Proprioceptive alignment of visual and somatosensory maps in the posterior parietal cortex

A touch on one hand can enhance the response to a visual stimulus delivered at a nearby location [1, 2], improving our interactions with the external world. In order to keep such visual-tactile spatial interactions effective, the brain updates the continuous postural changes, like those typically accompanying hand actions, through proprioception, thus maintaining the somatosensory and visual maps in spatial register [2, 3]. The posterior parietal cortex (PPC) might be critical for such a spatial remapping [4]; nevertheless, a direct causal demonstration of its involvement is lacking. Here, we found that unattended touches to one hand enhanced visual sensitivity for phosphenes induced by occipital trancranial magnetic stimulation (TMS) [5] when the touched hand was spatially coincident to the reported location of the phosphenes in external space. Notably, this spatially specific crossmodal facilitation was maintained after hand crossing, suggesting an efficient visual-tactile remapping. Critically, after 1 Hz repetitive TMS interference [6] over the PPC, but not over the primary somatosensory cortex, phosphene detection was still enhanced by spatially coincident touches with uncrossed hands, but it was enhanced by spatially noncoincident touches after hand crossing. This is the first causal evidence in humans that the PPC constantly updates the representation of the body in space in order to facilitate crossmodal interactions.     

Basal forebrain lesions with or without reserpine injection inhibit cortical reorganization in rat hindpaw primary somatosensory cortex following sciatic nerve section.

To test the hypothesis that cortical reorganization depends on acetylcholine and one or more of the monoamines, the hindpaw cortex was mapped in eight different groups of mature rats: (1) untreated; (2) after sciatic nerve transection; (3) after intraperitoneal injections of reserpine, to reduce the level of cortical monoamines; (4) after ibotenic acid lesion of the nucleus basalis of Meynert (NBM), to destroy cholinergic cells projecting to the cortex; (5) after reserpine treatment and transection; (6) after ibotenic acid lesion and transection; (7) after reserpine treatment and ibotenic acid lesion; and (8) after reserpine treatment, ibotenic acid lesion, and transection. Four days after transection, the cortex had reorganized in the transected group. However, this process of reorganization was prevented in transected animals with NBM lesions. Treatment with reserpine alone did not inhibit the process of reorganization, nor did it enhance the effect of NBM lesion. Nonetheless, the animals treated with reserpine and transected had higher response thresholds in the reorganized cortex than did the animals that were treated but not transected. These data suggest that acetylcholine plays an important role in the early reorganization that follows deafferentation, and that one or more of the monoamines may have other influences on reorganization of the primary somatosensory cortex of adult rats.     

Differential acetylation of core histones in rat cerebral cortex neurons during development and aging

Core histones can be modified by aceylation and this modification has been correlated with the modulation of chromatin condensation and histone deposition. We have now studied the levels of acetylation of the core histones in rat brain cortical neurons from the middle of the period of neuronal proliferation through postnatal development and aging. The results show that the level of acetylation of H4 decreases with age. The kinetics of H4 deacetylation show a perinatal fast phase followed by a much slower phase that spans the rest of the period examined. H4 deacetylation is accounted for by the decrease of the monoacetylated species, the proportions of the more highly acetylated species remaining essentially constant. By contrast to histone H4, the overall levels of acetylation and the proportions of the different acetylated species of H2A, H2B and H3 remain unchanged throughout the period examined. Furthermore, the variants belonging to a given histone class always show the same level of acetylation. The fact that in neurons the level of monoacetylated H4 decreases during development and aging, in sharp contrast with the constancy of the levels of all other acetylated histone species, raises the possibility that in interphase chromatin monoacetylated H4 may have a central role in the modulation of chromatin structure. The results also suggest that the slow decrease of the proportion of monoacetylated H4 may imply a gradual loss of chromatin structural plasticity and thus lead to aging.     

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.     

