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1.  Relating Hippocampus to Relational Memory Processing across Domains and Delays 
Journal of cognitive neuroscience  2015;27(2):234-245.
The hippocampus has been implicated in a diverse set of cognitive domains and paradigms, including cognitive mapping, long-term memory, and relational memory, at long or short study–test intervals. Despite the diversity of these areas, their association with the hippocampus may rely on an underlying commonality of relational memory processing shared among them. Most studies assess hippocampal memory within just one of these domains, making it difficult to know whether these paradigms all assess a similar underlying cognitive construct tied to the hippocampus. Here we directly tested the commonality among disparate tasks linked to the hippocampus by using PCA on performance from a battery of 12 cognitive tasks that included two traditional, long-delay neuropsychological tests of memory and two laboratory tests of relational memory (one of spatial and one of visual object associations) that imposed only short delays between study and test. Also included were different tests of memory, executive function, and processing speed. Structural MRI scans from a subset of participants were used to quantify the volume of the hippocampus and other subcortical regions. Results revealed that the 12 tasks clustered into four components; critically, the two neuropsychological tasks of long-term verbal memory and the two laboratory tests of relational memory loaded onto one component. Moreover, bilateral hippocampal volume was strongly tied to performance on this component. Taken together, these data emphasize the important contribution the hippocampus makes to relational memory processing across a broad range of tasks that span multiple domains.
doi:10.1162/jocn_a_00717
PMCID: PMC4336790  PMID: 25203273
2.  Implicit and Explicit Second Language Training Recruit Common Neural Mechanisms for Syntactic Processing 
Journal of cognitive neuroscience  2013;25(6):936-951.
In contrast to native language acquisition, adult second language (L2) acquisition occurs under highly variable learning conditions. While most adults acquire their L2 at least partially through explicit instruction, as in a classroom setting, many others acquire their L2 primarily through implicit exposure, as is typical of an immersion environment. Whether these differences in acquisition environment play a role in determining the neural mechanisms that are ultimately recruited to process L2 grammar has not been well characterized. The present study investigated this issue by comparing the event-related potential response to novel L2 syntactic rules acquired under conditions of implicit exposure and explicit instruction, using a novel laboratory language-learning paradigm. Native speakers tested on these stimuli showed a biphasic response to syntactic violations, consisting of an earlier negativity followed by a later P600 effect. After merely an hour of training, both implicitly- and explicitly-trained learners who were capable of detecting grammatical violations also elicited P600 effects. In contrast, learners who were unable to discriminate between grammatically correct and incorrect sentences did not show significant P600 effects. The magnitude of the P600 effect was found to correlate with learners’ behavioral proficiency. Behavioral measures revealed that successful learners from both the implicit and explicit groups gained explicit, verbalizable knowledge about the L2 grammar rules. Taken together, these results indicate that late, controlled mechanisms indexed by the P600 play a crucial role in processing a late-learned L2 grammar, regardless of training condition. These findings underscore the remarkable plasticity of later, attention-dependent processes and their importance in lifelong learning.
doi:10.1162/jocn_a_00354
PMCID: PMC4334462  PMID: 23631551
3.  The Impact of Early Amygdala Damage on Juvenile Rhesus Macaque Social Behavior 
Journal of cognitive neuroscience  2013;25(12):2124-2140.
The present experiments continue a longitudinal study of rhesus macaque social behavior following bilateral neonatal ibotenic acid lesions of the amygdala or hippocampus, or sham operations. Juvenile animals (approximately 1.5- 2.5 years of age) were tested in four different social contexts—alone, while interacting with one familiar peer, while interacting with one unfamiliar peer, and in their permanent social groups. During infancy, the amygdala-lesioned animals displayed more interest in conspecifics (indexed by increased affiliative signaling) and paradoxically demonstrated more submission or fear (Bauman, Lavenex, Mason, Capitanio, & Amaral, 2004a, this journal). When these animals were assessed as juveniles, differences were less striking. Amygdala-lesioned animals generated fewer aggressive and affiliative signals (e.g., vocalizations, facial displays) and spent less time in social interactions with familiar peers. When animals were observed alone or with an unfamiliar peer, amygdala-lesioned, compared with other subjects, spent more time being inactive and physically explored the environment less. Despite the subtle, lesion-based differences in the frequency and duration of specific social behaviors, there were lesion-based differences in the organization of behavior such that lesion groups could be identified based on the patterning of social behaviors in a discriminant function analysis. The findings indicate that, although overall frequencies of many of the observed behaviors do not differ between groups, the general patterning of social behavior may distinguish the amygdala-lesioned animals.
