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Cognitive control is a key factor in insight generation. However, the neurocognitive mechanisms underlying the generation of insight for different cognitive control remain poorly understood. This study developed a parametric fMRI design, wherein hints for solving Chinese idiom riddles were gradually provided in a stepwise manner (from the first hint, H1, to the final hint, H4). By classifying the step-specific items solved in different hint-uncovering steps/conditions, we could identify insightful responses for different levels of spontaneous or controlled processing. At the behavioral level, the number of insightful problem solving trials reached the maximum at a intermediate level of the cognitively controlled processing and the spontaneously idea generating in H3, while the bilateral insular cortex and thalamus showed the robust engagement, implying the function of these regions in making the optimal balance between external hint processing and internal generated ideas. In addition, we identified brain areas, including the dorsolateral prefrontal cortex (dlPFC), angular gyrus (AG), dorsal anterior cingulate cortex (dACC), and precuneus (PreC), whose activities were parametrically increased with the levels of controlled (from H1 to H4) insightful processing which were increasingly produced by the sequentially revealed hints. Further representational similarity analysis (RSA) found that spontaneous processing in insight featured greater within-condition representational variabilities in widely distributed regions in the executive, salience, and default networks. Altogether, the present study provided new evidence for the relationship between the process of cognitive control and that of spontaneous idea generation in insight problem solving and demystified the function of the insula and thalamus as an interactive interface for the optimal balance of these two processes.

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The relationship between language and thinking has long been a matter of debate and a research focus in studies on thinking and problem solving, including creativity. Previous behavioral studies have found that verbalization of one's internal thoughts does not participate in or even interfere with the creative insight process, thus suggesting that insight may take place nonverbally. In contrast to this hypothesis, the present study proposes a new one. That is, given that the basic categories or fundamental functions of key concepts or objects are critically changed or expanded during insightful thinking, the linguistic processing accompanying insight can be reflected as category-related representation and recategorization processes, which can be critically mediated by the posterior middle temporal gyrus and the angular gyrus (pMTG/AG). Using constraint-relaxation insight riddles as materials in a guided-insight experimental design with external hints to trigger the insightful representational change, this preliminary neuroimaging study of 11 participants found the involvement of pMTG/AG during moments of induced insight, but did not find the activation of left ventral frontal areas which are typically involved in verbalizing of one's internal thoughts. Although this observation still cannot exclude the possibility of internal verbalization in insightful restructuring, it implies that linguistic processing in insight may take the more fundamental form of category-related processing.

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Theoretical assessment of the function of the hippocampus has suggested that given certain physiological constraints at both the neuronal and cortical level, the hippocampus is best suited to associate discontiguous items that occur in different temporal or spatial positions. Conceptually, "discontiguous" refers to events that are to be associated with one another but do not temporally or spatially overlap. However, given that humans can actively maintain information "online" by rehearsing it, even when the information is no longer being presented to the sensory system, the right way to experimentally define "discontiguity" is still a question. Does it refer to a "gap" in the presentation of information (temporal discontiguity) or to an "interruption" of the active maintenance of working memory (WM) information (functional discontiguity)? To assess this, participants were imaged by functional magnetic resonance imaging (fMRI) when making judgments on whether two words were semantically related or not. In contrast with recognition memory that can be carried out through perceptual familiarity heuristics, judgments on semantic relatedness can only be accomplished through associative processing. To assess this experimentally, two words are either (1) presented at the same time (Event AB) or (2) one after the other with an unfilled, cross-viewing delay (Event A_B) (the uninterrupted discontiguity) or (3) presented one after the other, between which participants are required to perform a calculation task (Event A#B) (the interrupted discontiguity). Results of event-related fMRI analysis revealed that relative to Event AB, Event A_B was not associated with more hippocampal activity, whereas Event A#B was. The direct contrast of Event A#B relative to Event A_B also revealed significant hippocampal and parahippocampal activity. This result implied that functional discontiguity (the interruption of online maintenance of the inputted information) could be more apt at engaging the function of the hippocampus.

