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1.  Evidence for specificity of ERP abnormalities during response inhibition in ADHD children: A comparison with Reading Disorder children without ADHD 
Brain and cognition  2009;72(2):228-237.
Executive function and working memory deficits are not only present in ADHD, but also in Reading Disorder (RD). Here, high-density ERPs were recorded during the Stop Signal Task in 53 children and adolescents: An ADHD-combined type group, a group with RD, and a healthy control group. The ADHD-C group displayed unique abnormalities of the frontal N200. Both healthy controls and RD groups showed a success-related right frontal N200 modulation, which was absent in the ADHD group. Second, for Success Inhibition trials, the ADHD-C had smaller right frontal N200 waves relative to healthy controls, while the RD group didn't. In contrast, NoGo-P3 abnormalities were present both in the ADHD-C and RD groups. Impaired early response inhibition mechanisms, indexed by the frontal N200, appear to be limited to ADHD-C. In contrast, deficits in later cognitive control and error monitoring mechanisms, indexed by the NoGo-P3, appear to be present in both conditions.
doi:10.1016/j.bandc.2009.09.007
PMCID: PMC4321824  PMID: 19850394
ADHD; Reading Disorder; Event-Related Potentials; Stop Signal Task; Inhibitory Control; N200; NoGo-P3
2.  [No title available] 
PMCID: PMC3926098  PMID: 24355545
3.  Measuring Attention in the Hemispheres: The Lateralized Attention Network Test (LANT) 
Brain and cognition  2007;66(1):21-31.
The Attention Network Test (ANT) is a brief computerized battery measuring three independent behavioral components of attention: Conflict resolution (ability to overcome distracting stimuli), spatial Orienting (the benefit of valid spatial pre-cues), and Alerting (the benefit of temporal pre-cues). Imaging, clinical, and behavioral evidence demonstrate hemispheric asymmetries in these attentional networks. We constructed a lateralized version of the ANT (LANT), with brief targets flashed in one or the other visual hemifield. We also modified the tests by including invalid spatial cues in order to measure the cost component of Orienting. In a series of experiments, we investigated the efficiency of the attention networks separately in each hemisphere. Participants exhibited significant estimates of all networks measured by the LANT, comparable to the ANT. The three networks were represented in each hemisphere separately and were largely comparable across the two hemispheres. We suggest that the LANT is an informative extension of the original ANT, allowing for measurement of the three attention networks in each hemisphere separately.
doi:10.1016/j.bandc.2007.05.003
PMCID: PMC4283820  PMID: 17590491
attention; conflict; executive; orienting; vigilance; hemispheric specialization; hemispheric independence
4.  Neural processing of intentional biological motion in unaffected siblings of children with autism spectrum disorder: an fMRI study 
Brain and cognition  2013;83(3):10.1016/j.bandc.2013.09.007.
Despite often showing behaviorally typical levels of social cognitive ability, unaffected siblings of children with autism spectrum disorder have been found to show similar functional and morphological deficits within brain regions associated with social processing. They have also been reported to show increased activation to biological motion in these same regions, such as the posterior superior temporal sulcus (pSTS), relative to both children with autism and control children. It has been suggested that this increased activation may represent a compensatory reorganization of these regions as a result of the highly heritable genetic influence of autism. However, the response patterns of unaffected siblings in the domain of action perception are unstudied, and the phenomenon of compensatory activation has not yet been replicated. The present study used functional magnetic resonance imaging to determine the neural responses to intentional biological actions in 22 siblings of children with autism and 22 matched controls. The presented actions were either congruent or incongruent with the actor’s emotional cue. Prior studies reported that typically developing children and adults, but not children with autism, show increased activation to incongruent actions (relative to congruent), within the pSTS and dorsolateral prefrontal cortex. We report that unaffected siblings did not show a compensatory response, or a preference for incongruent over congruent trials, in any brain region. Moreover, interaction analyses revealed a sub-region of the pSTS in which control children showed an incongruency preference to a significantly greater degree than siblings, which suggests a localized deficit in siblings. A sample of children with autism also did not show differential activation in the pSTS, providing further evidence that it is an area of selective disruption in children with autism and siblings. While reduced activation to both conditions was unique to the autism sample, lack of differentiation to incongruent and congruent intentional actions was common to both children with ASD and unaffected siblings.
doi:10.1016/j.bandc.2013.09.007
PMCID: PMC3869032  PMID: 24128657
action perception; autism; endophenotype; superior temporal sulcus; fMRI
5.  Impact of negative affectively charged stimuli and response style on cognitive-control-related neural activation: An ERP study 
Brain and cognition  2013;83(2):10.1016/j.bandc.2013.07.012.
