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issn:0898-929
1.  Early visual responses predict conscious face perception within and between subjects during binocular rivalry 
Journal of cognitive neuroscience  2013;25(6):969-985.
Previous studies indicate that conscious face perception may be related to neural activity in a large time window around 170-800ms after stimulus presentation, yet in the majority of these studies changes in conscious experience are confounded with changes in physical stimulation. Using multivariate classification on MEG data recorded when participants reported changes in conscious perception evoked by binocular rivalry between a face and a grating, we showed that only MEG signals in the 120-320ms time range, peaking at the M170 around 180ms and the P2m at around 260ms, reliably predicted conscious experience. Conscious perception could not only be decoded significantly better than chance from the sensors that showed the largest average difference, as previous studies suggest, but also from patterns of activity across groups of occipital sensors that individually were unable to predict perception better than chance. Additionally, source space analyses showed that sources in the early and late visual system predicted conscious perception more accurately than frontal and parietal sites, although conscious perception could also be decoded there. Finally, the patterns of neural activity associated with conscious face perception generalized from one participant to another around the times of maximum prediction accuracy. Our work thus demonstrates that the neural correlates of particular conscious contents (here, faces) are highly consistent in time and space within individuals and that these correlates are shared to some extent between individuals.
doi:10.1162/jocn_a_00353
PMCID: PMC4060063  PMID: 23281780
2.  MEG activity related to conscious perception is stable within individuals across years but not between individuals 
Journal of cognitive neuroscience  2013;26(4):840-853.
Studies indicate that conscious perception is related to changes in neural activity within a time window that varies between 130-320ms after stimulus presentation, yet it is not known whether such neural correlates of conscious perception are stable across time. Here, we examined the generalization across time within individuals and across different individuals. We trained classification algorithms to decode conscious perception from neural activity recorded during binocular rivalry using MEG. The classifiers were then used to predict the perception of the same participants during different recording sessions either days or years later, as well as between different participants. No drop in decoding accuracy was observed when decoding across years compared to days, whereas a large drop in decoding accuracy was found for between-participant decoding. Furthermore, underlying percept-specific MEG signals remained stable in terms of latency, amplitude and sources within participants across years whereas differences were found in all of these domains between individuals. Our findings demonstrate that the neural correlates of conscious perception are stable across years for adults, but differ across individuals. Moreover, the study validates decoding based on MEG data as a method for further studies of correlations between individual differences in perceptual contents and between-participant decoding accuracies.
doi:10.1162/jocn_a_00525
PMCID: PMC4059382  PMID: 24236699
Ambiguous perception; awareness; consciousness; magnetoencephalography; plasticity
3.  T’ain’t what you say, it’s the way that you say it – left insula and inferior frontal cortex work in interaction with superior temporal regions to control the performance of vocal impersonations 
Journal of cognitive neuroscience  2013;25(11):1875-1886.
Historically, the study of human identity perception has focused on faces, but the voice is also central to our expressions and experiences of identity (P. Belin, Fecteau, & Bedard, 2004). Our voices are highly flexible and dynamic; talkers speak differently depending on their health, emotional state, and the social setting, as well as extrinsic factors such as background noise. However, to date, there have been no studies of the neural correlates of identity modulation in speech production. In the current fMRI experiment, we measured the neural activity supporting controlled voice change in adult participants performing spoken impressions. We reveal that deliberate modulation of vocal identity recruits the left anterior insula and inferior frontal gyrus, supporting the planning of novel articulations. Bilateral sites in posterior superior temporal/inferior parietal cortex and a region in right mid/anterior superior temporal sulcus showed greater responses during the emulation of specific vocal identities than for impressions of generic accents. Using functional connectivity analyses, we describe roles for these three sites in their interactions with the brain regions supporting speech planning and production. Our findings mark a significant step toward understanding the neural control of vocal identity, with wider implications for the cognitive control of voluntary motor acts.
doi:10.1162/jocn_a_00427
PMCID: PMC4050359  PMID: 23691984
4.  A Neural Measure of Precision in Visual Working Memory 
Journal of cognitive neuroscience  2013;25(5):754-761.
