Journal of Cognitive Neuroscience

Published by MIT Press

Online ISSN: 1530-8898


Print ISSN: 0898-929X


Figure 1. Study Task. Cues were presented in which the color of the diamond indicated whether a subject could control the length of the heat or not. In the controllable condition subjects were instructed that if they moved the joystick in the direction of the arrow and their response was faster than a threshold (based on their previous response times) they would receive the short, rather than long heat. Subjects were asked to respond on all trials, whether they had control or not. Pictured above are one controllable and one uncontrollable trial. Each subject received eight trials of each condition. Colors were counterbalanced between subjects, and the direction of the arrow varied randomly within condition. ITI = intertrial interval. 
Figure 2. Correlations between pain rating difference (UC À C) and mean parameter estimates from the (UC À C) contrast for clusters in the (A) ventral lateral prefrontal cortex (VLPFC) (anticipatory response), (B) pregenual anterior cingulate cortex (pACC) (pain response), (C) periaqueductal gray (PAG) (pain response), and (D) posterior insula/SII (pain response). All four clusters were significant in the main contrast (UC À C) ( z = 2.3, p = .01, corrected for whole-brain comparisons), VLPFC during anticipation, and pACC, PAG, and insula/SII during pain. Activations shown in the figure survived a higher threshold ( z = 3.09, p = .005), with the regions extracted outlined in blue. 
Table 2 . Regions with a Significant Activation Difference (UC À C) between Anticipation of Uncontrollable and Controllable Pain
Figure 3. Partial Correlation Plots. Correlation between activation difference (UC À C) in the two prefrontal regions and pain rating difference (UC À C) when both are included in a regression model predicting pain rating difference. (A) Correlation of VLPFC (anticipatory response) and pain rating difference with effects of pACC (pain response) partialled out. (B) Correlation of pACC (pain response) and pain rating difference with effects of VLPFC (anticipatory response) partialled out. 
Salomons TV, Johnstone T, Backonja MM, Shackman AJ, Davidson RJ. Individual differences in the effects of perceived controllability on pain perception: critical role of the prefrontal cortex. J Cogn Neurosci 19: 993-1003
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June 2007


530 Reads






The degree to which perceived controllability alters the way a stressor is experienced varies greatly among individuals. We used functional magnetic resonance imaging to examine the neural activation associated with individual differences in the impact of perceived controllability on self-reported pain perception. Subjects with greater activation in response to uncontrollable (UC) rather than controllable (C) pain in the pregenual anterior cingulate cortex (pACC), periaqueductal gray (PAG), and posterior insula/SII reported higher levels of pain during the UC versus C conditions. Conversely, subjects with greater activation in the ventral lateral prefrontal cortex (VLPFC) in anticipation of pain in the UC versus C conditions reported less pain in response to UC versus C pain. Activation in the VLPFC was significantly correlated with the acceptance and denial subscales of the COPE inventory [Carver, C. S., Scheier, M. F., & Weintraub, J. K. Assessing coping strategies: A theoretically based approach. Journal of Personality and Social Psychology, 56, 267-283, 1989], supporting the interpretation that this anticipatory activation was associated with an attempt to cope with the emotional impact of uncontrollable pain. A regression model containing the two prefrontal clusters (VLPFC and pACC) predicted 64% of the variance in pain rating difference, with activation in the two additional regions (PAG and insula/SII) predicting almost no additional variance. In addition to supporting the conclusion that the impact of perceived controllability on pain perception varies highly between individuals, these findings suggest that these effects are primarily top-down, driven by processes in regions of the prefrontal cortex previously associated with cognitive modulation of pain and emotion regulation.

Anderson DE, Bell TA, Awh E. Polymorphisms in the 5-HTTLPR gene mediate storage capacity of visual working memory. J Cogn Neurosci 24: 1069-1076

February 2012


64 Reads

By the request of the authors, the following two research articles will be retracted from the Journal of Cognitive Neuroscience: 1. Anderson, D. E., Ester, E. F., Klee, D., Vogel, E. K., & Awh, E. (2014). Electrophysiological evidence for failures of item individuation in crowded visual displays. Journal of Cognitive Neuroscience, 26(10), 2298– 2309. . 2. Anderson, D. E., Bell, T. A., & Awh, E. (2012). Polymorphisms in the 5-HTTLPR gene mediate storage capacity of visual working memory. Journal of Cognitive Neuroscience, 24(5), 1069–1076. https://dx.doi. org/10.1162/jocn_a_00207 . On August 1, 2015, the Office of Research Integrity (ORI) announced a settlement agreement with David E. Anderson, the Respondent ( case-summary-anderson-david ). On the basis of the Respondent’s admission and an analysis by the University of Oregon, ORI concluded that the Respondent had engaged in research misconduct by falsifying and/or fabricating data in four publications. Those publications were retracted immediately after the release of the ORI findings. Since that time, additional problems have been discovered with Article 1 above. Data points shown in Figure 8 were removed without justification and in contradiction to the analytic approach described in the methods and results. In light of this discovery and of the previous ORI findings, authors Bell and Awh no longer have confidence in the integrity of the data in Article 2. For these reasons, all authors on both articles (including the Respondent) have agreed to the retraction of Articles 1 and 2 above.

Figure 2. Mean proportion correct for short, medium and long time lags. Error bars indicate standard error of the mean.  
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St Jacques P, Rubin DC, LaBar KS, Cabeza R. The short and long of it: neural correlates of temporal-order memory for autobiographical events. J Cogn Neurosci 20: 1327-1341

March 2008


154 Reads

Previous functional neuroimaging studies of temporal-order memory have investigated memory for laboratory stimuli that are causally unrelated and poor in sensory detail. In contrast, the present functional magnetic resonance imaging (fMRI) study investigated temporal-order memory for autobiographical events that were causally interconnected and rich in sensory detail. Participants took photographs at many campus locations over a period of several hours, and the following day they were scanned while making temporal-order judgments to pairs of photographs from different locations. By manipulating the temporal lag between the two locations in each trial, we compared the neural correlates associated with reconstruction processes, which we hypothesized depended on recollection and contribute mainly to short lags, and distance processes, which we hypothesized to depend on familiarity and contribute mainly to longer lags. Consistent with our hypotheses, parametric fMRI analyses linked shorter lags to activations in regions previously associated with recollection (left prefrontal, parahippocampal, precuneus, and visual cortices), and longer lags with regions previously associated with familiarity (right prefrontal cortex). The hemispheric asymmetry in prefrontal cortex activity fits very well with evidence and theories regarding the contributions of the left versus right prefrontal cortex to memory (recollection vs. familiarity processes) and cognition (systematic vs. heuristic processes). In sum, using a novel photo-paradigm, this study provided the first evidence regarding the neural correlates of temporal-order for autobiographical events.

Figure 1. ERPs to all three word-types for the sixteen 14-month-olds over the anterior and posterior regions of the left and right hemispheres. ERPs to known words are shown in the solid lines, to phonemic contrasts in the dotted lines, and to phonetically dissimilar nonsense words in the dashed lines.
Figure 2. ERPs to all three word-types for the seventeen 20-month-olds over the anterior and posterior regions of the left and right hemispheres. ERPs to known words are shown in the solid lines, to phonemic contrasts in the dotted lines, and to phonetically dissimilar nonsense words in the dashed lines.  
Figure 3. For the 14-month-olds, ERP differences are directly compared to known words and nonsense words (top), known words and phonemic contrasts (middle), and phonemic contrasts and phonetically dissimilar nonsense words (bottom). Significant differences in N200–N400 mean amplitudes are shaded and enclosed in the rectangle. Left and temporal and parietal regions are shown on the left and right sides of the figure, respectively.  
Figure 4. For the 20-month-olds, ERP differences are directly compared to known words and nonsense words (top), known words and phonemic contrasts (middle), and phonemic contrasts and phonetically dissimilar nonsense words (bottom). Significant differences in N200–N400 mean amplitudes are shaded and enclosed in the rectangle. Left and temporal and parietal regions are shown on the left and right sides of the figure, respectively.  
Figure 5. N200–N400 mean areas averaged across temporal and parietal regions of the left and right hemispheres to the three word-types for the 14-and 20-month-olds.  
Language Experience and the Organization of Brain Activity to Phonetically Similar Words: ERP Evidence from 14- and 20-Month-Olds

November 2004


432 Reads

The ability to discriminate phonetically similar speech sounds is evident quite early in development. However, inexperienced word learners do not always use this information in processing word meanings [Stager & Werker (1997). Nature, 388, 381-382]. The present study used event-related potentials (ERPs) to examine developmental changes from 14 to 20 months in brain activity important in processing phonetic detail in the context of meaningful words. ERPs were compared to three types of words: words whose meanings were known by the child (e.g., ''bear''), nonsense words that differed by an initial phoneme (e.g., ''gare''), and nonsense words that differed from the known words by more than one phoneme (e.g., ''kobe''). These results supported the behavioral findings suggesting that inexperienced word learners do not use information about phonetic detail when processing word meanings. For the 14-month-olds, ERPs to known words (e.g., ''bear'') differed from ERPs to phonetically dissimilar nonsense words (e.g., ''kobe''), but did not differ from ERPs to phonetically similar nonsense words (e.g., ''gare''), suggesting that known words and similar mispronunciations were processed as the same word. In contrast, for experienced word learners (i. e., 20-month-olds), ERPs to known words (e.g., ''bear'') differed from those to both types of nonsense words (''gare'' and ''kobe''). Changes in the lateral distribution of ERP differences to known and unknown (nonce) words between 14 and 20 months replicated previous findings. The findings suggested that vocabulary development is an important factor in the organization of neural systems linked to processing phonetic detail within the context of word comprehension.

