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Over past decades, converging neuroimaging and electrophysiological findings have suggested a crucial role of posterior parietal cortex (PPC) in supporting the storage capacity of visual short-term memory (VSTM). Moreover, a few recent studies have shown that electrical stimulation over PPC can enhance VSTM capacity, making it a promising method for improving VSTM function. However, the reliability of these results is still in question because null findings have also been observed. Among studies that reported significant effects, some found increased VSTM capacity only in people with low capacity. Here, we hypothesized that subjects' encoding strategy might be a key source of these variable results. To directly test this hypothesis, we stimulated PPC using transcranial direct-current stimulation (tDCS) in male and female human subjects instructed to employ different encoding strategies during a VSTM recall task. We found that VSTM capacity was higher in subjects who were instructed to remember all items in the supra-capacity array of visual stimuli (i.e., the remember-all group), compared to subjects who were told to focus on a subset of these stimuli (i.e., the remember-subset group). As predicted, anodal tDCS over PPC significantly enhanced VSTM capacity only in the remember-subset group, but not in the remember-all group. Additionally, no effect of encoding strategy or its interaction with electrical stimulation was found on VSTM precision. Together, these results suggest that encoding strategy has a selective influence on VSTM capacity and this influence of encoding strategy mediates the effect of electrical stimulation over PPC on VSTM function.

The neural mechanisms associated with the limited capacity of working memory (WM) has long been studied, but it is still unclear which neural regions are associated with the precision of visual WM. Here, an orientation recall task for estimating the trial-wise precision of visual WM was performed and then repeated two weeks later in an fMRI scanner. Results showed that activity in frontal and parietal regions during WM maintenance scaled with WM load, but not with the precision of WM (i.e., recall error in radians). Conversely, activity in the lateral occipital complex (LOC) during WM maintenance was not affected by memory load, but rather, correlated with WM precision on a trial-by-trial basis. Moreover, activity in LOC also correlated with the individual participant's precision of WM from a separate behavioral experiment. Interestingly, a region within the prefrontal cortex, the inferior frontal junction (IFJ), exhibited greater functional connectivity with LOC when the WM load increased. Together, our findings provide unique evidence that the LOC supports visual WM precision, while communication between the IFJ and LOC varies based on WM load demands. These results suggest an intriguing possibility that distinct neural mechanisms may be associated with general content (load) or detailed information (precision) of WM.

Working memory (WM) deficits have been widely documented in schizophrenia (SZ), and almost all existing studies attributed the deficits to decreased capacity as compared to healthy control (HC) subjects. Recent developments in WM research suggest that other components, such as precision, also mediate behavioral performance. It remains unclear how different WM components jointly contribute to deficits in schizophrenia. We measured the performance of 60 SZ (31 females) and 61 HC (29 females) in a classical delay-estimation visual working memory (VWM) task and evaluated several influential computational models proposed in basic science of VWM to disentangle the effect of various memory components. We show that the model assuming variable precision (VP) across items and trials is the best model to explain the performance of both groups. According to the VP model, SZ exhibited abnormally larger variability of allocating memory resources rather than resources or capacity per se. Finally, individual differences in the resource allocation variability predicted variation of symptom severity in SZ, highlighting its functional relevance to schizophrenic pathology. This finding was further verified using distinct visual features and subject cohorts. These results provide an alternative view instead of the widely accepted decreased-capacity theory and highlight the key role of elevated resource allocation variability in generating atypical VWM behavior in schizophrenia. Our findings also shed new light on the utility of Bayesian observer models to characterize mechanisms of mental deficits in clinical neuroscience., Author summary Working memory is a core cognitive function related to a broad range of cognitive domains such as problem-solving, attention, executive control, and IQ. Although working memory deficits have been well-documented in schizophrenia, the underlying mechanisms remain unclear. Conventional working memory theories attribute working memory deficits in schizophrenia to their reduced memory capacity, overlooking the potential roles of other memory components, such as precision. In this study, we take the approach of computational psychiatry and use computational modeling to uncover the major determinants of working memory deficits. We assess working memory performance of a large cohort of participants (60 schizophrenia patients and 61 demographic matched healthy controls) and evaluate multiple mainstream computational models of visual working memory. The variable precision model turns out to be the best model for both groups. We further find that the poorer performance of schizophrenia patients arises from heterogeneous distribution of memory resources when encoding items in memory. This resource allocation variability can also predict symptom severity in schizophrenia. Our study highlights the use of computational models in psychiatric researches.

