N400-like Magnetoencephalography Responses Modulated by Semantic Context, Word Frequency, and Lexical Class in Sentences

Words have been found to elicit a negative potential at the scalp peaking at approximately 400 ms that is strongly modulated by semantic context. The current study used whole-head magnetoencephalography (MEG) as male subjects read sentences ending with semantically congruous or incongruous words. Compared with congruous words, sentence-terminal incongruous words consistently evoked a large magnetic field over the left hemisphere, peaking at approximately 450 ms. Source modeling at this latency with conventional equivalent current dipoles (ECDs) placed the N400 m generator in or near the left superior temporal sulcus. A distributed solution constrained to the cortical surface suggested a sequence of differential activation, beginning in Wernicke's area at approximately 250 ms, spreading to anterior temporal sites at approximately 270 ms, to Broca's area by approximately 300 ms, to dorsolateral prefrontal cortices by approximately 320 ms, and to anterior orbital and frontopolar cortices by approximately 370 ms. Differential activity was exclusively left-sided until >370 ms, and then involved right anterior temporal and orbital cortices. At the peak of the N400 m, activation in the left hemisphere was estimated to be widespread in the anterior temporal, perisylvian, orbital, frontopolar, and dorsolateral prefrontal cortices. In the right hemisphere, the orbital, as well as, weakly, the right anterior temporal cortices were activated. Similar but weaker field patterns were evoked by intermediate words in the sentences, especially to low-frequency words occurring in early sentence positions where there is little preceding context. The locations of the N400 m sources identified with the distributed solution correspond well with those previously demonstrated with direct intracranial recordings, and suggested by functional magnetic resonance imaging (fMRI). These results help identify a distributed cortical network that supports online semantic processing.

[1]  A. Puce,et al.  Visual recognition memory. Neurophysiological evidence for the role of temporal white matter in man. , 1991, Brain : a journal of neurology.

[2]  E. Halgren,et al.  Spatio-temporal stages in face and word processing. 2. Depth-recorded potentials in the human frontal and Rolandic cortices , 1994, Journal of Physiology-Paris.

[3]  M. Kutas,et al.  Reading senseless sentences: brain potentials reflect semantic incongruity. , 1980, Science.

[4]  T. Paine Common Sense , 1995 .

[5]  A. David,et al.  The planum temporale: a systematic, quantitative review of its structural, functional and clinical significance , 1999, Brain Research Reviews.

[6]  E. Halgren,et al.  Face-selective spectral changes in the human fusiform gyrus , 1999, Clinical Neurophysiology.

[7]  Klaus Lehnertz,et al.  Human temporal lobe potentials in verbal learning and memory processes , 1997, Neuropsychologia.

[8]  P. Holcomb,et al.  Event-Related Brain Potentials Reflect Semantic Priming in an Object Decision Task , 1994, Brain and Cognition.

[9]  C. Van Petten,et al.  Words and sentences: event-related brain potential measures. , 1995, Psychophysiology.

[10]  Ava J. Senkfor,et al.  Electrophysiological dissociation between verbal and nonverbal semantic processing in learning disabled adults , 2000, Neuropsychologia.

[11]  G. McCarthy,et al.  Language-related field potentials in the anterior-medial temporal lobe: II. Effects of word type and semantic priming , 1995, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[12]  G. McCarthy,et al.  Language-related field potentials in the anterior-medial temporal lobe: I. Intracranial distribution and neural generators , 1995, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[13]  E. Halgren,et al.  Dynamic Statistical Parametric Mapping Combining fMRI and MEG for High-Resolution Imaging of Cortical Activity , 2000, Neuron.

[14]  M S Gazzaniga,et al.  Processing of semantic anomaly by right and left hemispheres of commissurotomy patients. Evidence from event-related brain potentials. , 1988, Brain : a journal of neurology.

[15]  John W Belliveau,et al.  Monte Carlo simulation studies of EEG and MEG localization accuracy , 2002, Human brain mapping.

[16]  M. Kutas,et al.  Event-related potential asymmetries during the reading of sentences. , 1988, Electroencephalography and clinical neurophysiology.