Ehrlichman RS, Maxwell CR, Majumdar S, Siegel SJ. Deviance-elicited changes in event-related potentials are attenuated by ketamine in mice. J Cogn Neurosci 20: 1403-1414

September 2008


216 Reads

People with schizophrenia exhibit reduced ability to detect change in the auditory environment, which has been linked to abnormalities in N-methyl-D-aspartate (NMDA) receptor-mediated glutamate neurotransmission. This ability to detect changes in stimulus qualities can be measured with electroencephalography using auditory event-related potentials (ERPs). For example, reductions in the N100 and mismatch negativity (MMN), in response to pitch deviance, have been proposed as endophenotypes of schizophrenia. This study examines a novel rodent model of impaired pitch deviance detection in mice using the NMDA receptor antagonist ketamine. ERPs were recorded from unanesthetized mice during a pitch deviance paradigm prior to and following ketamine administration. First, N40 amplitude was evaluated using stimuli between 4 and 10 kHz to assess the amplitude of responses across the frequency range used. The amplitude and latency of the N40 were analyzed following standard (7 kHz) and deviant (5-9 kHz) stimuli. Additionally, we examined which portions of the ERP are selectively altered by pitch deviance to define possible regions for the mouse MMN. Mice displayed increased N40 amplitude that was followed by a later negative component between 50 and 75 msec in response to deviant stimuli. Both the increased N40 and the late N40 negativity were attenuated by ketamine. Ketamine increased N40 latency for both standard and deviant stimuli alike. The mouse N40 and a subsequent temporal region have deviance response properties similar to the human N100 and, possibly, MMN. Deviance responses were abolished by ketamine, suggesting that ketamine-induced changes in mice mimic deviance detection deficits in schizophrenia.

Table 1 . Standardized Factor Loadings for Executive Function Tests Executive Function Factor 
Table 2 . Unstandardized Means for All Cognitive Tasks 
Figure 2. Individuals with the Val/ Val genotype declined in executive functioning over a 5-year period. 
Catechol O-Methyltransferase Val 158 Met Polymorphism is Associated with Cognitive Performance in Nondemented Adults

August 2005


271 Reads

The catechol O-methyltransferase (COMT) gene is essential in the metabolic degradation of dopamine in the prefrontal cortex. In the present study, we examined the effect of a Val158Met polymorphism in the COMT gene on individual differences and changes in cognition (executive functions and visuospatial ability) in adulthood and old age. The participants were 292 nondemented men (initially aged 35-85 years) from a random sample of the population (i.e., the Betula study) tested at two occasions with a 5-year interval. Confirmatory factor analyses were used to test the underlying structure of three indicators of executive functions (verbal fluency, working memory, and Tower of Hanoi). Associations between COMT, age, executive functioning, and visuospatial (block design) tasks were examined using repeated-measures analyses of variance. Carriers of the Val allele (with higher enzyme activity) compared with carriers of the Met/Met genotype (with low enzyme activity) performed worse on executive functioning and visuospatial tasks. Individuals with the Val/Val genotype declined in executive functioning over the 5-year period, whereas carriers of the Met allele remained stable in performance. An Age x COMT interaction for visuospatial ability located the effect for middle-aged men only. This COMT polymorphism is a plausible candidate gene for executive functioning and fluid intelligence in nondemented middle-aged and older adults.

Individual Differences in Cerebral Blood Flow in Area 17 Predict the Time to Evaluate Visualized Letters

January 1996


87 Reads

Sixteen subjects closed their eyes and visualized uppercase letters of the alphabet at two sizes, as small as possible or as large as possible while remaining "visible." Subjects evaluated a shape characteristic of each letter (e.g., whether it has any curved lines), and responded as quickly as possible. Cerebral blood flow was normalized to the same value for each subject, and relative blood flow was computed for a set of regions of interest. The mean response time for each subject in the task was regressed onto the blood flow values. Blood flow in area 17 was negatively correlated with response time (r = -0.65), as was blood flow in area 19 (r = -0.66), whereas blood flow in the inferior parietal lobe was positively correlated with response time (r = 0.54). The first two effects persisted even when variance due to the other correlations was removed. These findings suggest that individual differences in the activation of specific brain loci are directly related to performance of tasks that rely on processing in those loci.

Barraclough, N. E., Xiao, D., Baker, C. I., Oram, M. W. & Perrett, D. I. Integration of visual and auditory information by superior temporal sulcus neurons responsive to the sight of actions. J. Cogn. Neurosci. 17, 377-391

April 2005


836 Reads

Processing of complex visual stimuli comprising facial movements, hand actions, and body movements is known to occur in the superior temporal sulcus (STS) of humans and nonhuman primates. The STS is also thought to play a role in the integration of multimodal sensory input. We investigated whether STS neurons coding the sight of actions also integrated the sound of those actions. For 23% of neurons responsive to the sight of an action, the sound of that action significantly modulated the visual response. The sound of the action increased or decreased the visually evoked response for an equal number of neurons. In the neurons whose visual response was increased by the addition of sound (but not those neurons whose responses were decreased), the audiovisual integration was dependent upon the sound of the action matching the sight of the action. These results suggest that neurons in the STS form multisensory representations of observed actions.

Hemisphere-specific Episodic Memory Networks in the Human Brain: A Correlation Study between Intracarotid Amobarbital Test and [ 18 F]FDG-PET

July 2008


46 Reads

The purpose of the present study was to explore the brain regions involved in human episodic memory by correlating unilateral memory performance estimated by the intracarotid amobarbital test (IAT) and interictal cerebral metabolism measured by [(18)F]fluorodeoxyglucose positron emission tomography ([(18)F]FDG-PET). Using this method, regional alterations of cerebral metabolism associated with epilepsy pathophysiology are used to predict hemisphere-specific episodic memory function, hence, investigate the differential distribution of memory in each hemisphere. Sixty-two patients with unilateral temporal lobe epilepsy (35 left and 27 right) were studied using [(18)F]FDG-PET with complementary voxel-based statistical parametric mapping (SPM) and region-of-interest (ROI) methods of analysis. Positive regression was analyzed in SPM with a series of different thresholds (p = .001, .01 or .05) with a correction to 100 voxels. IAT memory performance in which left hemisphere was tested by right-sided injection of amobarbital correlated with [(18)F]FDG uptake in left lateral and medial temporal regions, and in the left ventrolateral frontal cortex. Right IAT memory performance correlated with [(18)F]FDG uptake in the right inferior parietal lobule, right dorsolateral frontal cortex, right precentral gyrus, and caudal portion of the right anterior cingulate cortex. ROI analysis corroborated these results. Analyses carried out separately in patients with left (n = 50) and nonleft (n = 12) dominance for language showed that in the nonleft dominant group, right IAT scores correlated with right fronto-temporal regions, whereas left total memory scores correlated with left lateral and medial temporal regions. The findings indicate that (i) episodic memory is subserved by more widespread cortical regions beyond the core mesiotemporal lobe memory structures; (ii) there are different networks functional in the two hemispheres; and (iii) areas involved in memory may be different between patients with left and nonleft dominance for language, particularly in the right hemisphere.

Chikazoe J, Konishi S, Asari T, Jimura K, Miyashita Y. Activation of right inferior frontal gyrus during response inhibition across response modalities. J Cogn Neurosci 19: 69-80

February 2007


68 Reads

The go/no-go task, which effectively taps the ability to inhibit prepotent response tendency, has consistently activated the lateral prefrontal cortex, particularly the right inferior frontal gyrus (rIFG). On the other hand, rIFG activation has rarely been reported in the antisaccade task, seemingly an oculomotor version of the manual go/no-go task. One possible explanation for the variable IFG activation is the modality difference of the two tasks: The go/no-go task is performed manually, whereas the antisaccade task is performed in the oculomotor modality. Another explanation is that these two tasks have different task structures that require different cognitive processes: The traditional antisaccade task requires (i) configuration of a preparatory set prior to antisaccade execution and (ii) response inhibition at the time of antisaccade execution, whereas the go/no-go task requires heightened response inhibition under a minimal preparatory set. To test these possibilities, the traditional antisaccade task was modified in the present functional magnetic resonance imaging study such that it required heightened response inhibition at the time of antisaccade execution under a minimal preparatory set. Prominent activation related to response inhibition was observed in multiple frontoparietal regions, including the rIFG. Moreover, meta-analyses revealed that the rIFG activation in the present study was observed in the go/no-go tasks but not in the traditional antisaccade task, indicating that the rIFG activation was sensitive to the task structure difference, but not to the response modality difference. These results suggest that the rIFG is part of a network active during response inhibition across different response modalities.