The canonical AX-CPT task measures two forms of cognitive control: sustained goal-oriented control (“proactive” control) and transient changes in cognitive control following unexpected events (“reactive” control). We modified this task by adding negative and neutral International Affective Picture System (IAPS) pictures to assess the effects of negative emotion on these two forms of cognitive control. Proactive and reactive control styles were assessed based on measures of behavior and electrophysiology, including the N2 event-related potential component and source space activation (Low Resolution Tomography [LORETA]). We found slower reaction-times and greater DLPFC activation for negative relative to neutral stimuli. Additionally, we found that a proactive style of responding was related to less prefrontal activation (interpreted to reflect increased efficiency of processing) during actively maintained previously cued information and that a reactive style of responding was related to less prefrontal activation (interpreted to reflect increased efficiency of processing) during just-in-time environmentally triggered information. This pattern of results was evident in relatively neutral contexts, but in the face of negative emotion, these associations were not found, suggesting potential response style-by-emotion interaction effects on prefrontal neural activation
doi:10.1016/j.bandc.2013.07.012
PMCID: PMC3854958  PMID: 24021156
emotion; AX-CPT; reactive control; proactive control; N2; LORETA; prefrontal neural activation
6.  Musical experts recruit action-related neural structures in harmonic anomaly detection: Evidence for embodied cognition in expertise 
Brain and cognition  2013;83(2):10.1016/j.bandc.2013.07.002.
Humans are extremely good at detecting anomalies in sensory input. For example, while listening to a piece of Western-style music, an anomalous key change or an out-of-key pitch is readily apparent, even to the non-musician. In this paper we investigate differences between musical experts and non-experts during musical anomaly detection. Specifically, we analyzed the electroencephalograms (EEG) of five expert cello players and five non-musicians while they listened to excerpts of J.S. Bach’s Prelude from Cello Suite No.1. All subjects were familiar with the piece, though experts also had extensive experience playing the piece. Subjects were told that anomalous musical events (AMEs) could occur at random within the excerpts of the piece and were told to report the number of AMEs after each excerpt. Furthermore, subjects were instructed to remain still while listening to the excerpts and their lack of movement was verified via visual and EEG monitoring. Experts had significantly better behavioral performance (i.e. correctly reporting AME counts) than non-experts, though both groups had mean accuracies greater than 80%. These group differences were also reflected in the EEG correlates of key-change detection post-stimulus, with experts showing more significant, greater magnitude, longer periods of and earlier peaks in condition-discriminating EEG activity than novices. Using the timing of the maximum discriminating neural correlates, we performed source reconstruction and compared significant differences between cellists and non-musicians. We found significant differences that included a slightly right lateralized motor and frontal source distribution. The right lateralized motor activation is consistent with the cortical representation of the left hand – i.e. the hand a cellist would use, while playing, to generate the anomalous key-changes. In general, these results suggest that sensory anomalies detected by experts may in fact be partially a result of an embodied cognition, with a model of the action for generating the anomaly playing a role in its detection.
doi:10.1016/j.bandc.2013.07.002
PMCID: PMC3877747  PMID: 24056235
electroencephalography (EEG); expertise; single-trial analysis; pattern recognition; perceptual decision-making
7.  Aberrant neural mediation of verbal fluency in Autism Spectrum Disorders 
Brain and cognition  2013;83(2):10.1016/j.bandc.2013.08.003.
Objective:
Contrasts of verbal fluency and automatic speech provide an opportunity to evaluate the neural underpinnings of generativity and flexibility in autism spectrum disorders (ASD).
Method:
We used functional magnetic resonance imaging (fMRI) to contrast brain activity in high functioning ASD (n=17, mean verbal IQ=117) and neurotypical (NT; n=20, mean verbal IQ=112) adolescent and young adult males (12-23 years). Participants responded to three word generation conditions: automatic speech (reciting months), category fluency, and letter fluency.