[17]  E. Bullmore,et al.  Common and Distinct Neural Substrates for Pragmatic, Semantic, and Syntactic Processing of Spoken Sentences: An fMRI Study , 2000, Journal of Cognitive Neuroscience.

[18]  Ava J. Senkfor,et al.  Memory for words and novel visual patterns: repetition, recognition, and encoding effects in the event-related brain potential. , 1996, Psychophysiology.

[19]  A. Friederici,et al.  First-Pass versus Second-Pass Parsing Processes in a Wernicke's and a Broca's Aphasic: Electrophysiological Evidence for a Double Dissociation , 1998, Brain and Language.

[20]  Anders M. Dale,et al.  Cortical Surface-Based Analysis I. Segmentation and Surface Reconstruction , 1999, NeuroImage.

[21]  M I Posner,et al.  Topography of the N400: brain electrical activity reflecting semantic expectancy. , 1993, Electroencephalography and clinical neurophysiology.

[22]  B. N'Kaoua,et al.  Intracranial topography of event-related potentials (N400/P600) elicited during a continuous recognition memory task. , 1995, Psychophysiology.

[23]  E. Halgren,et al.  Spatiotemporal mapping of brain activity by integration of multiple imaging modalities , 2001, Current Opinion in Neurobiology.

[24]  A. Szentkuti,et al.  Differences in brain potentials to open and closed class words: class and frequency effects , 2001, Neuropsychologia.

[25]  C Van Petten,et al.  Word repetition in amnesia. Electrophysiological measures of impaired and spared memory. , 2000, Brain : a journal of neurology.

[26]  Ian Spence,et al.  Monte Carlo Simulation Studies , 1983 .

[27]  M. Hämäläinen,et al.  Realistic conductivity geometry model of the human head for interpretation of neuromagnetic data , 1989, IEEE Transactions on Biomedical Engineering.

[28]  M. Kutas,et al.  The lateral distribution of event-related potentials during sentence processing , 1982, Neuropsychologia.

[29]  Marta Kutas,et al.  Neural plasticity in the dynamics of human visual word recognition , 1998, Neuroscience Letters.

[30]  H G Smid,et al.  Event-related potentials of verbal encoding into episodic memory: dissociation between the effects of subsequent memory performance and distinctiveness. , 1998, Psychophysiology.

[31]  J. D. de Munck A linear discretization of the volume conductor boundary integral equation using analytically integrated elements (electrophysiology application) , 1992, IEEE Transactions on Biomedical Engineering.

[32]  A. Dale,et al.  High‐resolution intersubject averaging and a coordinate system for the cortical surface , 1999, Human brain mapping.

[33]  A. Dale,et al.  Improved Localizadon of Cortical Activity by Combining EEG and MEG with MRI Cortical Surface Reconstruction: A Linear Approach , 1993, Journal of Cognitive Neuroscience.

[34]  E. Halgren,et al.  LOCALISED FACE PROCESSING BY THE HUMAN PREFRONTAL CORTEX: FACE-SELECTIVE INTRACEREBRAL POTENTIALS AND POST-LESION DEFICITS , 2000, Cognitive neuropsychology.

[35]  C. Petten,et al.  Conceptual relationships between spoken words and environmental sounds: Event-related brain potential measures , 1995, Neuropsychologia.

[36]  E. Halgren,et al.  Human medial temporal lobe potentials evoked in memory and language tasks. , 1986, Electroencephalography and clinical neurophysiology.

[37]  D. Wegesin,et al.  Event-Related Potentials in Homosexual and Heterosexual Men and Women: Sex-Dimorphic Patterns in Verbal Asymmetries and Mental Rotation , 1998, Brain and Cognition.

[38]  L. Parkkonen,et al.  122-channel squid instrument for investigating the magnetic signals from the human brain , 1993 .

[39]  E. Halgren,et al.  Spatio-temporal stages in face and word processing. 1. Depth recorded potentials in the human occipital and parietal lobes , 1994, Journal of Physiology-Paris.

[40]  P. D. Eimas,et al.  Neurobiology of cognition , 1990 .

[41]  R. Salmelin,et al.  Semantic Cortical Activation in Dyslexic Readers , 1999, Journal of Cognitive Neuroscience.

