Second somatic sensory cortical area (SII) in a prosimian primate, Galago crassicaudatus

The cortex adjacent to and along the upper bank of the lateral sulcus (UB-LS) of a prosimian primate, Galago crassicaudatus, was explored to determine the topographical representation of low-threshold cutaneous inputs to this region. The somatic sensory projections to this cortex were considered homologous to those defined in other species as the second somatosensory cortical area (SII). Multiple and single neuron recordings were obtained with tungsten microelectrodes in animals anesthetized with sodium pentobarbital or ketamine hydrochloride; receptive fields were determined by means of manually applied tactile stimuli. The area of SII was located approximately 1-1.5 mm rostral to the posterior limit of LS, extended rostrally approximately 4 mm, and occupied nearly all of the upper bank of the sulcus throughout this region. Receptive fields (RFs) in SII were primarily contralateral except for some bilateral input in the cortex representing portions of the trunk, head, and face. The boundaries of RFs were well defined, especially where recordings were located in the middle layers of the cortex. The distribution of RFs across SII was somatotopically organized into a single, relatively erect representation of the body that involved inputs from the face rostral and medial (superficially along the UB-LS) surrounding an enlarged forelimb area; the latter, in turn, lies rostral and medial to the hindlimb zone. Projections from the tail and sacrum are located furthest caudal and lateral (deeper along UB-LS). Separate regions that were devoted to the glabrous skin surfaces of the distal limbs formed the rostral and lateral boundaries of the distal fore- and hindlimb representations, respectively. In the zone for the glabrous surfaces of the forelimb digits, individual digits dominated discrete components of the SII map, especially medially where digit 1 was represented. The glabrous tip of digit 5 was represented caudal and lateral to the tip of digit 1. A similar radial to ulnar medial to lateral sequence was noted in the area representing the palm. Except for a possible medially located toe 1 zone in the hindlimb representation, separated representations for the glabrous skin of individual toes were not noted. The dorsal hairy surfaces of the digits and toes were, respectively, amalgamated within the representations for the dorsal surfaces of the hand and foot. In these regions, which were found superficial and slightly caudal to their respective glabrous zones, some RFs were found that were devoted only to the distal extremities, but most RFs included more proximal portions of the hand or foot dorsum.(ABSTRACT TRUNCATED AT 400 WORDS)     

Early secondary somatosensory area (SII) SEPs. Data from intracerebral recordings in humans.

OBJECTIVE: To record somatosensory evoked potentials (SEPs) to median nerve stimulation by chronically implanted electrodes in the parieto-rolandic opercular area of 9 epileptic patients, in order to evaluate whether somatosensory evoked responses could be generated in the second somatosensory area (SII) earlier than 40 ms after stimulus. METHODS: Nine patients (4 males, 5 females) with drug-resistant partial epileptic seizures were investigated using stereotactically implanted electrodes in the parietal cortex, posterior to vertical anterior commissure plane and in the frontal opercular region rostral to vertical anterior commissure (VAC). RESULTS: The main finding of this study is the recording of an early somatosensory evoked potential, (N30op), by chronically implanted electrodes in the SII area of 8 epileptic patients. In 3 patients where SEPs were performed after ipsilateral median nerve (MN) stimulation, a N30op was recorded 5.8+/-2 ms later than contralateral one. CONCLUSIONS: This is the first report of early SEPs recorded by electrodes implanted in SII area. The N30op potential, even if less consistent than later potentials, confirmed the important role of the SII area in the early processing of somatosensory inputs.     