Cognitive processes require working memory (WM) that involves a brief period of memory retention known as the delay period. Elevated delay-period activity in the medial prefrontal cortex (mPFC) has been observed, but its functional role in WM tasks remains unclear. We optogenetically suppressed or enhanced activity of pyramidal neurons in mouse mPFC during the delay period. Behavioral performance was impaired during the learning phase but not after the mice were well trained. Delay-period mPFC activity appeared to be more important in memory retention than in inhibitory control, decision-making, or motor selection. Furthermore, endogenous delay-period mPFC activity showed more prominent modulation that correlated with memory retention and behavioral performance. Thus, properly regulated mPFC delay-period activity is critical for information retention during learning of a WM task.

Visual short-term memory (VSTM) provides an on-line mental space for incoming sensory information to be temporally maintained to carry out complex behavioral tasks. Despite its essential functions, the capacity at which VSTM could maintain sensory information is limited (i.e., VSTM can hold only about three to four visual items at once). Moreover, the quality of sensory representation (i.e., precision) degrades as more information has to be maintained in VSTM. Correlational evidence suggests that the level and the pattern of neural activity measured in the posterior parietal cortex (PPC) track both VSTM capacity and precision. However, the causal contributions of the PPC to these different VSTM operations are unclear. Here, we tested whether stimulating the PPC with transcranial direct current stimulation (tDCS) could increase VSTM capacity or precision. We found that stimulating the PPC in male and female human participants selectively enhanced VSTM capacity when the number of memory items exceeded capacity limit, without significant effects on VSTM precision. Moreover, this enhancement of VSTM capacity is region specific as stimulating the prefrontal cortex did not change VSTM capacity or precision. Null stimulation effects in the sensory memory condition confirmed that the tDCS-induced enhancement of VSTM capacity was not simply due to changes in sensory or attentional processes. Altogether, these results provide causal evidence suggesting that the PPC has a more dominant role in supporting the storage capacity of VSTM compared with maintaining the quality of sensory representations. Furthermore, tDCS could be used as a promising noninvasive method to enhance this PPC VSTM-related function.SIGNIFICANCE STATEMENTCorrelational evidence from neuroimaging and electrophysiology suggests that the posterior parietal cortex (PPC) supports the storage capacity of visual short-term memory (VSTM) and the precision of sensory representations maintained in VSTM. However, the causal contributions of the PPC to these different VSTM functions were unclear. Here, we found that electrical stimulation over the PPC selectively enhanced VSTM capacity without changing VSTM precision. Overall, our findings suggest that the PPC has a dominant and causal role in supporting the storage capacity of VSTM.

Neural activity in the dorsolateral prefrontal cortex (DLPFC) has been suggested to integrate information from distinct sensory areas. However, how the DLPFC interacts with the bilateral primary somatosensory cortices (SIs) in tactile-visual cross-modal working memory has not yet been established. In the present study, we applied single-pulse transcranial magnetic stimulation (sp-TMS) over the contralateral DLPFC and bilateral SIs of human participants at various time points, while they performed a tactile-visual delayed matching-to-sample task with a 2-second delay. sp-TMS over the contralateral DLPFC or the contralateral SI at either an sensory encoding stage [i.e. 100 ms after the onset of a vibrotactile sample stimulus (200-ms duration)] or an early maintenance stage (i.e. 300 ms after the onset), significantly impaired the accuracy of task performance; sp-TMS over the contralateral DLPFC or the ipsilateral SI at a late maintenance stage (1600 ms and 1900 ms) also significantly disrupted the performance. Furthermore, at 300 ms after the onset of the vibrotactile sample stimulus, there was a significant correlation between the deteriorating effects of sp-TMS over the contralateral SI and the contralateral DLPFC. These results imply that the DLPFC and the bilateral SIs play causal roles at distinctive stages during cross-modal working memory, while the contralateral DLPFC communicates with the contralateral SI in the early delay, and cooperates with the ipsilateral SI in the late delay.

