Mastication-related neurons in the orofacial first somatosensory cortex of awake cats

In the orofacial area of the first somatosensory cortex (SI), we recorded single unit activity from 699 neurons in 11 awake cats. Fifty-two percent (362/699) were mastication-related neurons (MRNs) showing activity related to some aspects of masticatory movements. MRNs were divided into three types by their activity patterns: (1) the rhythmical type, showing rhythmical bursts in pace with the masticatory rhythm; (2) the sustained type, showing a sustained firing during the period of taking food and (3) the transient (biting) type, showing intense discharges in coincidence with biting hard food. MRNs had mechanoreceptive fields in the perioral, tongue, periodontal and mandibular regions. The activities of perioral rhythmical-MRNs, mandibular transient-MRNs, tongue rhythmical-MRNs and periodontal transient-MRNs were correlated with food texture, while perioral rhythmical-MRNs, perioral sustained-MRNs and tongue sustained-MRMs were not. Both facial and intraoral MRNs were scattered throughout the facial and intraoral projection areas in SI. These findings provide evidence that the orofacial SI monitors masticatory movements for food ingestion.     

Deficits of masticatory movements caused by lesions in the orofacial somatosensory cortex of the awake cat

The purpose of this study was to determine the role of somatosensory cortex (SI) in the control of orofacial movements during eating.We identified perioral and tongue projection regions of the cat SI and destroyed cells in one region by injecting kainic acid. The effects on orofacial behavior were then studied over a period of 4-6 weeks. Cats with unilateral lesions in the perioral region (PL-cats) dropped food from the contralateral side of the mouth in the early phase. Failure in erection of the contralateral whisker hairs during masticatory movements and delay of the masticatory start were observed throughout the experimental period. Furthermore, in the late phase, PL-cats showed prolongations of the masticatory and food intake periods, which were accompanied by the increase in the number of swallows and chewing cycles. Cats with unilateral lesions in the tongue region (TL-cats) showed the prolongation of the masticatory period in the early phase, which was accompanied by the increase in the number of swallows and chewing cycles. TL-cats did not show the prolongation of the food intake period and failure in erection of the contralateral whisker hairs. In both PL- and TL-cats, masticatory and swallowing rhythms were normal.     

Deficits of masticatory movements caused by lesions in the orofacial somatosensory cortex of the awake cat

The purpose of this study was to determine the role of somatosensory cortex (SI) in the control of orofacial movements during eating. We identified perioral and tongue projection regions of the cat SI and destroyed cells in one region by injecting kainic acid. The effects on orofacial behavior were then studied over a period of 4-6 weeks. Cats with unilateral lesions in the perioral region (PL-cats) dropped food from the contralateral side of the mouth in the early phase. Failure in erection of the contralateral whisker hairs during masticatory movements and delay of the masticatory start were observed throughout the experimental period. Furthermore, in the late phase, PL-cats showed prolongations of the masticatory and food intake periods, which were accompanied by the increase in the number of swallows and chewing cycles. Cats with unilateral lesions in the tongue region (TL-cats) showed the prolongation of the masticatory period in the early phase, which was accompanied by the increase in the number of swallows and chewing cycles. TL-cats did not show the prolongation of the food intake period and failure in erection of the contralateral whisker hairs. In both PL- and TL-cats, masticatory and swallowing rhythms were normal.     

Spontaneous spiking and synaptic depression underlie noradrenergic control of feed-forward inhibition

Inhibitory interneurons across diverse brain regions commonly exhibit spontaneous spiking activity, even in the absence of external stimuli. It is not well understood how stimulus-evoked inhibition can be distinguished from background inhibition arising from spontaneous firing. We found that noradrenaline simultaneously reduced spontaneous inhibitory inputs and enhanced evoked inhibitory currents recorded from principal neurons of the mouse dorsal cochlear nucleus (DCN). Together, these effects produced a large increase in signal-to-noise ratio for stimulus-evoked inhibition. Surprisingly, the opposing effects on background and evoked currents could both be attributed to noradrenergic silencing of spontaneous spiking in glycinergic interneurons. During spontaneous firing, glycine release was decreased due to strong short-term depression. Elimination of background spiking relieved inhibitory synapses from depression and thereby enhanced stimulus-evoked inhibition. Our findings illustrate a simple yet powerful neuromodulatory mechanism to shift the balance between background and stimulus-evoked signals.     

C-fibres provide a source of masking inhibition to primary somatosensory cortex.