Recent studies suggest that the temporary storage of visual detail in working memory is mediated by sensory recruitment or sustained patterns of stimulus-specific activation within feature-selective regions of visual cortex. According to a strong version of this hypothesis, the relative “quality” of these patterns should determine the clarity of an individual’s memory. Here, we provide a direct test of this claim. We used fMRI and a forward encoding model to characterize population-level orientation-selective responses in visual cortex while human participants held an oriented grating in memory. This analysis, which enables a precise quantitative description of multivoxel, population-level activity measured during working memory storage, revealed graded response profiles whose amplitudes were greatest for the remembered orientation and fell monotonically as the angular distance from this orientation increased. Moreover, interparticipant differences in the dispersion—but not the amplitude—of these response profiles were strongly correlated with performance on a concurrent memory recall task. These findings provide important new evidence linking the precision of sustained population-level responses in visual cortex and memory acuity.
doi:10.1162/jocn_a_00357
PMCID: PMC4041615  PMID: 23469889
5.  The neural basis of inhibitory effects of semantic and phonological neighbors in spoken word production 
Journal of cognitive neuroscience  2013;25(9):1504-1516.
Theories of word production and word recognition generally agree that multiple word candidates are activated during processing. The facilitative and inhibitory effects of these “lexical neighbors” have been studied extensively using behavioral methods and have spurred theoretical development in psycholinguistics, but relatively little is known about the neural basis of these effects and how lesions may affect them. The present study used voxel-wise lesion overlap subtraction to examine semantic and phonological neighbor effects in spoken word production following left hemisphere stroke. Increased inhibitory effects of near semantic neighbors were associated with inferior frontal lobe lesions, suggesting impaired selection among strongly activated semantically-related candidates. Increased inhibitory effects of phonological neighbors were associated with posterior superior temporal and inferior parietal lobe lesions. In combination with previous studies, these results suggest that such lesions cause phonological-to-lexical feedback to more strongly activate phonologically-related lexical candidates. The comparison of semantic and phonological neighbor effects and how they are affected by left hemisphere lesions provides new insights into the cognitive dynamics and neural basis of phonological, semantic, and cognitive control processes in spoken word production.
doi:10.1162/jocn_a_00408
PMCID: PMC4038097  PMID: 23647518
lexical neighborhood; phonological neighborhood; semantic neighborhood; spoken word production; aphasia; lesion overlap
6.  Behavioral Choice-related Neuronal Activity in Monkey Primary Somatosensory Cortex in a Haptic Delay Task 
Journal of cognitive neuroscience  2012;24(7):1634-1644.
The neuronal activity in the primary somatosensory cortex was collected when monkeys performed a haptic–haptic DMS task. We found that, in trials with correct task performance, a substantial number of cells showed significant differential neural activity only when the monkeys had to make a choice between two different haptic objects. Such a difference in neural activity was significantly reduced in incorrect response trials. However, very few cells showed the choice-only differential neural activity in monkeys who performed a control task that was identical to the haptic–haptic task but did not require the animal to either actively memorize the sample or make a choice between two objects at the end of a trial. From these results, we infer that the differential activity recorded from cells in the primary somato-sensory cortex in correct performance reflects the neural process of behavioral choice, and therefore, it is a neural correlate of decision-making when the animal has to make a haptic choice.
doi:10.1162/jocn_a_00235
PMCID: PMC4034389  PMID: 22452554
7.  Robust Selectivity for Faces in the Human Amygdala in the Absence of Expressions 
Journal of cognitive neuroscience  2013;25(12):2086-2106.
There is a well-established posterior network of cortical regions that plays a central role in face processing and that has been investigated extensively. In contrast, although responsive to faces, the amygdala is not considered a core face-selective region, and its face selectivity has never been a topic of systematic research in human neuroimaging studies. Here, we conducted a large-scale group analysis of fMRI data from 215 participants. We replicated the posterior network observed in prior studies but found equally robust and reliable responses to faces in the amygdala. These responses were detectable in most individual participants, but they were also highly sensitive to the initial statistical threshold and habituated more rapidly than the responses in posterior face-selective regions. A multivariate analysis showed that the pattern of responses to faces across voxels in the amygdala had high reliability over time. Finally, functional connectivity analyses showed stronger coupling between the amygdala and posterior face-selective regions during the perception of faces than during the perception of control visual categories. These findings suggest that the amygdala should be considered a core face-selective region.
doi:10.1162/jocn_a_00469
PMCID: PMC4029947  PMID: 23984945
8.  Searching for something familiar or novel: ERP correlates of top-down attentional selection for specific items or categories 
Journal of cognitive neuroscience  2013;25(5):719-729.
