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Showing papers in "Journal of Cognitive Neuroscience in 2000"
Journal Article•10.1162/08989290051137585•
Imaging Cognition II: An Empirical Review of 275 PET and fMRI Studies

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Roberto Cabeza1, Lars Nyberg2•
University of Alberta1, Umeå University2
01 Jan 2000-Journal of Cognitive Neuroscience
TL;DR: Analysis of regional activations across cognitive domains suggested that several brain regions, including the cerebellum, are engaged by a variety of cognitive challenges.
Abstract: 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.

3,809 citations

Journal Article•10.1162/08989290051137549•
Mental Imagery of Faces and Places Activates Corresponding Stiimulus-Specific Brain Regions

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K. M. O'Craven, N. Kanwisher
01 Nov 2000-Journal of Cognitive Neuroscience
TL;DR: Evidence that imagery and perception share common processing mechanisms is strengthened, and it is demonstrated that the specific brain regions activated during mental imagery depend on the content of the visual image.
Abstract: What happens in the brain when you conjure up a mental image in your mind's eye? We tested whether the particular regions of extrastriate cortex activated during mental imagery depend on the content of the image. Using functional magnetic resonance imaging (fMRRI), we demonstrated selective activation within a region of cortex specialized for face perception during mental imagery of faces, and selective activation within a place-selective cortical region during imagery of places. In a further study, we compared the activation for imagery and perception in these regions, and found greater response magnitudes for perception than for imagery of the same items. Finally, we found that it is possible to determine the content of single cognitive events from an inspection of the fMRI data from individual imagery trials. These findings strengthen evidence that imagery and perception share common processing mechanisms, and demonstrate that the specific brain regions activated during mental imagery depend on the content of the visual image.

1,083 citations

Journal Article•10.1162/089892900562552•
Performance on Indirect Measures of Race Evaluation Predicts Amygdala Activation

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Elizabeth A. Phelps1, Kevin J. O'Connor2, William A. Cunningham3, E. Sumie Funayama3, J. Christopher Gatenby3, John C. Gore3, Mahzarin R. Banaji3 •
New York University1, Massachusetts Institute of Technology2, Yale University3
01 Sep 2000-Journal of Cognitive Neuroscience
TL;DR: The results suggest that amygdala and behavioral responses to Black-versus-White faces in White subjects reflect cultural evaluations of social groups modified by individual experience.
Abstract: We used fMRI to explore the neural substrates involved in the unconscious evaluation of Black and White social groups. Specifically, we focused on the amygdala, a subcortical structure known to play a role in emotional learning and evaluation. In Experiment 1, White American subjects observed faces of unfamiliar Black and White males. The strength of amygdala activation to Black-versus-White faces was correlated with two indirect (unconscious) measures of race evaluation (Implicit Association Test lIATr and potentiated startle), but not with the direct (conscious) expression of race attitudes. In Experiment 2, these patterns were not obtained when the stimulus faces belonged to familiar and positively regarded Black and White individuals. Together, these results suggest that amygdala and behavioral responses to Black-versus-White faces in White subjects reflect cultural evaluations of social groups modified by individual experience.

1,078 citations

Journal Article•10.1162/089892900561814•
Age Differences in the Frontal Lateralization of Verbal and Spatial Working Memory Revealed by PET

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Patricia A. Reuter-Lorenz1, John Jonides1, Edward E. Smith1, Alan A. Hartley2, Andrea Miller1, Christina Marshuetz1, Robert A. Koeppe1 •
University of Michigan1, Scripps College2
01 Jan 2000-Journal of Cognitive Neuroscience
TL;DR: Positron emission tomography was used to investigate verbal and spatial short-term storage in older and younger adults to consider several mechanisms that could account for these age differences including the possibility that bilateral activation reflects recruitment to compensate for neural decline.
Abstract: Age-related decline in working memory figures prominently in theories of cognitive aging. However, the effects of aging on the neural substrate of working memory are largely unknown. Positron emission tomography (PET) was used to investigate verbal and spatial short-term storage (3 sec) in older and younger adults. Previous investigations with younger subjects performing these same tasks have revealed asymmetries in the lateral organization of verbal and spatial working memory. Using volume of interest (VOI) analyses that specifically compared activation at sites identified with working memory to their homologous twin in the opposite hemisphere, we show pronounced age differences in this organization, particularly in the frontal lobes: In younger adults, activation is predominantly left lateralized for verbal working memory, and right lateralized for spatial working memory, whereas older adults show a global pattern of anterior bilateral activation for both types of memory. Analyses of frontal subregions indicate that several underlying patterns contribute to global bilaterality in older adults: most notably, bilateral activation in areas associated with rehearsal, and paradoxical laterality in dorsolateral prefrontal sites (DLPFC; greater left activation for spatial and greater right activation for verbal). We consider several mechanisms that could account for these age differences including the possibility that bilateral activation reflects recruitment to compensate for neural decline.

1,072 citations

Journal Article•10.1162/089892900562417•
Brain Areas Involved in Perception of Biological Motion

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Emily D. Grossman1, M. Donnelly1, Ronald R. Price1, David R. Pickens1, Victoria L. Morgan1, G. Neighbor1, Randolph Blake1 •
Vanderbilt University1
01 Sep 2000-Journal of Cognitive Neuroscience
TL;DR: Consistent with earlier neuroimaging and single-unit studies, this pattern of results points to the existence of neural mechanisms specialized for analysis of the kinematics defining biological motion.
Abstract: These experiments use functional magnetic resonance imaging (fMRI) to reveal neural activity uniquely associated with perception of biological motion. We isolated brain areas activated during the viewing of point-light figures, then compared those areas to regions known to be involved in coherent-motion perception and kinetic-boundary perception. Coherent motion activated a region matching previous reports of human MT/MST complex located on the temporo-parieto-occipital junction. Kinetic boundaries activated a region posterior and adjacent to human MT previously identified as the kinetic-occipital (KO) region or the lateral-occipital (LO) complex. The pattern of activation during viewing of biological motion was located within a small region on the ventral bank of the occipital extent of the superior-temporal sulcus (STS). This region is located lateral and anterior to human MT/MST, and anterior to KO. Among our observers, we localized this region more frequently in the right hemisphere than in the left. This was true regardless of whether the point-light figures were presented in the right or left hemifield. A small region in the medial cerebellum was also active when observers viewed biological-motion sequences. Consistent with earlier neuroimaging and single-unit studies, this pattern of results points to the existence of neural mechanisms specialized for analysis of the kinematics defining biological motion.