Capsaicin was applied to the exposed radial nerve of adult flying foxes (n = 5) and cats (n = 2) while recording in primary somatosensory cortex from a single neuron with a receptive field on digits 1 or 2. Within four minutes of application of capsaicin the borders of these receptive fields dramatically expanded. In a further four flying foxes it was shown, with subcutaneous delivery just proximal to the receptive fields, that capsaicin need affect only afferents from the region of a neuron's receptive field to induce expansion. Capsaicin applied directly to a nerve, or subcutaneously in high concentrations, is a selective neurotoxin that rapidly prevents the propagation of action potentials in most C-fibres. The result provides a partial explanation for experiments involving the specific and complete denervation of receptive fields of neurons in primary somatosensory cortex. Such denervation does not lead to unresponsiveness but to immediate sensitivity to stimulation of areas surrounding the original fields. Thus it appears that some subclass of capsaicin-sensitive C-fibres provides a primary source for the masking inhibition that normally limits the extent of the receptive fields of cortical neurons.     

Population coding in somatosensory cortex

Computational analyses have begun to elucidate which components of somatosensory cortical population activity may encode basic stimulus features. Recent results from rat barrel cortex suggest that the essence of this code is not synergistic spike patterns, but rather the precise timing of single neuron's first post-stimulus spikes. This may form the basis for a fast, robust population code.     

Habituation in the somatosensory cortex

Responses of neurons without spontaneous activity ("silent") to prolonged stimulation of the mesenteric nerves were studied in cats anesthetized with chloralose (65–70 mg/kg) and immobilized with flaxedil. The results showed that neurons of the association and primary projection areas are characterized by habituation to prolonged stimulation of visceral nerves. The rate of development of habituation depends on the parameters of stimulation and on the cortical region studied. Habituation developed more rapidly in the association area and was slower to develop at the focus of maximal activity of the second somatosensory area. The special features of cortical habituation during visceral stimulation are discussed.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 11, No. 5, pp. 412–418, September–October, 1979.     

Somatosensory integration controlled by dynamic thalamocortical feed-forward inhibition

The temporal features of tactile stimuli are faithfully represented by the activity of neurons in the somatosensory cortex. However, the cellular mechanisms that enable cortical neurons to report accurate temporal information are not known. Here, we show that in the rodent barrel cortex, the temporal window for integration of thalamic inputs is under the control of thalamocortical feed-forward inhibition and can vary from 1 to 10 ms. A single thalamic fiber can trigger feed-forward inhibition and contacts both excitatory and inhibitory cortical neurons. The dynamics of feed-forward inhibition exceed those of each individual synapse in the circuit and are captured by a simple disynaptic model of the thalamocortical projection. The variations in the integration window produce changes in the temporal precision of cortical responses to whisker stimulation. Hence, feed-forward inhibitory circuits, classically known to sharpen spatial contrast of tactile inputs, also increase the temporal resolution in the somatosensory cortex.     

Ipsilateral median somatosensory evoked potentials recorded from human somatosensory cortex

Somatosensory evoked potentials (SEP) to ipsilateral and contralateral median nerve stimulations were recorded from subdural electrode grids over the perirolandic areas in 41 patients with medically refractory focal epilepsies who underwent evaluation for epilepsy surgery. All patients showed clearly defined, high-amplitude contralateral median SEPs. In addition, four patients showed ipsilateral SEPs. Compared with the contralateral SEPs, ipsilateral SEPs were very localized, had a different spatial distribution, were of considerably lower amplitude, had a longer latency (1.2–17.8 ms), did not show an initial negativity, and were markedly attenuated during sleep. Stimulation of the subdural electrodes overlying the sensory hand area was associated with contralateral hand paresthesias, but no ipsilateral hand paresthesias occurred. It was concluded that subdurally recorded cortical SEPs to ipsilateral stimulation of the median nerve (M) reflect unconscious sensory input from the hand possibly serving fast bimanual hand control. The anatomical pathway of these ipsilateral short-latency MSEPs is not yet known. Transcallosal transmission seems unlikely because of the short delay between the ipsilateral and contralateral responses in selected cases. The infrequent occurrence of ipsilateral subdurally recorded SEPs and their low amplitude and limited distribution suggest that they contribute very little to the short-latency ipsilateral median SEPs recorded on the scalp.     

Neural generators of early cortical somatosensory evoked potentials in the awake monkey

Controversy continues to exist regarding the generators of the initial cortical components of the somatosensory evoked potential (SEP). This issue was explored by detailed epidural and intracortical mapping of somatosensory evoked activity in Old World monkeys. In depth recordings, 3 complementary procedures were utilized: (1) the intracortical and subcortical distribution of SEPs was determined from approximately 4000 locations; (2) concomitant profiles of multiple unit activity (MUA) were recorded as an estimate of local action potential profiles; (3) 1-dimensional calculations of current source density (CSD) were used to outline the timing and pattern of regional transmembrane current flow. Our analysis confirms the participation of multiple cortical areas, located on either side of the central sulcus, in the generation of the initial cortical SEP components. Earliest activity, P10, was localized to area 3, followed within milliseconds by activation of areas 1, 2 (P12), and 4 (P13). In SI (Brodmann's areas 3, 1 and 2), the initial SEP components reflect the depolarization of lamina 4 stellate cells and the subsequent activation of adjacent pyramidal cells in laminae 3 and 5. The genesis of later cortical components (P20, N45) represents the composite of activity distributed across multiple cortical laminae and the interaction of overlapping excitatory and inhibitory events. These findings have direct implications for the clinical interpretation of SEP waveforms.     