Visual search is often guided by top-down attentional templates that specify target-defining features. But search can also occur at the level of object categories. We measured the N2pc component, a marker of attentional target selection, in two visual search experiments where targets were defined either categorically (e.g., any letter), or at the item level (e.g., the letter C) by a prime stimulus. In both experiments, an N2pc was elicited during category search, in both familiar and novel contexts (Experiment 1) and with symbolic primes (Experiment 2), indicating that even when targets are only defined at the category level, they are selected at early sensory-perceptual stages. However, the N2pc emerged earlier and was larger during item-based search compared to category-based search, demonstrating the superiority of attentional guidance by item-specific templates. We discuss the implications of these findings for attentional control and category learning.
doi:10.1162/jocn_a_00352
PMCID: PMC3804645  PMID: 23281777
9.  Genetic polymorphisms regulating dopamine signaling in the frontal cortex interact to affect target detection under high working memory load 
Journal of cognitive neuroscience  2013;26(2):10.1162/jocn_a_00501.
Frontal-dependent task performance is typically modulated by dopamine (DA) according to an inverted-U pattern, whereby intermediate levels of DA signaling optimizes performance. Numerous studies implicate trait differences in DA signaling based on differences in the catechol-O-methyltransferase (COMT) gene in executive function task performance. However, little work has investigated genetic variations in DA signaling downstream from COMT. One candidate is the dopamine- and cAMP-regulated phosphoprotein of molecular weight 32 kDa (DARPP-32), which mediates signaling through the DA D1-type receptor, the dominant DA receptor in the frontal cortex. Using an n-back task, we used signal detection theory to measure performance in a healthy adult population (n=97) genotyped for single nucleotide polymorphisms in the COMT (rs4680) and DARPP-32 (rs907094) genes. Correct target detection (hits), and false alarms were used to calculate d' measures for each working memory load (0-, 2-, and 3-back). At the highest load (3-back) only, we observed a significant COMT×DARPP-32 interaction, such that the DARPP-32 T/T genotype enhanced target detection in COMTValVal individuals, but impaired target detection in COMTMet carriers. These findings suggest that enhanced dopaminergic signaling via the DARPP-32 T allele aids target detection in individuals with presumed low frontal DA (COMTValVal) but impairs target detection in those with putatively higher frontal DA levels (COMTMet carriers). Moreover, these data support an inverted-U model with intermediate levels of DA signaling optimizing performance on tasks requiring maintenance of mental representations in working memory.
doi:10.1162/jocn_a_00501
PMCID: PMC3877727  PMID: 24144248
COMT; DARPP-32; DRD1; executive function; n-back
10.  Love to win or hate to lose? Asymmetry of dopamine D2 receptor binding predicts sensitivity to reward vs. punishment 
Journal of cognitive neuroscience  2013;26(5):1039-1048.
Humans show consistent differences in the extent to which their behavior reflects a bias towards appetitive approach-related behavior or avoidance of aversive stimuli (Elliot, 2008). We examined the hypothesis that in healthy subjects this motivational bias (assessed by self-report and by a probabilistic learning task that allows direct comparison of the relative sensitivity to reward and punishment) reflects lateralization of dopamine signaling. Using [F-18]fallypride to measure D2/D3 binding , we found that self-reported motivational bias was predicted by the asymmetry of frontal D2 binding. Similarly, striatal and frontal asymmetries in D2 dopamine receptor binding, rather than absolute binding levels, predicted individual differences in learning from reward vs. punishment. These results suggest that normal variation in asymmetry of dopamine signaling may, in part, underlie human personality and cognition.
doi:10.1162/jocn_a_00544
PMCID: PMC3972269  PMID: 24345165
dopamine; asymmetry; reward; punishment; individual differences
11.  Conceptual and Data-based Investigation of Genetic Influences and Brain Asymmetry: A Twin Study of Multiple Structural Phenotypes 
Journal of cognitive neuroscience  2013;26(5):1100-1117.