Results:
Our paradigm closely mirrored behavioral fluency tasks by requiring overt, free recall word generation while controlling for differences in verbal output between the groups and systematically increasing the task demand. The ASD group showed reduced neural response compared to the NT participants during fluency tasks in multiple regions of left anterior and posterior cortices, and sub-cortical structures. Six of these regions fell in corticostriatal circuits previously linked to repetitive behaviors (Langen, et al, 2011), and activity in two of them (putamen and thalamus) was negatively correlated with autism repetitive behavior symptoms in the ASD group. In addition, response in left inferior frontal gyrus was differentially modulated in the ASD, relative to the NT, group as a function of task demand.
Conclusions:
These data indicate a specific, atypical brain response in ASD to demanding generativity tasks that may have relevance to repetitive behavior symptoms in ASD as well as to difficulties generating original verbal responses.
doi:10.1016/j.bandc.2013.08.003
PMCID: PMC3829693  PMID: 24056237
autism; verbal fluency; fMRI; executive function; left inferior frontal gyrus
8.  Functional connectivity abnormalities vary by amygdala subdivision and are associated with psychiatric symptoms in unilateral temporal epilepsy 
Brain and cognition  2013;83(2):10.1016/j.bandc.2013.08.001.
The amygdala has been described as a structure affected by mesial temporal lobe epilepsy (MTLE). Indeed, it is suggested that amygdala abnormalities are related to the co-morbid depression and anxiety reported in MTLE. In this context, we investigated the relation between functional connectivity (FC) emerging from this structure in fMRI and depression and anxiety levels reported in MTLE patients. We focused on resting-state BOLD activity and evaluated whether FC differences emerge from each of three amygdala subdivisions (laterobasal, centromedial and superficial) in left and right MTLE groups, compared with healthy controls. Results revealed significant differences between patient groups and controls. Specifically, the left MTLE group showed abnormal FC for the left-sided seeds only. Furthermore, regardless of the seed, we observed more reliable differences between the right MTLE group and controls. Further analysis of these results revealed correlations between these impaired connectivities and psychiatric symptoms in both MTLE groups. Opposite relations, however, were highlighted: the more depressed or anxious the right MTLE patients, the closer their FC values approached controls; whereas the less anxious the left MTLE patients, the closer their FC values were normative. These results highlight how MTLE alter FC emerging from the limbic system. Overall, our data demonstrate that right TLE has a more maladaptive impact on emotion-related networks, in ways specific to the amygdala region, and the emotion symptom involved, than left TLE.
doi:10.1016/j.bandc.2013.08.001
PMCID: PMC3864635  PMID: 24036129
resting state functional connectivity; amygdala; depression; anxiety; epilepsy
9.  Sex differences in the IQ-white matter microstructure relationship: A DTI study 
Brain and Cognition  2014;91:71-78.
Sex differences in the relationship between general intelligence and brain structure are a topic of increasing research interest. Early studies focused mainly on gray and white matter differences using voxel-based morphometry, while more recent studies investigated neural fiber tracts using diffusion tensor imaging (DTI) to analyze the white matter microstructure. In this study we used tract-based spatial statistics (TBSS) on DTI to test how intelligence is associated with brain diffusion indices and to see whether this relationship differs between men and women. 63 Men and women divided into groups of lower and higher intelligence were selected. Whole-brain DTI scans were analyzed using TBSS calculating maps of fractional anisotropy (FA), radial diffusivity (RD), and axial diffusivity (AD). The results reveal that the white matter microstructure differs between individuals as a function of intelligence and sex. In men, higher intelligence was related to higher FA and lower RD in the corpus callosum. In women, in contrast, intelligence was not related to the white matter microstructure. The higher values of FA and lower values of RD suggest that intelligence is associated with higher myelination and/or a higher number of axons particularly in men. This microstructural difference in the corpus callosum may increase cognitive functioning by reducing inter-hemispheric transfer time and thus account for more efficient brain functioning in men.
doi:10.1016/j.bandc.2014.08.006
PMCID: PMC4245721  PMID: 25238623
Corpus callosum; DTI; Intelligence; Sex; TBSS
10.  Differential roles of the frontal and parietal cortices in the control of saccades 
Brain and cognition  2013;83(1):1-9.