Studies have indicated that a cortical sensory system is capable of processing information from different sensory modalities. However, it still remains unclear when and how a cortical system integrates and retains information across sensory modalities during learning. Here we investigated the neural dynamics underlying crossmodal associations and memory by recording event-related potentials (ERPs) when human participants performed visuo-tactile (crossmodal) and visuo-visual (unimodal) paired-associate (PA) learning tasks. In a trial of the tasks, the participants were required to explore and learn the relationship (paired or non-paired) between two successive stimuli. EEG recordings revealed dynamic ERP changes during participants' learning of paired-associations. Specifically, (1) the frontal N400 component showed learning-related changes in both unimodal and crossmodal tasks but did not show any significant difference between these two tasks, while the central P400 displayed both learning changes and task differences; (2) a late posterior negative slow wave (LPN) showed the learning effect only in the crossmodal task; (3) alpha-band oscillations appeared to be involved in crossmodal working memory. Additional behavioral experiments suggested that these ERP components were not relevant to the participants' familiarity with stimuli per se. Further, by shortening the delay length (from 1300ms to 400ms or 200 ms) between the first and second stimulus in the crossmodal task, declines in participants' task performance were observed accordingly. Taken together, these results provide insights into the cortical plasticity (induced by PA learning) of neural networks involved in crossmodal associations in working memory.

Previous studies have consistently demonstrated that superior mnemonists (SMs) outperform normal individuals in domain-specific memory tasks. However, the neural correlates of memory-related processes remain unclear. In the current EEG study, SMs and control participants performed a digit memory task during which their brain activity was recorded. Chinese SMs used a digit-image mnemonic for encoding digits, in which they associated 2-digit groups with images immediately after the presentation of each even-position digit in sequences. Behaviorally, SMs' memory of digit sequences was better than the controls'. During encoding in the study phase, SMs showed an increased right central P2 (150-250ms post onset) and a larger right posterior high-alpha (10-14Hz, 500-1720ms) oscillation on digits at even-positions compared with digits at odd-positions. Both P2 and high-alpha oscillations in the study phase co-varied with performance in the recall phase, but only in SMs, indicating that neural dynamics during encoding could predict successful retrieval of digit memory in SMs. Our findings suggest that representation of a digit sequence in SMs using mnemonics may recruit both the early-stage attention allocation process and the sustained information preservation process. This study provides evidence for the role of dynamic and efficient neural encoding processes in mnemonists.

During the course of divergent thinking (DT), the number of generated ideas decreases while the originality of ideas increases. This phenomenon is labeled as serial order effect in DT. The present study investigated whether different executive processes (i.e., updating, shifting, and inhibition) specifically contribute to the serial order effect in DT. Participants' executive functions were measured by corresponding experimental tasks outside of the EEG lab. They were required to generate original uses of conventional objects (alternative uses task) during EEG recording. The behavioral results revealed that the originality of ideas was higher in later stage of DT (i.e., Epoch 2) than in its earlier stage (i.e., Epoch 1) for higher-shifting individuals, but showed no difference between two epochs for lower-shifting individuals. The EEG results revealed that lower-inhibition individuals showed stronger upper alpha (10-13Hz) synchronization in left frontal areas during Epoch 1 compared to during Epoch 2. For higher-inhibition individuals, no changes in upper alpha activity from Epoch 1 to Epoch 2 were found. These findings indicated that shifting and inhibition contributed to create a serial order effect in DT, perhaps because individuals suppress interference from obvious ideas and switch to new idea categories during DT, thus more original ideas appear as time passes by.