Right–left regional cerebral differences are a feature of the human brain linked to functional abilities, aging, and neuro-developmental and mental disorders. The role of genetic factors in structural asymmetry has been incompletely studied. We analyzed data from 515 individuals (130 monozygotic twin pairs, 97 dizygotic pairs, and 61 unpaired twins) from the Vietnam Era Twin Study of Aging to answer three questions about genetic determinants of brain structural asymmetry: First, does the magnitude of heritability differ for homologous regions in each hemisphere? Despite adequate power to detect regional differences, heritability estimates were not significantly larger in one hemisphere versus the other, except left > right inferior lateral ventricle heritability. Second, do different genetic factors influence left and right hemisphere size in homologous regions? Inter-hemispheric genetic correlations were high and significant; in only two subcortical regions (pallidum and accumbens) did the estimate statistically differ from 1.0. Thus, there was little evidence for different genetic influences on left and right hemisphere regions. Third, to what extent do genetic factors influence variability in left–right size differences? There was no evidence that variation in asymmetry (i.e., the size difference) of left and right homologous regions was genetically determined, except in pallidum and accumbens. Our findings suggest that genetic factors do not play a significant role in determining individual variation in the degree of regional cortical size asymmetries measured with MRI, although they may do so for volume of some subcortical structures. Despite varying interpretations of existing left–right, we view the present results as consistent with previous findings.
doi:10.1162/jocn_a_00531
PMCID: PMC3999438  PMID: 24283492
12.  Oscillatory coupling reveals the dynamic reorganization of large-scale neural networks as cognitive demands change 
Journal of cognitive neuroscience  2013;26(1):175-188.
Cognitive operations are thought to emerge from dynamic interactions between spatially distinct brain areas. Synchronization of oscillations has been proposed to regulate these interactions, but we do not know whether this large-scale synchronization can respond rapidly to changing cognitive demands. Here we show that as task demands change during a trial, multiple distinct networks are dynamically formed and reformed via oscillatory synchronization. Distinct frequency-coupled networks were rapidly formed to process reward value, maintain information in visual working memory, and deploy visual attention. Strong single-trial correlations showed that networks formed even before the presentation of imperative stimuli could predict the strength of subsequent networks, as well as the speed and accuracy of behavioral responses seconds later. These frequency-coupled networks better predicted single-trial behavior than either local oscillations or event-related potentials. Our findings demonstrate the rapid reorganization of networks formed by dynamic activity in response to changing task demands within a trial.
doi:10.1162/jocn_a_00470
PMCID: PMC3990735  PMID: 23984947
cross-frequency coupling; synchrony; reward; visual working memory; visual attention
13.  Attention-modulated alpha-band oscillations protect against intrusion of irrelevant information 
Journal of cognitive neuroscience  2013;25(9):1463-1476.
Combining high-density scalp EEG recordings with a sensitive analog measure of short-term memory’s fidelity, we characterized the temporal dynamics of intentional ignoring, and related those dynamics to the intrusion of task-irrelevant information. On each trial of the task, two study Gabors were briefly presented in succession. A green or red disc preceding each Gabor signified whether that Gabor should be remembered or ignored, respectively. With cue-stimulus intervals of 300, 600, or 900 ms presented in separate sessions, we found that the onset of posterior, pre-stimulus alpha oscillations varied with the length of the interval. Although stimulus onset time was entirely predictable, the longer the cue-stimulus interval, the earlier the increase in pre-stimulus alpha power. However, the alpha-band modulation was not simply locked to the cue offset. The temporal envelopes of posterior alpha-band modulation were strikingly similar for both cued attending and cued ignoring, and differed only in magnitude. This similarity suggests that cued attending includes suppression of task irrelevant, spatial processing. Supporting the view that alpha-band oscillations represent inhibition, our graded measure of recall revealed that when the stimulus to be ignored appears second in the sequence, peri-stimulus alpha power predicted the degree to which that irrelevant stimulus distorted subsequent recall of the stimulus that was to be remembered. These results demonstrate that timely deployment of attention-related alpha-band oscillations can aid short-term memory by filtering out task-irrelevant information.
doi:10.1162/jocn_a_00395
PMCID: PMC3987899  PMID: 23530921
14.  When memory is not enough: Electrophysiological evidence for goal-dependent use of working memory representations in guiding visual attention 
Journal of cognitive neuroscience  2011;23(10):2650-2664.