Rhythm generation through period concatenation in rat somatosensory cortex

Rhythmic voltage oscillations resulting from the summed activity of neuronal populations occur in many nervous systems. Contemporary observations suggest that coexistent oscillations interact and, in time, may switch in dominance. We recently reported an example of these interactions recorded from in vitro preparations of rat somatosensory cortex. We found that following an initial interval of coexistent gamma ( approximately 25 ms period) and beta2 ( approximately 40 ms period) rhythms in the superficial and deep cortical layers, respectively, a transition to a synchronous beta1 ( approximately 65 ms period) rhythm in all cortical layers occurred. We proposed that the switch to beta1 activity resulted from the novel mechanism of period concatenation of the faster rhythms: gamma period (25 ms)+beta2 period (40 ms) = beta1 period (65 ms). In this article, we investigate in greater detail the fundamental mechanisms of the beta1 rhythm. To do so we describe additional in vitro experiments that constrain a biologically realistic, yet simplified, computational model of the activity. We use the model to suggest that the dynamic building blocks (or motifs) of the gamma and beta2 rhythms combine to produce a beta1 oscillation that exhibits cross-frequency interactions. Through the combined approach of in vitro experiments and mathematical modeling we isolate the specific components that promote or destroy each rhythm. We propose that mechanisms vital to establishing the beta1 oscillation include strengthened connections between a population of deep layer intrinsically bursting cells and a transition from antidromic to orthodromic spike generation in these cells. We conclude that neural activity in the superficial and deep cortical layers may temporally combine to generate a slower oscillation.     

State-dependent effects of light-dark cycle on somatosensory and visual cortex EEG in rats

Somatosensory (SSctx) and visual cortex (Vctx) EEG were evaluated in rats under a 12:12-h light-dark (LD) cycle and under constant light (LL) or constant dark (DD) in each sleep or wake state. Under LD conditions during light period, relative Vctx EEG slow-wave activity (SWA) was higher than that of the SSctx, whereas during dark period, relative Vctx EEG SWA was lower than in the SSctx. These effects were state specific, occurring only during non-rapid eye movement sleep (NREMS). Under LL conditions, the duration of REMS and NREMS during the period that would have been dark if the LD cycle had continued (subjective dark period) was greater than under LD conditions. DD conditions had little effect on the duration of NREMS and REMS. SSctx and Vctx EEG SWA were suppressed by LL during the subjective dark period; however, the degree of Vctx SWA suppression was smaller than that of the SSctx. DD conditions during the subjective light period enhanced SSctx SWA, whereas Vctx SWA was suppressed. Under LL conditions during the subjective dark period, Vctx EEG power was higher than that of the SSctx across a broad frequency range during NREMS, REMS, and wakefulness. During DD, SSctx EEG power during NREMS was higher than that of the Vctx in the delta wave band, whereas SSctx power during REMS and wakefulness was higher than that of the Vctx in frequencies higher than 8 Hz. We concluded that the SSctx and Vctx EEGs are differentially affected by light during subsequent sleep. Results provide support for the notion that regional sleep intensity is dependent on prior regional afferent input.     

Ontogeny of corticocortical projections of the rat somatosensory cortex.

Rhodamine-coated microspheres (RCMs) were injected into the primary somatosensory cortex (SI) of rats ranging in age from postnatal (PN) day 1 to adulthood. Ipsilateral corticocortical and callosal projections within the SI were identified as early as PN day 1. At the end of the first PN week, ipsilaterally projecting neurons located in sublayer VIb were the first to assume an adult-like pattern of connectivity. Injections at subsequent postnatal ages revealed that an adult pattern of lamination of ipsilateral corticocortical projections within the SI is established between PN weeks 2 and 3, comprising projection neurons from layers II/III, layer V, and sublayer VIb. Therefore, local interactions in the rat SI are mediated not only by pyramidal neurons of layers III and V, derived from the cortical plate, but also by a subpopulation of ontogenetically older neurons located in the sublayer VIb, which may correspond to the subplate neurons of other species. Overall, these results suggest the existence of three independent short-range corticocortical systems of projections within the rat SI, which differ in terms of the laminar distribution and ontogenetic origin of their cells.     