The neural processes underlying pain memory are not well understood. To explore these processes, contact heat-evoked potentials (CHEPs) were recorded in humans with electroencephalography (EEG) technique during a delayed matching-to-sample task, a working memory task involving presentations of two successive painful heat stimuli (S-1 and S-2) with different intensities separated by a 2-s interval (the memorization period). At the end of the task, the subject was required to discriminate the stimuli by indicating which (S-1 or S-2) induced more pain. A control task was used, in which no active discrimination was required between stimuli. All event-related potential (ERP) analysis was aligned to the onset of S-1. EEG activity exhibited two successive CHEPs: an N2-P2 complex (∼400 ms after onset of S-1) and an ultralate component (ULC, ∼900 ms). The amplitude of the N2-P2 at vertex, but not the ULC, was significantly correlated with stimulus intensity in these two tasks, suggesting that the N2-P2 represents neural coding of pain intensity. A late negative component (LNC) in the frontal recording region was observed only in the memory task during a 500-ms period before onset of S-2. LNC amplitude differed between stimulus intensities and exhibited significant correlations with the N2-P2 complex. These indicate that the frontal LNC is involved in maintenance of intensity of pain in working memory. Furthermore, alpha-band oscillations observed in parietal recording regions during the late delay displayed significant power differences between tasks. This study provides in the temporal domain previously unidentified neural evidence showing the neural processes involved in working memory of painful stimuli.

The present study aimed to explore the neural correlates underlying the effects of idea evaluation on idea generation in creative thinking. Participants were required to generate original uses of conventional objects (alternative uses task) during EEG recording. A reflection task (mentally evaluating the generated ideas) or a distraction task (object characteristics task) was inserted into the course of idea generation. Behavioral results revealed that participants generated ideas with higher originality after evaluating the generated ideas than after performing the distraction task. The EEG results revealed that idea evaluation was accompanied with upper alpha (10-13 Hz) synchronization, most prominent at frontal cortical sites. Moreover, upper alpha activity in frontal cortices during idea generation was enhanced after idea evaluation. These findings indicate that idea evaluation may elicit a state of heightened internal attention or top-down activity that facilitates efficient retrieval and integration of internal memory representations.

Reactivation renders consolidated memory labile again, and the ensuing temporary reconsolidation process is highly susceptible to mnemonic modification. Here, we show that memories in such an unstable state could be influenced by sheer behavioral means, bypassing the need for pharmacological intervention. Across several experiments using a "face-location association" paradigm in which participants experienced a "Learning - New-learning - Final-test" procedure, we demonstrate that reactivated memory traces were hampered when the new learning was strategically administered at between 0-min and 20-min delay. Using fMRI, we further advance our theoretical understanding that this lability can be mechanistically explained by the differential activation in the hippocampal-amygdala memory system implicated by the post-activation new-learning whereas the mnemonic intrusion caused by newly learned memories is efficaciously reconciled by the left inferior frontal gyrus.

Previous studies have shown that the speed of identifying emotional words is affected by pre-cues of up or down arrows, called the space-valence congruency effect (Zhang, Hu, Zhang, & Wang, 2015). In the present study, we investigate whether this effect is influenced by hypnotic or non-hypnotic suggestions to ignore pre-cues in highly hypnotizable individuals (HHIs). In all conditions, target words (including positive words, negative words and neutral words) primed by up or down arrows were presented to pre-screened HHIs. They were asked to identify whether the target words had emotional valence. Behavioral results showed that the space-valence congruency effect was absent in the hypnotic and non-hypnotic suggestion groups, but present in the non-suggestion control group. Consistently, the amplitudes of P2 components elicited by negative words were significantly larger when primed by down than by up arrows in the non-suggestion control group, and this P2 effect was absent in the hypnotic and non-hypnotic suggestion groups. Moreover, the amplitudes of the late positive components (LPC) showed no significant arrow-priming difference for positive targets in the hypnotic-suggestion and control groups. However, in the non-hypnotic suggestion group, the amplitudes of LPC were significantly larger when primed by down than up arrows for positive targets. Our results showed that suggestions, even without a hypnotic induction, can de-automatize embodied emotional recognition processes and modulate early attentional processing.

