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1.  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
2.  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
3.  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
4.  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
5.  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
6.  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
7.  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
8.  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
9.  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
10.  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
11.  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
12.  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
13.  Meaningful Family Relationships: Neurocognitive Buffers of Adolescent Risk Taking 
Journal of cognitive neuroscience  2012;25(3):374-387.
Discordant development of brain regions responsible for cognitive control and reward processing may render adolescents susceptible to risk taking. Identifying ways to reduce this neural imbalance during adolescence can have important implications for risk taking and associated health outcomes. Accordingly, we sought to examine how a key family relationship—family obligation—can reduce this vulnerability. Forty-eight adolescents underwent an fMRI scan during which they completed a risk-taking and cognitive control task. Results suggest that adolescents with greater family obligation values show decreased activation in the ventral striatum when receiving monetary rewards and increased dorsolateral PFC activation during behavioral inhibition. Reduced ventral striatum activation correlated with less real-life risk-taking behavior and enhanced dorsolateral PFC activation correlated with better decision-making skills. Thus, family obligation may decrease reward sensitivity and enhance cognitive control, thereby reducing risk-taking behaviors.
doi:10.1162/jocn_a_00331
PMCID: PMC3932333  PMID: 23163412
14.  Generic Inhibition of the Selected Movement and Constrained Inhibition of Nonselected Movements during Response Preparation 
Journal of cognitive neuroscience  2013;26(2):269-278.
Previous studies have identified two inhibitory mechanisms that operate during action selection and preparation. One mechanism, competition resolution, is manifest in the inhibition of the nonselected response and attributed to competition between candidate actions. The second mechanism, impulse control, is manifest in the inhibition of the selected response and is presumably invoked to prevent premature response. To identify constraints on the operation of these two inhibitory mechanisms, we manipulated the effectors used for the response alternatives, measuring changes in corticospinal excitability with motor-evoked potentials to TMS. Inhibition of the selected response (impulse control) was independent of the task context, consistent with a model in which this form of inhibition is automatically triggered as part of response preparation. In contrast, inhibition of the nonselected response (competition resolution) was context-dependent. Inhibition of the nonselected response was observed when the response alternatives involved movements of the upper limbs but was absent when one response alternative involved an upper limb and the other involved a lower limb. Interestingly, competition resolution for pairs of upper limbs did not require homologous effectors, observed when a left index finger response was pitted with either a nonhomologous right index finger movement or a right arm movement. These results argue against models in which competition resolution is viewed as a generic or fully flexible process, as well as models based on strong anatomical constraints. Rather, they are consistent with models in which inhibition for action selection is constrained by the similarity between the potential responses, perhaps reflecting an experience-dependent mechanism sensitive to the past history of competitive interactions.
doi:10.1162/jocn_a_00492
PMCID: PMC3912843  PMID: 24047388
15.  Dynamic oscillatory processes governing cued orienting and allocation of auditory attention 
Journal of cognitive neuroscience  2013;25(11):1926-1943.
In everyday listening situations, we need to constantly switch between alternative sound sources and engage attention according to cues that match our goals and expectations. The exact neuronal bases of these processes are poorly understood. We investigated oscillatory brain networks controlling auditory attention using cortically constrained fMRI-weighted magnetoencephalography/ electroencephalography (MEG/EEG) source estimates. During consecutive trials, subjects were instructed to shift attention based on a cue, presented in the ear where a target was likely to follow. To promote audiospatial attention effects, the targets were embedded in streams of dichotically presented standard tones. Occasionally, an unexpected novel sound occurred opposite to the cued ear, to trigger involuntary orienting. According to our cortical power correlation analyses, increased frontoparietal/temporal 30–100 Hz gamma activity at 200–1400 ms after cued orienting predicted fast and accurate discrimination of subsequent targets. This sustained correlation effect, possibly reflecting voluntary engagement of attention after the initial cue-driven orienting, spread from the temporoparietal junction, anterior insula, and inferior frontal (IFC) cortices to the right frontal eye fields. Engagement of attention to one ear resulted in a significantly stronger increase of 7.5–15 Hz alpha in the ipsilateral than contralateral parieto-occipital cortices 200–600 ms after the cue onset, possibly reflecting crossmodal modulation of the dorsal visual pathway during audiospatial attention. Comparisons of cortical power patterns also revealed significant increases of sustained right medial frontal cortex theta power, right dorsolateral prefrontal cortex and anterior insula/IFC beta power, and medial parietal cortex and posterior cingulate cortex gamma activity after cued vs. novelty-triggered orienting (600–1400 ms). Our results reveal sustained oscillatory patterns associated with voluntary engagement of auditory spatial attention, with the frontoparietal and temporal gamma increases being best predictors of subsequent behavioral performance.