doi:10.1162/jocn_a_00483
PMCID: PMC4330965  PMID: 24047387
amygdala; social behavior; nonhuman primate; Macaca mulatta; Rhesus macaque
4.  Sequence Learning is Preserved in Individuals with Cerebellar Degeneration when the Movements are Directly Cued 
Journal of cognitive neuroscience  2009;21(7):1302-1310.
Cerebellar pathology is associated with impairments on a range of motor learning tasks including sequence learning. However, various lines of evidence are at odds with the idea that the cerebellum plays a central role in the associative processes underlying sequence learning. Behavioral studies indicate that sequence learning, at least with short periods of practice, involves the establishment of effector-independent, abstract spatial associations, a form of representation not associated with cerebellar function. Moreover, neuroimaging studies have failed to identify learning-related changes within the cerebellum. We hypothesize that the cerebellar contribution to sequence learning may be indirect, related to the maintenance of stimulus–response associations in working memory, rather than through processes directly involved in the formation of sequential predictions. Consistent with this hypothesis, individuals with cerebellar pathology were impaired in learning movement sequences when the task involved a demanding stimulus–response translation. When this translation process was eliminated by having the stimuli directly indicate the response location, the cerebellar ataxia group demonstrated normal sequence learning. This dissociation provides an important constraint on the functional domain of the cerebellum in motor learning.
doi:10.1162/jocn.2009.21102
PMCID: PMC4330994  PMID: 18752399
5.  Early Auditory Evoked Potential Is Modulated by Selective Attention and Related to Individual Differences in Visual Working Memory Capacity 
Journal of cognitive neuroscience  2014;26(12):2682-2690.
A growing body of research suggests that the predictive power of working memory (WM) capacity for measures of intellectual aptitude is due to the ability to control attention and select relevant information. Crucially, attentional mechanisms implicated in controlling access to WM are assumed to be domain-general, yet reports of enhanced attentional abilities in individuals with larger WM capacities are primarily within the visual domain. Here, we directly test the link between WM capacity and early attentional gating across sensory domains, hypothesizing that measures of visual WM capacity should predict an individual’s capacity to allocate auditory selective attention. To address this question, auditory ERPs were recorded in a linguistic dichotic listening task, and individual differences in ERP modulations by attention were correlated with estimates of WM capacity obtained in a separate visual change detection task. Auditory selective attention enhanced ERP amplitudes at an early latency (ca. 70–90 msec), with larger P1 components elicited by linguistic probes embedded in an attended narrative. Moreover, this effect was associated with greater individual estimates of visual WM capacity. These findings support the view that domain-general attentional control mechanisms underlie the wide variation of WM capacity across individuals.
doi:10.1162/jocn_a_00684
PMCID: PMC4327887  PMID: 25000526
6.  [No title available] 
PMCID: PMC4080218  PMID: 24650280
7.  [No title available] 
PMCID: PMC4291167  PMID: 24702458
8.  Functional Organization of the Orbitofrontal Cortex 
Journal of cognitive neuroscience  2014;26(7):1347-1362.