Timing and specificity of feed-forward inhibition within the LGN

Local interneurons provide feed-forward inhibition from retinal ganglion cells (RGCs) to thalamocortical (TC) neurons, but questions remain regarding the timing, magnitude, and functions of this inhibition. Here, we identify two types of inhibition that are suited to play distinctive roles. We recorded excitatory and inhibitory postsynaptic currents (EPSCs/IPSCs) in TC neurons in mouse brain slices and activated individual RGC inputs. In 34% of TC neurons, we identified EPSCs and IPSCs with identical thresholds that were tightly correlated, indicating activation by the same RGC. Such "locked" IPSCs occurred 1 ms after EPSC onset. The remaining neurons had only "nonlocked" inhibition, in which EPSCs and IPSCs had different thresholds, indicating activation by different RGCs. Nonlocked inhibition may refine receptive fields within the LGN by providing surround inhibition. In contrast, dynamic-clamp recordings suggest that locked inhibition improves the precision of synaptically evoked responses in individual TC neurons by eliminating secondary spikes.     

Afferent inhibition and functional properties of neurons in the vibrissae projection area of the cat somatosensory cortex]

The aim of this study was to investigate the role of inhibitory processes in S-1 cortex of cats. The inhibition was evoked by "natural" afferent stimulation of the fascial vibrissae. For this purpose, two neighboring vibrissae were sequentially stimulated by mechanical deflection; single unit activity was recorded simultaneously from the cortex. Results showed that conditioning by afferent stimulation significantly influenced the directional sensitivity of cortical neurons. These data and analysis of spatial pattern of stimulated vibrissa indicate that detector neurons could be quickly modified during sensory processing.     

Synaptic basis for developmental plasticity in somatosensory cortex

Sensory experience drives plasticity of the body map in developing and adult somatosensory cortex, but the synaptic mechanisms underlying such plasticity are not well understood. Recently, several mechanisms that are likely to contribute to map plasticity have been directly observed in response to altered experience in vivo. These mechanisms include long-term potentiation and long-term depression at specific excitatory synapses, competition between lemniscal (barrel) and non-lemniscal (septal) processing streams, and regulation of the number of inhibitory synapses.     

Electrophysiological actions of VIP in rat somatosensory cortex.

Electrophysiological and biochemical studies suggest that VIP may exert a facilitating action in the neocortical local circuitry. In the present study, we examined the actions of VIP and VIP + norepinephrine (NE) on somatosensory cortical neuron responses to direct application of the putative transmitters acetylcholine (ACh) and gamma-aminobutyric acid (GABA). Spontaneous and transmitter-induced discharges of cortical neurons from halothane-anesthetized rats were monitored before, during and after VIP, NE and VIP + NE iontophoresis. In 57 VIP-sensitive cells tested, VIP application (5-70 nA) increased (n = 18), decreased (n = 36) or had biphasic actions (n = 3) on background firing rate. In a group of 20 neurons tested for NE + VIP, the combined effect of both peptide and bioamine was predominantly (70%) inhibitory. On the other hand, inhibitory and excitatory responses of cortical neurons to GABA (11 of 15 cases) and ACh (10 of 18 cases), respectively, were enhanced during VIP iontophoresis. Concomitant application of VIP and NE produced additive (n = 2) or more than additive (n = 3) enhancing effects on GABA inhibition. NE administration reversed or enhanced further VIP modulatory actions on ACh-induced excitation. These findings provide electrophysiological evidence that NE and VIP afferents may exert convergent influences on cortical neuronal responses to afferent synaptic inputs such that modulatory actions are anatomically focused within the cortex.     

An examination of somatosensory area SIII in ferret cortex

A somatotopically organized region on the suprasylvian gyrus of the ferret was examined using multiunit recordings and anatomical tracer injections. This area, which contains a representation of the face, was bordered by the primary somatosensory area (SI), anteriorly, and by the visually responsive rostral posterior parietal cortex (PPr), posteriorly. Anatomical tracers revealed connections to this region from cortical areas MI, SI, MRSS, PPr, and the thalamic posterior nucleus. These results are consistent with previous work in ferrets as well as with the location, physiology, and connectivity of area SIII in cats. Given its associations, functional properties, location, and homology, it is proposed that this region represents the third cortical somatosensory area (SIII) in ferrets.