N400-like Semantic Incongruity Effect in 19-Month-Olds: Processing Known Words in Picture Contexts

November 2004


243 Reads

To understand mechanisms of early language acquisition, it is important to know whether the child's brain acts in an adult-like manner when processing words in meaningful contexts. The N400, a negative component in the eventrelated potential (ERP) of adults, is a sensitive index of semantic processing reflecting neural mechanisms of semantic integration into context. In the present study, we investigated whether the mechanisms indexed by the N400 are already working during early language acquisition. While 19-month-olds were looking at sequentially presented pictures, they were acoustically presented with words that were either congruous or incongruous to the picture content. The ERP averaged across the group of 55 children revealed an N400-like semantic incongruity effect in addition to an early phonological-lexical priming effect. The results suggest that both lexical expectations facilitating early phonological processing and mechanisms of semantic priming facilitating integration into semantic context are already present in 19-month-olds. The child's specific comprehension abilities are reflected in strength, latency, and hemispheric differences of the semantic incongruity effect. Spatio-temporal differences in that effect, thus, indicate changes in the organization of brain activity correlated with the child's behavioral development.

Face Familiarity Decisions Take 200 msec in the Human Brain: Electrophysiological Evidence from a Go/No-go Speeded Task

August 2013


572 Reads

Recognizing a familiar face rapidly is a fundamental human brain function. Here we used scalp EEG to determine the minimal time needed to classify a face as personally familiar or unfamiliar. Go (familiar) and no-go (unfamiliar) responses elicited clear differential waveforms from 210 msec onward, this difference being first observed at right occipito-temporal electrode sites. Similar but delayed (by about 40 msec) responses were observed when go response were required to the unfamiliar rather than familiar faces, in a second group of participants. In both groups, a small increase of amplitude was also observed on the right hemisphere N170 face-sensitive component for familiar faces. However, unlike the post-200 msec differential go/no-go effect, this effect was unrelated to behavior and disappeared with repetition of unfamiliar faces. These observations indicate that accumulation of evidence within the first 200 msec poststimulus onset is sufficient for the human brain to decide whether a person is familiar based on his or her face, a time frame that puts strong constraints on the time course of face processing.

Figure 2. Subjects’ ratings of perceived contradictions in statements by Bush, Kerry, and neutral figures (higher ratings indicate greater perceived contradictions). Democrats and Republicans reasoned to distinctly different conclusions about their preferred candidates, with mirror-image responses: Democrats readily identified the contradictions in Bush’s statements but not Kerry’s, whereas Republicans readily identified the contradictions in Kerry’s statements but not Bush’s. 2 As can be seen from the 
Figure 3. Three orthogonal views (axial, sagittal, coronal; at x = 0, y = 50, z = 6) of the areas of activation that differed when subjects were confronted with contradictory (threatening) information regarding their own party’s candidate versus a neutral target person. ACC = anterior cingulate; mPFC = medial prefrontal cortex; pCING = posterior cingulate; PCU = precuneus; vmPFC = ventromedial prefrontal cortex. 
Neural Bases of Motivated Reasoning: An fMRI Study of Emotional Constraints on Partisan Political Judgment in the 2004 U.S. Presidential Election

December 2006


4,886 Reads

Research on political judgment and decision-making has converged with decades of research in clinical and social psychology suggesting the ubiquity of emotion-biased motivated reasoning. Motivated reasoning is a form of implicit emotion regulation in which the brain converges on judgments that minimize negative and maximize positive affect states associated with threat to or attainment of motives. To what extent motivated reasoning engages neural circuits involved in "cold" reasoning and conscious emotion regulation (e.g., suppression) is, however, unknown. We used functional neuroimaging to study the neural responses of 30 committed partisans during the U.S. Presidential election of 2004. We presented subjects with reasoning tasks involving judgments about information threatening to their own candidate, the opposing candidate, or neutral control targets. Motivated reasoning was associated with activations of the ventromedial prefrontal cortex, anterior cingulate cortex, posterior cingulate cortex, insular cortex, and lateral orbital cortex. As predicted, motivated reasoning was not associated with neural activity in regions previously linked to cold reasoning tasks and conscious (explicit) emotion regulation. The findings provide the first neuroimaging evidence for phenomena variously described as motivated reasoning, implicit emotion regulation, and psychological defense. They suggest that motivated reasoning is qualitatively distinct from reasoning when people do not have a strong emotional stake in the conclusions reached.

On Phrase Structure and Brain Responses: A Comment on Bahlmann, Gunter, and Friederici (2006)

November 2007


25 Reads

Bahlmann et al. (2006) reported an experiment on event-related brain potentials of sequences of syllables obeying two rules, one defined by A(n)B(n) and the other by (AB)(n), where the As and Bs are different classes of syllables. They interpreted their findings on the assumption that A(n)B(n) are parsed according a center-embedded phrase-structure grammar. In fact, such sequences are much more likely to be parsed in terms of the repetition of element types, without reference to phrase structure. This raises a general issue about attempting to study syntactic processing independently of semantics.

“Do You See Yonder Cloud?”—On Priming Concepts, A New Test, and a Familiar Outcome. Reply to Lucas et al.: “Familiarity or Conceptual Priming? Good Question! Comment on Stenberg, Hellman, Johansson, and Rosén (2009)”

June 2009


54 Reads

Lucas, Voss, and Paller sympathize with our intentions but disagree with our findings. They argue that a relation between frequency and conceptual priming may have been obscured by methodological details in our second experiment, therefore failing to complete a bridge between conceptual priming and FN400 with name frequency as the mediator. However, renewed inspections of our experiment and a new additional experiment, designed to meet the objections, fail to find any role for name frequency in conceptual priming and therefore re-establish the dissociation of priming and the FN400. On closer inspection, our differing views seem to derive from different interpretations of the term "concept."

Familiarity or Conceptual Priming? Good Question! Comment on Stenberg, Hellman, Johansson, and Rosén (2009)

May 2009


74 Reads

Stenberg et al. argued that FN400 brain potentials index familiarity rather than conceptual priming. Their data from a test of name recognition showed that both familiarity and FN400s were influenced by frequency but not fame, whereas separate behavioral measures of priming were influenced by fame but not frequency. However, this apparent dissociation was gravely weakened by confounds in task demands and inadequate behavioral measures of priming. Although Stenberg et al. failed to provide evidence suitable for disentangling neural correlates of familiarity from those of conceptual priming, an analysis of their report can be used to highlight difficulties that remain to be surmounted to understand recognition and the neural events that signal distinct memory functions engaged during recognition.

Integrating the Behavioral and Neural Dynamics of Response Selection in a Dual-task Paradigm: A Dynamic Neural Field Model of Dux et al. (2009)

October 2013


224 Reads

People are typically slower when executing two tasks than when only performing a single task. These dual-task costs are initially robust but are reduced with practice. Dux et al. (2009) explored the neural basis of dual-task costs and learning using fMRI. Inferior frontal junction (IFJ) showed a larger hemodynamic response on dual-task trials compared with single-task trial early in learning. As dual-task costs were eliminated, dual-task hemodynamics in IFJ reduced to single-task levels. Dux and colleagues concluded that the reduction of dual-task costs is accomplished through increased efficiency of information processing in IFJ. We present a dynamic field theory of response selection that addresses two questions regarding these results. First, what mechanism leads to the reduction of dual-task costs and associated changes in hemodynamics? We show that a simple Hebbian learning mechanism is able to capture the quantitative details of learning at both the behavioral and neural levels. Second, is efficiency isolated to cognitive control areas such as IFJ, or is it also evident in sensory motor areas? To investigate this, we restrict Hebbian learning to different parts of the neural model. None of the restricted learning models showed the same reductions in dual-task costs as the unrestricted learning model, suggesting that efficiency is distributed across cognitive control and sensory motor processing systems.

Methodological Pitfalls in the “Objective” Approach to Consciousness: Comments on Busch et al. (2009)

November 2009


106 Reads

What is the observer's conscious experience in a change blindness task? Overgaard, Jensen, and Sandberg argue that subjective measures are required for any conclusions about conscious experience. We will lay out how the choice of subjective or objective ...

Some Regions within Broca's Area Do Respond More Strongly to Sentences than to Linguistically Degraded Stimuli: A Comment on Rogalsky and Hickok (2011)

May 2011


43 Reads

On the basis of their review of the literature, Rogalsky and Hickok [Rogalsky, C., & Hickok, G. The role of Broca's area in sentence comprehension. Journal of Cognitive Neuroscience, 23, 1664-1680, 2011] conclude that there is currently no strong evidence for the existence of "sentence-specific processing regions within Broca's area" (p. 1). Their argument is based, in part, on the observation that many previous studies have failed to detect an effect in the left inferior frontal regions for contrasts between sentences and linguistically degraded control conditions (e.g., lists of unconnected words, lists of nonwords, or acoustically degraded sentence stimuli). Our data largely replicate this lack of activation in inferior frontal regions when traditional random-effects group analyses are conducted but crucially show robust activations in the same data for the same contrasts in almost every subject individually. Thus, it is the use of group analyses in studies of language processing, not the idea that sentences robustly activate frontal regions, that needs to be reconsidered. This reconsideration has important methodological and theoretical implications.