996 citations

Journal Article•10.1162/089892900562165•
The Fusiform Face Area is Part of a Network that Processes Faces at the Individual Level

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Isabel Gauthier1, Michael J. Tarr2, J Moylan3, Pawel Skudlarski3, John C. Gore3, Adam W. Anderson3 •
Vanderbilt University1, Brown University2, Yale University3
01 May 2000-Journal of Cognitive Neuroscience
TL;DR: It is suggested that face- selective areas may be involved in the perception of faces at the individual level, whereas letter-selective regions may be tuning themselves to font information in order to recognize letters more efficiently.
Abstract: According to modular models of cortical organization, many areas of the extrastriate cortex are dedicated to object categories. These models often assume an early processing stage for the detection of category membership. Can functional imaging isolate areas responsible for detection of members of a category, such as faces or letters? We consider whether responses in three different areas (two selective for faces and one selective for letters) support category detection. Activity in these areas habituates to the repeated presentation of one exemplar more than to the presentation of different exemplars of the same category, but only for the category for which the area is selective. Thus, these areas appear to play computational roles more complex than detection, processing stimuli at the individual level. Drawing from prior work, we suggest that face-selective areas may be involved in the perception of faces at the individual level, whereas letter-selective regions may be tuning themselves to font information in order to recognize letters more efficiently.

884 citations

Journal Article•10.1162/089892900561959•
I. The Neurocognitive Profile of Williams Syndrome: A Complex Pattern of Strengths and Weaknesses

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Ursula Bellugi1, Liz Lichtenberger1, Wendy Jones1, Zona Lai1, Marie St. George2 •
Salk Institute for Biological Studies1, University of California, San Diego2
01 Mar 2000-Journal of Cognitive Neuroscience
TL;DR: The rare, genetically based disorder, Williams syndrome, produces a constellation of distinctive cognitive, neuroanatomical, and electrophysiological features which are explored through the series of studies reported here.
Abstract: The rare, genetically based disorder, Williams syndrome (WMS), produces a constellation of distinctive cognitive, neuroanatomical, and electrophysiological features which we explore through the series of studies reported here. In this paper, we focus primarily on the cognitive characteristics of WMS and begin to forge links among these characteristics, the brain, and the genetic basis of the disorder. The distinctive cognitive profile of individuals with WMS includes relative strengths in language and facial processing and profound impairment in spatial cognition. The cognitive profile of abilities, including what is ‘typical’ for individuals with WMS is discussed, but we also highlight areas of variability across the group of individuals with WMS that we have studied. Although the overall cognitive abilities (IQs) of individuals with WMS are typically in the mild-to-moderate range of mental retardation, the peaks and valleys within different cognitive domains make this syndrome especially intriguing to study across levels. Understanding the brain basis (and ultimately the genetic basis) for higher cognitive functioning is the goal we have begun to undertake with this line of interdisciplinary research.

742 citations

Journal Article•10.1162/089892900562075•
Visual Discrimination Task Improvement: A Multi-Step Process Occurring During Sleep

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Robert Stickgold1, Dana Whidbee1, Beth Schirmer1, Vipul Patel1, J. Allan Hobson1 •
Harvard University1
01 Mar 2000-Journal of Cognitive Neuroscience
TL;DR: The results suggest that, in the case of this visual discrimination task, both SWS and REM are required to consolidate experience-dependent neuronal changes into a form that supports improved task performance.
Abstract: Performance on a visual discrimination task shows long-term improvement after a single training session. When tested within 24 hr of training, improvement was not observed unless subjects obtained at least 6 hr of post-training sleep prior to retesting, in which case improvement was proportional to the amount of sleep in excess of 6 hr. For subjects averaging 8 hr of sleep, overnight improvement was proportional to the amount of slow wave sleep (SWS) in the first quarter of the night, as well as the amount of rapid eye movement sleep (REM) in the last quarter. REM during the intervening 4 hr did not appear to contribute to improvement. A two-step process, modeling throughput as the product of the amount of early SWS and late REM, accounts for 80 percent of intersubject variance. These results suggest that, in the case of this visual discrimination task, both SWS and REM are required to consolidate experience-dependent neuronal changes into a form that supports improved task performance.

604 citations

Journal Article•10.1162/089892900562183•
Brain Indices of Music Processing: Nonmusicians are Musical

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Stefan Koelsch1, Tomas Gunter1, Angela D. Friederici1, Erich Schröger2•
Max Planck Society1, Leipzig University2
01 May 2000-Journal of Cognitive Neuroscience
TL;DR: The amplitudes of both early and late negativities were found to be sensitive to the degree of musical expectancy induced by the preceding harmonic context, and to the probability for deviant acoustic events.
Abstract: Only little systematic research has examined event-related brain potentials (ERPs) elicited by the cognitive processing of music. The present study investigated how music processing is influenced by a preceding musical context, affected by the task relevance of unexpected chords, and influenced by the degree and the probability of violation. Four experiments were conducted in which "nonmusicians" listened to chord sequences, which infrequently contained a chord violating the sound expectancy of listeners. Integration of in-key chords into the musical context was reflected as a late negative-frontal deflection in the ERPs. This negative deflection declined towards the end of a chord sequence, reflecting normal buildup of musical context. Brain waves elicited by chords with unexpected notes revealed two ERP effects: an early right-hemispheric preponderant-anterior negativity, which was taken to reflect the violation of sound expectancy; and a late bilateral-frontal negativity. The late negativity was larger compared to in-key chords and taken to reflect the higher degree of integration needed for unexpected chords. The early right-anterior negativity (ERAN) was unaffected by the task relevance of unexpected chords. The amplitudes of both early and late negativities were found to be sensitive to the degree of musical expectancy induced by the preceding harmonic context, and to the probability for deviant acoustic events. The employed experimental design opens a new field for the investigation of music processing. Results strengthen the hypothesis of an implicit musical ability of the human brain.