Bilateral interaction in the second somatosensory area (SII) of the cat and contribution of the corpus callosum

There are indications in the literature that convergent ipsilateral and contralateral input to the second somatosensory area (SII) may interact. Single unit activity of SII bilateral cells was studied to evaluate the impact of simultaneous bilateral stimulation of the receptive fields (RF) on neural discharge. The cellular responses to unilateral ipsilateral and contralateral, as well as to bilateral stimulation were compared. 22% of bilateral cells showed interaction, usually facilitation. Bilaterally evoked responses were found to be as great as 250% of the strongest unilateral response. Only bilateral responses stronger or weaker than the dominant unilateral response by at least 50% were considered as interactive. The great majority of interactive cells had their RF on the forelimb and were responsive to deep stimulation. The corpus callosum appears to be responsible for part of the observed interaction since in callosotomized cats only 5% of bilateral cells were interactive. A non-callosal ipsilateral pathway must be postulated because both bilaterality and bilateral interaction persist to some degree after callosotomy. A putative role for bilateral interaction in sensory-motor integration is discussed.     

Spinothalamocortical projections to the secondary somatosensory cortex (SII) in squirrel monkey

Anterograde labeling of the cervical spinothalamic tract was combined with retrograde labeling of thalamocortical cells projecting to the hand region of the second somatosensory cortex (hSII) to identify likely sites in the thalamus for processing and transmitting nociceptive information to hSII. Anterograde labeling of terminals was done with 2% WGA-HRP injections in the cervical enlargement; thalamocortical cells were retrogradely labeled with fluorescent tracers. In one experiment, the contralateral primary somatosensory cortex hand region (hSI) was injected to provide a direct comparison with hSII thalamic label. Both labeled cells and terminal-like structures were visualized in single thalamic sections and their numbers and positions quantitatively analyzed. The number of labeled cells within 100 microns from the STT terminals were counted as overlapping cells. Four thalamic nuclei, ventroposterior inferior (VPI), ventroposterior lateral (VPL), posterior nucleus (PO) and centrolateral nucleus (CL) combined to contain 86.5% of all hSII-projecting overlapping cells. Of all hSII-projecting thalamic overlapping cells, VPI contained the largest number (36.4% of the total) followed by the anterior portion of the posterior nuclear complex (POa; 20.4%), VPL (18.3%) and CL (11.4%). Results of the hSI injection show a different pattern of overlap in agreement with our earlier study. The relative distribution of overlapping cells was dependent on the antero-posterior position of the SII injections. The most anterior injections resulted in small numbers of labeled cells, with the majority of overlapping cells located in PO and CL. The more posterior injections resulted in overlapping cells mainly in VPI and VPL. The results indicate that, in the squirrel monkey, VPI, VPL, POa and CL relay nociceptive information from the spinal cord to the second somatosensory cortex.     

Intracortical recordings of early pain-related CO2-laser evoked potentials in the human second somatosensory (SII) area.

We studied responses of the parieto-frontal opercular cortex to CO2-laser stimulation of A delta fiber endings, as recorded by intra-cortical electrodes during stereotactic-EEG (SEEG) presurgical assessment of patients with drug-resistant temporal lobe epilepsy. After CO2-laser stimulation of the skin at the dorsum of the hand, we consistently recorded in the upper bank of the sylvian fissure contralateral to stimulation, a negative response at a latency of 135 +/- 18 ms (N140), followed by a positivity peaking around 171 +/- 22 ms (P170). The stereotactic coordinates in the Talairach's atlas of the electrode contacts recording these early responses covered the pre- and post-rolandic part of the upper bank of the sylvian fissure (-27 < y < +12 mm; 31 < x < 57 mm; 4 < z < 23 mm), corresponding to the accepted localization of the SII area in man, possibly including the upper part of the insular cortex. The spatial distribution of these early contralateral responses in the SII-insular cortex fits wit that of the modeled sources of scalp CO2-laser evoked potentials (LEPs) and with PET data from pain activation studies. Moreover, this study showed the likely existence of dipolar sources radial to the scalp surface in SII, which are overlooked in magnetic recordings. Early responses also occurred in the SII area ipsilateral to stimulation peaking 15 ms later than in contralateral SII, suggesting a callosal transmission of nociceptive inputs between the two SII areas. Other pain responsive areas such as the anterior cingulate gyrus, the amygdala and the orbitofrontal cortex did not show early LEPs in the 200 ms post-stimulus. These findings suggest that activation of SII area contralateral to stimulation, possibly through direct thalamocortical projections, represents the first step in the cortical processing of peripheral A delta fiber pain inputs.     