Emerging evidence suggests that specific cognitive functions localize to different subregions of orbitofrontal cortex (OFC), but the nature of these functional distinctions remains unclear. One prominent theory, derived from human neuroimaging, proposes that different stimulus valences are processed in separate orbital regions, with medial and lateral OFC processing positive and negative stimuli respectively. Thus far, neurophysiology data have not supported this theory. We attempted to reconcile these accounts by recording neural activity from the full medial-lateral extent of the orbital surface in monkeys receiving rewards and punishments via gain or loss of secondary reinforcement. We found no convincing evidence for valence selectivity in any orbital region. Instead, we report differences between neurons in central OFC and those on the inferior-lateral orbital convexity (IC), in that they encoded different sources of value information provided by the behavioral task. Neurons in IC encoded the value of external stimuli, whereas those in OFC encoded value information derived from the structure of the behavioral task. We interpret these results in light of recent theories of OFC function and propose that these distinctions, not valence selectivity, may shed light on a fundamental organizing principle for value processing in orbital cortex.
doi:10.1162/jocn_a_00573
PMCID: PMC4091960  PMID: 24405106
9.  Feature diagnosticity affects representations of novel and familiar objects 
Journal of cognitive neuroscience  2014;26(12):2735-2749.
Many features can describe a concept, but only some features define a concept in that they enable discrimination of items that are instances of a concept from (similar) items that are not. We refer to this property of some features as feature diagnosticity. Previous work has described the behavioral effects of feature diagnosticity, but there has been little work on explaining why and how these effects arise. In this study, we aimed to understand the impact of feature diagnosticity on concept representations across two complementary experiments. In Experiment 1, we manipulated the diagnosticity of one feature, color, for a set of novel objects that human subjects learned over the course of one week. We report behavioral and neural evidence that diagnostic features are likely to be automatically recruited during remembering. Specifically, individuals activated color-selective regions of ventral temporal cortex (specifically, left fusiform gyrus and left inferior temporal gyrus) when thinking about the novel objects, even though color information was never explicitly probed during the task. Moreover, multiple behavioral and neural measures of the effects of feature diagnosticity were correlated across subjects. In Experiment 2, we examined relative color association in familiar object categories, which varied in feature diagnosticity (fruits and vegetables, household items). Taken together, these results offer novel insights into the neural mechanisms underlying concept representations by demonstrating that automatic recruitment of diagnostic information gives rise to behavioral effects of feature diagnosticity.
doi:10.1162/jocn_a_00661
PMCID: PMC4216244  PMID: 24800630
10.  Experience-Dependent Hemispheric Specialization of Letters and Numbers is Revealed in Early Visual Processing 
Journal of cognitive neuroscience  2014;26(10):2239-2249.
Recent functional magnetic resonance imaging research has demonstrated that letters and numbers are preferentially processed in distinct regions and hemispheres in the visual cortex. In particular, the left visual cortex preferentially processes letters compared to numbers, while the right visual cortex preferentially processes numbers compared to letters. Because letters and numbers are cultural inventions and are otherwise physically arbitrary, such a double dissociation is strong evidence for experiential effects on neural architecture. Here, we use the high temporal resolution of event-related potentials (ERPs) to investigate the temporal dynamics of the neural dissociation between letters and numbers. We show that the divergence between ERP traces to letters and numbers emerges very early in processing. Letters evoked greater N1 waves (latencies 140–170 ms) than did numbers over left occipital channels, while numbers evoked greater N1s than letters over the right, suggesting letters and numbers are preferentially processed in opposite hemispheres early in visual encoding. Moreover, strings of letters, but not single letters, elicited greater P2 ERP waves, (starting around 250 ms) than numbers did over the left hemisphere, suggesting that the visual cortex is tuned to selectively process combinations of letters, but not numbers, further along in the visual processing stream. Additionally, the processing of both of these culturally defined stimulus types differentiated from similar but unfamiliar visual stimulus forms (false fonts) even earlier in the processing stream (the P1 at 100 ms). These findings imply major cortical specialization processes within the visual system driven by experience with reading and mathematics.
doi:10.1162/jocn_a_00621
PMCID: PMC4261939  PMID: 24669789
Letter processing; number processing; ERP; hemispheric specialization
11.  The Necessity of the Medial Temporal Lobe for Statistical Learning 
Journal of cognitive neuroscience  2014;26(8):1736-1747.