561 citations

Journal Article•10.1162/089892900562606•
Hemispheric Asymmetries for Whole-Based and Part-Based Face Processing in the Human Fusiform Gyrus

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Bruno Rossion1, Laurence Dricot1, Anne Devolder1, J.-M Bodart1, Marc Crommelinck1, Beatrice de Gelder1, Richard Zoontjes2 •
Université catholique de Louvain1, Tilburg University2
01 Sep 2000-Journal of Cognitive Neuroscience
TL;DR: The present PET study identifies the anatomical localization of these effects in well-defined regions of the middle fusiform gyri of both hemispheres as a double dissociation between two modes of face processing.
Abstract: Behavioral studies indicate a right hemisphere advantage for processing a face as a whole and a left hemisphere superiority for processing based on face features. The present PET study identifies the anatomical localization of these effects in well-defined regions of the middle fusiform gyri of both hemispheres. The right middle fusiform gyrus, previously described as a face-specific region, was found to be more activated when matching whole faces than face parts whereas this pattern of activity was reversed in the left homologous region. These lateralized differences appeared to be specific to faces since control objects processed either as wholes or parts did not induce any change of activity within these regions. This double dissociation between two modes of face processing brings new evidence regarding the lateralized localization of face individualization mechanisms in the human brain.

475 citations

Journal Article•10.1162/08989290051137521•
fMRI Studies of Stroop Tasks Reveal Unique Roles of Anterior and Posterior Brain Systems in Attentional Selection

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Marie T. Banich1, Michael P. Milham1, Ruthann Atchley1, Neal J. Cohen1, Andrew G. Webb1, Tracey Wszalek1, Arthur F. Kramer1, Zhei-Pei Liang1, Alexander Wright1, Joel I. Shenker1, Richard L. Magin1 •
University of Illinois at Urbana–Champaign1
01 Nov 2000-Journal of Cognitive Neuroscience
TL;DR: The results of this second experiment indicate that attentional selection in tasks such as the Stroop task, which contain multiple potential sources of relevant information, acts more by modulating the processing of task-irrelevant information than by modulated processing oftask-relevant information.
Abstract: The brain's attentional system identifies and selects information that is task-relevant while ignoring information that is task-irrelevant. In two experiments using functional magnetic resonance imaging, we examined the effects of varying task-relevant information compared to task-irrelevant information. In the first experiment, we compared patterns of activation as attentional demands were increased for two Stroop tasks that differed in the task-relevant information, but not the task-irrelevant information: a color–word task and a spatial–word task. Distinct subdivisions of dorsolateral prefrontal cortex and the precuneus became activated for each task, indicating differential sensitivity of these regions to task-relevant information (e.g., spatial information vs. color). In the second experiment, we compared patterns of activation with increased attentional demands for two Stroop tasks that differed in task-irrelevant information, but not task-relevant information: a color–word task and color–object task. Little differentiation in activation for dorsolateral prefrontal and precuneus regions was observed, indicating a relative insensitivity of these regions to task-irrelevant information. However, we observed a differentiation in the pattern of activity for posterior regions. There were unique areas of activation in parietal regions for the color–word task and in occipitotemporal regions for the color–object task. No increase in activation was observed in regions responsible for processing the perceptual attribute of color. The results of this second experiment indicate that attentional selection in tasks such as the Stroop task, which contain multiple potential sources of relevant information (e.g., the word vs. its ink color), acts more by modulating the processing of task-irrelevant information than by modulating processing of task-relevant information.
Journal Article•10.1162/089892900562273•
Neuroanatomical Substrates of Arabic Number Processing, Numerical Comparison, and Simple Addition: A PET Study

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Mauro Pesenti1, Marc Thioux1, Xavier Seron1, Anne De Volder1•
Université catholique de Louvain1
01 May 2000-Journal of Cognitive Neuroscience
TL;DR: Comparison and simple addition fact retrieval revealed a fronto-parietal network involving mainly the left intraparietal sulcus, the superior parietal lobule and the precentral gyrus, whereas addition also activated the orbito-frontal areas and the anterior insula in the right hemisphere.
Abstract: Positron emission tomography was used to localize the cerebral networks specifically involved in three basic numerical processes: arabic numeral processing, numerical magnitude comparison, and retrieval of simple addition facts. Relative cerebral blood flow changes were measured while normal volunteers were resting with eyes closed, making physical judgment on nonnumerical characters or arabic digits, comparing, or adding the same digits. Processing arabic digits bilaterally produced a large nonspecific activation of occipito-parietal areas, as well as a specific activation of the right anterior insula. Comparison and simple addition fact retrieval revealed a fronto-parietal network involving mainly the left intraparietal sulcus, the superior parietal lobule and the precentral gyrus. Comparison also activated, but to a lesser extent, the right superior parietal lobe, whereas addition also activated the orbito-frontal areas and the anterior insula in the right hemisphere. Implications for current anatomo-functional models of numerical cognition are drawn.
Journal Article•10.1162/089892900564055•
The Representation of Objects in the Human Occipital and Temporal Cortex

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Alumit Ishai1, Leslie G. Ungerleider1, Alex Martin1, James V. Haxby1•
National Institutes of Health1
01 Nov 2000-Journal of Cognitive Neuroscience
TL;DR: It is proposed that the representation of objects in the ventral visual pathway, including both occipital and temporal regions, is not restricted to small, highly selective patches of cortex but, instead, is a distributed representation of information about object form within this distributed system.
Abstract: Recently, we identified, using fMRI, three bilateral regions in the ventral temporal cortex that responded preferentially to faces, houses, and chairs [Ishai, A., Ungerleider, L. G., Martin, A., Schouten, J. L., & Haxby, J. V. (1999). Distributed representation of objects in the human ventral visual pathway. Proceedings of the National Academy of Sciences, U.S.A., 96, 9379-9384]. Here, we report differential patterns of activation, similar to those seen in the ventral temporal cortex, in bilateral regions of the ventral occipital cortex. We also found category-related responses in the dorsal occipital cortex and in the superior temporal sulcus. Moreover, rather than activating discrete, segregated areas, each category was associated with its own differential pattern of response across a broad expanse of cortex. The distributed patterns of response were similar across tasks (passive viewing, delayed matching) and presentation formats (photographs, line drawings). We propose that the representation of objects in the ventral visual pathway, including both occipital and temporal regions, is not restricted to small, highly selective patches of cortex but, instead, is a distributed representation of information about object form. Within this distributed system, the representation of faces appears to be less extensive as compared to the representations of nonface objects.
Journal Article•10.1162/089892900562228•
Sex Steroids Modify Working Memory