The effects of unilateral or bilateral removals of the second somatosensory cortex (area SII): a profound tactile disorder in monkeys.

Three groups of monkeys were studied; one with bilateral removals of the second somatosensory projection cortex (SII); another with similar unilateral removals; and an unoperated control group. Both lesion groups were significantly impaired on tasks of tactile learning and tactile retention. However, there were no differences between groups on the great majority of tasks of motor learning, nor on spatial alternation in the dark. The animals with unilateral removals were generally impaired whether using the hand contralateral to or ispilateral to the removal.     

Broca's area plays a causal role in morphosyntactic processing

Although there is strong evidence that Broca's area is important for syntax, this may simply be a by-product of greater working memory and/or cognitive control demands for more complex syntactic structures. Here we report an experiment with event-related transcranial magnetic stimulation (TMS) to investigate whether Broca's area plays a causal role in morphosyntactic processing when both working memory and cognitive control demands are low. Participants were presented with word pairs that could either agree or disagree in grammatical number or gender while receiving stimulation to Broca's area or to the right intraparietal sulcus (a control site). Stimulation of Broca's area significantly reduced the advantage for grammatical relative to ungrammatical word pairs. In contrast, stimulation of control site left this grammaticality advantage unchanged. The interaction between grammaticality and stimulation was specific to Broca's area, suggesting a clear involvement of the region in morphosyntactic processing.     

Distribution of GAD-immunoreactive neurons in the first (SI) and second (SII) somatosensory cortex of the monkey

The distribution of neurons immunoreactive for glutamic acid decar☐ylase (GAD), the synthesizing enzyme of γ-aminobutyric acid (GABA), was examined in the first (SI) and second (SII) somatosensory cortex of monkeys. GAD-like immunoreactive (GAD- LI) somata and puncta were present in all layers of SI and SII. All GAD-LI somata were identified as non-pyramidal neurons and were most numerous in layer IV of SI and in layer II of SII. Layer IV of SI also contained the highest density of GAD-LI puncta. In SII, GAD-LI puncta were distributed more homogeneously and did not show a dense band of labelled puncta in layer IV. The major and minor diameters of GAD-LI somata in SII ranged from 6.9 to 26.2 μm and from 6.2 to 19.0 μm, respectively. The major diameters of GAD-LI somata in SII were significantly smaller than those in SI in layers I, III and IV. Differences between the distributions of GAD-LI puncta and somata in SI and SII may be accounted for by differences in the number and/or distribution of different types of GABAergic neurons. Functional differences of neurons in SI and SII may be related to the differences in GABAergic inhibitory mechanisms and reflected in the distribution of GABAergic neurons.     

Mapping study of somatosensory evoked potentials during selective spatial attention.

We have investigated the effects of selective spatial attention on early and middle-latency SEPs. Baseline control responses to electrical stimulation of 2 digits of the hand were recorded first in conditions of mental relaxation, in the absence of any cognitive task, to obtain truly 'neutral' responses uncontaminated by cognitive components. Then, during a 'task condition,' identical stimuli were applied to the same two fingers, but the subject's attention was driven towards the stimulated territory by the bias of mechanical taps delivered to the same digits. The earliest effect of directing attention towards the territory stimulated was a positive shift on contralateral somatosensory responses, with onset at 27.4 +/- 4 msec post stimulus. This SEP modification: (a) did not entail any change in the scalp distribution of components, as assessed by topographic mapping, and (b) was not present when attention was directed towards the hand contralateral to that receiving electrical stimuli. A second effect was represented by a parieto-central negativity in the 60-80 msec latency range; this feature could also be observed during contralaterally driven attention and was associated with topographical changes in SEP scalp distribution. Finally, a late centro-frontal negativity beginning at 90-100 msec (N140) appeared during ipsilateral attention, while P100 was not enhanced. Subcortical P14 and primary cortical N20 were not significantly affected by the tasks. We conclude that the 'early positive shift' is linked to the spatial aspects of selective attention and represents in part modulation of obligatory components (P25 through P45) existing in control SEPs; it probably corresponds with the deflections with similar polarity and time-course that have been described by others in response to somatosensory target stimuli. Conversely, 60-80 msec negative enhancement is less spatially selective and may represent non-specific arousal effects. The late negative component (N140) shares several features with the 'processing negativity' described in auditory paradigms and could represent the equivalent of this effect in the somatosensory system.     