The sensory input that we experience is highly patterned, and we are experts at detecting these regularities. Although the extraction of such regularities, or statistical learning (SL), is typically viewed as a cortical process, recent studies have implicated the medial temporal lobe (MTL), including the hippocampus. These studies have employed fMRI, leaving open the possibility that the MTL is involved but not necessary for SL. Here, we examined this issue in a case study of LSJ, a patient with complete bilateral hippocampal loss and broader MTL damage. In Experiments 1 and 2, LSJ and matched control participants were passively exposed to a continuous sequence of shapes, syllables, scenes, or tones containing temporal regularities in the co-occurrence of items. In a subsequent test phase, the control groups exhibited reliable SL in all conditions, successfully discriminating regularities from recombinations of the same items into novel foil sequences. LSJ, however, exhibited no SL, failing to discriminate regularities from foils. Experiment 3 ruled out more general explanations for this failure, such as inattention during exposure or difficulty following test instructions, by showing that LSJ could discriminate which individual items had been exposed. These findings provide converging support for the importance of the MTL in extracting temporal regularities.
doi:10.1162/jocn_a_00578
PMCID: PMC4264662  PMID: 24456393
12.  When Events Change Their Nature: The Neurocognitive Mechanisms underlying Aspectual Coercion 
Journal of cognitive neuroscience  2014;26(9):1905-1917.
The verb ‘pounce’ describes a single, near-instantaneous event. Yet, we easily understand that, “For several minutes the cat pounced…” describes a situation in which multiple pounces occurred, even though this interpretation is not overtly specified by the sentence’s syntactic structure or by any of its individual words—a phenomenon known as ‘aspectual coercion’. Previous psycholinguistic studies have reported processing costs in association with aspectual coercion, but the neurocognitive mechanisms giving rise to these costs remain contentious. Additionally, there is some controversy about whether readers commit to a full interpretation of the event when the aspectual information becomes available, or whether they leave it temporarily underspecified until later in the sentence. Using event-related potentials (ERPs), we addressed these questions in a design that fully crossed context type (punctive, durative, frequentative) with verb type (punctive, durative). We found a late, sustained negativity to punctive verbs in durative contexts, but not in frequentative (e.g. explicitly iterative) contexts. This effect was distinct from the N400 in both its time course and scalp distribution, suggesting that it reflected a different underlying neurocognitive mechanism. We also found that ERPs to durative verbs were unaffected by context type. Together, our results provide strong evidence that neural activity associated with aspectual coercion is driven by the engagement of a morphosyntactically unrealized semantic operator rather than by violations of real-world knowledge, more general shifts in event representation, or event iterativity itself. More generally, our results add to a growing body of evidence that a set of late-onset sustained negativities reflect elaborative semantic processing that goes beyond simply combining the meaning of individual words with syntactic structure to arrive at a final representation of meaning.
doi:10.1162/jocn_a_00638
PMCID: PMC4204801  PMID: 24702457
13.  Electrophysiological Evidence for the Involvement of the Approximate Number System in Preschoolers’ Processing of Spoken Number Words 
Journal of cognitive neuroscience  2014;26(9):1891-1904.
Little is known about the neural underpinnings of number-word comprehension in young children. Here we investigated the neural processing of these words during the crucial developmental window in which children learn their meanings and whether such processing relies on the Approximate Number System (ANS). Event-related potentials (ERPs) were recorded as 3- to 5-year-old children heard the words one, two, three or six while looking at pictures of 1, 2, 3 or 6 objects. The number word was incongruent with the number of visual objects on half the trials, and congruent on the other half. Children’s number-word comprehension predicted their ERP incongruency effects. Specifically, children with the least number-word knowledge did not show any ERP incongruency effects, whereas those with intermediate and high number-word knowledge showed an enhanced, negative-polarity ERP incongruency response (Ninc) over centro-parietal sites from 200–500 ms after the number-word onset. This negativity was followed by an enhanced, positive- polarity incongruency effect (Pinc) that emerged bilaterally over parietal sites at about 700 ms. Moreover, children with the most number-word knowledge showed ratio dependence in the Pinc (larger for greater compared to smaller numerical mismatches), a hallmark of the ANS. A similar modulation of the Pinc from 700–800 ms for number words “one”, “two” and “three” was found in children with intermediate number-word knowledge. These results provide the first neural correlates of spoken number-word comprehension in preschoolers and are consistent with the view that children map number words onto approximate-number representations before they master the verbal count list.
doi:10.1162/jocn_a_00631
PMCID: PMC4228473  PMID: 24702455
14.  Exploring the Roles of Spectral Detail and Intonation Contour in Speech Intelligibility: An fMRI Study 
Journal of cognitive neuroscience  2014;26(8):1748-1763.