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Jeri S. Janowsky1, Bambi Chavez1, Eric S. Orwoll1•
Oregon Health & Science University1
01 May 2000-Journal of Cognitive Neuroscience
TL;DR: The results suggest that sex steroids can modulate working memory in men and can act as modulators of cognition throughout life.
Abstract: In the last ten years, numerous mechanisms by which sex steroids modify cortical function have been described. For example, estrogen replacement improves verbal memory in women, and animal studies have shown effects of estrogen on hippocampal synaptogenesis and function. Little is known about sex steroid effects on other aspects of memory, such as frontal lobe-mediated working memory. We examined the relationships between working memory and sex steroid concentrations and whether sex steroid supplementation would modify age-related loss of working memory in older men and women. Before hormone supplementation, working memory, tested with the Subject Ordered Pointing Test (SOP), was worse in older subjects than younger subjects, and there was no evidence of gender differences at either age. Testosterone supplementation improved working memory in older men, but a similar enhancement of working memory was not found in older women supplemented with estrogen. In men, testosterone and estrogen effects were reciprocal - with better working memory related to a higher testosterone to estrogen ratio. These results suggest that sex steroids can modulate working memory in men and can act as modulators of cognition throughout life.
Journal Article•10.1162/08989290051137558•
Imaginal, Semantic, and Surface-Level Processing of Concrete and Abstract Words: An Electrophysiological Investigation

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W. Caroline West1, Phillip J. Holcomb2•
Harvard University1, Tufts University2
01 Nov 2000-Journal of Cognitive Neuroscience
TL;DR: The results of this study support the extended dual-coding hypothesis that superior associative connections and the use of mental imagery both contribute to processing advantages for concrete words over abstract words.
Abstract: Words representing concrete concepts are processed more quickly and efficiently than words representing abstract concepts. Concreteness effects have also been observed in studies using event-related brain potentials (ERPs). The aim of this study was to examine concrete and abstract words using both reaction time (RT) and ERP measurements to determine (1) at what point in the stream of cognitive processing concreteness effects emerge and (2) how different types of cognitive operations influence these concreteness effects. Three groups of subjects performed a sentence verification task in which the final word of each sentence was concrete or abstract. For each group the truthfulness judgment required either (1) image generation, (2) semantic decision, or (3) evaluation of surface characteristics. Concrete and abstract words produced similar RTs and ERPs in the surface task, suggesting that postlexical semantic processing is necessary to elicit concreteness effects. In both the semantic and imagery tasks, RTs were shorter for concrete than for abstract words. This difference was greatest in the imagery task. Also, in both of these tasks concrete words elicited more negative ERPs than abstract words between 300 and 550 msec (N400). This effect was widespread across the scalp and may reflect activation in a linguistic semantic system common to both concrete and abstract words. ERPs were also more negative for concrete than abstract words between 550 and 800 msec. This effect was more frontally distributed and was most evident in the imagery task. We propose that this later anterior effect represents a distinct ERP component (N700) that is sensitive to the use of mental imagery. The N700 may reflect the a access of specific characteristics of the imaged item or activation in a working memory system specific to mental imagery. These results also support the extended dual-coding hypothesis that superior associative connections and the use of mental imagery both contribute to processing advantages for concrete words over abstract words.
Journal Article•10.1162/089892900562138•
Common and Distinct Neural Substrates for Pragmatic, Semantic, and Syntactic Processing of Spoken Sentences: An fMRI Study

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Gina R. Kuperberg, Philip McGuire, Edward T. Bullmore1, Mick Brammer, Sophia Rabe-Hesketh, Ian C. Wright, David J. Lythgoe, Steven Williams, A.S. David •
University of Cambridge1
01 Mar 2000-Journal of Cognitive Neuroscience
TL;DR: It is suggested that the left-inferior-temporal/fusiform gyrus may play a key role in using pragmatic, semantic, and subcategorical information to construct a higher representation of meaning of sentences.
Abstract: Extracting meaning from speech requires the use of pragmatic, semantic, and syntactic information. A central question is: Does the processing of these different types of linguistic information have common or distinct neuroanatomical substrates? We addressed this issue using functional magnetic resonance imaging (fMRI) to measure neural activity when subjects listened to spoken normal sentences contrasted with sentences that had either (A) pragmatical, (B) semantic (selection restriction), or (C) syntactic (subcategorical) violations sentences. All three contrasts revealed robust activation of the left-inferior-temporal/fusiform gyrus. Activity in this area was also observed in a combined analysis of all three experiments, suggesting that it was modulated by all three types of linguistic violation. Planned statistical comparisons between the three experiments revealed (1) a greater difference between conditions in activation of the left-superior-temporal gyrus for the pragmatic experiment than the semantic/syntactic experiments; (2) a greater difference between conditions in activation of the right-superior and middle-temporal gyrus in the semantic experiment than in the syntactic experiment; and (3) no regions activated to a greater degree in the syntactic experiment than in the semantic experiment. These data show that, while left- and right-superior-temporal regions may be differentially involved in processing pragmatic and lexico-semantic information within sentences, the left-inferior-temporal/fusiform gyrus is involved in processing all three types of linguistic information. We suggest that this region may play a key role in using pragmatic, semantic (selection restriction), and subcategorical information to construct a higher representation of meaning of sentences.
Journal Article•10.1162/089892900562390•
Electrophysiological Correlates of Phonological Processing: A Cross-linguistic Study

[...]