Biased competition theory proposes that representations in working memory drive visual attention to select similar inputs. However, behavioral tests of this hypothesis have led to mixed results. These inconsistent findings could be due to the inability of behavioral measures to reliably detect the early, automatic effects on attentional deployment that the memory representations exert. Alternatively, executive mechanisms may govern how working memory representations influence attention based on higher-level goals. In the present study, we tested these hypotheses using the N2pc component of participants’ event-related potentials (ERPs) to directly measure the early deployments of covert attention. Participants searched for a target in an array that sometimes contained a memory-matching distractor. In Experiments 1–3, we manipulated the difficulty of the target discrimination and the proximity of distractors, but consistently observed that covert attention was deployed to the search targets and not the memory-matching distractors. In Experiment 4, we showed that when participants’ goal involved attending to memory-matching items that these items elicited a large and early N2pc. Our findings demonstrate that working memory representations alone are not sufficient to guide early deployments of visual attention to matching inputs and that goal-dependent executive control mediates the interactions between working memory representations and visual attention.
doi:10.1162/jocn.2011.21602
PMCID: PMC3981747  PMID: 21254796
Event related potentials; Attention: Visual; Memory: Working memory; Executive functions
15.  High Proficiency in a Second Language is Characterized by Greater Involvement of the First Language Network: Evidence from Chinese Learners of English 
Journal of cognitive neuroscience  2013;25(10):1649-1663.
The assimilation hypothesis argues that second language learning recruits the brain network for processing the native language, whereas the accommodation hypothesis argues that learning a second language recruits brain structures not involved in native language processing. This study tested these hypotheses by examining brain activation of a group of native Chinese speakers, who were late bilinguals with varying levels of proficiency in English, when they performed a rhyming judgment to visually presented English word pairs (CE group) during fMRI. Assimilation was examined by comparing the CE group to native Chinese speakers performing the rhyming task in Chinese (CC group), and accommodation was examined by comparing the CE group to native English speakers performing the rhyming task in English (EE group). The CE group was very similar in activation to the CC group, supporting the assimilation hypothesis. Additional support for the assimilation hypothesis was the finding that higher proficiency in the CE group was related to increased activation in the Chinese network (as defined by the CC > EE), including the left middle frontal gyrus, the right inferior parietal lobule, and the right precuneus, and decreased activation in the English network (as defined by the EE > CC), including the left inferior frontal gyrus and the left inferior temporal gyrus. Although most of the results support assimilation, there was some evidence for accommodation as the CE group showed less activation in the Chinese network including the right middle occipital gyrus, which has been argued to be involved in holistic visuospatial processing of Chinese characters.
doi:10.1162/jocn_a_00414
PMCID: PMC3979436  PMID: 23654223
16.  The Biology of Linguistic Expression Impacts Neural Correlates for Spatial Language 
Journal of cognitive neuroscience  2012;25(4):517-533.
Biological differences between signed and spoken languages may be most evident in the expression of spatial information. PET was used to investigate the neural substrates supporting the production of spatial language in American Sign Language as expressed by classifier constructions, in which handshape indicates object type and the location/motion of the hand iconically depicts the location/motion of a referent object. Deaf native signers performed a picture description task in which they overtly named objects or produced classifier constructions that varied in location, motion, or object type. In contrast to the expression of location and motion, the production of both lexical signs and object type classifier morphemes engaged left inferior frontal cortex and left inferior temporal cortex, supporting the hypothesis that unlike the location and motion components of a classifier construction, classifier handshapes are categorical morphemes that are retrieved via left hemisphere language regions. In addition, lexical signs engaged the anterior temporal lobes to a greater extent than classifier constructions, which we suggest reflects increased semantic processing required to name individual objects compared with simply indicating the type of object. Both location and motion classifier constructions engaged bilateral superior parietal cortex, with some evidence that the expression of static locations differentially engaged the left intraparietal sulcus. We argue that bilateral parietal activation reflects the biological underpinnings of sign language. To express spatial information, signers must transform visual–spatial representations into a body-centered reference frame and reach toward target locations within signing space.
doi:10.1162/jocn_a_00339
PMCID: PMC3715382  PMID: 23249348
17.  Putting the pieces together: The role of dorsolateral prefrontal cortex in relational memory encoding 
Journal of cognitive neuroscience  2011;23(1):257-265.