Figure 1. Left: Selected sagittal slices where high FA is associated with high delay AUC. Right: Selected sagittal slices where low MD is associated with high delay AUC. Signed numbers are x-coordinates in standard MNI space. 
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White Matter Integrity Predicts Delay Discounting Behavior in 9- to 23-Year-Olds: A Diffusion Tensor Imaging Study

October 2008


121 Reads

Healthy participants (n = 79), ages 9-23, completed a delay discounting task assessing the extent to which the value of a monetary reward declines as the delay to its receipt increases. Diffusion tensor imaging (DTI) was used to evaluate how individual differences in delay discounting relate to variation in fractional anisotropy (FA) and mean diffusivity (MD) within whole-brain white matter using voxel-based regressions. Given that rapid prefrontal lobe development is occurring during this age range and that functional imaging studies have implicated the prefrontal cortex in discounting behavior, we hypothesized that differences in FA and MD would be associated with alterations in the discounting rate. The analyses revealed a number of clusters where less impulsive performance on the delay discounting task was associated with higher FA and lower MD. The clusters were located primarily in bilateral frontal and temporal lobes and were localized within white matter tracts, including portions of the inferior and superior longitudinal fasciculi, anterior thalamic radiation, uncinate fasciculus, inferior fronto-occipital fasciculus, corticospinal tract, and splenium of the corpus callosum. FA increased and MD decreased with age in the majority of these regions. Some, but not all, of the discounting/DTI associations remained significant after controlling for age. Findings are discussed in terms of both developmental and age-independent effects of white matter organization on discounting behavior.

Imaging cognition II: An empirical review of 275 PET and fMRI studies

February 2000


10,261 Reads

Positron emission tomography (PET) and functional magnetic resonance imaging (fMRI) have been extensively used to explore the functional neuroanatomy of cognitive functions. Here we review 275 PET and fMRI studies of attention (sustained, selective, Stroop, orientation, divided), perception (object, face, space/motion, smell), imagery (object, space/motion), language (written/spoken word recognition, spoken/no spoken response), working memory (verbal/numeric, object, spatial, problem solving), semantic memory retrieval (categorization, generation), episodic memory encoding (verbal, object, spatial), episodic memory retrieval (verbal, nonverbal, success, effort, mode, context), priming (perceptual, conceptual), and procedural memory (conditioning, motor, and nonmotor skill learning). To identify consistent activation patterns associated with these cognitive operations, data from 412 contrasts were summarized at the level of cortical Brodmann's areas, insula, thalamus, medial-temporal lobe (including hippocampus), basal ganglia, and cerebellum. For perception and imagery, activation patterns included primary and secondary regions in the dorsal and ventral pathways. For attention and working memory, activations were usually found in prefrontal and parietal regions. For language and semantic memory retrieval, typical regions included left prefrontal and temporal regions. For episodic memory encoding, consistently activated regions included left prefrontal and medial temporal regions. For episodic memory retrieval, activation patterns included prefrontal, medial temporal, and posterior midline regions. For priming, deactivations in prefrontal (conceptual) or extrastriate (perceptual) regions were consistently seen. For procedural memory, activations were found in motor as well as in non-motor brain areas. Analysis of regional activations across cognitive domains suggested that several brain regions, including the cerebellum, are engaged by a variety of cognitive challenges. These observations are discussed in relation to functional specialization as well as functional integration.

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The Brain Network for Deductive Reasoning: A Quantitative Meta-analysis of 28 Neuroimaging Studies

May 2011


629 Reads

Over the course of the past decade, contradictory claims have been made regarding the neural bases of deductive reasoning. Researchers have been puzzled by apparent inconsistencies in the literature. Some have even questioned the effectiveness of the methodology used to study the neural bases of deductive reasoning. However, the idea that neuroimaging findings are inconsistent is not based on any quantitative evidence. Here, we report the results of a quantitative meta-analysis of 28 neuroimaging studies of deductive reasoning published between 1997 and 2010, combining 382 participants. Consistent areas of activations across studies were identified using the multilevel kernel density analysis method. We found that results from neuroimaging studies are more consistent than what has been previously assumed. Overall, studies consistently report activations in specific regions of a left fronto-parietal system, as well as in the left BG. This brain system can be decomposed into three subsystems that are specific to particular types of deductive arguments: relational, categorical, and propositional. These dissociations explain inconstancies in the literature. However, they are incompatible with the notion that deductive reasoning is supported by a single cognitive system relying either on visuospatial or rule-based mechanisms. Our findings provide critical insight into the cognitive organization of deductive reasoning and need to be accounted for by cognitive theories.

Contextual Priming in Grapheme–Color Synesthetes and Yoked Controls: 400 msec in the Life of a Synesthete

March 2010


341 Reads

Grapheme-color synesthesia is a heritable trait where graphemes ("2") elicit the concurrent perception of specific colors (red). Researchers have questioned whether synesthetic experiences are meaningful or simply arbitrary associations and whether these associations are perceptual or conceptual. To address these fundamental questions, ERPs were recorded as 12 synesthetes read statements such as "The Coca-Cola logo is white and 2," in which the final grapheme induced a color that was either contextually congruous (red) or incongruous ("...white and 7," for a synesthetes who experienced 7 as green). Grapheme congruity was found to modulate the amplitude of the N1, P2, N300, and N400 components in synesthetes, suggesting that synesthesia impacts perceptual as well as conceptual aspects of processing. To evaluate whether observed ERP effects required the experience of colored graphemes versus knowledge of grapheme-color pairings, we ran three separate groups of controls on a similar task. Controls trained to a synesthete's associations elicited N400 modulation, indicating that knowledge of grapheme-color mappings was sufficient to modulate this component. Controls trained to synesthetic associations and given explicit visualization instructions elicited both N300 and N400 modulations. Lastly, untrained controls who viewed physically colored graphemes ("2" printed in red) elicited N1 and N400 modulations. The N1 grapheme congruity effect began earlier in synesthetes than colored grapheme controls but had similar scalp topography. Data suggest that, in synesthetes, achromatic graphemes engage similar visual processing networks as colored graphemes in nonsynesthetes and are in keeping with models of synesthesia that posit early feed-forward connections between form and color processing areas in extrastriate cortex. The P2 modulation was unique to the synesthetes and may reflect neural activity that underlies the conscious experience of the synesthetic induction.

Lane RD, Reiman EM, Axelrod B, Yun LS, Holmes A, Schwartz GE. Neural correlates of levels of emotional awareness. Evidence of an interaction between emotion and attention in the anterior cingulate cortex. J Cogn Neurosci 10: 525-535

July 1998


215 Reads

Recent functional imaging studies have begun to identify the neural correlates of emotion in healthy volunteers. However, studies to date have not differentially addressed the brain areas associated with the perception, experience, or expression of emotion during emotional arousal. To explore the neural correlates of emotional experience, we used positron emission tomography (PET) and 15-water to measure cerebral blood flow (CBF) in 12 healthy women during film- and recall-induced emotion and correlated CBF changes attributable to emotion with subjects' scores on the Levels of Emotional Awareness Scale (LEAS), a measure of individual differences in the capacity to experience emotion in a differentiated and complex way. A conjunction analysis revealed that the correlations between LEAS and CBF during film- and recall-induced emotion overlapped significantly (z = 3.74, p < 0. 001) in Brodmann's area 24 of the anterior cingulate cortex (ACC). This finding suggests that individual differences in the ability to accurately detect emotional signals interoceptively or exteroceptively may at least in part be a function of the degree to which the ACC participates in the experiential processing and response to emotion cues. To the extent that this finding is consistent with the functions of the ACC involving attention and response selection, it suggests that this neural correlate of conscious emotional experience is not exclusive to emotion.

Mostofsky SH, Simmonds DJ. Response inhibition and response selection: two sides of the same coin. J Cogn Neurosci 20: 751-761

June 2008


273 Reads

Response inhibition refers to the suppression of actions that are inappropriate in a given context and that interfere with goal-driven behavior. Studies using a range of methodological approaches have implicated executive control processes mediated by frontal-subcortical circuits as being critical to response inhibition; however, localization within the frontal lobe has been inconsistent. In this review, we present evidence from behavioral, lesion, neuroimaging, electrophysiology, and neurological population studies. The findings lay the foundation for a construct in which response inhibition is akin to response selection, such that pre-SMA circuits are critical to selection of appropriate behavior, including both selecting to engage appropriate motor responses and selecting to withhold (inhibit) inappropriate motor responses. Recruitment of additional prefrontal and posterior cortical circuits, necessary to guide response selection, varies depending on the cognitive and behavioral demands of the task.