Ghislaine Dehaene-Lambertz1, Emmanuel Dupoux1, Ariel Gout1•
School for Advanced Studies in the Social Sciences1
01 Jul 2000-Journal of Cognitive Neuroscience
TL;DR: The results suggest that the impact of phonotactics takes place early in speech processing and support models of speech perception, which postulate that the input signal is directly parsed into the native language phonological format.
Abstract: It is well known that speech perception is deeply affected by the phoneme categories of the native language. Recent studies have found that phonotactics, i.e., constraints on the cooccurrence of phonemes within words, also have a considerable impact on speech perception routines. For example, Japanese does not allow (nonnasal) coda consonants. When presented with stimuli that violate this constraint, as in / ebzo/, Japanese adults report that they hear a /u/ between consonants, i.e., /ebuzo/. We examine this phenomenon using event-related potentials (ERPs) on French and Japanese participants in order to study how and when the phonotactic properties of the native language affect speech perception routines. Trials using four similar precursor stimuli were presented followed by a test stimulus that was either identical or different depending on the presence or absence of an epenthetic vowel /u/ between two consonants (e.g., "ebuzo ebuzo ebuzo- ebzo"). Behavioral results confirm that Japanese, unlike French participants, are not able to discriminate between identical and deviant trials. In ERPs, three mismatch responses were recorded in French participants. These responses were either absent or significantly weaker for Japanese. In particular, a component similar in latency and topography to the mismatch negativity (MMN) was recorded for French, but not for Japanese participants. Our results suggest that the impact of phonotactics takes place early in speech processing and support models of speech perception, which postulate that the input signal is directly parsed into the native language phonological format. We speculate that such a fast computation of a phonological representation should facilitate lexical access, especially in degraded conditions.
Journal Article•10.1162/08989290051137567•
Auditory Cortex Accesses Phonological Categories: An MEG Mismatch Study

[...]

Colin Phillips1, Thomas Pellathy1, Alec Marantz2, Elron Yellin2, Kenneth Wexler2, David Poeppel3, Martha McGinnis4, Timothy P.L. Roberts5 •
University of Delaware1, Massachusetts Institute of Technology2, University of Maryland, College Park3, University of Calgary4, University of California, San Francisco5
01 Nov 2000-Journal of Cognitive Neuroscience
TL;DR: The studies presented here use an adapted oddball paradigm to show evidence that representations of discrete phonological categories are available to the human auditory cortex, and demonstrate the all-or-nothing property of phonological category membership.
Abstract: The studies presented here use an adapted oddball paradigm to show evidence that representations of discrete phonological categories are available to the human auditory cortex. Brain activity was recorded using a 37-channel biomagnetometer while eight subjects listened passively to synthetic speech sounds. In the phonological condition, which contrasted stimuli from an acoustic /dae/-/tae/ continuum, a magnetic mismatch field (MMF) was elicited in a sequence of stimuli in which phonological categories occurred in a many-to-one ratio, but no acoustic many-to-one ratio was present. In order to isolate the contribution of phonological categories to the MMF responses, the acoustic parameter of voice onset time, which distinguished standard and deviant stimuli, was also varied within the standard and deviant categories. No MMF was elicited in the acoustic condition, in which the acoustic distribution of stimuli was identical to the first experiment, but the many-to-one distribution of phonological categories was removed. The design of these studies makes it possible to demonstrate the all-or-nothing property of phonological category membership. This approach contrasts with a number of previous studies of phonetic perception using the mismatch paradigm, which have demonstrated the graded property of enhanced acoustic discrimination at or near phonetic category boundaries.
Journal Article•10.1162/089892900562002•
VI. Genome Structure and Cognitive Map of Williams Syndrome

[...]

Julie R. Korenberg1, Xiao-Ning Chen1, Hamao Hirota1, Zona Lai2, Ursula Bellugi2, Dennis Burian3, Bruce A. Roe3, Rumiko Matsuoka •
University of California, Los Angeles1, Salk Institute for Biological Studies2, University of Oklahoma3
01 Mar 2000-Journal of Cognitive Neuroscience
TL;DR: An integrated physical, genetic, and transcriptional map of the WMS and flanking regions is generated using multicolor metaphase and interphase fluorescence in situ hybridization of bacterial artificial chromosomes and P1 artificial chromosomes, BAC end sequencing, PCR gene marker and microsatellite, large-scale sequencing, cDNA library, and database analyses, which establish regions and consequent gene candidates for WMS features including mental retardation, hypersociability, and facial features.
Abstract: Williams syndrome (WMS) is a most compelling model of human cognition, of human genome organization, and of evolution. Due to a deletion in chromosome band 7q11.23, subjects have cardiovascular, connective tissue, and neurodevelopmental deficits. Given the striking peaks and valleys in neurocognition including deficits in visual-spatial and global processing, preserved language and face processing, hypersociability, and heightened affect, the goal of this work has been to identify the genes that are responsible, the cause of the deletion, and its origin in primate evolution. To do this, we have generated an integrated physical, genetic, and transcriptional map of the WMS and flanking regions using multicolor metaphase and interphase fluorescence in situ hybridization (FISH) of bacterial artificial chromosomes (BACs) and P1 artificial chromosomes (PACs), BAC end sequencing, PCR gene marker and microsatellite, large-scale sequencing, cDNA library, and database analyses. The results indicate the genomic organization of the WMS region as two nested duplicated regions flanking a largely single-copy region. There are at least two common deletion breakpoints, one in the centromeric and at least two in the telomeric repeated regions. Clones anchoring the unique to the repeated regions are defined along with three new pseudogene families. Primate studies indicate an evolutionary hot spot for chromosomal inversion in the WMS region. A cognitive phenotypic map of WMS is presented, which combines previous data with five further WMS subjects and three atypical WMS subjects with deletions; two larger (deleted for D7S489L) and one smaller, deleted for genes telomeric to FZD9, through LIMK1, but not WSCR1 or telomeric. The results establish regions and consequent gene candidates for WMS features including mental retardation, hypersociability, and facial features. The approach provides the basis for defining pathways linking genetic underpinnings with the neuroanatomical, functional, and behavioral consequences that result in human cognition.
Journal Article•10.1162/089892900561832•
Age-Related Differences in Neural Activity during Item and Temporal-Order Memory Retrieval: A Positron Emission Tomography Study

[...]