The possible role of selective attention in acquired dyslexia

Literal dyslexia, the inability to read letters within a word which itself could be read, was analysed in two patients. Their impairment was found not to be specific to letters but to stimuli in which more than one item of the same category was simultaneously present in the visual field. Thus a letter surrounded by numbers could be read more easily than if surrounded by other letters. Explanations in terms of visual disorientation or implicit response interference were rejected. It is suggested that their impairment arises from a specific defect at the level at which visual input is selected for meaningful analysis.     

Supplementary motor area plays a causal role in automatic inhibition of motor responses

BackgroundThe masked-priming paradigm is used to test unconscious inhibitory processes of the brain. A tendency towards responses that are incompatible with the prime, designated as negative compatibility effect (NCE), emerges when the perception of a priming visual stimulus is “masked” afterwards. This effect presumably stems from a subliminal inhibitory process against the masked-prime. Prior lesions as well as activation studies suggest a key role of SMA in this effect.ObjectiveThis study was conducted to elucidate a causal role of SMA in the subliminal response inhibition represented by the NCE.MethodsUsing a repeated-measures pre–post design with a group of healthy people, physiological measures (resting and active motor thresholds and motor evoked potential (MEP) amplitude) and behavioral ones (choice reaction time (CRT), positive compatibility effect (PCE) and NCE) were obtained before and after three quadripulse stimulation (QPS), namely sham, M1-QPS, and SMA-QPS, on different days. CRT and PCE served as indices for different aspects of motor execution.ResultsMotor thresholds were not altered after any QPS, although the M1-QPS increased MEP amplitude. Neither CRT nor PCE was altered significantly after QPS protocols. NCE was abolished after the SMA-QPS.ConclusionsAbolished NCE after the SMA-QPS in the absence of MEP changes suggests that (1) SMA plays a cardinal role in the NCE, and (2) the network involved in NCE is different from that of MEP generation.     

Medial prefrontal cortex plays a critical and selective role in 'feeling of knowing' meta-memory judgments

The frontal lobes are thought to play a role in the monitoring of memory performance, or 'meta-memory', but the specific circuits involved have yet to be definitively established. Medial prefrontal cortex in general and the anterior cingulate cortex in particular, have been implicated in other forms of monitoring, such as error and conflict monitoring. Here, we tested the hypothesis that medial prefrontal cortex plays a critical role in memory monitoring, aiming to determine whether this region contributed to all, or only some classes of meta-memory judgments. We also investigated the relationship between these judgments and memory performance itself. Three types of meta-memory judgment were measured in 5 subjects with focal damage to medial prefrontal cortex, with maximal overlap in dorsal anterior cingulate cortex, compared to 19 healthy, demographically matched control subjects performing a face-name episodic memory task. Judgment-of-learning accuracy was not affected by such damage. In contrast, both recall confidence and feeling-of-knowing judgments were impaired. Memory performance was itself impaired in the patient group, so we performed a second experiment to examine the relationship between memory and meta-memory deficits. In an easier memory task, where patients performed as well as controls, recall confidence accuracy improved to within the control range despite medial prefrontal damage. In contrast, feeling-of-knowing judgments remained less accurate in the patient group. These results argue that medial prefrontal cortex plays a critical role in generating accurate recall confidence and feeling-of-knowing judgments, but is not necessary for judgment-of-learning. The role of this region in feeling-of-knowing seems to be, at least in part, independent of its role in memory itself.     