Differential effects of aging on estrogen negative and positive feedback

Recent studies have demonstrated an age-related decline in gonadotropins and a decrease in pituitary responsiveness to GnRH, indicating that aging influences the neuroendocrine components of the female reproductive axis independently of changes in ovarian function. To determine whether aging might also affect the luteinizing hormone (LH) negative and positive feedback responses to gonadal steroids, we administered a controlled, graded sex steroid infusion to 11 younger (45-56 yr) and nine older (70-80 yr) postmenopausal women (PMW) in whom endogenous ovarian steroids and peptides are uniformly low. The doses of estradiol (E(2)) and progesterone (P) were chosen to mimic levels across the normal follicular phase and have been shown previously to induce negative followed by positive feedback on LH. Similar E(2) and P levels were achieved in younger and older PMW (P = 0.4 and 0.3, respectively) and produced a biphasic LH response in all subjects. The early decline in LH to 53% of baseline was not different in older vs. younger PMW. However, the positive feedback effect was attenuated in older compared with younger PMW (peak LH 144.4 ± 19.5 vs. 226.8 ± 22.3 IU/l, respectively, P = 0.01). In conclusion, these studies in PMW demonstrate preservation of short-term steroid negative and positive feedback in response to exogenous E(2) and P with aging. Attenuation of positive feedback in older compared with younger PMW is consistent with previous reports of declining GnRH responsiveness with aging.     

Acute effects of total or partial digit denervation on raccoon somatosensory cortex

The immediate effects of total or partial denervation of single digits (0-16 hr after nerve transection) on primary somatosensory cortex were studied electrophysiologically. Comparisons of response properties and cortical somatotopy were made between intact raccoons and four groups of raccoons with transection of some or all of the nerves innervating the fourth or fifth digit. Animals with all four digital nerves cut (amputation of the digit) were most different from normal. Approximately half of the penetrations in the affected cortical region showed inhibitory responses to stimulation of adjacent skin regions. These consisted of a strong response to stimulus offset and/or a suppression of spontaneous activity during indentation. Since these responses were substantially different from those recorded several months after digit amputation, additional changes in connectivity and synaptic strength must occur with chronic denervation. These inhibitory responses were not seen in animals with one, two, or three nerves cut per digit. In the animals with partial denervation of a digit, the greatest disruption occurred when both ventral nerves to the glabrous skin were transected. This yielded cell clusters with abnormally large receptive fields, disruptions in somatotopic organization, and a decreased occurrence of low-threshold responses. If only one nerve to glabrous skin was transected, there was less change, even if it was combined with transection of both nerves to hairy skin. These results suggest that the release of inhibitory responses in a cortical digital region by amputation is prevented by the retention of even one ventral nerve. None of the denervation conditions produced large nonresponsive areas of cortex, which would have indicated a loss of all inputs.     

Population coding of stimulus location in rat somatosensory cortex.

This study explores the nature of population coding in sensory cortex by applying information theoretic analyses to neuron pairs recorded simultaneously from rat barrel cortex. We quantified the roles of individual spikes and spike patterns in encoding whisker stimulus location. 82%-85% of the total information was contained in the timing of individual spikes: first spike time was particularly crucial. Spike patterns within neurons accounted for the remaining 15%-18%. Neuron pairs located in the same barrel column coded redundantly, whereas pairs in neighboring barrel columns coded independently. The barrel cortical population code for stimulus location appears to be the time of single neurons' first poststimulus spikes-a fast, robust coding mechanism that does not rely on "synergy" in crossneuronal spike patterns.     

Infrared thermal imaging of rat somatosensory cortex with whisker stimulation

The present study aims to validate the applicability of infrared (IR) thermal imaging for the study of brain function through experiments on the rat barrel cortex. Regional changes in neural activity within the brain produce alterations in local thermal equilibrium via increases in metabolic activity and blood flow. We studied the relationship between temperature change and neural activity in anesthetized rats using IR imaging to visualize stimulus-induced changes in the somatosensory cortex of the brain. Sensory stimulation of the vibrissae (whiskers) was given for 10 s using an oscillating whisker vibrator (5-mm deflection at 10, 5, and 1 Hz). The brain temperature in the observational region continued to increase significantly with whisker stimulation. The mean peak recorded temperature changes were 0.048 ± 0.028, 0.054 ± 0.036, and 0.097 ± 0.015°C at 10, 5, and 1 Hz, respectively. We also observed that the temperature increase occurred in a focal spot, radiating to encompass a larger region within the contralateral barrel cortex region during single-whisker stimulation. Whisker stimulation also produced ipsilateral cortex temperature increases, which were localized in the same region as the pial arterioles. Temperature increase in the barrel cortex was also observed in rats treated with a calcium channel blocker (nimodipine), which acts to suppress the hemodynamic response to neural activity. Thus the location and area of temperature increase were found to change in accordance with the region of neural activation. These results indicate that IR thermal imaging is viable as a functional quantitative neuroimaging technique.     