High-resolution 7T fMRI of Human Hippocampal Subfields during Associative Learning

December 2014


256 Reads

Examining the function of individual human hippocampal subfields remains challenging because of their small sizes and convoluted structures. Previous human fMRI studies at 3 T have successfully detected differences in activation between hippocampal cornu ammonis (CA) field CA1, combined CA2, CA3, and dentate gyrus (DG) region (CA23DG), and the subiculum during associative memory tasks. In this study, we investigated hippocampal subfield activity in healthy participants using an associative memory paradigm during high-resolution fMRI scanning at 7 T. We were able to localize fMRI activity to anterior CA2 and CA3 during learning and to the posterior CA2 field, the CA1, and the posterior subiculum during retrieval of novel associations. These results provide insight into more specific human hippocampal subfield functions underlying learning and memory and a unique opportunity for future investigations of hippocampal subregion function in healthy individuals as well as those suffering from neurodegenerative diseases.

Figure 1. Assessment of CC efficiency using the "Animal Stroop task." (Left) A child performing the "Animal Stroop task" in the classroom. (Right) Example of "conflict" and "no-conflict" stimuli used in
Figure 2. Morphological patterns of ACC. The two ACC sulcal patterns: "single" type, with only the cingulate sulcus, and "double parallel" type, with an additional PCS. ACC sulci (blue) are represented on the cortical surface (gray/white interface).
Figure 3. Interindividual variability of ACC sulcal pattern. Superimposition of the 3-D mesh-based reconstructions of the cingulate sulcus (turquoise) and PCS (blue) for all children included in the study. Sulci were represented on the cortical surface (gray/white interface). The reconstructions of the sulci of each child were linearly aligned in a common referential (MNI space) for visualization purpose. 
The Shape of the AACC Contributes to Cognitive Control Efficiency in Preschoolers.

August 2013


844 Reads

Cognitive success at school and later in life is supported by executive functions including cognitive control (CC). The pFC plays a major role in CC, particularly the dorsal part of ACC or midcingulate cortex. Genes, environment (including school curricula), and neuroplasticity affect CC. However, no study to date has investigated whether ACC sulcal pattern, a stable brain feature primarily determined in utero, influences CC efficiency in the early stages of cognitive and neural development. Using anatomical MRI and three-dimensional reconstruction of cortical folds, we investigated the effect that ACC sulcal pattern may have on the Stroop score, a classical behavioral index of CC efficiency, in 5-year-old preschoolers. We found higher CC efficiency, that is, lower Stroop interference scores for both RTs and error rates, in children with asymmetrical ACC sulcal pattern (i.e., different pattern in each hemisphere) compared with children with symmetrical pattern (i.e., same pattern in both hemispheres). Critically, ACC sulcal pattern had no effect on performance in the forward and backward digit span tasks suggesting that ACC sulcal pattern contributes to the executive ability to resolve conflicts but not to the ability to maintain and manipulate information in working memory. This finding provides the first evidence that preschoolers' CC efficiency is likely associated with ACC sulcal pattern, thereby suggesting that the brain shape could result in early constraints on human executive ability.

Working Memory Delay Activity Predicts Individual Differences in Cognitive Abilities

December 2014


140 Reads

A great deal of prior research has examined the relation between estimates of working memory and cognitive abilities. Yet, the neural mechanisms that account for these relations are still not very well understood. The current study explored whether individual differences in working memory delay activity would be a significant predictor of cognitive abilities. A large number of participants performed multiple measures of capacity, attention control, long-term memory, working memory span, and fluid intelligence, and latent variable analyses were used to examine the data. During two working memory change detection tasks, we acquired EEG data and examined the contralateral delay activity. The results demonstrated that the contralateral delay activity was significantly related to cognitive abilities, and importantly these relations were because of individual differences in both capacity and attention control. These results suggest that individual differences in working memory delay activity predict individual differences in a broad range of cognitive abilities, and this is because of both differences in the number of items that can be maintained and the ability to control access to working memory.

When Less Is More: Evidence for a Facilitative Cathodal tDCS Effect in Attentional Abilities

May 2012


80 Reads

Many previous studies reported that the hyperpolarization of cortical neurons following cathodal stimulation (in transcranial direct current stimulation) has resulted in cognitive performance degradation. Here, we challenge this assumption by showing that cathodal stimulation will not always degrade cognitive performance. We used an attentional load paradigm in which irrelevant stimuli are processed only under low but not under high attentional load. Thirty healthy participants were randomly allocated into three interventional groups with different brain stimulation parameters (active anodal posterior parietal cortex [PPC], active cathodal PPC, and sham). Cathodal but not anodal stimulation enabled flanker processing even in high-loaded scenes. A second experiment was carried out to assert whether the improved flanker processing under cathodal stimulation is because of altered attention allocation between center and surround or, alternatively, enhanced attentional resources. In this experiment, the flanker was presented centrally. The results of Experiment 2 replicated Experiment 1's finding of improved flanker processing. We interpret the results from these two experiments as evidence for the ability of cathodal stimulation to enhance attentional resources rather than simply change attention allocation between center and periphery. Cathodal stimulation in high-loaded scenes can act like a noise filter and may in fact enhance cognitive performance. This study contributes to understanding the way the PPC is engaged with attentional functions and explains the cathodal effects, which thus might lead to more efficient brain stimulation protocols.

Extensive Left Temporal Pole Damage Does Not Impact on Theory of Mind Abilities

September 2013


227 Reads

The temporal poles (TPs) are among the brain regions that are often considered as the brain network sustaining our ability to understand other people's mental states or "Theory of Mind" (ToM). However, so far the functional role of the left and right TPs in ToM is still debated, and it is even not clear yet whether these regions are necessary for ToM. In this study, we tested whether the left TP is necessary for ToM by assessing the mentalizing abilities of a patient (C.M.) diagnosed with semantic dementia. Converging evidence from detailed MRI and (18)F-fluoro-2-deoxy-d-glucose PET examinations showed a massive atrophy of the left TP with the right TP being relatively unaffected. Furthermore, C.M.'s atrophy encompassed most regions of the left TP usually activated in neuroimaging studies investigating ToM. Given C.M.'s language impairments, we used a battery of entirely nonverbal ToM tasks. Across five tasks encompassing 100 trials, which probed the patient's ability to attribute various mental states (intentions, knowledge, and beliefs), C.M. showed a totally spared performance. This finding suggests that, despite its consistently observed activation in neuroimaging studies involving ToM tasks, the left TP is not necessary for ToM reasoning, at least in nonverbal conditions and as long as its right counterpart is preserved. Implications for understanding the social abilities of patients with semantic dementia are discussed.

Multiple Cognitive Abilities from a Single Cortical Algorithm

May 2012


176 Reads

One strong claim made by the representational-hierarchical account of cortical function in the ventral visual stream (VVS) is that the VVS is a functional continuum: the basic computations carried out in service of a given cognitive function, such as recognition memory or visual discrimination, might be the same at all points along the VVS. Here, we use a single-layer computational model with a fixed learning mechanism and set of parameters to simulate a variety of cognitive phenomena from different parts of the functional continuum of the VVS: recognition memory, categorization of perceptually related stimuli, perceptual learning of highly similar stimuli, and development of retinotopy and orientation selectivity. The simulation results indicate--consistent with the representational-hierarchical view--that the simple existence of different levels of representational complexity in different parts of the VVS is sufficient to drive the emergence of distinct regions that appear to be specialized for solving a particular task, when a common neurocomputational learning algorithm is assumed across all regions. Thus, our data suggest that it is not necessary to invoke computational differences to understand how different cortical regions can appear to be specialized for what are considered to be very different psychological functions.

Development of Elementary Numerical Abilities: A Neuronal Model

October 1993


1,041 Reads

Abstract Despite their lack of language, human infants and several animal species possess some elementary abilities for numerical processing. These include the ability to recognize that a given numerosity is being presented visually or auditorily, and, at a later stage of development, the ability to compare two nume-rosities and to decide which is larger. We propose a model for the development of these abilities in a formal neuronal network. Initially, the model is equipped only with unordered numerosity detectors. It can therefore detect the numerosity of an input set and can be conditioned to react accordingly. In a later stage, the addition of a short-term memory network is shown to be sufficient for number comparison abilities to develop. Our computer simulations account for several phenomena in the numerical domain, including the distance effect and Fechner's law for numbers. They also demonstrate that infants' numerosity detection abilities may be explained without assuming that infants can count. The neurobiological bases of the critical components of the model are discussed.