In the present study, we searched for causal evidence linking activity in the bilateral primary somatosensory cortex (SI), posterior parietal cortex (PPC), and prefrontal cortex (PFC) with behavioral performance in vibrotactile working memory. Participants performed a vibrotactile delayed matching-to-sample task, while single-pulse transcranial magnetic stimulation (sp-TMS) was applied over these cortical areas at 100, 200, 300, 600, 1600, and 1900 ms after the onset of vibrotactile stimulation (200 ms duration). In our experiments, sp-TMS over the contralateral SI at the early delay (100 and 200 ms) deteriorated the accuracy of task performance, and over the ipsilateral SI at the late delay (1600 and 1900 ms) also induced such deteriorating effects. Furthermore, deteriorating effects caused by sp-TMS over the contralateral DLPFC at the same maintenance stage (1600 ms) were correlated with the effects caused by sp-TMS over the ipsilateral SI, indicating that information retained in the ipsilateral SI during the late delay may be associated with the DLPFC. Taken together, these results suggest that both the contralateral and ipsilateral SIs are involved in tactile WM, and the contralateral DLPFC bridges the contralateral SI and ipsilateral SI for goal-directed action.

Laboratory studies have investigated how individuals with normal memory spans attained digit spans over 80 digits after hundreds of hours of practice. Experimental analyses of their memory skills suggested that their attained memory spans were constrained by the encoding time, for the time needed will increase if the length of digit sequences to be memorised becomes longer. These constraints seemed to be violated by a world-class memorist, Feng Wang (FW), who won the World Memory Championship by recalling 300 digits presented at 1 digit/s. In several studies we examined FW's memory skills underlying his exceptional performance. First FW reproduced his superior memory span of 200 digits under laboratory condition, and we obtained his retrospective reports describing his encoding/retrieval processes (Experiment 1). Further experiments used self-paced memorisation to identify temporal characteristics of encoding of digits in 4-digit clusters (Experiment 2), and explored memory encoding at presentation speeds much faster than 1 digit/s (Experiment 3). FW's superiority over previous digit span experts is explained by his acquisition of well-known mnemonic techniques and his training that focused on rapid memorisation. His memory performance supports the feasibility of acquiring memory skills for improved working memory based on storage in long-term memory.

Spatial attention has been postulated to facilitate perceptual processing via several different mechanisms. For instance, attention can amplify neural responses in sensory areas (sensory gain), mediate neural variability (noise modulation), or alter the manner in which sensory signals are selectively read out by postsensory decision mechanisms (efficient readout). Even in the context of simple behavioral tasks, it is unclear how well each of these mechanisms can account for the relationship between attention-modulated changes in behavior and neural activity because few studies have systematically mapped changes between stimulus intensity, attentional focus, neural activity, and behavioral performance. Here, we used a combination of psychophysics, event-related potentials (ERPs), and quantitative modeling to explicitly link attention-related changes in perceptual sensitivity with changes in the ERP amplitudes recorded from human observers. Spatial attention led to a multiplicative increase in the amplitude of an early sensory ERP component (the P1, peaking ∼80–130 ms poststimulus) and in the amplitude of the late positive deflection component (peaking ∼230–330 ms poststimulus). A simple model based on signal detection theory demonstrates that these multiplicative gain changes were sufficient to account for attention-related improvements in perceptual sensitivity, without a need to invoke noise modulation. Moreover, combining the observed multiplicative gain with a postsensory readout mechanism resulted in a significantly poorer description of the observed behavioral data. We conclude that, at least in the context of relatively simple visual discrimination tasks, spatial attention modulates perceptual sensitivity primarily by modulating the gain of neural responses during early sensory processing