Differential effects of aging on EEG after baclofen administration

Baclofen is a selective gamma-aminobutyric acid (GABA) type B agonist that may have important medicinal uses, such as in analgesics and drug addiction treatment. In addition, evidence is accumulating that suggests GABAergic-mediated neurotransmission is altered during aging. This study investigated whether baclofen administration (5 mg kg⁻¹) induces differential effects on cortical electrical activity with age. Electroencephalograms (EEGs) were recorded from young (3–4 months) and aged (15–17 months) rats, and both the absolute and relative powers in five frequency bands (delta: 2–4 Hz; theta: 4–8 Hz; alpha: 8–12 Hz; beta: 12–20 Hz; gamma: 20–100 Hz) were analyzed. Before administration of baclofen, we found that the EEG relative power in the beta band was higher in the aged than that in the young rats. After administration of baclofen, there was a slower increase in the relative power in the delta band in the aged than that in the young rats. Moreover, there was no significant difference between the two age groups in absolute power in any frequency band. These findings indicate that baclofen treatment appears to differentially modify cortical EEG activity as a function of age. Our data further elucidate the relationship between GABA_B receptor-mediated neurotransmission and aging.     

Non-NMDA receptor-mediated vibrotactile responses of neurons from the hindpaw representation in the rat SI cortex

Non-NMDA receptor-mediated vibrotactile responses of neurons from the hindpaw representation were investigated in the rat SI cortex. We recorded single-unit spikes evoked by sinusoidal (duration: 500?ms; frequency: 5, 40, and 250?Hz; amplitude: 100?μm peak-to-peak) stimulation of the glabrous skin. The responses were obtained with microinjection of aCSF (sham), bicuculline, and AMPA near the isolated neurons in anaesthetized rats. Blocking most of the NMDA receptors by ketamine revealed local dynamics differentially modulated by each drug. The responses were generally suppressed after the initial 100-ms period of the 40- and 250-Hz stimulus, but not at 5?Hz. Both drugs increased average firing rates (AFRs) only during vibrotactile stimulation, and increased entrainment as measured by the vector strength (VS) of spike phases. However, bicuculline was more effective on the AFR in the late period particularly at 40?Hz. Complex interactions were found with AMPA; late activity increased only for fast spiking neurons at 40?Hz, and more for regular spiking neurons at 5?Hz. The increase of VS by bicuculline was much higher in layer IV. In addition to thalamocortical feed-forward inhibition, vibrotactile information seems to be suppressed after 100?ms by longer-latency inhibitory networks tuned to mid-frequency inputs. Combined with the presumed AMPA-receptor desensitization, those two inhibitory factors could limit the excitatory flow mostly to lower frequencies. The frequency dependence of the drug effects highlights the role of local cortical dynamics in the hindpaw area.     

Phase resetting curves and oscillatory stability in interneurons of rat somatosensory cortex

Synchronous oscillations in neural activity are found over wide areas of the cortex. Specific populations of interneurons are believed to play a significant role in generating these synchronized oscillations through mutual synaptic and gap-junctional interactions. Little is known, though, about the mechanism of how oscillations are maintained stably by particular types of interneurons and by their local networks. To obtain more insight into this, we measured membrane-potential responses to small current-pulse perturbations during regular firing, to construct phase resetting curves (PRCs) for three types of interneurons: nonpyramidal regular-spiking (NPRS), low-threshold spiking (LTS), and fast-spiking (FS) cells. Within each cell type, both monophasic and biphasic PRCs were observed, but the proportions and sensitivities to perturbation amplitude were clearly correlated to cell type. We then analyzed the experimentally measured PRCs to predict oscillation stability, or firing reliability, of cells for a complex stochastic input, as occurs in vivo. To do this, we used a method from random dynamical system theory to estimate Lyapunov exponents of the simplified phase model on the circle. The results indicated that LTS and NPRS cells have greater oscillatory stability (are more reliably entrained) in small noisy inputs than FS cells, which is consistent with their distinct types of threshold dynamics.     