Figure 1. Lesion overlaps. Lesions of participants in the vmPFC group are shown in warmer colors, where the maximum overlap (15) is within ventromedial prefrontal sector. Lesions in the BDC group cover a much broader swath of cortical territory with a maximum overlap of 4. Purple represents regions of overlap between the vmPFC and BDC groups and covers bilateral dorsal pFC. 
Figure 2. Difference in proportional investment by group. Bars represent the difference in the proportion of money invested when given dispositional information minus situational information. Participants with vmPFC damage displayed a greater difference than the other groups. 
Figure 3. Proportional investment by information type. Bars represent the proportion of money invested with dispositional or situational information types. Participants with vmPFC damage invested more with dispositional information and less with situational information compared with the other groups. 
Figure 4. Investments for catch trials. Bars represent the amount invested for dispositional and situational information types on trials where only one of these types of information was given. There are no differences between groups. 
Figure 5. Specificity of vmPFC subregions. Each voxel represents the group average difference in the proportion of money invested with dispositional and situational information (dispositional – situational) for participants whose lesion includes that voxel. Warmer colors represent a larger difference. Only voxels where the number of lesion overlaps is greater than 5 are included. 
Abnormal Causal Attribution Leads to Advantageous Economic Decision-making: A Neuropsychological Approach

April 2013


88 Reads

People tend to assume that outcomes are caused by dispositional factors, for example, a person's constitution or personality, even when the actual cause is due to situational factors, for example, luck or coincidence. This is known as the "correspondence bias." This tendency can lead normal, intelligent persons to make suboptimal decisions. Here, we used a neuropsychological approach to investigate the neural basis of the correspondence bias, by studying economic decision-making in patients with damage to the ventromedial pFC (vmPFC). Given the role of the vmPFC in social cognition, we predicted that vmPFC is necessary for the normal correspondence bias. In our experiment, consistent with expectations, healthy (n = 46) and brain-damaged (n = 30) comparison participants displayed the correspondence bias during economic decision-making and invested no differently when given dispositional or situational information. By contrast, vmPFC patients (n = 17) displayed a lack of correspondence bias and invested more when given dispositional than situational information. The results support the conclusion that vmPFC is critical for normal social inference and the correspondence bias. The findings help clarify the important (and sometimes disadvantageous) role of social inference in economic decision-making.

Abnormal Auditory Cortical Activation in Dyslexia 100 msec after Speech Onset

June 2002


32 Reads

Reading difficulties are associated with problems in processing and manipulating speech sounds. Dyslexic individuals seem to have, for instance, difficulties in perceiving the length and identity of consonants. Using magnetoencephalography (MEG), we characterized the spatio-temporal pattern of auditory cortical activation in dyslexia evoked by three types of natural bisyllabic pseudowords (/ata/, /atta/, and /a a/), complex nonspeech sound pairs (corresponding to /atta/ and /a a/) and simple 1-kHz tones. The most robust difference between dyslexic and non-reading-impaired adults was seen in the left supratemporal auditory cortex 100 msec after the onset of the vowel /a/. This N100m response was abnormally strong in dyslexic individuals. For the complex nonspeech sounds and tone, the N100m response amplitudes were similar in dyslexic and nonimpaired individuals. The responses evoked by syllable /ta/ of the pseudoword /atta/ also showed modest latency differences between the two subject groups. The responses evoked by the corresponding nonspeech sounds did not differ between the two subject groups. Further, when the initial formant transition, that is, the consonant, was removed from the syllable /ta/, the N100m latency was normal in dyslexic individuals. Thus, it appears that dyslexia is reflected as abnormal activation of the auditory cortex already 100 msec after speech onset, manifested as abnormal response strengths for natural speech and as delays for speech sounds containing rapid frequency transition. These differences between the dyslexic and nonimpaired individuals also imply that the N100m response codes stimulus-specific features likely to be critical for speech perception. Which features of speech (or nonspeech stimuli) are critical in eliciting the abnormally strong N100m response in dyslexic individuals should be resolved in future studies.

Abnormal Functional Activation During a Simple Word Repetition Task: A PET Study of Adult Dyslexics

October 2000


272 Reads

Eight dyslexic subjects, impaired on a range of tasks requiring phonological processing, were matched for age and general ability with six control subjects. Participants were scanned using positron emission tomography (PET) during three conditions: repeating real words, repeating pseudowords, and rest. In both groups, speech repetition relative to rest elicited widespread bilateral activation in areas associated with auditory processing of speech; there were no significant differences between words and pseudowords. However, irrespective of word type, the dyslexic group showed less activation than the control group in the right superior temporal and right post-central gyri and also in the left cerebellum. Notably, the right anterior superior temporal cortex (Brodmann's area 22 [BA 22]) was less activated in each of the eight dyslexic subjects, compared to each of the six control subjects. This deficit appears to be specific to auditory repetition as it was not detected in a previous study of reading which used the same sets of stimuli (Brunswick, N., McCrory, E., Price, C., Frith, C.D., & Frith, U. [1999]. Explicit and implicit processing of words and pseudowords by adult developmental dyslexics: A search for Wernicke's Wortschatz? Brain, 122, 1901-1917). This implies that the observed neural manifestation of developmental dyslexia is task-specific (i.e., functional rather than structural). Other studies of normal subjects indicate that attending to the phonetic structure of speech leads to a decrease in right-hemisphere processing. Lower right hemisphere activation in the dyslexic group may therefore indicate less processing of non-phonetic aspects of speech, allowing greater salience to be accorded to phonological aspects of attended speech.

Abnormal Processing of Social Information from Faces in Autism

March 2001


787 Reads

Autism has been thought to be characterized, in part, by dysfunction in emotional and social cognition, but the pathology of the underlying processes and their neural substrates remain poorly understood. Several studies have hypothesized that abnormal amygdala function may account for some of the impairments seen in autism, specifically, impaired recognition of socially relevant information from faces. We explored this issue in eight high-functioning subjects with autism in four experiments that assessed recognition of emotional and social information, primarily from faces. All tasks used were identical to those previously used in studies of subjects with bilateral amygdala damage, permitting direct comparisons. All subjects with autism made abnormal social judgments regarding the trustworthiness of faces; however, all were able to make normal social judgments from lexical stimuli, and all had a normal ability to perceptually discriminate the stimuli. Overall, these data from subjects with autism show some parallels to those from neurological subjects with focal amygdala damage. We suggest that amygdala dysfunction in autism might contribute to an impaired ability to link visual perception of socially relevant stimuli with retrieval of social knowledge and with elicitation of social behavior.

Functional Magnetic Resonance Imaging Evidence for Abnormalities in Response Selection in Attention Deficit Hyperactivity Disorder: Differences in Activation Associated with Response Inhibition but Not Habitual Motor Response

April 2008


204 Reads

Impaired response inhibition is thought to be a core deficit in attention deficit hyperactivity disorder (ADHD). Prior imaging studies investigating response inhibition in children with ADHD have used tasks involving different cognitive resources, thereby complicating the interpretation of their findings. In this study, a classical go/no-go task with a well-ingrained stimulus-response association (green = go; red = no-go) was used in order to minimize extraneous cognitive demands. Twenty-five children with ADHD and 25 typically developing (TD) children between the ages of 8 and 13 years and group-matched for IQ and performance on the go/no-go task were studied using event-related functional magnetic resonance imaging (fMRI). Analyses were used to examine differences in activation between the ADHD and TD groups for "go" (habitual motor response) and "no-go" (requiring inhibition of the motor response) events. Region-of-interest analyses revealed no between-group difference in activation in association with "go" events. For "no-go" events, the children with ADHD demonstrated significantly less activation than did TD controls within a network important for inhibiting a motor response to a visual stimulus, with frontal differences localized to the pre-supplementary motor area. Although blood oxygenation level-dependent fMRI data show no differences between children with ADHD and TD children in association with a habituated motor "go" response, during "no-go" events, which require selecting not to respond, children with ADHD show diminished recruitment of networks important for response inhibition. The findings suggest that abnormalities in circuits important for motor response selection contribute to deficits in response inhibition in children with ADHD and lend support to the growing awareness of ADHD-associated anomalies in medial frontal regions important for the control of voluntary actions.

Figure 1.  
Figure 2. Lesion overlap of vmPFC patients. Mesial and frontal views of the overlap map of lesions for the 9 vmPFC patients. The color bar indicates the number of overlapping lesions at each voxel. The area of maximal overlap lies in the ventromedial prefrontal cortex.  
Harming Kin to Save Strangers: Further Evidence for Abnormally Utilitarian Moral Judgments after Ventromedial Prefrontal Damage

October 2010


272 Reads

The ventromedial PFC (vmPFC) has been implicated as a critical neural substrate mediating the influence of emotion on moral reasoning. It has been shown that the vmPFC is especially important for making moral judgments about "high-conflict" moral dilemmas involving direct personal actions, that is, scenarios that pit compelling utilitarian considerations of aggregate welfare against the highly emotionally aversive act of directly causing harm to others [Koenigs, M., Young, L., Adolphs, R., Tranel, D., Cushman, F., Hauser, M., et al. Damage to the prefrontal cortex increases utilitarian moral judgments. Nature, 446, 908-911, 2007]. The current study was designed to elucidate further the role of the vmPFC in high-conflict moral judgments, including those that involve indirect personal actions, such as indirectly causing harm to one's kin to save a group of strangers. We found that patients with vmPFC lesions were more likely than brain-damaged and healthy comparison participants to endorse utilitarian outcomes on high-conflict dilemmas regardless of whether the dilemmas (1) entailed direct versus indirect personal harms and (2) were presented from the Self versus Other perspective. In addition, all groups were more likely to endorse utilitarian outcomes in the Other perspective as compared with the Self perspective. These results provide important extensions of previous work, and the findings align with the proposal that the vmPFC is critical for reasoning about moral dilemmas in which anticipating the social-emotional consequences of an action (e.g., guilt or remorse) is crucial for normal moral judgments [Greene, J. D. Why are VMPFC patients more utilitarian?: A dual-process theory of moral judgment explains. Trends in Cognitive Sciences, 11, 322-323, 2007; Koenigs, M., Young, L., Adolphs, R., Tranel, D., Cushman, F., Hauser, M., et al. Damage to the prefrontal cortex increases utilitarian moral judgments. Nature, 446, 908-911, 2007].