Monocarboxylate transporter 4 plays a significant role in the neuroprotective mechanism of ischemic preconditioning in transient cerebral ischemia

Monocarboxylate transporters(MCTs), which carry monocarboxylates such as lactate across biological membranes, have been associated with cerebral ischemia/reperfusion process. In this study, we studied the effect of ischemic preconditioning(IPC) on MCT4 immunoreactivity after 5 minutes of transient cerebral ischemia in the gerbil. Animals were randomly designated to four groups(sham-operated group, ischemia only group, IPC + sham-operated group and IPC + ischemia group). A serious loss of neuron was found in the stratum pyramidale of the hippocampal CA1 region(CA1), not CA2/3, of the ischemia-only group at 5 days post-ischemia; however, in the IPC + ischemia groups, neurons in the stratum pyramidale of the CA1 were well protected. Weak MCT4 immunoreactivity was found in the stratum pyramidale of the CA1 in the sham-operated group. MCT4 immunoreactivity in the stratum pyramidale began to decrease at 2 days post-ischemia and was hardly detected at 5 days post-ischemia; at this time point, MCT4 immunoreactivity was newly expressed in astrocytes. In the IPC + sham-operated group, MCT4 immunoreactivity in the stratum pyramidale of the CA1 was increased compared with the sham-operated group, and, in the IPC + ischemia group, MCT4 immunoreactivity was also increased in the stratum pyramidale compared with the ischemia only group. Briefly, present findings show that IPC apparently protected CA1 pyramidal neurons and increased or maintained MCT4 expression in the stratum pyramidale of the CA1 after transient cerebral ischemia. Our findings suggest that MCT4 appears to play a significant role in the neuroprotective mechanism of IPC in the gerbil with transient cerebral ischemia.     

Magnetic source imaging of tactile input shows task-independent attention effects in SII

We investigated whether attention to different stimulus attributes (location, intensity) has different effects on the activity of the secondary (SII) somatosensory cortex. Tactile stimuli were applied to the left index finger and somatosensory evoked fields (SEFs) were recorded using a whole-head magnetoencephalography (MEG) system. Two oddball paradigms with stimuli varying in location or intensity were performed in an ignore and an attend condition. Brain sources were estimated by magnetic source imaging. No attention effect was observed for the primary SI area. However, attention enhanced SII activity bilaterally from 55 to 130 ms by 52% in the spatial and 64% in the intensity discrimination task. SII attentional enhancement was very similar in both paradigms and occurred both for deviants and standards.     

The role of selective attention in matching observed and executed actions

Substantial evidence suggests that observed actions can engage their corresponding motor representations within the observer. It is currently believed that this process of observation-execution matching occurs relatively automatically, without the need for top-down control. In this study we tested the susceptibility of the observation-execution matching process to selective attention. We used a Go/NoGo paradigm to investigate the phenomenon of ‘automatic imitation’, in which participants are faster to initiate a hand movement that is congruent with a concurrently observed action, relative to one that is incongruent. First, we replicated previous findings of automatic imitation, and excluded the possibility that spatial compatibility effects might explain these results (Experiment 1). We then presented participants with the same goal-directed actions while directing their attention to an imperative stimulus that spatially overlapped, but was distinct from, the observed actions (Experiment 2). Crucially, automatic imitation no longer occurred when participants directed their attention away from the displayed actions and towards the spatially overlapping stimulus. In a final experiment, we examined whether the automatic imitation of grasp persists when participants attend to an irrelevant feature of the observed action, such as whether it is performed by a left or right hand (Experiment 3). Here we found that automatic imitation is contingent on participants attending to the feature of the observed hand that was relevant to their responses. Together these findings demonstrate the importance of selective mechanisms in the filtering of task-irrelevant actions, and indicate a role for top-down control in limiting the motoric simulation of observed actions.     

