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1.  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.
PMCID: PMC3667979  PMID: 23665948
aging; cerebellum; cognitive training; plasticity; vascular risk; longitudinal; MRI
2.  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.
PMCID: PMC3756498  PMID: 23665947
Crossmodal matching; Audio–visual processing; pSTS; Magnetoencephalography
3.  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.
PMCID: PMC4067454  PMID: 22935543
Anterior cingulate cortex; Prefrontal cortex; Executive function; Event-related potentials; Cognitive control; Novelty P3
4.  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.
PMCID: PMC4067484  PMID: 23561915
Body expression of emotion; Nonverbal behavior; Emotion regulation; Embodiment; Simulation; Motor imagery
5.  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.
PMCID: PMC3625470  PMID: 23523717
7.  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.
PMCID: PMC3652620  PMID: 23511846
Adolescence; working memory; fMRI; brain; lateralization
8.  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.
PMCID: PMC3683872  PMID: 23434916
Rhythm; ERP; Time perception; Omission elicited potential (OEP); Meter
9.  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.
PMCID: PMC3955709  PMID: 19942334
10.  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.
PMCID: PMC3596875  PMID: 23291265
Attention; dorsolateral prefrontal cortex; fusiform gyrus; pre-supplementary motor area; posterior cingulate; distraction
11.  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.
PMCID: PMC3926862  PMID: 23485023 CAMSID: cams3892
N170; Faces; Cars; Eyes; Inversion; Specificity
12.  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.
PMCID: PMC3808183  PMID: 23164732
fear conditioning; computational model; hippocampus; amygdala; ventromedial prefrontal cortex; extinction
13.  Cortical Integration of Audio-Visual Information 
Brain and cognition  2010;74(2):10.1016/j.bandc.2010.07.002.
We investigated the neural basis of audio-visual processing in speech and non-speech stimuli. Physically identical auditory stimuli (speech and sinusoidal tones) and visual stimuli (animated circles and ellipses) were used in this fMRI experiment. Relative to unimodal stimuli, each of the multimodal conjunctions showed increased activation in largely non-overlapping areas. The conjunction of Ellipse and Speech, which most resembles naturalistic audiovisual speech, showed higher activation in the right inferior frontal gyrus, fusiform gyri, left posterior superior temporal sulcus, and lateral occipital cortex. The conjunction of Circle and Tone, an arbitrary audio-visual pairing with no speech association, activated middle temporal gyri and lateral occipital cortex. The conjunction of Circle and Speech showed activation in lateral occipital cortex, and the conjunction of Ellipse and Tone did not show increased activation relative to unimodal stimuli. Further analysis revealed that middle temporal regions, although identified as multimodal only in the Circle-Tone condition, were more strongly active to Ellipse-Speech or Circle-Speech, but regions that were identified as multimodal for Ellipse-Speech were always strongest for Ellipse-Speech. Our results suggest that combinations of auditory and visual stimuli may together be processed by different cortical networks, depending on the extent to which speech or non-speech percepts are evoked.
PMCID: PMC3869029  PMID: 20709442
audio-visual; fMRI; multi-modal processing
14.  Efficiency of route selection as a function of adult age 
Brain and cognition  2006;63(3):10.1016/j.bandc.2006.09.006.
Two tasks hypothesized to assess the efficiency of route selection were administered to 328 adults ranging from 18 to 93 years of age. Increased age was associated with slower completion of mazes, even after adjusting for differences in perceptual-motor speed, and with longer and less accurate routes in a task in which participants were asked to visit designated exhibits in a zoo. The route selection measures were correlated with measures hypothesized to represent executive functioning, such as the number of categories in the Wisconsin card sorting test and the number of words generated in a category fluency test. However, most of the age-related influences on the measures from the route selection tasks were shared with age-related effects on established cognitive abilities, which implies that the same mechanisms may account for the relations of age on both sets of variables.