doi:10.1162/jocn_a_00452
PMCID: PMC3788846  PMID: 23915050
16.  Cortical Spatio-temporal Dynamics Underlying Phonological Target Detection in Humans 
Journal of cognitive neuroscience  2010;23(6):1437-1446.
Selective processing of task-relevant stimuli is critical for goal-directed behavior. We used electrocorticography to assess the spatio-temporal dynamics of cortical activation during a simple phonological target detection task, in which subjects press a button when a prespecified target syllable sound is heard. Simultaneous surface potential recordings during this task revealed a highly ordered temporal progression of high gamma (HG, 70– 200 Hz) activity across the lateral hemisphere in less than 1 sec. The sequence demonstrated concurrent regional sensory processing of speech syllables in the posterior superior temporal gyrus (STG) and speech motor cortex, and then transitioned to sequential task-dependent processing from prefrontal cortex (PFC), to the final motor response in the hand sensorimotor cortex. STG activation was modestly enhanced for target over nontarget sounds, supporting a selective gain mechanism in early sensory processing, whereas PFC was entirely selective to targets, supporting its role in guiding response behavior. These results reveal that target detection is not a single cognitive event, but rather a process of progressive target selectivity that involves large-scale rapid parallel and serial processing in sensory, cognitive, and motor structures to support goal-directed human behavior.
doi:10.1162/jocn.2010.21466
PMCID: PMC3895406  PMID: 20465359
17.  How Does Reward Expectation Influence Cognition in the Human Brain? 
Journal of cognitive neuroscience  2008;20(11):10.1162/jocn.2008.20140.
The prospect of reward changes how we think and behave. We investigated how this occurs in the brain using a novel continuous performance task in which fluctuating reward expectations biased cognitive processes between competing spatial and verbal tasks. Critically, effects of reward expectancy could be distinguished from induced changes in task-related networks. Behavioral data confirm specific bias toward a reward-relevant modality. Increased reward expectation improves reaction time and accuracy in the relevant dimension while reducing sensitivity to modulations of stimuli characteristics in the irrelevant dimension. Analysis of functional magnetic resonance imaging data shows that the proximity to reward over successive trials is associated with increased activity of the medial frontal cortex regardless of the modality. However, there are modality-specific changes in brain activity in the lateral frontal, parietal, and temporal cortex. Analysis of effective connectivity suggests that reward expectancy enhances coupling in both early visual pathways and within the prefrontal cortex. These distributed changes in task-related cortical networks arise from subjects’ representations of future events and likelihood of reward.
doi:10.1162/jocn.2008.20140
PMCID: PMC3886193  PMID: 18416677
18.  Prestimulus subsequent memory effects for auditory and visual events 
Journal of cognitive neuroscience  2010;22(6):10.1162/jocn.2009.21298.