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Although the function of the hippocampus and adjacent medial temporal lobe (MTL) areas in forming associations is generally recognized, how MTL contributes to form creative associations that could result in novel and appropriate functions or meanings remains unclear. In this study, we compared highly creative combinations (HCCs) of two objects (e.g., that of "lifejacket" and "distress signal device") that resulted in genuine innovative designs comprising additionally unprecedented functions (the "1 + 1 > 2" effects) with the lowly creative combinations (LCCs, e.g., the combination of "set-top box" and "jewelry box") that resulted in nothing more than simple "1 + 1 = 2" effects. The event-related functional magnetic resonance imaging (fMRI) study found that during the "early binding phase," when the combinations of the two objects were initially encoded, the right parahippocampus was more intensively activated during the encoding of HCC relative to LCC trials. However, during the "late integration phase," when participants finally formed a holistic mental representation of new products based on the two-object combinations, both HCCs and LCCs were found to be associated with significantly increased hippocampal and parahippocampal activation relative to the baseline condition, but at a similar level. In this "late integration phase," the functional areas appeared to be more intensively activated in HCCs relative to LCCs located in the posterior middle temporal gyrus (pMTG), the area known to mediate category-related processing. Consistently, our supplementary behavioral study found that, relative to LCCs, HCCs had a higher possibility of resulting in some new conceptual expansions that differed from each of the original two objects that constituted the combinations. These findings indicate that the formation of creative combinations not only require MTL-based novel association-formation, but also pMTG-based novel concept-expansion.

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Total sleep deprivation (TSD) has been consistently found to impair encoding of information during ensuing wakefulness, probably through suppressing NonREM (non-rapid eye movement) sleep. However, a possible contribution of missing REM sleep to this encoding impairment after TSD has so far not been systematically examined in humans, although such contribution might be suspected in particular for emotional information. Here, in two separate experiments in young healthy men, we compared effects of TSD and of selective REM sleep deprivation (REMD), relative to respective control conditions of undisturbed sleep, on the subsequent encoding of neutral and emotional pictures. The pictures were presented in conjunction with colored frames to also assess related source memory. REMD was achieved by tones presented contingently upon initial signs of REM sleep. Encoding capabilities were examined in the evening (18:00h) after the experimental nights, by a picture recognition test right after encoding. TSD significantly decreased both the rate of correctly recognized pictures and of recalled frames associated with the pictures. The TSD effect was robust and translated into an impaired long term memory formation, as it was likewise observed on a second recognition testing one week after the encoding phase. Contrary to our expectation, REMD did not affect encoding in general, or particularly of emotional pictures. Also, REMD did not affect valence ratings of the encoded pictures. However, like TSD, REMD distinctly impaired vigilance at the time of encoding. Altogether, these findings indicate an importance of NonREM rather than REM sleep for the encoding of information that is independent of the emotionality of the materials.

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BACKGROUND: The purpose of this study was to examine the effect of sleep deprivation on flow experience. METHODS: Sixteen healthy male volunteers of mean age 21.4±1.59 (21-24) years participated in two experimental conditions, ie, sleep-deprivation and normal sleep. In the sleep-deprived condition, participants stayed awake at home for 36 hours (from 8 am until 10 pm the next day) beginning on the day prior to an experimental day. In both conditions, participants carried out a simple reaction time (psychomotor vigilance) task and responded to a questionnaire measuring flow experience and mood status. RESULTS: Flow experience was reduced after one night of total sleep deprivation. Sleep loss also decreased positive mood, increased negative mood, and decreased psychomotor performance. CONCLUSION: Sleep deprivation has a strong impact on mental and behavioral states associated with the maintenance of flow, namely subjective well-being.

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We investigated how the structure of the brain network relates to the stability of perceptual alternation in binocular rivalry. Historically, binocular rivalry has provided important new insights to our understandings in neuroscience. Although various relationships between the local regions of the human brain structure and perceptual switching phenomena have been shown in previous researches, the global organization of the human brain structural network relating to this phenomenon has not yet been addressed. To approach this issue, we reconstructed fiber-tract bundles using diffusion tensor imaging and then evaluated the correlations between the speeds of perceptual alternation and fractional anisotropy (FA) values in each fiber-tract bundle integrating among 84 brain regions. The resulting comparison revealed that the distribution of the global organization of the structural brain network showed positive or negative correlations between the speeds of perceptual alternation and the FA values. First, the connections between the subcortical regions stably were negatively correlated. Second, the connections between the cortical regions mainly showed positive correlations. Third, almost all other cortical connections that showed negative correlations were located in one central cluster of the subcortical connections. This contrast between the contribution of the cortical regions to destabilization and the contribution of the subcortical regions to stabilization of perceptual alternation provides important information as to how the global architecture of the brain structural network supports the phenomenon of binocular rivalry.