PMCID: PMC3838961  PMID: 17079064
Route selection; Aging; Executive functioning
15.  The amygdala is involved in affective priming effect for fearful faces 
Brain and cognition  2012;80(1):15-22.
The object of this study was to investigate whether the amygdala is involved in affective priming effect after stimuli are encoded unconsciously and consciously. During the encoding phase, each masked face (fearful or neutral) was presented to participants six times for 17 ms each, using a backward masking paradigm. During the retrieval phase, participants made a fearful/neutral judgment for each face. Half of the faces had the same valence as that seen during encoding (congruent condition) and the other half did not (incongruent condition). Participants were divided into unaware and aware groups based on their subjective and objective awareness assessments. The fMRI results showed that during encoding, the amygdala elicited stronger activation for fearful faces than neutral faces but differed in the hemisphere according to the awareness level. During retrieval, the amygdala showed a significant repetition priming effect, with the congruent faces producing less activation than the incongruent faces, especially for fearful faces. These data suggest that the amygdale is important in unconscious retrieving of memories for emotional faces whether they are encoded consciously or unconsciously.
PMCID: PMC3408776  PMID: 22626918
amygdala; unconscious; emotional memory; affective priming
16.  Hemispheric Asymmetries in the Activation and Monitoring of Memory Errors 
Brain and cognition  2012;80(1):7-14.
Previous research on the lateralization of memory errors suggests that the right hemisphere’s tendency to produce more memory errors than the left hemisphere reflects hemispheric differences in semantic activation. However, all prior research that has examined the lateralization of memory errors has used self-paced recognition judgments. Because activation occurs early in memory retrieval, with more time to make a decision, other memory processes, like strategic monitoring processes, may affect memory errors. By manipulating the time subjects were given to make memory decisions, this study separated the influence of automatic memory processes (activation) from strategic memory processes (monitoring) on the production of false memories. The results indicated that when retrieval was fast, the right hemisphere produced more memory errors than the left hemisphere. However, when retrieval was slow, the left hemisphere’s error-proneness increased compared to the fast retrieval condition, while the right hemisphere’s error-proneness remained the same. These results suggest that the right hemisphere’s errors are largely due to activation, while the left hemisphere’s errors are influenced by both activation and monitoring.
PMCID: PMC3408826  PMID: 22626917
false memory; lateralization; semantic activation; source monitoring
17.  Spatial analysis after perinatal stroke: patterns of neglect and exploration in extra-personal space 
Brain and Cognition  2012;79(2):107-116.
This study was conducted to determine whether school-aged children who had experienced a perinatal stroke demonstrate evidence of persistent spatial neglect, and if such neglect was specific to the visual domain or was more generalized. Two studies were carried out. In the first, 38 children with either left hemisphere (LH) or right hemisphere (RH) damage and 50 age-matched controls were given visual cancellation tasks varying in two factors: target stimuli and stimulus array. In the second study, tactile neglect was evaluated in 41 children with LH or RH damage and 72 age-matched controls using a blindfolded manual exploration task. On the visual cancellation task, LH subjects omitted more target stimuli on the right, but also on the left, compared with controls. Children with RH lesions also produced a larger number of omissions on both the left and right sides than controls, but with poorer performance on the left. On the manual exploration task, LH children required significantly longer times to locate the target on both sides of the board than did controls. RH children had significantly prolonged search times on the left side, but not on the right, compared with controls. In both tasks, LH subjects employed unsystematic search strategies more often than both control and RH children. The search strategy of RH children also tended to be erratic when compared to controls, but only in the random arrays of the visual cancellation tasks; structure of the target stimuli improved their organization. These results demonstrate that children with early LH brain damage display bilateral difficulties in visual and tactile modalities; a pattern that is in contrast to that seen in adults with LH damage. This may result from disorganized search strategies or other subtle spatial or attentional deficits. Results of performance of RH children suggests the presence of contralateral neglect in both the visual and tactile modalities; a finding that is similar to the neglect in adult stroke patients with RH lesions. The fact that deficits in spatial attention and organizational strategies are present after very early focal damage to either the LH or the RH broadens our understanding of the differences in functional lateralization between the immature and mature brain. These results also add to evidence for limitations to plasticity in the developing brain. Our findings may have therapeutic and rehabilitative implications for the management of children with early focal brain lesions.