It has been assumed that the effective encoding of information into memory primarily depends on neural activity elicited when an event is initially encountered. Recently, it has been shown that memory formation also relies on neural activity just before an event. The precise role of such activity in memory is currently unknown. Here, we address whether pre-stimulus activity affects the encoding of auditory and visual events, is set up on a trial-by-trial basis, and varies as a function of the type of recognition judgment an item later receives. Electrical brain activity was recorded from the scalps of 24 healthy young adults while they made semantic judgments on randomly intermixed series of visual and auditory words. Each word was preceded by a cue signalling the modality of the upcoming word. Auditory words were preceded by auditory cues and visual words by visual cues. A recognition memory test with remember/know judgments followed after a delay of about 45 min. As observed previously, a negative-going, frontally-distributed modulation just before visual word onset predicted later recollection of the word. Crucially, the same effect was found for auditory words and observed on stay as well as switch trials. These findings emphasize the flexibility and general role of pre-stimulus activity in memory formation, and support a functional interpretation of the activity in terms of semantic preparation. At least with an unpredictable trial sequence, the activity is set up anew on each trial.
doi:10.1162/jocn.2009.21298
PMCID: PMC3881065  PMID: 19583467
Memory encoding; recognition memory; EEG; pre-stimulus activity; input modality; task switching
19.  Electrophysiological correlates of stimulus-driven reorienting deficits after interference with right parietal cortex during a spatial attention task: a TMS-EEG study 
Journal of cognitive neuroscience  2012;24(12):10.1162/jocn_a_00287.
Transcranial magnetic stimulation (TMS) interference over right intraparietal sulcus (IPS) causally disrupts behaviorally and electroencephalographic (EEG) rhythmic correlates of endogenous spatial orienting prior to visual target presentation (Capotosto et al. 2009; 2011). Here we combine data from our previous studies to examine whether right parietal TMS during spatial orienting also impairs stimulus-driven re-orienting or the ability to efficiently process unattended stimuli, i.e. stimuli outside the current focus of attention. Healthy subjects (N=24) performed a Posner spatial cueing task while their EEG activity was being monitored. Repetitive TMS (rTMS) was applied for 150 milliseconds (ms) simultaneously to the presentation of a central arrow directing spatial attention to the location of an upcoming visual target.
Right IPS-rTMS impaired target detection, especially for stimuli presented at unattended locations; it also caused a modulation of the amplitude of parieto-occipital positive ERPs peaking at about 480 ms (P3) post-target. The P3 significantly decreased for unattended targets, and significantly increased for attended targets after right IPS-rTMS as compared to Sham stimulation. Similar effects were obtained for left IPS stimulation albeit in a smaller group of subjects. We conclude that disruption of anticipatory processes in right IPS has prolonged effects that persist during target processing. The P3 decrement may reflect interference with post-decision processes that are part of stimulus-driven re-orienting. Right IPS is a node of functional interaction between endogenous spatial orienting and stimulus-driven re-orienting processes in human vision.
doi:10.1162/jocn_a_00287
PMCID: PMC3879779  PMID: 22905824
Visuospatial attention; visual cortex; rTMS; EEG; P30
20.  Dorsal and ventral attention systems underlie social and symbolic cueing 
Journal of cognitive neuroscience  2013;26(1):10.1162/jocn_a_00461.
Eye gaze is a powerful cue for orienting attention in space. Studies examining whether gaze and symbolic cues recruit the same neural mechanisms have found mixed results. We tested whether there is a specialized attentional mechanism for social cues. We separately measured BOLD activity during orienting and reorienting attention following predictive gaze and symbolic cues. Results showed that gaze and symbolic cues exerted their influence through the same neural networks, but also produced some differential modulations. Dorsal fronto-parietal regions in left IPS and bilateral MT+/lateral occipital cortex only showed orienting effects for symbolic cues while right pIPS showed larger validity effects following gaze cues. Both exceptions may reflect the greater automaticity of gaze cues: symbolic orienting may require more effort, while disengaging attention during reorienting may be more difficult following gaze cues. Face-selective regions, identified with a face localizer, showed selective activations for gaze cues reflecting sensory processing but no attentional modulations. Therefore, no evidence was found linking face-selective regions to a hypothetical, specialized mechanism for orienting attention to gaze cues. However, a functional connectivity analysis showed greater connectivity between face-selective regions and right pIPS, pSTS and IFG during gaze cueing, consistent with proposals that face-selective regions may send gaze signals to parts of the DAN and VAN. Finally, although the default-mode network is thought to be involved in social cognition, this role does not extend to gaze orienting as these regions were more deactivated following gaze cues and showed less functional connectivity with face-selective regions during gaze cues.