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We investigated the key anatomical structures mediating interhemispheric integration during the perception of apparent motion across the retinal midline. Previous studies of commissurotomized patients suggest that subcortical structures mediate interhemispheric transmission but the specific regions involved remain unclear. Here, we exploit interindividual variations in the propensity of normal subjects to perceive horizontal motion, in relation to vertical motion. We characterize these differences psychophysically using a Dynamic Dot Quartet (an ambiguous stimulus that induces illusory motion). We then tested for correlations between a tendency to perceive horizontal motion and fractional anisotropy (FA) (from structural diffusion tensor imaging), over subjects. FA is an indirect measure of the orientation and integrity of white matter tracts. Subjects who found it easy to perceive horizontal motion showed significantly higher FA values in the pulvinar. Furthermore, fiber tracking from an independently identified (subject-specific) visual motion area converged on the pulvinar nucleus. These results suggest that the pulvinar is an anatomical hub and may play a central role in interhemispheric integration.

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The present study addressed the hypothesis that emotional stimuli relevant to survival or reproduction (biologically emotional stimuli) automatically affect cognitive processing (e.g., attention, memory), while those relevant to social life (socially emotional stimuli) require elaborative processing to modulate attention and memory. Results of our behavioral studies showed that (1) biologically emotional images hold attention more strongly than do socially emotional images, (2) memory for biologically emotional images was enhanced even with limited cognitive resources, but (3) memory for socially emotional images was enhanced only when people had sufficient cognitive resources at encoding. Neither images' subjective arousal nor their valence modulated these patterns. A subsequent functional magnetic resonance imaging study revealed that biologically emotional images induced stronger activity in the visual cortex and greater functional connectivity between the amygdala and visual cortex than did socially emotional images. These results suggest that the interconnection between the amygdala and visual cortex supports enhanced attention allocation to biological stimuli. In contrast, socially emotional images evoked greater activity in the medial prefrontal cortex (MPFC) and yielded stronger functional connectivity between the amygdala and MPFC than did biological images. Thus, it appears that emotional processing of social stimuli involves elaborative processing requiring frontal lobe activity.

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In the present study, we examined whether and how brief viewing of positive and negative images influences subsequent understanding of solutions to insight problems. For each trial, participants were first presented with an insight problem and then briefly viewed a task-irrelevant positive, negative, or neutral image (660 ms), which was followed by the solution to the problem. In our behavioral study (Study 1), participants were faster to report that they understood the solutions following positive images, and were slower to report it following negative images. A subsequent fMRI study (Study 2) revealed enhanced activity in the angular gyrus and medial prefrontal cortex (MPFC) while viewing solutions following positive, as compared with negative, images. In addition, greater activation of the angular gyrus was associated with more rapid understanding of the solutions. These results suggest that brief viewing of positive images enhances activity in the angular gyrus and MPFC, which results in facilitation of understanding solutions to insight problems.

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In life, we must often learn new associations to people, places, or things we already know. The current fMRI study investigated the neural mechanisms underlying emotional memory updating. Nineteen participants first viewed negative and neutral pictures and learned associations between those pictures and other neutral stimuli, such as neutral objects and encoding tasks. This initial learning phase was followed by a memory updating phase, during which participants learned picture-location associations for old pictures (i.e., pictures previously associated with other neutral stimuli) and new pictures (i.e., pictures not seen in the first phase). There was greater frontopolar/orbito-frontal (OFC) activity when people learned picture-location associations for old negative pictures than for new negative pictures, but frontopolar OFC activity did not significantly differ during learning locations of old versus new neutral pictures. In addition, frontopolar activity was more negatively correlated with the amygdala when participants learned picture-location associations for old negative pictures than for new negative or old neutral pictures. Past studies revealed that the frontopolar OFC allows for updating the affective values of stimuli in reversal learning or extinction of conditioning [e.g., Izquierdo, A., & Murray, E. A. Opposing effects of amygdala and orbital PFC lesions on the extinction of instrumental responding in macaque monkeys. European Journal of Neuroscience, 22, 2341-2346, 2005]; our findings suggest that it plays a more general role in updating associations to emotional stimuli.