PMCID: PMC3381944  PMID: 22475578
spatial neglect; perinatal stroke; extrapersonal space; tactile neglect; visual neglect
18.  Exploring the role of space-defining objects in constructing and maintaining imagined scenes 
Brain and Cognition  2013;82(1):100-107.
•Some objects evoke a sense of surrounding space (space-defining; SD).•We explored their role in constructing and maintaining imagined scenes.•SD objects enjoyed a privileged role in scene construction and maintenance.•SD objects appear to be an essential building block of scenes.
It has recently been observed that certain objects, when viewed or imagined in isolation, evoke a strong sense of three-dimensional local space surrounding them (space-defining (SD) objects), while others do not (space-ambiguous (SA) objects), and this is associated with engagement of the parahippocampal cortex (PHC). But activation of the PHC is classically associated with scene stimuli. The comparable neural response within PHC to both full scenes and single SD objects, led us to hypothesise that SD objects might play a more critical role in the construction and maintenance of scene representations than SA objects. To test this we used scene construction and deconstruction paradigms, where participants gradually built and maintained scenes using SD, SA and background (wall, floors) items. By examining the order in which each item was added (and later removed) to (and from) a scene, we could estimate the significance of each item type. In two different experiments, participants chose SD over SA objects and background items as the first and most critical item in their constructed scenes and, more generally, selected SD objects earlier than SA objects across the scene construction process. When deconstructing scenes, participants retained significantly more SD objects than SA objects, and the last remaining object across all scenes was highly likely to be an SD object. SD objects therefore enjoy a privileged role in scene construction and maintenance, and appear to be an essential building block of scenes.
PMCID: PMC3661989  PMID: 23548838
Scenes; Objects; Space-defining; Parahippocampal cortex; Scene construction; Memory
19.  Dual-tasking alleviated sleep deprivation disruption in visuomotor tracking: An fMRI study 
Brain and Cognition  2012;78(3):248-256.
Effects of dual-responding on tracking performance after 49-hr of sleep deprivation (SD) were evaluated behaviorally and with functional magnetic resonance imaging (fMRI). Continuous visuomotor tracking was performed simultaneously with an intermittent color-matching visual detection task in which a pair of color-matched stimuli constituted a target and non-matches were non-targets. Tracking error means were binned time-locked to stimulus onset of the detection task in order to observe changes associated with dual-responding by comparing the error during targets and non-targets. Similar comparison was made with fMRI data. Our result showed that despite a significant increase in the overall tracking error post SD, from 20 pixels pre SD to 45 pixels post SD, error decreased to a minimum of about 25 pixels 0 to 6 s after dual-response. Despite an overall reduced activation post SD, greater activation difference between targets and non-targets was found post SD in task-related regions, such as the left cerebellum, the left somatosensory cortex, the left extrastriate cortex, bilateral precuneus, the left middle frontal gyrus, and the left motor cortex. Our results suggest that dual-response helps to alleviate performance impairment usually associated with SD. The duration of the alleviation effect was on the order of seconds after dual-responding.
PMCID: PMC3278524  PMID: 22305924
Visuomotor tracking; Visual detection; Continuous wakefulness; Dual responses; Dual tasks; Prefrontal cortex
Brain and cognition  2012;78(3):257-267.