Roberto Cabeza1, Nicole D. Anderson2, Sylvain Houle2, Jennifer A. Mangels3, Lars Nyberg4 •
University of Alberta1, University of Toronto2, Columbia University3, Umeå University4
01 Jan 2000-Journal of Cognitive Neuroscience
TL;DR: The results suggest that age-related changes in brain activity are rather process- and region-specific, and that they involve increases as well as decreases in neural activity.
Abstract: Positron emission tomography (PET) was used to investigate the hypothesis that older adults’ difficulties with temporal-order memory are related to deficits in frontal function. Young (mean 24.7 years) and old (mean 68.6 years) participants studied a list of words, and were then scanned while retrieving information about what words were in the list (item retrieval) or when they occurred within the list (temporal-order retrieval). There were three main results. First, whereas the younger adults engaged right prefrontal regions more during temporal-order retrieval than during item retrieval, the older adults did not. This result is consistent with the hypothesis that context memory deficits in older adults are due to frontal dysfunction. Second, ventromedial temporal activity during item memory was relatively unaffected by aging. This finding concurs with evidence that item memory is relatively preserved in old adults and with the notion that medial temporal regions are involved in automatic retrieval operations. Finally, replicating the result of a previous study (Cabeza, R., Grady, C. L., Nyberg, L., McIntosh, A. R., Tulving, E., Kapur, S., Jennings, J. M., Houle, S., & Craik, F. I. M., 1997), the old adults showed weaker activations than the young adults in the right prefrontal cortex but stronger activations in the left prefrontal cortex. The age-related increase in left prefrontal activity may be interpreted as compensatory. Taken together, the results suggest that age-related changes in brain activity are rather process- and region-specific, and that they involve increases as well as decreases in neural activity.
Journal Article•10.1162/089892900563975•
Neural Mechanisms of Visual Attention: Object-Based Selection of a Region in Space

[...]

Catherine M. Arrington1, Thomas H. Carr1, Andrew R. Mayer2, Stephen M. Rao2•
Michigan State University1, Medical College of Wisconsin2
01 Nov 2000-Journal of Cognitive Neuroscience
TL;DR: In this article, the authors used event-related functional magnetic resonance imaging (ER-fMRI) to measure brain activity during cued discrimination tasks requiring subjects to orient attention either to a region bounded by an object or to an unbounded region of space (location-based spatial attention).
Abstract: Objects play an important role in guiding spatial attention through a cluttered visual environment. We used event-related functional magnetic resonance imaging (ER-fMRI) to measure brain activity during cued discrimination tasks requiring subjects to orient attention either to a region bounded by an object (object-based spatial attention) or to an unbounded region of space (location-based spatial attention) in anticipation of an upcoming target. Comparison between the two tasks revealed greater activation when attention selected a region bounded by an object. This activation was strongly lateralized to the left hemisphere and formed a widely distributed network including (a) attentional structures in parietal and temporal cortex and thalamus, (b) ventral-stream object processing structures in occipital, inferior-temporal, and parahippocampal cortex, and (c) control structures in medial- and dorsolateral-prefrontal cortex. These results suggest that object-based spatial selection is achieved by imposing additional constraints over and above those processes already operating to achieve selection of an unbounded region. In addition, ER-fMRI methodology allowed a comparison of validly versus invalidly cued trials, thereby delineating brain structures involved in the reorientation of attention after its initial deployment proved incorrect. All areas of activation that differentiated between these two trial types resulted from greater activity during the invalid trials. This outcome suggests that all brain areas involved in attentional orienting and task performance in response to valid cues are also involved on invalid trials. During invalid trials, additional brain regions are recruited when a perceiver recovers from invalid cueing and reorients attention to a target appearing at an uncued location. Activated brain areas specific to attentional reorientation were strongly right-lateralized and included posterior temporal and inferior parietal regions previously implicated in visual attention processes, as well as prefrontal regions that likely subserve control processes, particularly related to inhibition of inappropriate responding.
Journal Article•10.1162/089892900562372•
Activation Timecourse of Ventral Visual Stream Object-recognition Areas: High Density Electrical Mapping of Perceptual Closure Processes

[...]

Glen M. Doniger1, John J. Foxe2, Micah M. Murray2, Beth A. Higgins2, Charles E. Schroeder3, Daniel C. Javitt3 •
New York University1, Albert Einstein College of Medicine2, Nathan Kline Institute for Psychiatric Research3
01 Jul 2000-Journal of Cognitive Neuroscience
TL;DR: Scalp-current density mapping of the Ncl revealed bilateral occipito-temporal scalp foci, which are consistent with generators in the human ventral visual stream, and specifically the lateral-occipital or LO complex as defined by hemodynamic studies of object recognition.
Abstract: Object recognition is achieved even in circumstances when only partial information is available to the observer. Perceptual closure processes are essential in enabling such recognitions to occur. We presented successively less fragmented images while recording high-density event-related potentials (ERPs), which permitted us to monitor brain activity during the perceptual closure processes leading up to object recognition. We reveal a bilateral ERP component (Ncl) that tracks these processes (onsets ~230 msec, maximal at ~290 msec). Scalp-current density mapping of the Ncl revealed bilateral occipito-temporal scalp foci, which are consistent with generators in the human ventral visual stream, and specifically the lateral-occipital or LO complex as defined by hemodynamic studies of object recognition.
Journal Article•10.1162/089892900563984•
Neural Activation During Response Competition

[...]

Eliot Hazeltine1, Russell A. Poldrack2, John D. E. Gabrieli3•
Ames Research Center1, Harvard University2, Stanford University3
01 Nov 2000-Journal of Cognitive Neuroscience
TL;DR: This study measured brain activity using functional magnetic resonance imaging (fMRI) during performance of the flanker task to suggest that the frontal foci may be related to response inhibition processes whereas the posterior foci is related to the activation of representations of the inappropriate responses.
Abstract: The flanker task, introduced by Eriksen and Eriksen lEriksen, B. A., & Eriksen, C. W. (1974). Effects of noise letters upon the identification of a target letter in a nonsearch task. Perception & Psychophysics, 16, 143–149r, provides a means to selectively manipulate the presence or absence of response competition while keeping other task demands constant. We measured brain activity using functional magnetic resonance imaging (fMRI) during performance of the flanker task. In accordance with previous behavioral studies, trials in which the flanking stimuli indicated a different response than the central stimulus were performed significantly more slowly than trials in which all the stimuli indicated the same response. This reaction time effect was accompanied by increases in activity in four regions: the right ventrolateral prefrontal cortex, the supplementary motor area, the left superior parietal lobe, and the left anterior parietal cortex. The increases were not due to changes in stimulus complexity or the need to overcome previously learned associations between stimuli and responses. Correspondences between this study and other experiments manipulating response interference suggest that the frontal foci may be related to response inhibition processes whereas the posterior foci may be related to the activation of representations of the inappropriate responses.
Journal Article•10.1162/089892900562444•
The Role of Spatial Selective Attention in Working Memory for Locations: Evidence from Event-Related Potentials

[...]