Although externally as well as internally-guided eye movements allow us to flexibly explore the visual environment, their differential neural mechanisms remain elusive. A better understanding of these neural mechanisms will help us to understand the control of action and to elucidate the nature of cognitive deficits in certain psychiatric populations (e.g. schizophrenia) that show increased latencies in volitional but not visually-guided saccades. Both the superior precentral sulcus (sPCS) and the intraparietal sulcus (IPS) are implicated in the control of eye movements. However, it remains unknown what differential contributions the two areas make to the programming of visually-guided and internally-guided saccades. In this study we tested the hypotheses that sPCS and IPS distinctly encode internally-guided saccades and visually-guided saccades. We scanned subjects with fMRI while they generated visually-guided and internally-guided delayed saccades. We used multi-voxel pattern analysis to test whether patterns of cue related, preparatory and saccade related activation could be used to predict the direction of the planned eye movement. Results indicate that patterns in the human sPCS predicted internally-guided saccades but not visually-guided saccades in all trial periods and patterns in the IPS predicted internally-guided saccades and visually-guided saccades equally well. The results support the hypothesis that the human sPCS and IPS make distinct contributions to the control of volitional eye movements.
doi:10.1016/j.bandc.2013.06.005
PMCID: PMC3954743  PMID: 23867736
action control; fMRI; MVPA; saccades; volition
11.  Lifespan changes in working memory in fragile X premutation males 
Brain and cognition  2008;69(3):551-558.
Fragile X syndrome is the world’s most common hereditary cause of developmental delay in males and is now well characterized at the biological, brain and cognitive levels. The disorder is caused by the silencing of a single gene on the X chromosome, the FMR1 gene. The premutation (carrier) status, however, is less well documented but has an emerging literature that highlights a more subtle profile of executive cognitive deficiencies that mirror those reported in fully affected males. Rarely, however, has the issue of age-related declines in cognitive performance in premutation males been addressed. In the present study we focus specifically on the cognitive domain of working memory and its sub-components (verbal, spatial and central executive memory) and explore performance across a broad sample of premutation males aged 18–69 years matched on age and IQ to unaffected comparison males. We further tease apart the premutation status into those males with symptoms of the newly identified neurodegenerative disorder, the fragile X-associated tremor/ataxia syndrome (FXTAS) and those males currently symptom-free. Our findings indicate a specific vulnerability in premutation males on tasks that require simultaneous manipulation and storage of new information, so-called executive control of memory. Furthermore, this vulnerability appears to exist regardless of the presence of FXTAS symptoms. Males with FXTAS symptoms demonstrated a more general impairment encompassing phonological working memory in addition to central executive working memory. Among asymptomatic premutation males, we observed the novel finding of a relationship between increased CGG repeat size and impairment to central executive working memory.
doi:10.1016/j.bandc.2008.11.006
PMCID: PMC4158922  PMID: 19114290
Fragile X syndrome; Fragile X tremor and ataxia syndrome; premutation status; working memory; central executive; phonological loop; visual-spatial sketchpad; development; aging
12.  Cognitive Control and Attentional Functions 
Brain and cognition  2013;82(3):301-312.
Cognitive control is essential to flexible, goal-directed behavior under uncertainty, yet its underlying mechanisms are not clearly understood. Because attentional functions are known to allocate mental resources and prioritize the information to be processed by the brain, we propose that the attentional functions of alerting, orienting, and executive control and the interactions among them contribute to cognitive control in the service of uncertainty reduction. To test this hypothesis, we examined the relationship between cognitive control and attentional functions. We used the Majority Function Task (MFT) to manipulate uncertainty in order to evoke cognitive control along with the Revised Attention Network Test (ANT-R) to measure the efficiency and the interactions of attentional functions. A backwards, stepwise regression model revealed that performance on the MFT could be significantly predicted by attentional functions and their interactions as measured by the ANT-R. These results provide preliminary support for our theory that the attentional functions may be involved in the implementation of cognitive control as required to reduce uncertainty, though further investigation is needed.
doi:10.1016/j.bandc.2013.05.004
PMCID: PMC3722267  PMID: 23792472
attention; cognitive control; executive control; uncertainty
13.  The relation between electroencephalogram asymmetry and attention biases to threat at baseline and under stress 
Brain and cognition  2013;82(3):337-343.