The melodic contour of speech forms an important perceptual aspect of tonal and nontonal languages and an important limiting factor on the intelligibility of speech heard through a cochlear implant. Previous work exploring the neural correlates of speech comprehension identified a left-dominant pathway in the temporal lobes supporting the extraction of an intelligible linguistic message, whereas the right anterior temporal lobe showed an overall preference for signals clearly conveying dynamic pitch information. The current study combined modulations of overall intelligibility (through vocoding and spectral inversion) with a manipulation of pitch contour (normal vs. falling) to investigate the processing of spoken sentences in functional MRI. Our overall findings replicate and extend those of Scott et al., whereas greater sentence intelligibility was predominately associated with increased activity in the left STS, the greatest response to normal sentence melody was found right superior temporal gyrus. These data suggest a spatial distinction between brain areas associated with intelligibility and those involved in the processing of dynamic pitch information in speech. By including a set of complexity-matched unintelligible conditions created by spectral inversion, this is additionally the first study reporting a fully factorial exploration of spectrotemporal complexity and spectral inversion as they relate to the neural processing of speech intelligibility. Perhaps surprisingly, there was no evidence for an interaction between the two factors—we discuss the implications for the processing of sound and speech in the dorsolateral temporal lobes.
doi:10.1162/jocn_a_00583
PMCID: PMC4238060  PMID: 24568205
15.  Regions of mid-level human visual cortex sensitive to the global coherence of local image patches 
Journal of cognitive neuroscience  2014;26(8):1764-1774.
The global structural arrangement and spatial layout of the visual environment must be derived from the integration of local signals represented in the lower tiers of the visual system. This interaction between the spatially local and global properties of visual stimulation underlies many of our visual capacities, and how this is achieved in the brain is a central question for visual and cognitive neuroscience. Here, we examine the sensitivity of regions of the posterior human brain to the global coordination of spatially displaced naturalistic image patches. We presented observers with image patches in two circular apertures to the left and right of central fixation, with the patches drawn from either the same (coherent condition) or different (non-coherent condition) extended image. Using functional magnetic resonance imaging (fMRI) at 7T (n = 5), we find that global coherence affected signal amplitude in regions of dorsal mid-level cortex. Furthermore, we find that extensive regions of mid-level visual cortex contained information in their local activity pattern that could discriminate coherent and non-coherent stimuli. These findings indicate that the global coordination of local naturalistic image information has important consequences for the processing in human mid-level visual cortex.
doi:10.1162/jocn_a_00588
PMCID: PMC4074231  PMID: 24564470
16.  Individual Differences in the Balance of GABA to Glutamate in pFC Predict the Ability to Select among Competing Options 
Journal of cognitive neuroscience  2014;26(11):2490-2502.
Individuals vary greatly in their ability to select one item or response when presented with a multitude of options. Here we investigate the neural underpinnings of these individual differences. Using magnetic resonance spectroscopy, we found that the balance of inhibitory versus excitatory neurotransmitters in pFC predicts the ability to select among task-relevant options in two language production tasks. The greater an individual’s concentration of GABA relative to glutamate in the lateral pFC, the more quickly he or she could select a relevant word from among competing options. This outcome is consistent with our computational modeling of this task [Snyder, H. R., Hutchison, N., Nyhus, E., Curran, T., Banich, M. T., O’Reilly, R. C., et al. Neural inhibition enables selection during language processing. Proceedings of the National Academy of Sciences, U.S.A., 107, 16483–16488, 2010], which predicts that greater net inhibition in pFC increases the efficiency of resolving competition among task-relevant options. Moreover, the association with the GABA/glutamate ratio was specific to selection and was not observed for executive function ability in general. These findings are the first to link the balance of excitatory and inhibitory neural transmission in pFC to specific aspects of executive function.
doi:10.1162/jocn_a_00655
PMCID: PMC4182145  PMID: 24742191
17.  White Matter Integrity Predicts Delay Discounting Behavior in 9- to 23-Year-Olds: A Diffusion Tensor Imaging Study 
Journal of cognitive neuroscience  2009;21(7):1406-1421.