doi:10.1162/jocn_a_00461
PMCID: PMC3879788  PMID: 23937692
21.  Youthful Memory Capacity in Old Brains: Anatomic and Genetic Clues from the Northwestern SuperAging Project 
The Northwestern University SuperAging Project recruits community dwellers over the age of 80 who have unusually high performance on tests of episodic memory. In a previous report, a small cohort of SuperAgers was found to have higher cortical thickness on structural MRI than a group of age-matched but cognitively average peers. SuperAgers also displayed a patch of ACC where cortical thickness was higher than in 50- to 60-year-old younger cognitively healthy adults. In additional analyses, some SuperAgers had unusually low densities of age-related Alzheimer pathology and unusually high numbers of von Economo neurons in the anterior cingulate gyrus. SuperAgers were also found to have a lower frequency of the ε4 allele of apolipoprotein E than the general population. These preliminary results show that above-average memory capacity can be encountered in advanced age. They also offer clues to potential biological factors that may promote resistance to age-related involutional changes in the structure and function of the brain.
doi:10.1162/jocn_a_00300
PMCID: PMC3541673  PMID: 23198888
22.  The Neural Correlates of Cognitive Reappraisal during Emotional Autobiographical Memory Recall 
We used fMRI to investigate the neural processes engaged as individuals down- and up-regulated the emotions associated with negative autobiographical memories (AMs) using cognitive reappraisal strategies. Our analyses examined neural activity during 3 separate phases, as participants: (a) viewed a reappraisal instruction (i.e., Decrease, Increase, Maintain), (b) searched for an AM referenced by a self-generated cue, and (c) elaborated upon the details of the AM being held in mind. Decreasing emotional intensity primarily engaged activity in regions previously implicated in cognitive control (e.g., dorsal and ventral lateral prefrontal cortex), emotion generation and processing (e.g., amygdala, insula), and visual imagery (e.g., precuneus) as participants searched for and retrieved events. In contrast, increasing emotional intensity engaged similar regions during the instruction phase (i.e., before a memory cue was presented) and again as individuals later elaborated upon the details of the events they had recalled. These findings confirm that reappraisal can modulate neural activity during the recall of personally-relevant events, though the time course of this modulation appears to depend on whether individuals are attempting to down- or up-regulate their emotions.
doi:10.1162/jocn_a_00289
PMCID: PMC3631011  PMID: 22905826
23.  The role of transverse occipital sulcus in scene perception and its relationship to object individuation in inferior intraparietal sulcus 
Journal of cognitive neuroscience  2013;25(10):1711-1722.
The parietal cortex has been functionally divided into various subregions; however, very little is known about how these areas relate to each other. Two such regions are the transverse occipital sulcus (TOS) scene area and inferior intraparietal sulcus (IPS). TOS exhibits similar activation patterns to the scene selective parahippocampal place area (PPA), suggesting its role in scene perception. Inferior IPS, in contrast, has been shown to participate in object individuation and selection via location. Interestingly, both regions have been localized to the same general area of the brain. If these two were actually the same brain region, it would have important implications regarding these regions’ role in cognition. To explore this, we first localized TOS and inferior IPS in individual participants and examined the degree of overlap between these regions in each participant. We found that TOS showed only a minor degree of overlap with inferior IPS (∼10%). We then directly explored the role of TOS and inferior IPS in object individuation and scene perception by examining their responses to furnished rooms, empty rooms, isolated furniture, and multiple isolated objects. If TOS and inferior IPS were the same region, we would expect to see similar response patterns in both. Instead, the response of TOS was predominantly scene selective, while activity in inferior IPS was primarily driven by the number of objects present in the display, regardless of scene context. These results show that TOS and inferior IPS are nearby, but distinct regions, with different functional roles in visual cognition.