Electroencephalogram (EEG) asymmetry in the alpha frequency band has been implicated in emotion processing and broad approach-withdrawal motivation systems. Questions remain regarding the cognitive mechanisms that may help elucidate the observed links between EEG asymmetry and patterns of socioemotional functioning. The current study observed frontal EEG asymmetry patterns at rest and under social threat among young adults (N=45, M=21.1 years). Asymmetries were, in turn, associated with performance on an emotion-face dot-probe attention bias task. Attention biases to threat have been implicated as potential causal mechanisms in anxiety and social withdrawal. Frontal EEG asymmetry at baseline did not predict attention bias patterns to angry or happy faces. However, increases in right frontal alpha asymmetry from baseline to the stressful speech condition were associated with vigilance to angry faces and avoidance of happy faces. The findings may reflect individual differences in the pattern of response (approach or withdrawal) with the introduction of a mild stressor. Comparison analyses with frontal beta asymmetry and parietal alpha asymmetry did not find similar patterns. Thus, the data may reflect the unique role of frontal regions, particularly the dorsolateral prefrontal cortex, in cognitive control and threat detection, coupled with ruminative processes associated with alpha activity.
doi:10.1016/j.bandc.2013.05.009
PMCID: PMC3722560  PMID: 23807238
EEG Asymmetry; Attention Bias; Speech Task
14.  Differential Brain Shrinkage Over Six Months Shows Limited Association with Cognitive Practice 
Brain and cognition  2013;82(2):171-180.
The brain shrinks with age, but the timing of this process and the extent of its malleability are unclear. We measured changes in regional brain volumes in younger (age 20–31) and older (age 65–80) adults twice over a six months period, and examined the association between changes in volume, history of hypertension, and cognitive training. Between two MRI scans, 49 participants underwent intensive practice in three cognitive domains for 100 consecutive days, whereas 23 control group members performed no laboratory cognitive tasks. Regional volumes of seven brain structures were measured manually and adjusted for intracranial volume. We observed significant mean shrinkage in the lateral prefrontal cortex, the hippocampus, the caudate nucleus, and the cerebellum, but no reliable mean change of the prefrontal white matter, orbital-frontal cortex, and the primary visual cortex. Individual differences in change were reliable in all regions. History of hypertension was associated with greater cerebellar shrinkage. The cerebellum was the only region in which significantly reduced shrinkage was apparent in the experimental group after completion of cognitive training. Thus, in healthy adults, differential brain shrinkage can be observed in a narrow time window, vascular risk may aggravate it, and intensive cognitive activity may have a limited effect on it.
doi:10.1016/j.bandc.2013.04.002
PMCID: PMC3667979  PMID: 23665948
aging; cerebellum; cognitive training; plasticity; vascular risk; longitudinal; MRI
15.  Assessing crossmodal matching of abstract auditory and visual stimuli in posterior superior temporal sulcus with MEG 
Brain and cognition  2013;82(2):161-170.
Associating crossmodal auditory and visual stimuli is an important component of perception, with the posterior superior temporal sulcus (pSTS) hypothesized to support this. However, recent evidence has argued that the pSTS serves to associate two stimuli irrespective of modality. To examine the contribution of pSTS to crossmodal recognition, participants (N = 13) learned 12 abstract, non-linguistic pairs of stimuli over 3 weeks. These paired associates comprised four types: auditory–visual (AV), auditory–auditory (AA), visual–auditory (VA), and visual–visual (VV). At week four, participants were scanned using magnetoencephalography (MEG) while performing a correct/incorrect judgment on pairs of items. Using an implementation of synthetic aperture magnetometry that computes real statistics across trials (SAMspm), we directly contrasted crossmodal (AV and VA) with unimodal (AA and VV) pairs from stimulus-onset to 2 s in theta (4–8 Hz), alpha (9–15 Hz), beta (16–30 Hz), and gamma (31–50 Hz) frequencies. We found pSTS showed greater desynchronization in the beta frequency for crossmodal compared with unimodal trials, suggesting greater activity during the crossmodal pairs, which was not influenced by congruency of the paired stimuli. Using a sliding window SAM analysis, we found the timing of this difference began in a window from 250 to 750 ms after stimulus-onset. Further, when we directly contrasted all sub-types of paired associates from stimulus-onset to 2 s, we found that pSTS seemed to respond to dynamic, auditory stimuli, rather than crossmodal stimuli per se. These findings support an early role for pSTS in the processing of dynamic, auditory stimuli, and do not support claims that pSTS is responsible for associating two stimuli irrespective of their modality.
doi:10.1016/j.bandc.2013.03.004
PMCID: PMC3756498  PMID: 23665947
Crossmodal matching; Audio–visual processing; pSTS; Magnetoencephalography
16.  Anterior cingulate cortex and cognitive control: Neuropsychological and electrophysiological findings in two patients with lesions to dorsomedial prefrontal cortex 
Brain and cognition  2012;80(2):237-249.