[42]  Klaus Lehnertz,et al.  Limbic ERPs predict verbal memory after left‐sided hippocampectomy , 1998, Neuroreport.

[43]  A. Dale,et al.  Cortical Surface-Based Analysis II: Inflation, Flattening, and a Surface-Based Coordinate System , 1999, NeuroImage.

[44]  Steven M. J. Hunt,et al.  MacProbe: A Macintosh-based experimenter’s workstation for the cognitive sciences , 1994 .

[45]  E Halgren,et al.  Dissociation of recognition memory components following temporal lobe lesions. , 1989, Journal of experimental psychology. Learning, memory, and cognition.

[46]  Colin M. Brown,et al.  Lexical-semantic event-related potential effects in patients with left hemisphere lesions and aphasia, and patients with right hemisphere lesions without aphasia. , 1996, Brain : a journal of neurology.

[47]  F. Rösler,et al.  Individual cortical current density reconstructions of the semantic N400 effect: Using a generalized minimum norm model with different constraints (L1 and L2 norm) , 2000, Human brain mapping.

[48]  J. D. Munck A linear discretization of the volume conductor boundary integral equation using analytically integrated elements (electrophysiology application) , 1992 .

[49]  Kara D. Federmeier,et al.  Electrophysiology reveals semantic memory use in language comprehension , 2000, Trends in Cognitive Sciences.

[50]  M. Kutas,et al.  Event-related brain potentials to grammatical errors and semantic anomalies , 1983, Memory & cognition.

[51]  H. Kucera,et al.  Computational analysis of present-day American English , 1967 .

[52]  F. Guillem,et al.  Short-and Long-Delay Intracranial ERP Repetition Effects Dissociate Memory Systems in the Human Brain , 1999, Journal of Cognitive Neuroscience.

[53]  E Halgren,et al.  Event-related potentials during lexical decision: effects of repetition, word frequency, pronounceability, and concreteness. , 1987, Electroencephalography and clinical neurophysiology. Supplement.

[54]  R. Ilmoniemi,et al.  Magnetoencephalography-theory, instrumentation, and applications to noninvasive studies of the working human brain , 1993 .

[55]  R L Buckner,et al.  Spatiotemporal Maps of Brain Activity Underlying Word Generation and Their Modification during Repetition Priming , 2001, The Journal of Neuroscience.

[56]  M. Kutas,et al.  The Search for Common Sense: An Electrophysiological Study of the Comprehension of Words and Pictures in Reading , 1996, Journal of Cognitive Neuroscience.

[57]  Kenji Kansaku,et al.  Imaging studies on sex differences in the lateralization of language , 2001, Neuroscience Research.

[58]  C Van Petten,et al.  Time course of word identification and semantic integration in spoken language. , 1999, Journal of experimental psychology. Learning, memory, and cognition.

[59]  Andrew C Papanicolaou,et al.  Source localization of the N400 response in a sentence-reading paradigm using evoked magnetic fields and magnetic resonance imaging , 1997, Brain Research.

[60]  M. Kutas,et al.  Influences of semantic and syntactic context on open- and closed-class words , 1991, Memory & cognition.

[61]  U. Bellugi,et al.  Neural Systems Mediating American Sign Language: Effects of Sensory Experience and Age of Acquisition , 1997, Brain and Language.

[62]  K. Lehnertz,et al.  Differential Involvement of Left Temporolateral and Temporomesial Structures in Verbal Declarative Learning and Memory: Evidence from Temporal Lobe Epilepsy , 1997, Brain and Cognition.

[63]  M. Torrens Co-Planar Stereotaxic Atlas of the Human Brain—3-Dimensional Proportional System: An Approach to Cerebral Imaging, J. Talairach, P. Tournoux. Georg Thieme Verlag, New York (1988), 122 pp., 130 figs. DM 268 , 1990 .

[64]  T. Allison,et al.  Word recognition in the human inferior temporal lobe , 1994, Nature.

[65]  M. Kutas,et al.  Interactions between sentence context and word frequencyinevent-related brainpotentials , 1990, Memory & cognition.