Anxiety is typically considered an impediment to cognition. We propose anxiety-related impairments in cognitive-behavioral performance are the consequences of enhanced stimulus-driven attention. Accordingly, reflexive, habitual behaviors that rely on stimulus-driven mechanisms should be facilitated in an anxious state, while novel, flexible behaviors that compete with the former should be impaired. To test these predictions, healthy adults (N=17) performed a mixed-saccade task, which pits habitual actions (pro-saccades) against atypical ones (anti-saccades), under anxiety-inducing threat of shock and safe conditions. Whole-head magnetoencephalography (MEG) captured oscillatory responses in the preparatory interval preceding target onset and saccade execution. Results showed threat-induced anxiety differentially impacted response times based on the type of saccade initiated, slowing anti-saccades but facilitating erroneous pro-saccades on anti-saccade trials. MEG source analyses revealed that successful suppression of reflexive pro-saccades and correct initiation of anti-saccades during threat was marked by increased theta power in right ventrolateral prefrontal cortical and midbrain regions (superior colliculi) implicated in stimulus-driven attention. Theta activity may delay stimulus-driven processes to enable generation of an anti-saccade. Moreover, compared to safety, threat reduced beta desynchronization in inferior parietal cortices during anti-saccade preparation but increased it during pro-saccade preparation. Differential effects in inferior parietal cortices indicate a greater readiness to execute anti-saccades during safety and to execute pro-saccades during threat. These findings suggest that, in an anxiety state, reduced cognitive-behavioral flexibility may stem from enhanced stimulus-driven attention, which may serve the adaptive function of optimizing threat detection.
PMCID: PMC3448553  PMID: 22289426
anxiety; beta oscillation; magnetoencephalography; saccade; stimulus-driven attention; theta oscillation
Brain and Cognition  2012;78(2):85-93.
Prior work has related sentence processing to executive deficits in non-demented patients with Parkinson’s disease (PD). We extended this investigation to patients with dementia with Lewy bodies (DLB) and PD dementia (PDD) by examining grammatical and working memory components of sentence processing in the full range of patients with Lewy body spectrum disorder (LBSD). Thirty-three patients with LBSD were given a two-alternative, forced-choice sentence-picture matching task. Sentence type, working memory, and grammatical structure were systematically manipulated in the sentences. We found that patients with PDD and DLB were significantly impaired relative to non-demented PD patients and healthy controls. The deficit in PDD/DLB was most pronounced for sentences lengthened by the strategic placement of an additional prepositional phrase and for sentences with an additional proposition due to a center-embedded clause. However, there was no effect for subject-relative versus object-relative grammatical structure. An MRI voxel-based morphometry analysis in a subset of patients showed significant gray matter thinning in the frontal lobe bilaterally, and this extended to temporal, parietal and occipital regions. A regression analysis related sentence processing difficulty in LBSD to frontal neocortex, including inferiorprefrontal, premotor, and dorsolateral prefrontal regions, as well as right superior temporal cortex. These findings are consistent with the hypothesis that patients with PDD and DLB have difficulty processing sentences with increased working memory demands and that this deficit is related in part to their frontal disease.
PMCID: PMC3265703  PMID: 22218297
Lewy body; Parkinson’s; sentence processing; working memory; MRI; prefrontal
22.  Speech Presentation Cues Moderate Frontal EEG Asymmetry in Socially Withdrawn Young Adults 
Brain and Cognition  2011;78(2):156-162.
Socially withdrawn individuals display solitary behavior across wide contexts with both unfamiliar and familiar peers. This tendency to withdraw may be driven by either past or anticipated negative social encounters. In addition, socially withdrawn individuals often exhibit right frontal electroencephalogram (EEG) asymmetry at baseline and when under stress. In the current study we examined shifts in frontal EEG activity in young adults (N=41) at baseline, as they viewed either an anxiety-provoking or a benign speech video, and as they subsequently prepared for their own speech. Results indicated that right frontal EEG activity increased, relative to the left, only for socially withdrawn participants exposed to the anxious video. These results suggest that contextual affective cues may prime an individual’s response to stress, particularly if they illustrate or substantiate an anticipated negative event.