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Cooperation among genetically unrelated individuals is a fundamental aspect of society, but it has been a longstanding puzzle in biological and social sciences. Recently, theoretical studies in biology and economics showed that conditional cooperation-cooperating only with those who have exhibited cooperative behavior-can spread over a society. Furthermore, experimental studies in psychology demonstrated that people are actually conditional cooperators. In this study, we used functional magnetic resonance imaging to investigate the neural system underlying conditional cooperation by scanning participants during interaction with cooperative, neutral and non-cooperative opponents in prisoner's dilemma games. The results showed that: (i) participants cooperated more frequently with both cooperative and neutral opponents than with non-cooperative opponents; and (ii) a brain area related to cognitive inhibition of pre-potent responses (right dorsolateral prefrontal cortex) showed greater activation, especially when participants confronted non-cooperative opponents. Consequently, we suggest that cognitive inhibition of the motivation to cooperate with non-cooperators drives the conditional behavior.

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Chunk decomposition is the decomposing of familiar patterns into their component elements so that they can be regrouped in another meaningful manner. Such a regrouping is sometimes critically required in problem solving because during initial encoding the problem elements become automatically grouped into familiar chunks and this may prohibit finding a novel or efficient solution to problems [G. Knoblich, S. Ohlsson, H. Haider, D. Rhenius, Constraint relaxation and chunk decomposition in insight problem solving, J. Exp. Psychol. Learn. Mem. Cogn. 25 (1999) 1534-1556]. In order to elucidate the brain mechanisms underlying the process of chunk decomposition, we developed a task that uses Chinese character as materials. Chinese characters are ideal examples of perceptual chunks. They are composed of radicals, which in turn, are composed of strokes. Because radicals are meaningful chunks themselves but strokes are not meaningful in isolation, it is much easier to separate a character by its radicals than to separate a character by its strokes. By comparing the stroke-level decomposition and the radical-level decomposition, we observed activities in occipital, frontal, and parietal lobes. Most importantly, during the moment of chunk decomposition, we found the early visual cortex showed a tendency of negative activation whereas the higher visual cortex showed a tendency of positive activation. This suggests that in order to successfully decompose a chunk, the higher visual areas must at least partly be 'disconnected' from the input provided by early visual processing in order to allow simple features to be rearranged into a different perceptual chunk. We conclude that early perceptual processes can crucially affect thinking and problem solving.

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Previous work identified bilateral regions in the occipitoparietal lobe sensitive to the complexity of relational information [Phillips, S., Niki, K., 2003. Increased bilateral occipitoparietal activity for retention of binary versus unary indexed lists in pair recognition. NeuroImage 20 (2), 1226-1235]. Here, we investigate the effect of learning on sensitivity to relational complexity. Eight subjects were scanned on a pair recognition task before, during and after a 2-week training period when subjects learned to recognize a set of shape pairs. For each trial of the pair recognition task, subjects determined whether a probe pair appeared in a list of learned or novel pairs. In the low/high relational complexity condition, every pair in list AB CD EF/AB AD CB was uniquely identifiable by an item in either/both the first or/and second position. Whole-brain and region of interest contrasts revealed a significant interaction between complexity and learning in the occipitoparietal lobe. The increase in activity for the retention of high versus low complexity lists was greater for learned than novel pair lists. Subjects were more likely to respond to low complexity lists as though they were high complexity prior to training. The results suggest that this region provides a window into effective relational complexity, that is, complexity of relational information as processed by the subject, not as presumed by task design.

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To examine the hypothesis that the hippocampus is necessary to overcome temporal or spatial "discontiguity" (Wallenstein et al., Trends Neurosci 1998; 21:317-323), subjects were imaged by functional magnetic resonance imaging (fMRI) when they were making judgments as to whether two words were semantically related. Two words were presented, either at the same time (the Simultaneous Presentation Condition) or one after the other with a short unfilled rest period (the Delayed Presentation Condition). The latter condition, relative to the former, was proposed to involve the process of "discontiguity association." Event-related fMRI results of eight subjects showed that, relative to the binding of simultaneously presented words, the binding of delay presented words was associated with left hippocampus activity. This result provided direct neuroimaging evidence for the role of the hippocampus in "discontiguity association."

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An 'Aha! reaction' is a brief moment of exceptional thinking where an unexpected change in one's mental perspective reveals the solution to an otherwise intractable problem. In this event-related fMRI study, subjects read incomprehensible sentences followed by solution cues that were used to evoke such a reaction by triggering an alternative interpretation of the critical concepts. For 73% of the trials, subjects attributed their failure in the initial stage of sentence presentation to "having thought about it in another direction". This behavior implies that the breaking of mental impasse is a critical component of the Aha! reaction phenomenon. During the Aha! reaction we observed anterior cingulate and left lateral prefrontal cortical activation, which are two areas known to mediate cognitive conflict.