Results from functional magnetic resonance imaging (fMRI) have strongly supported the idea that the ventrolateral prefrontal cortex (VLPFC) contributes to successful memory formation, but the role the dorsolateral prefrontal cortex (DLPFC) in memory encoding is more controversial. Some findings suggest that the DLPFC is recruited when one is processing relationships between items in working memory, and this processing specifically promotes subsequent memory for these relationships. However, previous studies could not rule out the possibility that DLPFC promotes memory during all elaborative encoding conditions and contributes to memory on all subsequent associative memory tests. To address this question directly, we used functional magnetic resonance imaging (fMRI) to examine activity during two encoding tasks which prompted participants to encode either relational or item-specific information. On relational trials, participants imagined pairs of items interacting, whereas on item-specific trials, participants imagined the items spatially separated and in different sizes. After scanning we examined memory for relational information and item-specific information. FMRI results showed that DLPFC activity specifically promoted memory for relational information during relational encoding and not memory for item-specific information during item-specific encoding. In contrast, activity in the VLPFC predicted memory for both relational and item-specific information. The present results are consistent with the idea that the DLPFC specifically contributes to successful memory formation through its role in building relationships amongst items.
doi:10.1162/jocn.2010.21459
PMCID: PMC3970078  PMID: 20146616
prefrontal; encoding; long-term memory; DLPFC; relational; item-specific; cognitive control; executive processing
18.  Perceptual demand modulates activation of human auditory cortex in response to task-irrelevant sounds 
Journal of cognitive neuroscience  2013;25(9):1553-1562.
In the visual modality, perceptual demand on a goal-directed task have been shown to modulate the extent to which irrelevant information can be disregarded at a sensory-perceptual stage of processing. In the auditory modality the effect of perceptual demand on neural representations of task-irrelevant sounds is unclear. We compared simultaneous event-related potentials (ERP) and functional magnetic resonance imaging (fMRI) responses associated with task-irrelevant sounds across parametrically modulated perceptual task demands in a dichotic-listening paradigm. Participants performed a signal detection task in one ear (Attend ear) while ignoring task-irrelevant syllable sounds in the other ear (Ignore ear). Results revealed modulation of syllable processing by auditory perceptual demand in a region of interest in middle left superior temporal gyrus and in negative ERP activity 130–230 ms post stimulus onset. Increasing the perceptual demand in the Attend ear was associated with a reduced neural response in both fMRI and ERP to task-irrelevant sounds. These findings are in support of a selection model whereby ongoing perceptual demands modulate task-irrelevant sound processing in auditory cortex.
doi:10.1162/jocn_a_00416
PMCID: PMC3970782  PMID: 23647558
Attention; Auditory; Perceptual demand/load; fMRI; ERP
19.  Neural Correlates of People’s Hypercorrection of Their False Beliefs 
Journal of cognitive neuroscience  2012;24(7):1571-1583.
Despite the intuition that strongly held beliefs are particularly difficult to change, the data on error correction indicate that general information errors that people commit with a high degree of belief are especially easy to correct. This finding is called the hypercorrection effect. The hypothesis was tested that the reason for hypercorrection stems from enhanced attention and encoding that results from a metacognitive mismatch between the person’s confidence in their responses and the true answer. This experiment, which is the first to use imaging to investigate the hyper-correction effect, provided support for this hypothesis, showing that both metacognitive mismatch conditions—that in which high confidence accompanies a wrong answer and that in which low confidence accompanies a correct answer—revealed anterior cingulate and medial frontal gyrus activations. Only in the high confidence error condition, however, was an error that conflicted with the true answer mentally present. And only the high confidence error condition yielded activations in the right TPJ and the right dorsolateral pFC. These activations suggested that, during the correction process after error commission, people (1) were entertaining both the false belief as well as the true belief (as in theory of mind tasks, which also manifest the right TPJ activation) and (2) may have been suppressing the unwanted, incorrect information that they had, themselves, produced (as in think/no-think tasks, which also manifest dorsolateral pFC activation). These error-specific processes as well as enhanced attention because of metacognitive mismatch appear to be implicated.
doi:10.1162/jocn_a_00228
PMCID: PMC3970786  PMID: 22452558
20.  Infection, Incest, and Iniquity: Investigating the Neural Correlates of Disgust and Morality 
Journal of cognitive neuroscience  2008;20(9):1529-1546.