PMCID: PMC3268053  PMID: 22169714
Social Withdrawal; EEG; Frontal Asymmetry; Presentation Task; Affective Cue
23.  Feedback and Stimulus-Offset Timing Effects in Perceptual Category Learning 
Brain and cognition  2013;81(2):283-293.
We examined how feedback delay and stimulus offset timing affected declarative, rule-based and procedural, information-integration category-learning. We predicted that small feedback delays of several hundred milliseconds would lead to the best information-integration learning based on a highly regarded neurobiological model of learning in the striatum. In Experiment 1 information-integration learning was best with feedback delays of 500ms compared to delays of 0 and 1,000ms. This effect was only obtained if the stimulus offset following the response. Rule-based learning was unaffected by the length of feedback delay, but was better when the stimulus was present throughout feedback than when it offset following the response. In Experiment 2 we found that a large variance (SD=150ms) in feedback delay times around a mean delay of 500ms attenuated information-integration learning, but a small variance (SD=75ms) did not. In Experiment 3 we found that the delay between stimulus offset and feedback is more critical to information-integration learning than the delay between the response and feedback. These results demonstrate the importance of feedback timing in category-learning situations where a declarative, verbalizable rule cannot easily be used as a heuristic to classify members into their correct category.
PMCID: PMC3560315  PMID: 23313835
Category-learning; Information-integration; Feedback delay, Dopamine, Calcium
24.  Attention Training and the Threat Bias: An ERP Study 
Brain and cognition  2011;78(1):63-73.
Anxiety is characterized by exaggerated attention to threat. Several studies suggest that this threat bias plays a causal role in the development and maintenance of anxiety disorders. Furthermore, although the threat bias can be reduced in anxious individuals and induced in non-anxious individual, the attentional mechanisms underlying these changes remain unclear. To address this issue, 49 non-anxious adults were randomly assigned to either attentional training toward or training away from threat using a modified version of the dot probe task. Behavioral measures of attentional biases were also generated pre- and post-training using the dot probe task. Event-related potentials (ERPs) were generated to threat and non-threat face pairs and probes during pre- and post-training assessments. Effects of training on behavioral measures of the threat bias were significant, but only for those participants showing pre-training biases. Attention training also influenced early spatial attention, as measured by post-training P1 amplitudes to cues. Results illustrate the importance of taking pre-training attention biases in non-anxious individuals into account when evaluating the effects of attention training and tracking physiological changes in attention following training.
PMCID: PMC3233611  PMID: 22083026
attention training; dot probe; ERP
25.  An fMRI investigation of posttraumatic flashbacks 
Brain and Cognition  2013;81(1):151-159.
► Flashbacks in PTSD are associated with decreased rather than increased MTL activation. ► Flashbacks in PTSD are associated with activation in the insula and in motor and sensory areas. ► Flashbacks in PTSD may correspond to familiarity rather than recollection responses.
Flashbacks are a defining feature of posttraumatic stress disorder (PTSD), but there have been few studies of their neural basis. We tested predictions from a dual representation model of PTSD that, compared with ordinary episodic memories of the same traumatic event, flashbacks would be associated with activity in dorsal visual stream and related areas rather than in the medial temporal lobe. Participants with PTSD, with depression but not PTSD, and healthy controls were scanned during a recognition task with personally relevant stimuli. The contrast of flashbacks versus ordinary episodic trauma memories in PTSD was associated with increased activation in sensory and motor areas including the insula, precentral gyrus, supplementary motor area, and mid-occipital cortex. The same contrast was associated with decreased activation in the midbrain, parahippocampal gyrus, and precuneus/posterior cingulate cortex. The results were discussed in terms of theories of PTSD and dual-process models of recognition.
PMCID: PMC3549493  PMID: 23207576
PTSD; Flashbacks; Memory; Dual-process; Familiarity

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