Millisecond-timescale local network coding in the rat primary somatosensory cortex

Correlation among neocortical neurons is thought to play an indispensable role in mediating sensory processing of external stimuli. The role of temporal precision in this correlation has been hypothesized to enhance information flow along sensory pathways. Its role in mediating the integration of information at the output of these pathways, however, remains poorly understood. Here, we examined spike timing correlation between simultaneously recorded layer V neurons within and across columns of the primary somatosensory cortex of anesthetized rats during unilateral whisker stimulation. We used bayesian statistics and information theory to quantify the causal influence between the recorded cells with millisecond precision. For each stimulated whisker, we inferred stable, whisker-specific, dynamic bayesian networks over many repeated trials, with network similarity of 83.3±6% within whisker, compared to only 50.3±18% across whiskers. These networks further provided information about whisker identity that was approximately 6 times higher than what was provided by the latency to first spike and 13 times higher than what was provided by the spike count of individual neurons examined separately. Furthermore, prediction of individual neurons' precise firing conditioned on knowledge of putative pre-synaptic cell firing was 3 times higher than predictions conditioned on stimulus onset alone. Taken together, these results suggest the presence of a temporally precise network coding mechanism that integrates information across neighboring columns within layer V about vibrissa position and whisking kinetics to mediate whisker movement by motor areas innervated by layer V.     

Hemodynamics evoked by microelectrical direct stimulation in rat somatosensory cortex

The aim of this study was to estimate the timing (latency) of the increase in red blood cell (RBC) velocity and RBC concentration, and the magnitude of response in local cerebral blood flow (LCBF) for neuronal activation. We measured LCBF change during activation of the somatosensory cortex by direct microelectrical stimulation. Electrical stimuli of 5, 10 and 50 Hz of 1 ms pulse with 10-15 microA, were given for 5 s. LCBF, RBC velocity and RBC concentration were monitored by laser-Doppler flowmetry (LDF) in alpha-chloralose anesthetized rats (n = 7). LCBF, RBC velocity and RBC concentration increased nearly proportionally to stimulus frequency, i.e. neuronal activity. LCBF rose approximately 0.5 s after the onset of stimulation, and there was no significant time lag of the latencies among LCBF, RBC velocity and RBC concentration at the same stimulus frequency. We interpret these results to mean that the onset of LCBF increase on cortical activation is reflected by a rapid change in arteriole (resistance vessel) dilation and capillary volume. The data also elucidate the linear relationship between LCBF increase and cortical activity.     

Chronic effects of total or partial digit denervation on raccoon somatosensory cortex.

Electrophysiological recordings were made in the primary somatosensory cortex of anesthetized raccoons 14 to 169 days following digit amputation or 60 to 129 days after transection of the two nerves innervating the ventral surface of the fourth digit. The incidence of inhibitory responses decreased from 50% of the penetrations immediately after amputation to 35% over the first 3 weeks and to almost zero after 2 months. The number of sites with low-threshold excitatory responses increased from 4% to 14% to 50% during these same intervals. Initially, the excitatory fields were small and located over the nerve stumps, and were therefore probably due to direct stimulation of the damaged nerves. At 2 months after amputation, the excitatory receptive fields were large and diffuse. Although the size of receptive fields decreased during the later period (when the thresholds were also decreasing), there was no recovery of any precise somatotopic organization in the deafferented cortex. The reorganization process in the raccoon thus consists of at least two stages: The early stage is dominated by inhibitory connections, whereas the second involves a recovery and restructuring of excitatory inputs. From 2 to 4 months after partial digit denervation, there were only minor changes in response properties or somatotopic organization in the deafferented cortex as compared to immediately after nerve transection. Thus, few of the characteristics of reorganization induced by digit amputation were elicited by this treatment, which leaves some of the digit innervation intact. There was, however, an unexpected increase in the portion of the ventral digit that was able to activate the cortex, suggesting complexities in the peripheral innervation of the digit that need to be resolved.     