The role of selective attention on academic foundations: a cognitive neuroscience perspective

To the extent that selective attention skills are relevant for academic foundations and amenable to training, they represent an important focus for the field of education. Here, drawing on research on the neurobiology of attention, we review hypothesized links between selective attention and processing across three domains important to early academic skills. First, we provide a brief review of the neural bases of selective attention, emphasizing the effects of selective attention on neural processing, as well as the neural systems important to deploying selective attention and managing response conflict. Second, we examine the developmental time course of selective attention. It is argued that developmental differences in selective attention are related to the neural systems important for deploying selective attention and managing response conflict. In contrast, once effectively deployed, selective attention acts through very similar neural mechanisms across ages. In the third section, we relate the processes of selective attention to three domains important to academic foundations: language, literacy, and mathematics. Fourth, drawing on recent literatures on the effects of video-game play and mind-brain training on selective attention, we discuss the possibility of training selective attention. The final section examines the application of these principles to educationally-focused attention-training programs for children.     

Electrophysiological evidence against a neurotransmitter role of corticotropin-releasing hormone (CRH) in primary somatosensory cortex

The possible neurotransmitter role of corticotropin-releasing hormone (CRH) was studied in the primary somatosensory cortex of the rat. Electrical activity of single neurones was recorded in layers II–VI of cortex, and in the region of the locus coeruleus. Iontophoresis and pressure ejection were employed to locally apply CRH, and changes in spontaneous, synaptically driven and iontophoretically driven firing were examined. In the cortex, of 62 neurones recorded most (51) were completely unaffected by high and prolonged current/pressure ejections of CRH. Depression of firing was occasionally seen (8 of 62), while a very few (3) were weakly excited. Of 25 cells studied with vibrissal stimulation to evoke excitatory synaptic responses, responses in two cells were depressed and in two they were enhanced. Activity that was evoked by iontophoretic ejection of excitatory amino acids, such as glutamate, was depressed in 6 of 40 cells (none were enhanced). Such effects as were seen were weak and often difficult to reproduce. The effect of CRH on depressions produced by GABA was also tested in four experiments. No effects on the amplitude or duration of the depressions were observed. In contrast recordings made in the midbrain, in the region of the locus coeruleus, resulted in over half the neurones (11 of 20) showing clear reproducible excitatory responses to CRH applications. Solutions used in the experiments were analysed using chromatography, radioimmunoassay and bioassay, and no significant degradation of the peptide was found compared with the synthetic standard (CRH (1–41)). The data provide evidence against CRH acting as a neurotransmitter or modulator in primary cortex, suggesting that the CRH which is localized in certain types of cortical cells is involved in other processes.     

Evidence against a role of the rat's dorsal noradrenergic bundle in selective attention and place memory

In Experiment 1,6-hydroxydopamine-induced lesions of the dorsal noradrenergic bundle (DNB) in rats did not impair either acquisition of non-delayed alternation, retention of non-delayed alternation, or performance of alternation with delays in a T-maze, whether or not the goal arms of the maze were visually distinctive. These results were in contrast with those of a previous report indicating that DNB lesions impair learning of spatial alternation. In Experiment 2, the lack of a reliable effect of DNB lesions on learning performance of spatial alternation was confirmed. However, the rats with DNB lesions showed an impairment of spontaneous alternation. The negative results of the present study do not support either the hypothesis that depletion of forebrain noradrenaline impairs selective attention or the hypothesis that such depletion induces an amnesia for past places. On the other hand, the finding of impaired spontaneous alternation is consistent with previous observations suggesting that depletion of forebrain noradrenaline impairs habituation of fear reactions.