Whereas neuroimaging studies of healthy subjects have demonstrated an association between the anterior cingulate cortex (ACC) and cognitive control functions, including response monitoring and error detection, lesion studies are sparse and have produced mixed results. Due to largely normal behavioral test results in two patients with medial prefrontal lesions, a hypothesis has been advanced claiming that the ACC is not involved in cognitive operations. In the current study, two comparably rare patients with unilateral lesions to dorsal medial prefrontal cortex (MPFC) encompassing the ACC were assessed with neuropsychological tests as well as Event-Related Potentials in two experimental paradigms known to engage prefrontal cortex (PFC). These included an auditory Novelty Oddball task and a visual Stop-signal task. Both patients performed normally on the Stroop test but showed reduced performance on tests of learning and memory. Moreover, altered attentional control was reflected in a diminished Novelty P3, whereas the posterior P3b to target stimuli was present in both patients. The error-related negativity, which has been hypothesized to be generated in the ACC, was present in both patients, but alterations of inhibitory behavior were observed. Although interpretative caution is generally called for in single case studies, and the fact that the lesions extended outside the ACC, the findings nevertheless suggest a role for MPFC in cognitive control that is not restricted to error monitoring.
doi:10.1016/j.bandc.2012.07.008
PMCID: PMC4067454  PMID: 22935543
Anterior cingulate cortex; Prefrontal cortex; Executive function; Event-related potentials; Cognitive control; Novelty P3
17.  Emotion regulation through execution, observation, and imagery of emotional movements 
Brain and cognition  2013;82(2):219-227.
According to Damasio’s somatic marker hypothesis, emotions are generated by conveying the current state of the body to the brain through interoceptive and proprioceptive afferent input. The resulting brain activation patterns represent unconscious emotions and correlate with subjective feelings. This proposition implies a corollary that the deliberate control of motor behavior could regulate feelings. We tested this possibility, hypothesizing that engaging in movements associated with a certain emotion would enhance that emotion and/or the corresponding valence. Furthermore, because motor imagery and observation are thought to activate the same mirror-neuron network engaged during motor execution, they might also activate the same emotional processing circuits, leading to similar emotional effects. Therefore, we measured the effects of motor execution, motor imagery and observation of whole-body dynamic expressions of emotions (happiness, sadness, fear) on affective state. All three tasks enhanced the corresponding affective state, indicating their potential to regulate emotions.
doi:10.1016/j.bandc.2013.03.001
PMCID: PMC4067484  PMID: 23561915
Body expression of emotion; Nonverbal behavior; Emotion regulation; Embodiment; Simulation; Motor imagery
18.  Language Dysfluencies in Females with the FMR1 Premutation 
Brain and cognition  2013;82(1):84-89.
Recent evidence suggests that there are age-related neurocognitive implications for fragile X premutation carriers, including deficits in executive function, and that such deficits are more common in male than female premutation carriers. The purpose of the current study is to examine one aspect of executive function, language dysfluencies, in a group of 193 women with the premutation, and to contrast them with a comparison group (mothers of children with autism spectrum disorders). Our results demonstrate a linguistic profile in the female premutation carriers characterized by dysfluencies associated with deficits in organization and planning, with a clear impact of age. The comparison group, matched on both age and education level, did not demonstrate the age effect. Our results suggest dysfluencies could be an early indicator of cognitive aging in some female premutation carriers, and could be used to target early intervention.
doi:10.1016/j.bandc.2013.02.009
PMCID: PMC3625470  PMID: 23523717
20.  Hemispheric lateralization of verbal and spatial working memory during adolescence 
Brain and cognition  2013;82(1):58-68.