Medial PFC Damage Abolishes the Self-reference Effect

September 2011


133 Reads

Functional neuroimaging studies suggest that the medial PFC (mPFC) is a key component of a large-scale neural system supporting a variety of self-related processes. However, it remains unknown whether the mPFC is critical for such processes. In this study, we used a human lesion approach to examine this question. We administered a standard trait judgment paradigm [Kelley, W. M., Macrae, C. N., Wyland, C. L., Caglar, S., Inati, S., & Heatherton, T. F. Finding the self? An event-related fMRI study. Journal of Cognitive Neuroscience, 14, 785-794, 2002] to patients with focal brain damage to the mPFC. The self-reference effect (SRE), a memory advantage conferred by self-related processing, served as a measure of intact self-processing ability. We found that damage to the mPFC abolished the SRE. The results demonstrate that the mPFC is necessary for the SRE and suggest that this structure is important for self-referential processing and the neural representation of self.

Robust Selectivity for Faces in the Human Amygdala in the Absence of Expressions

August 2013


32 Reads

There is a well-established posterior network of cortical regions that plays a central role in face processing and that has been investigated extensively. In contrast, although responsive to faces, the amygdala is not considered a core face-selective region, and its face selectivity has never been a topic of systematic research in human neuroimaging studies. Here, we conducted a large-scale group analysis of fMRI data from 215 participants. We replicated the posterior network observed in prior studies but found equally robust and reliable responses to faces in the amygdala. These responses were detectable in most individual participants, but they were also highly sensitive to the initial statistical threshold and habituated more rapidly than the responses in posterior face-selective regions. A multivariate analysis showed that the pattern of responses to faces across voxels in the amygdala had high reliability over time. Finally, functional connectivity analyses showed stronger coupling between the amygdala and posterior face-selective regions during the perception of faces than during the perception of control visual categories. These findings suggest that the amygdala should be considered a core face-selective region.

Inhibition-related Activation in the Right Inferior Frontal Gyrus in the Absence of Inhibitory Cues

March 2011


395 Reads

The right inferior frontal gyrus (rIFG) has been hypothesized to mediate response inhibition. Typically response inhibition is signaled by an external stop cue, which provides a top-down signal to initiate the process. However, recent behavioral findings suggest that response inhibition can also be triggered automatically by bottom-up processes. In the present study, we evaluated whether rIFG activity would also be observed during automatic inhibition, in which no stop cue was presented and no motor inhibition was actually required. We measured rIFG activation in response to stimuli that were previously associated with stop signals but which required a response on the current trial (reversal trials). The results revealed an increase in rIFG (pars triangularis) activity, suggesting that it can be activated by associations between stimuli and stopping. Moreover, its role in inhibition tasks is not contingent on the presence of an external stop cue. We conclude that rIFG involvement in stopping is consistent with a role in reprogramming of action plans, which may comprise inhibition, and its activity can be triggered through automatic, bottom-up processing.

Figure 2. The eight options presented in the multiple choice question after Experimental Runs 1 – 3 and the control run. For the NIB group, option A was chosen by all participants after Runs 1 – 3 and the control run. For the IB group, option A was never chosen after Runs 1 – 3. Instead, option B was chosen five times, option C was chosen two times, option F was chosen two times, option H was chosen three times, and options D, E, and G were never chosen. After the control run, all participants chose option A. 
Figure 3. ROIs. Each individualsʼ cortex was inflated and a retinotopic map (A, depicted here on a representative participant), eccentricity map, study-specific localizer, and a mapping for LOC (also shown here) were projected onto the inflated surface. On the basis of these mappings, six ROIs ( V1, V2, V3, V3A/B, V4, LOC) were identified for each participant (B, depicted here on the representative participant in A). The gray line in A represents the border between the central visual region and the periphery where the inducers were presented.
Figure 6. A whole-brain analysis was performed for the NIB (A) and IB groups (B), in which the Kanizsa figure was contrasted with the three control figures (Kanizsa > Controls). Activity is in lower and higher visual areas and does not differ between the two groups. Note that the scans did not cover the whole brain but only recorded visual cortex and extended into a part of the temporal and parietal cortex. Dotted lines indicate upper and lower limits of epi acquisition.
Figure 7. Univariate results for the control run. Activity (in normalized arbitrary units) associated with the four figures in each ROI for the NIB group (A) and the IB group (B). Error bars denote SE.
Seeing without Knowing: Neural Signatures of Perceptual Inference in the Absence of Report

November 2013


769 Reads

Everyday, we experience a rich and complex visual world. Our brain constantly translates meaningless fragmented input into coherent objects and scenes. However, our attentional capabilities are limited, and we can only report the few items that we happen to attend to. So what happens to items that are not cognitively accessed? Do these remain fragmentary and meaningless? Or are they processed up to a level where perceptual inferences take place about image composition? To investigate this, we recorded brain activity using fMRI while participants viewed images containing a Kanizsa figure, an illusion in which an object is perceived by means of perceptual inference. Participants were presented with the Kanizsa figure and three matched nonillusory control figures while they were engaged in an attentionally demanding distractor task. After the task, one group of participants was unable to identify the Kanizsa figure in a forced-choice decision task; hence, they were "inattentionally blind." A second group had no trouble identifying the Kanizsa figure. Interestingly, the neural signature that was unique to the processing of the Kanizsa figure was present in both groups. Moreover, within-subject multivoxel pattern analysis showed that the neural signature of unreported Kanizsa figures could be used to classify reported Kanizsa figures and that this cross-report classification worked better for the Kanizsa condition than for the control conditions. Together, these results suggest that stimuli that are not cognitively accessed are processed up to levels of perceptual interpretation.

Figure 1. (A) The conditional association learning paradigm. After a cue is briefly presented at the center of gaze, monkeys maintain fixation during a memory delay before making a saccade to one of two targets presented on the right and left of the extinguished fixation spot. Only a saccade to the target associated with the cue generates a reward. (B) The block structure of the conditional association learning task without reversals. On any given block of trials, each of two novel cues is associated with a single saccade direction. Once a block of trials is completed, the old images are discarded, and two novel cues are again presented and must be learned.  
Figure 2. (A) Percentage of correct performance averaged across blocks and cues for all trials (correct plus error trials). At the start of a block (Trial 1), average performance is not above chance levels (50%) but quickly jumps above chance by the second presentation of each cue and continues to gradually improve throughout the block. (B) RTs averaged across blocks and cues for correct trials to match neural data from the two monkeys. RTs are slowest at the start of new blocks but rapidly improve over the first four trials and then continue to gradually improve over the remainder of the block. Error bars show SEM.  
Figure 3. (A) Change in peri-cue direction selectivity during association learning without reversals. Population percentage of variance explained by direction (PEV dir , color scale) shown as a function of correct trials and time from cue onset, averaged across blocks and cues. Black dots indicate the half-maximal PEV dir or rise times. (B) Rise times replotted and fit with a sigmoid curve show bistable learning, with initial late activity (Trials 1–4) followed by an increase in early trial direction selectivity starting with Trial 5. (C) Mean PEV dir from the second half of the cue period (250–500 msec after cue onset) also shows a jump by Trial 5 but continues to increase with learning. Error bars show standard deviation of the mean.  
Rapid Association Learning in the Primate Prefrontal Cortex in the Absence of Behavioral Reversals

July 2011


127 Reads

The PFC plays a central role in our ability to learn arbitrary rules, such as "green means go." Previous experiments from our laboratory have used conditional association learning to show that slow, gradual changes in PFC neural activity mirror monkeys' slow acquisition of associations. These previous experiments required monkeys to repeatedly reverse the cue-saccade associations, an ability known to be PFC-dependent. We aimed to test whether the relationship between PFC neural activity and behavior was due to the reversal requirement, so monkeys were trained to learn several new conditional cue-saccade associations without reversing them. Learning-related changes in PFC activity now appeared earlier and more suddenly in correspondence with similar changes in behavioral improvement. This suggests that learning of conditional associations is linked to PFC activity regardless of whether reversals are required. However, when previous learning does not need to be suppressed, PFC acquires associations more rapidly.