Disgust, an emotion related to avoiding harmful substances, has been linked to moral judgments in many behavioral studies. However, the fact that participants report feelings of disgust when thinking about feces and a heinous crime does not necessarily indicate that the same mechanisms mediate these reactions. Humans might instead have separate neural and physiological systems guiding aversive behaviors and judgments across different domains. The present interdisciplinary study used functional magnetic resonance imaging (n = 50) and behavioral assessment to investigate the biological homology of pathogen-related and moral disgust. We provide evidence that pathogen-related and sociomoral acts entrain many common as well as unique brain networks. We also investigated whether morality itself is composed of distinct neural and behavioral subdomains. We provide evidence that, despite their tendency to elicit similar ratings of moral wrongness, incestuous and nonsexual immoral acts entrain dramatically separate, while still overlapping, brain networks. These results (i) provide support for the view that the biological response of disgust is intimately tied to immorality, (ii) demonstrate that there are at least three separate domains of disgust, and (iii) suggest strongly that morality, like disgust, is not a unified psychological or neurological phenomenon.
doi:10.1162/jocn.2008.20109
PMCID: PMC3969035  PMID: 18345982
21.  Investigating the Time Course of Spoken Word Recognition: Electrophysiological Evidence for the Influences of Phonological Similarity 
Journal of cognitive neuroscience  2009;21(10):1893-1906.
Behavioral and modeling evidence suggests that words compete for recognition during auditory word identification, and that phonological similarity is a driving factor in this competition. The present study used event-related potentials [ERPs] to examine the temporal dynamics of different types of phonological competition (i.e., cohort and rhyme). ERPs were recorded during a novel picture-word matching task, where a target picture was followed by an auditory word that either matched the target (CONE-cone), or mismatched in one of three ways: rhyme (CONE-bone), cohort (CONE-comb), and unrelated (CONE-fox). Rhymes and cohorts differentially modulated two distinct ERP components, the Phonological Mismatch Negativity [PMN] and the N400, revealing the influences of pre-lexical and lexical processing components in speech recognition. Cohort mismatches resulted in late increased negativity in the N400, reflecting disambiguation of the later point of miscue and the combined influences of top-down expectations and misleading bottom-up phonological information on processing. In contrast, we observed a reduction in the N400 for rhyme mismatches, reflecting lexical activation of rhyme competitors. Moreover, the observed rhyme effects suggest that there is interaction between phoneme-level and lexical-level information in the recognition of spoken words. The results support the theory that both levels of information are engaged in parallel during auditory word recognition in a way that permits both bottom-up and top-down competition effects.
doi:10.1162/jocn.2008.21142
PMCID: PMC3965566  PMID: 18855555 CAMSID: cams939
Perception; Auditory processing; Event related potentials
22.  Neural Responses to Ambiguity Involve Domain-general and Domain-specific Emotion Processing Systems 
Journal of cognitive neuroscience  2013;25(4):547-557.
Extant research has examined the process of decision making under uncertainty, specifically in situations of ambiguity. However, much of this work has been conducted in the context of semantic and low-level visual processing. An open question is whether ambiguity in social signals (e.g., emotional facial expressions) is processed similarly or whether a unique set of processors come on-line to resolve ambiguity in a social context. Our work has examined ambiguity using surprised facial expressions, as they have predicted both positive and negative outcomes in the past. Specifically, whereas some people tended to interpret surprise as negatively valenced, others tended toward a more positive interpretation. Here, we examined neural responses to social ambiguity using faces (surprise) and nonface emotional scenes (International Affective Picture System). Moreover, we examined whether these effects are specific to ambiguity resolution (i.e., judgments about the ambiguity) or whether similar effects would be demonstrated for incidental judgments (e.g., nonvalence judgments about ambiguously valenced stimuli). We found that a distinct task control (i.e., cingulo-opercular) network was more active when resolving ambiguity. We also found that activity in the ventral amygdala was greater to faces and scenes that were rated explicitly along the dimension of valence, consistent with findings that the ventral amygdala tracks valence. Taken together, there is a complex neural architecture that supports decision making in the presence of ambiguity: (a) a core set of cortical structures engaged for explicit ambiguity processing across stimulus boundaries and (b) other dedicated circuits for biologically relevant learning situations involving faces.
doi:10.1162/jocn_a_00363
PMCID: PMC3961008  PMID: 23363410
23.  Domain-general signals in the cingulo-opercular network for visuospatial attention and episodic memory 
Journal of cognitive neuroscience  2013;26(3):551-568.