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.
doi:10.1162/jocn.2009.21107
PMCID: PMC4203708  PMID: 18767918
18.  Selective Development of Anticorrelated Networks in the Intrinsic Functional Organization of the Human Brain 
Journal of cognitive neuroscience  2013;26(3):501-513.
We examined the normal development of intrinsic functional connectivity of the default network (brain regions typically deactivated for attention-demanding tasks) as measured by resting-state fMRI in children, adolescents, and young adults ages 8–24 years. We investigated both positive and negative correlations and employed analysis methods that allowed for valid interpretation of negative correlations and that also minimized the influence of motion artifacts that are often confounds in developmental neuroimaging. As age increased, there were robust developmental increases in negative correlations, including those between medial pFC (MPFC) and dorsolateral pFC (DLPFC) and between lateral parietal cortices and brain regions associated with the dorsal attention network. Between multiple regions, these correlations reversed from being positive in children to negative in adults. Age-related changes in positive correlations within the default network were below statistical threshold after controlling for motion. Given evidence in adults that greater negative correlation between MPFC and DLPFC is associated with superior cognitive performance, the development of an intrinsic anticorrelation between MPFC and DLPFC may be a marker of the large growth of working memory and executive functions that occurs from childhood to young adulthood.
doi:10.1162/jocn_a_00517
PMCID: PMC4175987  PMID: 24188367
19.  Distinct Contributions of the Magnocellular and Parvocellular Visual Streams to Perceptual Selection 
Journal of cognitive neuroscience  2011;24(1):246-259.
During binocular rivalry, conflicting images presented to the two eyes compete for perceptual dominance, but the neural basis of this competition is disputed. In interocular switch (IOS) rivalry, rival images periodically exchanged between the two eyes generate one of two types of perceptual alternation: 1) a fast, regular alternation between the images that is time-locked to the stimulus switches and has been proposed to arise from competition at lower levels of the visual processing hierarchy, or 2) a slow, irregular alternation spanning multiple stimulus switches that has been associated with higher levels of the visual system. The existence of these two types of perceptual alternation has been influential in establishing the view that rivalry may be resolved at multiple hierarchical levels of the visual system. We varied the spatial, temporal, and luminance properties of IOS rivalry gratings and found, instead, an association between fast, regular perceptual alternations and processing by the magnocellular stream and between slow, irregular alternations and processing by the parvocellular stream. The magnocellular and parvocellular streams are two early visual pathways that are specialized for the processing of motion and form, respectively. These results provide a new framework for understanding the neural substrates of binocular rivalry that emphasizes the importance of parallel visual processing streams, and not only hierarchical organization, in the perceptual resolution of ambiguities in the visual environment.
doi:10.1162/jocn_a_00121
PMCID: PMC4176614  PMID: 21861685
20.  Roles of Medial Prefrontal Cortex and Orbitofrontal Cortex in Self-evaluation 
Journal of cognitive neuroscience  2010;22(9):2108-2119.
Empirical investigations of the relation of frontal lobe function to self-evaluation have mostly examined the evaluation of abstract qualities in relation to self versus other people. The present research furthers our understanding of frontal lobe involvement in self-evaluation by examining two processes that have not been widely studied by neuroscientists: on-line self-evaluations and correction of systematic judgment errors that influence self-evaluation. Although people evaluate their abstract qualities, it is equally important that perform on-line evaluations to assess the success of their behavior in a particular situation. In addition, self-evaluations of task performance are sometimes overconfident because of systematic judgment errors. What role do the neural regions associated with abstract self-evaluations and decision bias play in on-line evaluation and self-evaluation bias? In this fMRI study, self-evaluation in two reasoning tasks was examined; one elicited overconfident self-evaluations of performance because of salient but misleading aspects of the task and the other was free from misleading aspects. Medial PFC (mPFC), a region associated with self-referential processing, was generally involved in on-line self-evaluations but not specific to accurate or overconfident evaluation. Orbitofrontal cortex (OFC) activity, a region associated with accurate nonsocial judgment, negatively predicted individual differences in overconfidence and was negatively associated with confidence level for incorrect trials.