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AIM: To examine whether precuneus subserves the attentive control of finger movement or whether it mediates the movement preparation and motor inhibition. METHODS: In the Preparation Stage, subjects were shown with a 5-number string in which each number stood for a key-pressing response, the number strings included a complex pattern (eg, 4-1-4-2-3), or a simple one (eg, 2-2-2-2-2), or a null one (ie, x-x-x-x-x). In the Execution Stage, five reaction signs were presented one by one and subjects were required to press the corresponding key to each sign sequentially (eg, in the 4-1-4-2-3 preparation example, subjects press key 4 to the first sign, press key 1 to the second sign, key 4 to the third sign and so on). For the null preparation pattern, five numbers, rather than the reaction signs, were shown at the same pace as in the other two conditions and subjects were to press the corresponding keys. RESULTS: Left medial frontal gyrus (BA 6) and precentral gyrus (BA 6) were involved in both of the Preparation Stage and the Execution Stage, whereas left precuneus (BA 7) was activated only in the Execution Stage. CONCLUSION: Precuneus mediates the attentive control of finger movement, but not the movement preparation or motor inhibition.

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Cognitive complexity has been characterized by relations processed, rather than items stored. Separating these factors is difficult, because processing more complex relations often involves holding more items in memory. Previous research, NeuroImage, 17, 1031-1055) identified parietal lobes with more item relationships, but not more items by varying index length-fewest number of positions having a unique combination of items. For example, AB CD EF is a unary (length one) indexed list of three pairs, because all items are unique at the first (or second) position; AB AD CB is a binary indexed list, because only pairs of items are unique. But, these lists also differ in number of associates. In this experiment, index length was varied independently of the numbers of items and associates. Subjects were asked to make a recognition judgment for each three-pair list: Was the test pair in the previous list? Random effects analysis contrasting two binary indexed lists (AB AC CB and AB AD CB) minus two unary indexed lists (AB BC CA and AB BC CD) revealed increased occipital and parietal activity (bilaterally) during the retention period for both binary indexed list types. This result is explained by index length, but not by item load or item fan, because the numbers of items and item associates were the same for the corresponding unary and binary list types. For peak voxels in left and right precuneus, activity during retention for both binary list types was also greater than for a third unary indexed list (AB CD EF). Because binary indexes require more positions (roles) to individuate pairs, we suggest that the increased activity in precuneus relates to spatial rehearsal in that more attention is directed to both positions to maintain the integrity of the memory trace.

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Since the work of Wolfgang Kohler, the process of "insight" in problem solving has been the subject of considerable investigation. Yet, the neural correlates of "insight" remain unknown. Theoretically, "insight" means the reorientation of one's thinking, including breaking of the unwarranted "fixation" and forming of novel, task-related associations among the old nodes of concepts or cognitive skills. Processes closely related to these aspects have been implicated in the hippocampus. In this research, the neural correlates of "insight" were investigated using Japanese riddles, by imaging the answer presentation and comprehension events, just after participants failed to resolve them. The results of event-related functional magnetic resonance imaging (fMRI) analysis demonstrated that the right hippocampus was critically highlighted and that a wide cerebral cortex was also involved in this "insight" event. To the best of our knowledge, this work is the first neuroimaging study to have investigated the neural correlates of "insight" in problem solving.

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Working memory is affected by items stored and the relations between them. However, separating these factors has been difficult, because increased items usually accompany increased associations/relations. Hence, some have argued, relational effects are reducible to item effects. We overcome this problem by manipulating index length: the fewest number of item positions at which there is a unique item, or tuple of items (if length >1), for every instance in the relational (memory) set. Longer indexes imply greater similarity (number of shared items) between instances and higher load on encoding processes. Subjects were given lists of study pairs and asked to make a recognition judgement. The number of unique items and index length in the three list conditions were: (1) AB, CD: four/one; (2) AB, CD, EF: six/one; and (3) AB, AD, CB: four/two, respectively. Japanese letters were used in Experiments 1 (kanji-ideograms) and 2 (hiragana-phonograms); numbers in Experiment 3; and shapes generated from Fourier descriptors in Experiment 4. Across all materials, right dominant temporoparietal and middle frontal gyral activity was found with increased index length, but not items during study. In Experiment 5, a longer delay was used to isolate retention effects in the absence of visual stimuli. Increased left hemispheric activity was observed in the precuneus, middle frontal gyrus, and superior temporal gyrus with increased index length for the delay period. These results show that relational load is not reducible to item load.

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