Edward Awh1, Lourdes Anllo-Vento2, Steven A. Hillyard2•
University of Oregon1, University of California, San Diego2
01 Sep 2000-Journal of Cognitive Neuroscience
TL;DR: Investigation of the hypothesis that the covert focusing of spatial attention mediates the on-line maintenance of location information in spatial working memory found that early ERP components were enlarged in response to probes that appeared at memorized locations.
Abstract: We investigated the hypothesis that the covert focusing of spatial attention mediates the on-line maintenance of location information in spatial working memory. During the delay period of a spatial working-memory task, behaviorally irrelevant probe stimuli were flashed at both memorized and nonmemorized locations. Multichannel recordings of event-related potentials (ERPs) were used to assess visual processing of the probes at the different locations. Consistent with the hypothesis of attention-based rehearsal, early ERP components were enlarged in response to probes that appeared at memorized locations. These visual modulations were similar in latency and topography to those observed after explicit manipulations of spatial selective attention in a parallel experimental condition that employed an identical stimulus display.
Journal Article•10.1162/089892900562101•
A Parametric Approach to Orthographic Processing in the Brain: An fMRI Study

[...]

Malle A. Tagamets1, Jared M. Novick2, Maria L. Chalmers2, Rhonda B. Friedman2•
University of Maryland, Baltimore1, Georgetown University Medical Center2
01 Mar 2000-Journal of Cognitive Neuroscience
TL;DR: There is a widespread network of brain regions that form a common network for the processing of all orthographic string types and differences among these string types were mainly expressed as graded changes in the balance of activations among the regions.
Abstract: Brain activation studies of orthographic stimuli typically start with the premise that different types of orthographic strings (e.g., words, pseudowords) differ from each other in discrete ways, which should be reflected in separate and distinct areas of brain activation. The present study starts from a different premise: Words, pseudowords, letterstrings, and false fonts vary systematically across a continuous dimension of familiarity to English readers. Using a one-back matching task to force encoding of the stimuli, the four types of stimuli were visually presented to healthy adult subjects while fMRI activations were obtained. Data analysis focused on parametric comparisons of fMRI activation sites. We did not find any region that was exclusively activated for real words. Rather, differences among these string types were mainly expressed as graded changes in the balance of activations among the regions. Our results suggest that there is a widespread network of brain regions that form a common network for the processing of all orthographic string types.
Journal Article•10.1162/089892900562318•
Tickling Expectations: Neural Processing in Anticipation of a Sensory Stimulus

[...]

Katrina Carlsson1, Predrag Petrovic1, Stefan Skare1, Karl Magnus Petersson1, Martin Ingvar1 •
Karolinska Institutet1
01 Jul 2000-Journal of Cognitive Neuroscience
TL;DR: The overlapping pattern of change, during the somatosensory stimulation and the anticipation, furthers the idea that predictions are subserved by a neuronal network similar to that which subserves the processing of actual sensory input.
Abstract: Predictions of the near future can optimize the accuracy and speed of sensory processing as well as of behavioral responses. Previous experience and contextual cues are essential elements in the generation of a subjective prediction. Using a blocked fMRI paradigm, we investigated the pattern of neural activation in anticipation of a sensory stimulus and during the processing of the somatosensory stimulus itself. Tickling was chosen as the somatosensory stimulus rather than simple touch in order to increase the probability to get a high degree of anticipation. The location and nature of the stimulus were well defined to the subject. The state of anticipation was initiated by attributing an uncertainty regarding the time of stimulus onset. The network of activation and deactivation during anticipation of the expected stimulus was similar to that engaged during the actual sensory stimulation. The areas that were activated during both states included the contralateral primary sensory cortex, bilateral areas in the inferior parietal lobules, the putative area SII, the right anterior cingulate cortex and areas in the right prefrontal cortex. Similarly, common decreases were observed in areas of sensorimotor cortex located outside the area representing the target of stimulus, i.e., areas that process information which is irrelevant to the attended process. The overlapping pattern of change, during the somatosensory stimulation and the anticipation, furthers the idea that predictions are subserved by a neuronal network similar to that which subserves the processing of actual sensory input. Moreover, this study indicates that activation of primary somatosensory cortex can be obtained without intra-modal sensory input. These findings suggest that anticipation may invoke a tonic top-down regulation of neural activity.
Journal Article•10.1162/089892900562264•
Cerebral Specialization for Speech and Non-Speech Stimuli in Infants

[...]

Ghislaine Dehaene-Lambertz1•
Centre national de la recherche scientifique1
01 May 2000-Journal of Cognitive Neuroscience
TL;DR: Early cerebral specialization and lateralization for auditory processing in 4-month-old infants was studied by recording high-density evoked potentials to acoustical and phonetic changes in a series of repeated stimuli indicating specialized modules are present within the auditory cortex very early in development.
Abstract: Early cerebral specialization and lateralization for auditory processing in 4-month-old infants was studied by recording high-density evoked potentials to acoustical and phonetic changes in a series of repeated stimuli (either tones or syllables). Mismatch responses to these stimuli exhibit a distinct topography suggesting that different neural networks within the temporal lobe are involved in the perception and representation of the different features of an auditory stimulus. These data confirm that specialized modules are present within the auditory cortex very early in development. However, both for syllables and continuous tones, higher voltages were recorded over the left hemisphere than over the right with no significant interaction of hemisphere by type of stimuli. This suggests that there is no greater left hemisphere involvement in phonetic processing than in acoustic processing during the first months of life.
Journal Article•10.1162/089892900564019•
The Emotional Modulation of Cognitive Processing: An fMRI Study

[...]