doi:10.1162/jocn.2009.21359
PMCID: PMC4159715  PMID: 19925187
21.  The corollary discharge in humans is related to synchronous neural oscillations 
Journal of cognitive neuroscience  2010;23(10):2892-2904.
How do animals distinguish between sensations coming from external sources and those resulting from their own actions? A corollary discharge system has evolved that involves the transmission of a copy of motor commands to sensory cortex, where the expected sensation is generated. Through this mechanism, sensations are tagged as coming from self, and responsiveness to them is minimized. The present study investigated whether neural phase synchrony between motor command and auditory cortical areas is related to the suppression of the auditory cortical response. We recorded electrocorticograms from the human brain during a vocalizing/listening task. Neural phase synchrony between Broca’s area and auditory cortex in the gamma band (35 Hz to ~50 Hz) in the 50 ms time window preceding speech onset was greater during vocalizing than listening to a playback of the same spoken sounds. Because pre-speech neural synchrony was correlated (r = −0.83, p = 0.006) with the subsequent suppression of the auditory cortical response to the spoken sound, we hypothesize that phase synchrony in the gamma band between Broca’s area and auditory cortex is the neural instantiation of the transmission of a copy of motor commands. We suggest that neural phase synchrony of gamma frequencies may contribute to transmission of corollary discharges in humans.
doi:10.1162/jocn.2010.21589
PMCID: PMC4155919  PMID: 20946054
22.  Anticipating conflict facilitates controlled stimulus-response selection 
Journal of cognitive neuroscience  2009;21(8):1461-1472.
Cognitive control can be triggered in reaction to previous conflict, as suggested by the finding of sequential effects in conflict tasks. Can control also be triggered proactively by presenting cues predicting conflict (‘proactive control’)? We exploited the high temporal resolution of event-related potentials (ERPs) and controlled for sequential effects to ask whether proactive control based on anticipating conflict modulates neural activity related to cognitive control, as may be predicted from the conflict-monitoring model. ERPs associated with conflict detection (N2) were measured during a cued flanker task. Symbolic cues were either informative or neutral with respect to whether the target involved conflicting or congruent responses. Sequential effects were controlled by analysing the congruency of the previous trial. The results showed that cuing conflict facilitated conflict resolution and reduced the N2 latency. Other potentials (frontal N1 and P3) were also modulated by cuing conflict. Cuing effects were most evident after congruent than after incongruent trials. This interaction between cuing and sequential effects suggests neural overlap between the control networks triggered by proactive and reactive signals. This finding clarifies why previous neuroimaging studies, in which reactive sequential effects were not controlled, have rarely found anticipatory effects upon conflict-related activity. Finally, the high temporal resolution of ERPs was critical to reveal a temporal modulation of conflict detection by proactive control. This novel finding suggests that anticipating conflict speeds up conflict detection and resolution. Recent research suggests that this anticipatory mechanism may be mediated by pre-activation of the ACC during the preparatory interval.
doi:10.1162/jocn.2009.21136
PMCID: PMC4152723  PMID: 18823248
cognitive control; cuing conflict; event-related potential (ERP); N2; error-related negativity (ERN); anterior cingulate cortex (ACC)
23.  Long-Term Memories Bias Sensitivity and Target Selection in Complex Scenes 
Journal of cognitive neuroscience  2012;24(12):2281-2291.