Differential effects of phencyclidine application on secretogranin II expression in organotypic slices of rat prefrontal cortex

Phencyclidine (PCP) is a non-competitive NMDA glutamate receptor antagonist that induces psychotomimetic effects in humans and experimental animals. Chronic PCP exposure elicits signs of persistently altered frontal brain activity and related behaviors which are also seen in patients with schizophrenia. Secretogranin II (sg II) belongs to the chromogranin family of proteins that exist in large dense core vesicles in nervous tissue. In the brain, 90% of sg II is processed to the small peptide secretoneurin. We previously detected differential effects of single-dose and subchronic PCP administration on sg II expression in the rat prefrontal cortex (PFC). In the present study, we applied PCP to organotypic PFC slices. PCP application for 28 h induced decreased tissue and culture medium secretoneurin content. In contrast, incubation with the adenylate cyclase activator forskolin caused significantly increased secretoneurin levels after 8 h. PCP for 4 h followed by 24 h without PCP resulted in increased culture medium secretoneurin content but no change in tissue levels. sg II mRNA expression was decreased after 28 h PCP application in cortical neurons. Immunohistochemical and TUNEL staining profiles indicated that the alterations were not due to neurodegeneration. PCP for 5 days changed neither the secretoneurin tissue or culture medium levels, nor the sg II mRNA expression. These results demonstrate that PCP modulates sg II expression in PFC tissue in the absence of afferent inputs and that the nature of these changes is dependent upon the duration of exposure to and/or withdrawal from PCP.     

Mapping the effects of SI cortex stimulation on somatosensory relay neurons in the rat thalamus: direct responses and afferent modulation

We have used single-unit recording techniques to map the spatial distribution of the primary somatosensory (SI) cortical influences on thalamic somatosensory relay nuclei in the rat. A total of 193 microelectrode penetrations were made to record single neurons in tracks through the medial and lateral ventroposterior (VPL and VPM), ventrolateral (VL), posterior (Po), and reticular (nRt) thalamic nuclei. Single units were classified according to their (1) location within the nuclei, (2) receptive fields, and (3) response to standardized microstimulation in deep layers of the SI cortical forepaw areas. The SI stimulation produced short-latency (1- to 7-msec) excitatory responses in different percentages of neurons recorded in the following thalamic nuclei: VPL, 42.0%; Po, 25.0%; nRt, 16.4%; VL, 13.6%; and VPM, 9.9%. Within the VPL, the highest proportion of responsive neurons was found in the anterior region. Although most of the VL region was unresponsive, the caudal subregion bordering the rostral VPL showed some responsiveness (13.6% of neurons). In general, the spatial pattern of corticothalamic influences appeared to reciprocate the known thalamocortical connection patterns, but with a heterogeneity that was unpredicted. The same parameters of SI cortical stimulation were used in studies of corticofugal modulation of afferent transmission through the VPL thalamus. A condition-test (C-T) paradigm was implemented in which the cortical stimulation (C) was delivered at a range of time intervals before test (T) mechanical vibratory stimulation was applied to digit 4 of the contralateral forepaw. The time course of cortical effects was analyzed by measuring the averaged evoked unit responses of thalamic neurons to the T stimuli, and plotting them as a function of C-T intervals from 5 to 50 msec. Of the 20 VPL neurons tested during SI stimulation, the average response to T stimulation was decreased a mean of 36%, with the suppression peaking (at 49% inhibition of the afferent response) about 15 msec after the C stimulus. Considerable rostrocaudal variation was observed, however. Whereas neurons in the rostral VPL (near VL) were strongly inhibited (-69%), neurons in the middle and caudal VPL exhibited facilitations at long and short C-T intervals, respectively. This study establishes a specific projection system from the forepaw region of SI cortex to different subregions of the VPL thalamus, producing specific temporal patterns of sensory modulation.