Joseph R. Simpson1, Dost Ouml1, ngür1, Erbil Akbudak1, Thomas E. Conturo1, John M. Ollinger1, Abraham Z. Snyder1, Debra A. Gusnard1, Marcus E. Raichle1 •
Washington University in St. Louis1
01 Nov 2000-Journal of Cognitive Neuroscience
TL;DR: Results provide evidence that the emotional valence and arousing nature of stimuli used during the performance of an attention-demanding cognitive task are reflected in discernable, quantitative changes in the functional anatomy associated with task performance.
Abstract: The functional neuroanatomy of visual processing of surface features of emotionally valenced pictorial stimuli was examined in normal human subjects using functional magnetic resonance imaging (fMRI). Pictorial stimuli were of two types: emotionally negative and neutral pictures. Task performance was slower for the negatively valenced than for the neutral pictures. Significant blood oxygen level dependent (BOLD) increases occurred in the medial and dorsolateral prefrontal cortex, midbrain, substantia innominata, and/or amygdala, and in the posterior cortical visual areas for both stimulus types. Increases were greater for the negatively valenced stimuli. While there was a small but significant BOLD decrease in the subgenual prefrontal cortex, which was larger in response to the negatively valenced pictures, there was an almost complete absence of other decreases prominently seen during the performance of demanding cognitive tasks [Shulman, G. L., Fiez, J. A., Corbetta, M., Buckner, R. L., Miezin, F. M., Raichle, M. E., & Petersen, S. E. (1997). Common blood flow changes across visual tasks: II. Decreases in cerebral cortex. Journal of Cognitive Neuroscience, 9, 648-663]. These results provide evidence that the emotional valence and arousing nature of stimuli used during the performance of an attention-demanding cognitive task are reflected in discernable, quantitative changes in the functional anatomy associated with task performance.
Journal Article•10.1162/089892900562147•
Language Processing Modulated by Literacy: A Network Analysis of Verbal Repetition in Literate and Illiterate Subjects

[...]

Karl Magnus Petersson1, Alexandra Reis1, Simon Askelöf1, Alexandre Castro-Caldas, Martin Ingvar1 •
Karolinska Institutet1
01 May 2000-Journal of Cognitive Neuroscience
TL;DR: The hypothesis that learning to read and write during childhood influences the functional architecture of the adult human brain is supported and the parallel interactive processing characteristics of the underlying language-processing brain network differ in literate and illiterate subjects is suggested.
Abstract: Previous behavioral and functional neuroimaging data indicate that certain aspects of phonological processing may not be acquired spontaneously, but are modulated by learning an alphabetic written language, that is, learning to read and write. It appears that learning an alphabetic written language modifies the auditory-verbal (spoken) language processing competence in a nontrivial way. We have previously suggested, based on behavioral and functional neuroimaging data, that auditory-verbal and written language interact not only during certain language tasks, but that learning and developing alphabetic written language capacities significantly modulates the spoken language system. Specifically, the acquisition of alphabetic orthographic knowledge has a modulatory influence on sublexical phonological processing and the awareness of sublexical phonological structure. We have suggested that developing an orthographic representation system for an alphabetic written language, and integrating a phoneme-grapheme correspondence with an existing infrastructure for auditory-verbal language processing, will result in a modified language network. Specifically, we suggest that the parallel interactive processing characteristics of the underlying language-processing brain network differ in literate and illiterate subjects. Therefore, the pattern of interactions between the regions of a suitably defined large-scale functional-anatomical network for language processing will differ between literate and illiterate subjects during certain language tasks. In order to investigate this hypothesis further, we analyzed the observed covariance structure in a PET data set from a simple auditory-verbal repetition paradigm in literate and illiterate subjects, with a network approach based on structural equation modeling (SEM). Based on a simple network model for language processing, the results of the present network analysis indicate that the network interactions during word and pseudoword repetition in the illiterate group differ, while there were no significant differences in the literate group. The differences between the two tasks in the illiterate group may reflect differences in attentional modulation of the language network, executive aspects of verbal working memory and the articulatory organization of verbal output. There were no significant differences between the literate and illiterate group during word repetition. In contrast, the network interactions differed between the literate and illiterate group during pseudoword repetition. In addition to differences similar to those observed in the illiterate group between word and pseudoword repetition, there were differences related to the interactions of the phonological loop between the groups. In particular, these differences related to the interaction between Broca's area and the inferior parietal cortex as well as the posterior-midinsula bridge between Wernicke's and Broca's area. In conclusion, the results of this network analysis are consistent with our previously presented results and support the hypothesis that learning to read and write during childhood influences the functional architecture of the adult human brain. In particular, the basic auditory-verbal language network in the human brain is modified as a consequence of acquiring orthographic language skills.
Journal Article•10.1162/089892900562291•
Electrophysiological Evidence for the Modulation of Retrieval Orientation by Depth of Study Processing

[...]

Michael D. Rugg1, Kevin Allan1, Claire S. Birch2•
University College London1, University of St Andrews2
01 Jul 2000-Journal of Cognitive Neuroscience
TL;DR: The findings indicate that the depth of study processing influences two kinds of memory-related neural activity, associated with memory search operations, and the processing of retrieved information, respectively.
Abstract: Event-related potentials (ERPs) were employed to investigate whether brain activity elicited by retrieval cues in a memory test varies according to the encoding task undertaken at study. Two recognition memory test blocks were administered, preceded, in one case, by a "shallow" study task (alphabetic judgement) and, in the other case, by a "deep" task (sentence generation). ERPs elicited by the new words in each test block differed, the ERPs elicited in the block following the shallow study task exhibiting the more positive-going waveforms. This finding was taken as evidence that subjects adopt different "retrieval sets" when attempting to retrieve items that had been encoded in terms of alphabetic versus semantic attributes. Differences between the ERPs elicited by correctly classified old and new words (old/new effects) also varied with encoding task. The effects for deeply studied words resembled those found in previous ERP studies of recognition memory, whereas old/new effects for shallowly studied words were confined to a late-onsetting, right frontal positivity. Together, the findings indicate that the depth of study processing influences two kinds of memory-related neural activity, associated with memory search operations, and the processing of retrieved information, respectively.
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