doi:10.1162/jocn_a_00422
PMCID: PMC3758388  PMID: 23662863
Inferior intraparietal sulcus; transverse occipital sulcus; scene processing; multiple object individuation; functional MRI
24.  Objects and categories: Feature statistics and object processing in the ventral stream 
Journal of cognitive neuroscience  2013;25(10):1723-1735.
Recognising an object involves more than just visual analyses; its meaning must also be decoded. Extensive research has shown that processing the visual properties of objects relies on a hierarchically organised stream in ventral occipitotemporal cortex, with increasingly more complex visual features being coded from posterior to anterior sites culminating in the perirhinal cortex (PRC) in the anteromedial temporal lobe (aMTL). The neurobiological principles of the conceptual analysis of objects remain more controversial. Much research has focussed on two neural regions - the fusiform gyrus and aMTL, both of which show semantic category differences, but of different types. fMRI studies show category differentiation in the fusiform gyrus, based on clusters of semantically similar objects, whereas category-specific deficits, specifically for living things, are associated with damage to the aMTL. These category-specific deficits for living things have been attributed to problems in differentiating between highly similar objects, a process which involves the PRC. To determine whether the PRC and the fusiform gyri contribute to different aspects of an object’s meaning, with differentiation between confusable objects in the PRC and categorisation based on object similarity in the fusiform, we carried out an fMRI study of object processing based on a feature-based model which characterises the degree of semantic similarity and difference between objects and object categories. Participants saw 388 objects for which feature statistic information was available, and named the objects at the basic-level while undergoing fMRI scanning. After controlling for the effects of visual information, we found that feature statistics that capture similarity between objects formed category clusters in fusiform gyri, such that objects with many shared features (typical of living things) were associated with activity in the lateral fusiform gyri while objects with fewer shared features (typical of nonliving things) were associated with activity in the medial fusiform gyri. Significantly, a feature statistic reflecting differentiation between highly similar objects, enabling object-specific representations, was associated with bilateral PRC activity. These results confirm that the statistical characteristics of conceptual object features are coded in the ventral stream, supporting a conceptual feature-based hierarchy, and integrating disparate findings of category responses in fusiform gyri and category deficits in aMTL into a unifying neurocognitive framework.
doi:10.1162/jocn_a_00419
PMCID: PMC3767967  PMID: 23662861
25.  RESTING-STATE MODULATION OF ALPHA RHYTHMS BY INTERFERENCE WITH ANGULAR GYRUS ACTIVITY 
Journal of cognitive neuroscience  2013;26(1):10.1162/jocn_a_00460.
The Default-Mode Networks (DMN) is active during restful wakefulness and suppressed during goal-driven behavior. We hypothesize that inhibitory interference with spontaneous ongoing, i.e. not task-driven, activity in the Angular Gyrus (AG), one of the core regions of the DMN, will enhance the dominant idling electroencephalographic (EEG) alpha rhythms observed in the resting state. Fifteen right-handed healthy adult volunteers underwent to this study. Compared to sham stimulation, magnetic stimulation (1 Hz for 1 min) over both left and right AG, but not over frontal eye field (FEF) or intra-parietal sulcus (IPS), core regions of the dorsal attention network (DAN), enhanced the dominant alpha power density (8–10 Hz) in occipito-parietal cortex. Furthermore, right AG-rTMS enhanced intra-hemispheric alpha coherence (8–10 Hz). These results suggest that AG plays a causal role in the modulation of dominant low-frequency alpha rhythms in the resting state condition.
doi:10.1162/jocn_a_00460
PMCID: PMC3861401  PMID: 23937690
Alpha rhythms; EEG; rTMS; Resting state

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