The study aimed to evaluate the effects of noninvasive brain stimulation on cognitive function in healthy older adults and patients with Alzheimer's disease. A comprehensive literature search was performed on noninvasive stimulation studies published from January 1990 to November 2014 in Pubmed and Web of Science. Fourteen articles with a total of 331 participants were identified as studies with healthy older adults, and the mean effect size and 95% confidence interval were estimated. A significant effect size of 0.42 was found for the cognitive outcome. Further subgroup analyses demonstrated more prominent effects for studies delivering the stimulation before the execution of the task and studies applying multiple sessions of stimulation. To assess the effects of stimulation on Alzheimer's disease patients, 11 studies with a total of 200 patients were included in the analysis. A significant effect size of 1.35 was found for the cognitive outcomes. Subgroup analyses indicated more pronounced effects for studies applying the stimulation during the execution of the task compared with studies delivering the stimulation before the execution of the task. Noninvasive brain stimulation has a positive effect on cognitive function in physiological and pathological aging.

In the present study, causal roles of both the primary somatosensory cortex (SI) and the posterior parietal cortex (PPC) were investigated in a tactile unimodal working memory (WM) task. Individual magnetic resonance imaging-based single-pulse transcranial magnetic stimulation (spTMS) was applied, respectively, to the left SI (ipsilateral to tactile stimuli), right SI (contralateral to tactile stimuli) and right PPC (contralateral to tactile stimuli), while human participants were performing a tactile-tactile unimodal delayed matching-to-sample task. The time points of spTMS were 300, 600 and 900 ms after the onset of the tactile sample stimulus (duration: 200 ms). Compared with ipsilateral SI, application of spTMS over either contralateral SI or contralateral PPC at those time points significantly impaired the accuracy of task performance. Meanwhile, the deterioration in accuracy did not vary with the stimulating time points. Together, these results indicate that the tactile information is processed cooperatively by SI and PPC in the same hemisphere, starting from the early delay of the tactile unimodal WM task. This pattern of processing of tactile information is different from the pattern in tactile-visual cross-modal WM. In a tactile-visual cross-modal WM task, SI and PPC contribute to the processing sequentially, suggesting a process of sensory information transfer during the early delay between modalities.

The face inversion effect is a defection in performance in recognizing inverted faces compared with faces presented in their usual upright orientation typically believed to be specific for facial stimuli. McLaren (1997) was able to demonstrate that (a) an inversion effect could be obtained with exemplars drawn from a familiar category, such that upright exemplars were better discriminated than inverted exemplars; and (b) that the inversion effect required that the familiar category be prototype-defined. In this article, we replicate and extend these findings. We show that the inversion effect can be obtained in a standard old/new recognition memory paradigm, demonstrate that it is contingent on familiarization with a prototype-defined category, and establish that the effect is made up of two components. We confirm the advantage for upright exemplars drawn from a familiar, prototype-defined category, and show that there is a disadvantage for inverted exemplars drawn from this category relative to suitable controls. We also provide evidence that there is an N170 event-related potential signature for this effect. These results allow us to integrate a theory of perceptual learning originally proposed by McLaren, Kaye, and Mackintosh (1989) with explanations of the face inversion effect, first reported by Yin.

Abacus experts are able to mentally calculate multi-digit numbers rapidly. Some behavioral and neuroimaging studies have suggested a visuospatial and visuomotor strategy during abacus mental calculation. However, no study up to now has attempted to dissociate temporally the visuospatial neural process from the visuomotor neural process during abacus mental calculation. In the present study, an abacus expert performed the mental addition tasks (8-digit and 4-digit addends presented in visual or auditory modes) swiftly and accurately. The 100% correct rates in this expert’s task performance were significantly higher than those of ordinary subjects performing 1-digit and 2-digit addition tasks. ERPs, EEG source localizations, and fMRI results taken together suggested visuospatial and visuomotor processes were sequentially arranged during the abacus mental addition with visual addends and could be dissociated from each other temporally. The visuospatial transformation of the numbers, in which the superior parietal lobule was most likely involved, might occur first (around 380 ms) after the onset of the stimuli. The visuomotor processing, in which the superior/middle frontal gyri were most likely involved, might occur later (around 440 ms). Meanwhile, fMRI results suggested that neural networks involved in the abacus mental addition with auditory stimuli were similar to those in the visual abacus mental addition. The most prominently activated brain areas in both conditions included the bilateral superior parietal lobules (BA 7) and bilateral middle frontal gyri (BA 6). These results suggest a supra-modal brain network in abacus mental addition, which may develop from normal mental calculation networks.