In everyday situations we often rely on our memories to find what we are looking for in our cluttered environment. Recently, we developed a new experimental paradigm to investigate how long-term memory (LTM) can guide attention, and showed how the pre-exposure to a complex scene in which a target location had been learned facilitated the detection of the transient appearance of the target at the remembered location (Summerfield, Lepsien, Gitelman, Mesulam, & Nobre, 2006; Summerfield, Rao, Garside, & Nobre, 2011). The present study extends these findings by investigating whether and how LTM can enhance perceptual sensitivity to identify targets occurring within their complex scene context. Behavioral measures showed superior perceptual sensitivity (d′) for targets located in remembered spatial contexts. We used the N2pc event-related potential to test whether LTM modulated the process of selecting the target from its scene context. Surprisingly, in contrast to effects of visual spatial cues or implicit contextual cueing, LTM for target locations significantly attenuated the N2pc potential. We propose that the mechanism by which these explicitly available LTMs facilitate perceptual identification of targets may differ from mechanisms triggered by other types of top-down sources of information.
doi:10.1162/jocn_a_00294
PMCID: PMC4152726  PMID: 23016670
24.  A Surface-based Analysis of Language Lateralization and Cortical Asymmetry 
Journal of cognitive neuroscience  2013;25(9):1477-1492.
Among brain functions, language is one of the most lateralized. Cortical language areas are also some of the most asymmetrical in the brain. An open question is whether the asymmetry in function is linked to the asymmetry in anatomy. To address this question, we measured anatomical asymmetry in 34 participants shown with fMRI to have language dominance of the left hemisphere (LLD) and 21 participants shown to have atypical right hemisphere dominance (RLD). All participants were healthy and left-handed, and most (80%) were female. Gray matter (GM) volume asymmetry was measured using an automated surface-based technique in both ROIs and exploratory analyses. In the ROI analysis, a significant difference between LLD and RLD was found in the insula. No differences were found in planum temporale (PT), pars opercularis (POp), pars triangularis (PTr), or Heschl’s gyrus (HG). The PT, POp, insula, and HG were all significantly left lateralized in both LLD and RLD participants. Both the positive and negative ROI findings replicate a previous study using manually labeled ROIs in a different cohort [Keller, S. S., Roberts, N., Garcia-Finana, M., Mohammadi, S., Ringelstein, E. B., Knecht, S., et al. Can the language-dominant hemisphere be predicted by brain anatomy? Journal of Cognitive Neuroscience, 23, 2013–2029, 2011]. The exploratory analysis was accomplished using a new surface-based registration that aligns cortical folding patterns across both subject and hemisphere. A small but significant cluster was found in the superior temporal gyrus that overlapped with the PT. A cluster was also found in the ventral occipitotemporal cortex corresponding to the visual word recognition area. The surface-based analysis also makes it possible to disentangle the effects of GM volume, thickness, and surface area while removing the effects of curvature. For both the ROI and exploratory analyses, the difference between LLD and RLD volume laterality was most strongly driven by differences in surface area and not cortical thickness. Overall, there were surprisingly few differences in GM volume asymmetry between LLD and RLD indicating that gross morphometric asymmetry is only subtly related to functional language laterality.
doi:10.1162/jocn_a_00405
PMCID: PMC3767398  PMID: 23701459
25.  Prefrontal cortex neurons reflect categorical decisions about ambiguous stimuli 
Journal of cognitive neuroscience  2014;26(6):1283-1291.
We examined whether prefrontal cortex (PFC) neuron activity reflects categorical decisions in monkeys categorizing ambiguous stimuli. A morphing system was used to systematically vary stimulus shape and precisely define category boundaries. Ambiguous stimuli were centered on a category boundary, i.e., they were a mix of 50% of two prototypes and therefore had no category information, so monkeys guessed at their category membership. We found that the monkey's trial-by-trial decision about the category membership of an ambiguous image was reflected in PFC activity. Activity to the same ambiguous image differed significantly depending on which category the monkey had assigned it to. This effect only occurred when that scheme was behaviorally relevant. These indicate that PFC activity reflects categorical decisions.
doi:10.1162/jocn_a_00568
PMCID: PMC4034266  PMID: 24405188
prefrontal cortex; categorization; monkey; flexibility; goal directed; object vision

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