Figure 2. Behavioral data. (A) Overall response times. Response times in the search condition were significantly longer than those in the no-search condition. (B) Overall accuracy. Response accuracy was significantly higher in the no-search condition than in the search condition. (C). Response times in the singleton-present/search condition, plotted as function of the relative position of the color singleton to the target. For about half the participants, responses were significantly slower when the target was presented to the location opposite to that of the distractor (opposite-slow group; black lines). For approximately the other half of the participants, responses were actually faster when the target was presented opposite to the distractor location (opposite-fast group; red lines. For each group, the single-subject response patterns are superimposed using dashed lines.). (D) Response times of the opposite-slow group. Note that in the no-search condition the response times to singleton-present and singleton-absent trials are similar. (E). Response time of the opposite-fast group. Notice that in contrast to the opposite-slow group, responses in the no-search condition are significantly faster here when a color singleton is present than when a color singleton is not present.
Table 2 . Areas Activated by the Color Singleton
Figure 3. Overview of brain areas that were more active in the search condition than in the no-search condition. (A) Brain areas showing significant activation. (B) Time courses of BOLD response in key areas activated by the top – down control. The activated areas are mainly located in frontal and parietal cortex. 
Figure 4. Brain areas activated by the color singleton. (A) Brain areas showing significant singleton related activation. (B) Time courses of the BOLD response in key areas activated by the singleton. Notice that in contrast to the search related activations (see Figure 2), the color singleton activated predominantly areas in parietal and occipital brain regions. 
Figure 5. Relation between brain activity and attentional control. (A) Correlation between the strength of attentional control and activation in the right insula. Top – down control strength is defined as the response time difference between responses to targets presented opposite to a color singleton and responses to targets that were presented at locations adjacent to the color singleton. (B) Interaction between group and singleton presence. Left: One area in the left precentral gyrus yielded a significant interaction between singleton presence and group, suggesting that this area was more strongly activated by color singletons in the opposite-slow group than in the opposite-fast group. Right: A post hoc analysis confirmed that this area was indeed significantly activated in the opposite-slow group alone (right), but not in the opposite-fast group (data not shown here). 
Brain Structures Involved in Visual Search in the Presence and Absence of Color Singletons

April 2009


124 Reads

It is still debated to what degree top-down and bottom-up driven attentional control processes are subserved by shared or by separate mechanisms. Interactions between these attentional control forms were investigated using a rapid event-related fMRI design, using an attentional search task. Following a prestimulus mask, target stimuli (consisting of a letter C or a mirror image of the C, enclosed in a diamond outline) were presented either at one unique location among three nontarget items (consisting of a random letter, enclosed in a circle outline; 50% probability), or at all four possible target locations (also 50% probability). On half the trials, irrelevant color singletons were presented, consisting of a color change of one of the four prestimulus masks, just prior to target appearance. Participants were required to search for a target letter inside the diamond and report its orientation. Results indicate that, in addition to a common network of parietal areas, medial frontal cortex is uniquely involved in top-down orienting, whereas bottom-up control is mainly subserved by a network of occipital and parietal areas. Additionally, we found that participants who were better able to suppress orienting to the color singleton showed middle frontal gyrus activation, and that the degree of top-down control correlated with insular activity. We conclude that, in addition to a common set of parietal areas, separate brain areas are involved in top-down and bottom-up driven attentional control, and that frontal areas play a role in the suppression of attentional capture by an irrelevant color singleton.

Figure 2. Mean median reaction times on the LVF-RH and RVF-LH trials as a function of spatial relation task. The results for the bright and contrast-balanced (CB) stimuli are presented in the left and right panels, respectively. CATE and COOR stand for categorical and coordinate, respectively. 
Hemispheric Processing of Categorical and Coordinate Spatial Relations in the Absence of Low Spatial Frequencies

March 2002


115 Reads

Right-handed participants performed the categorical and coordinate spatial relation judgments on stimuli presented to either the left visual field - right hemisphere (LVF-RH) or the right visual field - left hemisphere (RVF-LH). The stimulus patterns were formulated either by bright dots or contrast-balanced dots. When the stimuli were bright, an RVF-LH advantage was observed for the categorical task, whereas an LVF-RH advantage was observed for the coordinate task. When the stimuli were contrast balanced, the RVF-LH advantage was observed for the categorical task, but the LVF-RH advantage was eliminated for the coordinate task. Because the contrast-balanced dots are largely devoid by of low spatial frequency content, these results suggest that processing of low spatial frequency is responsible for the right hemisphere advantage for the coordinate spatial processing.

Persistent Neuronal Firing in Primary Somatosensory Cortex in the Absence of Working Memory of Trial-specific Features of the Sample Stimuli in a Haptic Working Memory Task

November 2011


174 Reads

Previous studies suggested that primary somatosensory (SI) neurons in well-trained monkeys participated in the haptic-haptic unimodal delayed matching-to-sample (DMS) task. In this study, 585 SI neurons were recorded in monkeys performing a task that was identical to that in the previous studies but without requiring discrimination and active memorization of specific features of a tactile or visual memorandum. A substantial number of those cells significantly changed their firing rate in the delay compared with the baseline, and some of them showed differential delay activity. These firing changes are similar to those recorded from monkeys engaged in active (working) memory. We conclude that the delay activity is not necessarily only observed as was generally thought in the situation of active memorization of different features between memoranda after those features have been actively discriminated. The delay activity observed in this study appears to be an intrinsic property of SI neurons and suggests that there exists a neural network in SI (the primary sensory cortex) for haptic working memory no matter whether the difference in features of memoranda needs to be memorized in the task or not. Over 400 SI neurons were also recorded in monkeys well-trained to discriminate two memoranda in the haptic-haptic DMS task for comparison of delay firing of SI neurons between the two different working memory tasks used in this study. The similarity observed in those two situations suggests that working memory uses already-existing memory apparatus by activating it temporarily. Our data also suggest that, through training (repetitive exposure to the stimulus), SI neurons may increase their involvement in the working memory of the memorandum.

Figure 5. Grand-averaged ERPs from the rare intra stimuli and rare inter stimuli for the control subjects (A), the acallosal subject S.G. with AC (B), and the acallosal subject S.Pe. without AC (C) at Po7, Po8, P7, P8, Tp7, Tp8, T7, T8, Ft7, Ft8, F7, and F8 recording sites. These sites were chosen because their lateral scalp positions allowed for clear interhemispheric comparisons.
Figure 5. (continued )
Inter- and Intra-hemispheric Processing of Visual Event-related Potentials in the Absence of the Corpus Callosum

April 2004


76 Reads

Interhemispheric differences of the N100 latency ill visual evoked potentials have been used to estimate interhemispheric transfer time (e.g., Saron & Davidson, 1989). Recent work has also suggested that the P300 component could reflect the efficacy of interhemispheric transmission (Polich & 1998). The purpose of the present study was to study the differential role of the corpus callosum (CC) and anterior commissure (AC) in the interhemispheric propagation of these two electrophysiological components. Thus, the amplitude and latency distribution of the N100 and P300 components were analyzed using high-density electrical mapping in a subject with agenesis of CC but preservation of AC, a subject with agenesis Of both CC and AC, and 10 neurologically intact Control subjects. The task consisted of a Modified visual oddball paradigm comprising one frequent and two rare stimuli, one presented on the same and the other on the opposite side of the frequent stimulus. Interhemispheric differences in latency were found for the N100 component in controls. However, in the acallosal subjects, this component was not identifiable in the indirectly stimulated hemisphere. In controls, no interhemispheric differences were observed in the distribution of the P300 latency and amplitude to rare and frequent stimuli. The distribution of the P300 amplitude in the acallosal subject with an AC was identical to that of the controls, whereas in the acallosal subject lacking the AC, the amplitude was greater in the hemisphere receiving the rare stimuli were presented. In both acallosal subjects, hemispheric differences in the P300 latency were observed, the latencies being shorter in the hemisphere directly stimulated for all categories of stimuli. These results suggest that the interhemispheric transfer of both the N100 and P300 components relies on the integrity of cortical commissures. Possible P300 generator sources are discussed.

Absence of Face-specific Cortical Activity in the Complete Absence of Awareness: Converging Evidence from Functional Magnetic Resonance Imaging and Event-related Potentials

September 2011


501 Reads

In this study, we explored the neural correlates of perceptual awareness during a masked face detection task. To assess awareness more precisely than in previous studies, participants employed a 4-point scale to rate subjective visibility. An event-related fMRI and a high-density ERP study were carried out. Imaging data showed that conscious face detection was linked to activation of fusiform and occipital face areas. Frontal and parietal regions, including the pre-SMA, inferior frontal sulcus, anterior insula/frontal operculum, and intraparietal sulcus, also responded strongly when faces were consciously perceived. In contrast, no brain area showed face-selective activity when participants reported no impression of a face. ERP results showed that conscious face detection was associated with enhanced N170 and also with the presence of a second negativity around 300 msec and a slow positivity around 415 msec. Again, face-related activity was absent when faces were not consciously perceived. We suggest that, under conditions of backward masking, ventral stream and fronto-parietal regions show similar, strong links of face-related activity to conscious perception and stress the importance of a detailed assessment of awareness to examine activity related to unseen stimulus events.

Global Versus Local Processing in the Absence of Low Spatial Frequencies

July 1990


36 Reads

When observers are presented with hierarchical visual stimuli that contain incongruous coarse ("global") and fine ("local") pattern attributes, the global structure interferes with local pattern processing more than local structure interferes with global pattern processing. This effect is referred to as "global precedence." The present experiments tested the hypothesis that global precedence depends on the presence of low spatial frequencies using stimuli constructed from "contrast balanced dots." Stimuli composed of contrast balanced dots are largely devoid of low-frequency content. Choice reaction time to identify either the local or global pattern information was the dependent measure. Global precedence was found only for control stimuli that contained low spatial frequencies. In the absence of low-frequency information, local precedence was obtained. These findings suggest that global precedence is heavily dependent on the low spatial frequency content of the patterns.

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