Functional neuroimaging data have shown that mental calculation involves fronto-parietal areas that are composed of different subsystems shared with other cognitive functions such as working memory and language. Event-related potential (ERP) analysis has also indicated sequential information changes during the calculation process. However, little is known about the dynamic properties of oscillatory networks in this process. In the present study, we applied both ERP and event-related (de-)synchronization (ERS/ERD) analyses to EEG data recorded from normal human subjects performing tasks for sequential visual/auditory mental addition. Results in the study indicate that the late positive components (LPCs) can be decomposed into two separate parts. The earlier element LPC1 (around 360 ms) reflects the computing attribute and is more prominent in calculation tasks. The later element LPC2 (around 590 ms) indicates an effect of number size and appears larger only in a more complex 2-digit addition task. The theta ERS and alpha ERD show modality-independent frontal and parietal differential patterns between the mental addition and control groups, and discrepancies are noted in the beta ERD between the 2-digit and 1-digit mental addition groups. The 2-digit addition (both visual and auditory) results in similar beta ERD patterns to the auditory control, which may indicate a reliance on auditory-related resources in mental arithmetic, especially with increasing task difficulty. These results coincide with the theory of simple calculation relying on the visuospatial process and complex calculation depending on the phonological process.

In the present study, we examined the neural mechanisms underlying cross-modal working memory by analyzing scalp-recorded event-related potentials (ERPs) from normal human subjects performing tactile-tactile unimodal or tactile-auditory cross-modal delay tasks that consisted of stimulus-1 (S-1, tactile), interval (delay), and stimulus-2 (S-2, tactile or auditory). We hypothesized that there would be sequentially discrete task-correlated changes in ERPs representing neural processes of tactile working memory, and in addition, significant differences would be observed in ERPs between the unimodal task and the cross-modal task. In comparison to the ERP components in the unimodal task, two late positive ERP components (LPC-1 and LPC-2) evoked by the tactile S-1 in the delay of the cross-modal task were enhanced by expectation of the associated auditory S-2 presented at the end of the delay. Such enhancement might represent neural activities involved in cross-modal association between the tactile stimulus and the auditory stimulus. Later in the delay, a late negative component (LNC) was observed. The amplitude of LNC depended on information retained during the delay, and when the same information was retained, this amplitude was not influenced by modality or location of S-2 (auditory S-2 through headphones, or tactile S-2 on the left index finger). LNC might represent the neural activity involved in working memory. The above results suggest that the sequential ERP changes in the present study represent temporally distinguishable neural processes, such as the cross-modal association and cross-modal working memory.

Our previous studies on scalp-recorded event-related potentials (ERPs) showed that somatosensory N140 evoked by a tactile vibration in working memory tasks was enhanced when human subjects expected a coming visual stimulus that had been paired with the tactile stimulus. The results suggested that such enhancement represented the cortical activities involved in tactile-visual crossmodal association. In the present study, we further hypothesized that the enhancement represented the neural activities in somatosensory and frontal cortices in the crossmodal association. By applying independent component analysis (ICA) to the ERP data, we found independent components (ICs) located in the medial prefrontal cortex (around the anterior cingulate cortex, ACC) and the primary somatosensory cortex (SI). The activity represented by the IC in SI cortex showed enhancement in expectation of the visual stimulus. Such differential activity thus suggested the participation of SI cortex in the task-related crossmodal association. Further, the coherence analysis and the Granger causality spectral analysis of the ICs showed that SI cortex appeared to cooperate with ACC in attention and perception of the tactile stimulus in crossmodal association. The results of our study support with new evidence an important idea in cortical neurophysiology: higher cognitive operations develop from the modality-specific sensory cortices (in the present study, SI cortex) that are involved in sensation and perception of various stimuli.