We investigated the functional properties of a previously described cingulo-opercular network (CON) putatively involved in cognitive control. Analyses of common fMRI task-evoked activity during perceptual and episodic memory search tasks that differently recruited the dorsal attention (DAN) and default mode network (DMN) established the generality of this network. Regions within the CON (anterior insula/frontal operculum and anterior cingulate/presupplementary cortex) displayed sustained signals during extended periods in which participants searched for behaviourally relevant information in a dynamically changing environment or from episodic memory in the absence of sensory stimulation. The CON was activated during all phases of both tasks, which involved trial initiation, target detection, decision and response, indicating its consistent involvement in a broad range of cognitive processes. Functional connectivity analyses showed that the CON flexibly linked with the DAN or DMN regions during perceptual or memory search, respectively. Aside from the CON, only a limited number of regions, including the lateral prefrontal cortex, showed evidence of domain-general, sustained activity, although in some cases the common activations may have reflected the functional-anatomical variability of domain-specific regions rather than a true domain-generality. These additional regions also showed task-dependent functional connectivity with the DMN and DAN, suggesting that this feature is not a specific marker of cognitive control. Finally, multivariate clustering analyses separated the CON from other fronto-parietal regions previously associated with cognitive control, indicating a unique fingerprint. We conclude that the CON’s functional properties and interactions with other brain regions support a broad role in cognition, consistent with its characterization as a task-control network.
doi:10.1162/jocn_a_00504
PMCID: PMC3947512  PMID: 24144246
24.  Contextual Processing of Abstract Concepts Reveals Neural Representations of Non-Linguistic Semantic Content 
Journal of cognitive neuroscience  2013;25(6):920-935.
Concepts develop for many aspects of experience, including abstract internal states and abstract social activities that do not refer to concrete entities in the world. The current study assessed the hypothesis that, like concrete concepts, distributed neural patterns of relevant, non-linguistic semantic content represent the meanings of abstract concepts. In a novel neuroimaging paradigm, participants processed two abstract concepts (convince, arithmetic) and two concrete concepts (rolling, red) deeply and repeatedly during a concept-scene matching task that grounded each concept in typical contexts. Using a catch trial design, neural activity associated with each concept word was separated from neural activity associated with subsequent visual scenes to assess activations underlying the detailed semantics of each concept. We predicted that brain regions underlying mentalizing and social cognition (e.g., medial prefrontal cortex, superior temporal sulcus) would become active to represent semantic content central to convince, whereas brain regions underlying numerical cognition (e.g., bilateral intraparietal sulcus) would become active to represent semantic content central to arithmetic. The results supported these predictions, suggesting that the meanings of abstract concepts arise from distributed neural systems that represent concept-specific content.
doi:10.1162/jocn_a_00361
PMCID: PMC3947606  PMID: 23363408
25.  Effects of Acute Dopamine Precusor Depletion on Immediate Reward Selection Bias and Working Memory Depend on Catechol-O-methyltransferase Genotype 
Journal of cognitive neuroscience  2013;25(12):10.1162/jocn_a_00464.
Little agreement exists as to acute dopamine (DA) manipulation effects on intertemporal choice in humans. We previously found that catechol-O-methyltransferase (COMT) Val158Met genotype predicts individual differences in immediate reward selection bias among adults. Moreover, we and others have shown that the relationship between COMT genotype and immediate reward bias is inverted in adolescents. No previous pharmacology studies testing DA manipulation effects on intertemporal choice have accounted for COMT genotype, and many have included participants in the adolescent age range (18–21) as adults. Moreover, many studies have included female subjects without strict cycle phase control, although recent evidence demonstrates that cyclic estradiol elevations interact with COMT genotype to affect DA-dependent cognition. These factors may have interacted with DA manipulations in past studies, potentially occluding detection of effects. Therefore, we predicted that among healthy adult males (ages 22–40), frontal DA tone, as indexed by COMT genotype, would interact with acute changes in DA signaling to affect intertemporal choice. In a double-blind, placebo-controlled design, we decreased central DA via administration of an amino acid beverage deficient in the DA precursors, phenylalanine and tyrosine (P/T[−]), and tested effects on immediate reward bias in a delay-discounting (DD) task and working memory (WM) in an n-back task. We found no main effect of beverage on DD or WM performance, but did find significant beverage*genotype effects. These results suggest that the effect of DA manipulations on DD depends on individual differences in frontal DA tone, which may have impeded some past efforts to characterize DA’s role in immediate reward bias in humans.
doi:10.1162/jocn_a_00464
PMCID: PMC3816120  PMID: 23937688
delay discounting; executive function; frontal; immediate reward bias; tyrosine

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