Adult functional magnetic resonance imaging (fMRI) literature suggests that a left-right hemispheric dissociation may exist between verbal and spatial working memory (WM), respectively. However, investigation of this type has been obscured by incomparable verbal and spatial WM tasks and/or visual inspection at arbitrary thresholds as means to assess lateralization. Furthermore, it is unclear whether this hemispheric lateralization is present during adolescence, a time in which WM skills are improving, and whether there is a developmental association with laterality of brain functioning. This study used comparable verbal and spatial WM n-back tasks during fMRI and a bootstrap analysis approach to calculate lateralization indices (LI) across several thresholds to examine the potential of a left-right WM hemispheric dissociation in healthy adolescents. We found significant left hemispheric lateralization for verbal WM, most notably in the frontal and parietal lobes, as well as right hemisphere lateralization for spatial WM, seen in frontal and temporal cortices. Although no significant relationships were observed between LI and age or LI and performance, significant age-related patterns of brain activity were demonstrated during both verbal and spatial WM. Specifically, increased adolescent age was associated with less activity in the default mode brain network during verbal WM. In contrast, increased adolescent age was associated with greater activity in task-positive posterior parietal cortex during spatial working memory. Our findings highlight the importance of utilizing non-biased statistical methods and comparable tasks for determining patterns of functional lateralization. Our findings also suggest that, while a left-right hemispheric dissociation of verbal and spatial WM is apparent by early adolescence, age-related changes in functional activation during WM are also present.
doi:10.1016/j.bandc.2013.02.007
PMCID: PMC3652620  PMID: 23511846
Adolescence; working memory; fMRI; brain; lateralization
21.  To the Beat of Your Own Drum: Cortical Regularization of Non-Integer Ratio Rhythms Toward Metrical Patterns 
Brain and cognition  2013;81(3):329-336.
Humans perceive a wide range of temporal patterns, including those rhythms that occur in music, speech, and movement; however, there are constraints on the rhythmic patterns that we can represent. Past research has shown that sequences in which sounds occur regularly at non-metrical locations in a repeating beat period (non-integer ratio subdivisions of the beat, e.g. sounds at 430 ms in a 1000 ms beat) are represented less accurately than sequences with metrical relationships, where events occur at even subdivisions of the beat (integer ratios, e.g. sounds at 500 ms in a 1000 ms beat). Why do non-integer ratio rhythms present cognitive challenges? An emerging theory is that non-integer ratio sequences are represented incorrectly, “regularized” in the direction of the nearest metrical pattern, and the present study sought evidence of such perceptual regularization toward integer ratio relationships. Participants listened to metrical and non-metrical rhythmic auditory sequences during electroencephalogram recording, and sounds were pseudorandomly omitted from the stimulus sequence. Cortical responses to these omissions (omission elicited potentials; OEPs) were used to estimate the timing of expectations for omitted sounds in integer ratio and non-integer ratio locations. OEP amplitude and onset latency measures indicated that expectations for non-integer ratio sequences are distorted toward the nearest metrical location in the rhythmic period. These top-down effects demonstrate metrical regularization in a purely perceptual context, and provide support for dynamical accounts of rhythm perception.
doi:10.1016/j.bandc.2013.01.005
PMCID: PMC3683872  PMID: 23434916
Rhythm; ERP; Time perception; Omission elicited potential (OEP); Meter
22.  Pubertal Development and Behavior: Hormonal Activation of Social and Motivational Tendencies 
Brain and cognition  2009;72(1):66-72.
Adolescence is a time of dramatic changes including rapid physical growth, the onset of sexual maturation, the activation of new drives and motivations, and a wide array of social and affective changes and challenges. This review focuses on behavioral changes in this interval and is organized by the claim that a key set of these adolescent changes are part of a more general re-orientation of social behavior. More specifically we hypothesize that pubertal maturation is associated with the activation of social and motivational tendencies, which in turn influence behavior and emotion in adolescence depending upon interactions with social context. We focus on evidence for two examples of these motivational changes: 1) increases in sensation seeking (motivational tendency to want to experience high-intensity, exciting experiences) and 2) stronger natural interest in—and pursuit of—contact with peers and potential romantic partners. We consider how these motivational changes contribute to the broader social re-orientation of adolescence, including exploration of social experiences, the development of skills and knowledge relevant to taking on adult social roles, individuation from family, and the establishment of an individual identity, all of which represent core developmental tasks during this period in the life span (Blakemore, 2008; Dahl & Spear, 2004; Steinberg & Morris, 2000). The paper also emphasizes the importance of investigating and understanding the direct influences of puberty on behavior and disentangling these from the broader set of changes during adolescent development.
doi:10.1016/j.bandc.2009.10.007
PMCID: PMC3955709  PMID: 19942334
23.  Neuronal effects of auditory distraction on visual attention 
Brain and cognition  2013;81(2):263-270.
