50 Hz but little effect at higher frequencies. In summary, the results reveal, first, a striking similarity in vibrotactile discriminative performance in hairy and glabrous skin despite marked differences in detection thresholds for the two sites, and, second, the results confirm that vibrotactile detection and discrimination in hairy skin depend on superficial receptors at low frequencies but depend on deep, probably Pacinian corpuscle, receptors for high frequencies.

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 disappearan...

Hz with a progressive decline at higher frequencies. Above 400 Hz, impulse activity occurred almost randomly throughout the vibratory stimulus cycle and therefore carried little further signal of vibratory frequency. The decline, with increasing frequency, in the ability of cuneate neurones to signal information about vibratory frequency parallels the known subjective capacities for frequency discrimination.5. A switch-over occurred at approximately 80 Hz in the population of cuneate neurones able to provide the more reliable signal of vibratory frequency; above 80 Hz, the Pacinian neurones; below 80 Hz, the neurones receiving intradermal, rapidly adapting receptor input from the pads. P. R. DOUGLAS, D. G. FERRINGTON AND M. ROWE 6. The observed properties of cuneate neurones are compatible with a role in signalling information which could contribute to subjective tactile abilities.

The organization of sensory and motor regions of the cerebral cortex has been studied in the platypus (Omithorhynchus anatinus), of the order Monotremata. Comparisons were made with the organization found in the other representative of this order, the echidna, and with primitive species of eutherian and metatherian mammals. Evoked potential and single neuron studies revealed a large single area of somatosensory representation in the posterior region of the hemisphere, extending from an approximately mid-sagittal position around to the region of the rhinal sulcus on the ventrolateral surface of the hemisphere. The mediolateral representation of contralateral body parts was consistent with the pattern in the primary somatosensory area of other mammalian species. No evidence of a second somatosensory area was found. Neurons with similar receptive fields were grouped in columns normal to the cortical surface and a highly ordered pattern of somatotopic representation was found. Within the large area of bill representation individual neurons had receptive fields which were often punctate and no more than 1 mm in diameter. They responded to dynamic components of tactile stimuli delivered to their receptive fields on the bill.Movements on the contralateral side of the body could be elicited by bipolar electrical stimulation over an area on the dorsal surface of the hemisphere which largely overlapped the somatosensory area, but extended further anteriorly towards the frontal pole of the hemisphere. Visual and auditory projection regions were found overlapping the somatosensory area in the posterior part of the hemisphere. The auditory area overlapped the visual area and appeared to be displaced posteromedially in relation to its position in other species, a displacement which may be a consequence of the large expanse of cortical area associated with the bill.The observation that a large proportion of cortical area is devoted to specific sensory and motor function in platypus corresponds with earlier findings in primitive eutherian and metatherian species. The platypus neocortex appears to represent a more primitive stage of cortical development than that found in the other member of the order Monotremata, the echidna.

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)

SUMMARY1. Responses were recorded in decereberate, unanaesthetized cats from individual cuneate neurones in order to determine firstly, the afferent sources of inhibition on cuneate neurones and secondly, the influence of afferent-induced inhibition on those response features of dynamically sensitive tactile neurones which determine their capacity to code information about parameters of tactile stimuli.2. For all cuneate neurones which displayed afferent-induced inhibition from areas surrounding or within their excitatory receptive field (71 % of the sample) it was consistently found that 300 Hz vibration at low amplitudes (<25-50 um) which selectively engages Pacinian corpuscles was an effective source of inhibition. In contrast, steady indentation which activates slowly adapting tactile afferents was quite ineffective, as was low frequency vibration (30 Hz) at amplitudes of < 50-100 jsm. The latter stimulus can be used to engage rapidly adapting receptors either within glabrous skin (presumed to be Meissners corpuscles) or in association with hair follicles. It is concluded that afferents from Pacinian corpuscles are the dominant or exclusive source of afferent-induced inhibition of cuneate neurones.3. For dynamically sensitive neurones responsive to low frequency cutaneous vibration (30 Hz) there was a reduction in the slope of stimulusresponse relations with afferent-induced inhibition, but no expansion of the range of stimulus amplitudes over which the neurone responded.4. The influence of afferent-induced inhibition on the phase-locking of impulse activity to a cutaneous vibratory wave form was examined by constructing post-stimulus time histograms and cycle histograms. Measures of dispersion of impulse activity around the preferred point of firing in the vibratory waveform indicated that the capacity of individual cuneate neuroses to code information about the frequency of the cutaneous 252 E. BYSTRZYCKA, B. S. NAIL AND MARK ROWE vibration was not systematically changed in the presence of afferentinduced inhibition.

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SUMMARY1. 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 see 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 counterparts. They displayed graded stimulus-response relations which, over the steepest segment of the curves, had mean slopes of 15-7 impulses/100um 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.