Selective attention in the presence of distraction is a key aspect of healthy cognition. The underlying neurobiological processes, have not, however, been functionally well characterized. In the present study, we used functional magnetic resonance imaging to determine how ecologically relevant distracting noise affects cortical activity in 27 healthy adults during two versions of the visual sustained attention to response task (SART) that differ in difficulty (and thus attentional load). A significant condition (noise or silence) by task (easy or difficult) interaction was observed in several areas, including dorsolateral prefrontal cortex (DLPFC), fusiform gyrus (FG), posterior cingulate (PCC), and pre-supplementary motor area (PreSMA). Post-hoc analyses of interaction effects revealed deactivation of DLPFC, PCC, and PreSMA during distracting noise under conditions of low attentional load, and activation of FG and PCC during distracting noise under conditions of high attentional load. These results suggest that distracting noise may help alert subjects to task goals and reduce demands on cortical resources during tasks of low difficulty and attentional load. Under conditions of higher load, however, additional cognitive resources may be required in the presence of noise.
doi:10.1016/j.bandc.2012.11.008
PMCID: PMC3596875  PMID: 23291265
Attention; dorsolateral prefrontal cortex; fusiform gyrus; pre-supplementary motor area; posterior cingulate; distraction
24.  Combined effects of inversion and feature removal on N170 responses elicited by faces and car fronts 
Brain and cognition  2013;81(3):321-328.
The face-sensitive N170 is typically enhanced for inverted compared to upright faces. Itier, Alain, Sedore, and McIntosh (2007) recently suggested that this N170 inversion effect is mainly driven by the eye region which becomes salient when the face configuration is disrupted. Here we tested whether similar effects could be observed with non-face objects that are structurally similar to faces in terms of possessing a homogeneous within-class first-order feature configuration. We presented upright and inverted pictures of intact car fronts, car fronts without lights, and isolated lights, in addition to analogous face conditions. Upright cars elicited substantial N170 responses of similar amplitude to those evoked by upright faces. In strong contrast to face conditions however, the car-elicited N170 was mainly driven by the global shape rather than the presence or absence of lights, and was dramatically reduced for isolated lights. Overall, our data confirm a differential influence of the eye region in upright and inverted faces. Results for car fronts do not suggest similar interactive encoding of eye-like features and configuration for non-face objects, even when these objects possess a similar feature configuration as faces.
doi:10.1016/j.bandc.2013.01.002
PMCID: PMC3926862  PMID: 23485023 CAMSID: cams3892
N170; Faces; Cars; Eyes; Inversion; Specificity
25.  A model of amygdala-hippocampal-prefrontal interaction in fear conditioning and extinction in animals 
Brain and cognition  2012;81(1):10.1016/j.bandc.2012.10.005.
Empirical research has shown that the amygdala, hippocampus, and ventromedial prefrontal cortex (vmPFC) are involved in fear conditioning. However, the functional contribution of each brain area and the nature of their interactions are not clearly understood. Here, we extend existing neural network models of the functional roles of the hippocampus in classical conditioning to include interactions with the amygdala and prefrontal cortex. We apply the model to fear conditioning, in which animals learn physiological (e.g. heart rate) and behavioral (e.g. freezing) responses to stimuli that have been paired with a highly aversive event (e.g. electrical shock). The key feature of our model is that learning of these conditioned responses in the central nucleus of the amygdala is modulated by two separate processes, one from basolateral amygdala and signaling a positive prediction error, and one from the vmPFC, via the intercalated cells of the amygdala, and signaling a negative prediction error. In addition, we propose that hippocampal input to both vmPFC and basolateral amygdala is essential for contextual modulation of fear acquisition and extinction. The model is sufficient to account for a body of data from various animal fear conditioning paradigms, including acquisition, extinction, reacquisition, and context specificity effects. Consistent with studies on lesioned animals, our model shows that damage to the vmPFC impairs extinction, while damage to the hippocampus impairs extinction in a different context (e.g., a different conditioning chamber from that used in initial training in animal experiments). We also discuss model limitations and predictions, including the effects of number of training trials on fear conditioning.
doi:10.1016/j.bandc.2012.10.005
PMCID: PMC3808183  PMID: 23164732
fear conditioning; computational model; hippocampus; amygdala; ventromedial prefrontal cortex; extinction

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