Thirty years and counting: finding meaning in the N400 component of the event-related brain potential (ERP).

We review the discovery, characterization, and evolving use of the N400, an event-related brain potential response linked to meaning processing. We describe the elicitation of N400s by an impressive range of stimulus types--including written, spoken, and signed words or pseudowords; drawings, photos, and videos of faces, objects, and actions; sounds; and mathematical symbols--and outline the sensitivity of N400 amplitude (as its latency is remarkably constant) to linguistic and nonlinguistic manipulations. We emphasize the effectiveness of the N400 as a dependent variable for examining almost every aspect of language processing and highlight its expanding use to probe semantic memory and to determine how the neurocognitive system dynamically and flexibly uses bottom-up and top-down information to make sense of the world. We conclude with different theories of the N400's functional significance and offer an N400-inspired reconceptualization of how meaning processing might unfold.

[1]  G. McCarthy,et al.  Augmenting mental chronometry: the P300 as a measure of stimulus evaluation time. , 1977, Science.

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

[3]  Salim Roukos,et al.  Brain potentials related to stages of sentence verification. , 1983, Psychophysiology.

[4]  M. Kutas,et al.  Electrophysiology of cognitive processing. , 1983, Annual review of psychology.

[5]  E. Donchin.  Cognitive Psychophysiology: Event-Related Potentials and the Study of Cognition , 2022 .

[6]  F. Macar,et al.  An event-related potential analysis of incongruity in music and other non-linguistic contexts. , 1987, Psychophysiology.

[7]  P. Holcomb.  Automatic and attentional processing: An event-related brain potential analysis of semantic priming , 1988, Brain and Language.

[8]  M. Kutas.  Event related brain potential studies of language , 1988 .

[9]  J. Connolly,et al.  The effects of processing requirements on neurophysiological responses to spoken sentences , 1990, Brain and Language.

[10]  W. Ritter,et al.  The relationship between N2 and N400: scalp distribution, stimulus probability, and task relevance. , 1991, Psychophysiology.

[11]  M. Smith,et al.  Decomposition of recognition memory event-related potentials yields target, repetition, and retrieval effects. , 1993, Electroencephalography and clinical neurophysiology.

[12]  M. Kutas.  In the company of other words: Electrophysiological evidence for single-word and sentence context effects , 1993 .

[13]  G McCarthy,et al.  Modulation of semantic processing by spatial selective attention. , 1993, Electroencephalography and clinical neurophysiology.

[14]  C. Petten.  A comparison of lexical and sentence-level context effects in event-related potentials , 1993 .

[15]  Peter Hagoort,et al.  The Processing Nature of the N400: Evidence from Masked Priming , 1993, Journal of Cognitive Neuroscience.

[16]  P. Holcomb,et al.  Cross-modal semantic priming: A time-course analysis using event-related brain potentials , 1993 .

[17]  M. Kutas,et al.  What Is Who Violating? A Reconsideration of Linguistic Violations in Light of Event-Related Brain Potentials , 1994 .

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

[19]  L. Osterhout,et al.  Event-Related Brain Potentials Elicited by Failure to Agree , 1995 .

[20]  Colin M. Brown,et al.  The N400 as a function of the level of processing. , 1995, Psychophysiology.

[21]  B. Grözinger,et al.  Evoked brain potentials and memory: more positivity in response to forgotten items , 1995, Neuroreport.

[22]  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.

[23]  W. Ritter,et al.  Lexical processing of visually and auditorily presented nouns and verbs: evidence from reaction time and N400 priming data. , 1997, Brain research. Cognitive brain research.

[24]  E. Vogel,et al.  Electrophysiological Evidence for a Postperceptual Locus of Suppression during the Attentional Blink Time-based Attention and the Attentional Blink , 1998 .

[25]  R. Johnson,et al.  A spatio-temporal analysis of recognition-related event-related brain potentials. , 1998, International journal of psychophysiology : official journal of the International Organization of Psychophysiology.

[26]  J. Kounios,et al.  Dual-coding, context-availability, and concreteness effects in sentence comprehension: an electrophysiological investigation. , 1999, Journal of experimental psychology. Learning, memory, and cognition.

[27]  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.

[28]  Frank Rösler,et al.  N400 Effects Reflect Activation Spread During Retrieval of Arithmetic Facts , 1999 .

[29]  E I Olivares,et al.  Searching for face-specific long latency ERPs: a topographic study of effects associated with mismatching features. , 1999, Brain research. Cognitive brain research.

[30]  F Rösler,et al.  Processing of incongruous mental calculation problems: evidence for an arithmetic N400 effect. , 1999, Psychophysiology.

[31]  E. Plante,et al.  Time course of word identification and semantic integration in spoken language. , 1999, Journal of experimental psychology. Learning, memory, and cognition.

[32]  G. Woodman,et al.  Event-related potential studies of attention , 2000, Trends in Cognitive Sciences.

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

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

[35]  D Friedman,et al.  Event‐related potential (ERP) studies of memory encoding and retrieval: A selective review , 2000, Microscopy research and technique.

[36]  D Deacon,et al.  How automatically is meaning accessed: a review of the effects of attention on semantic processing. , 2000, Frontiers in bioscience : a journal and virtual library.

[37]  A. A. Wijers,et al.  Visual semantic features are activated during the processing of concrete words: event-related potential evidence for perceptual semantic priming. , 2000, Brain research. Cognitive brain research.

[38]  A. Friederici,et al.  Syntactic Gender and Semantic Expectancy: ERPs Reveal Early Autonomy and Late Interaction , 2000, Journal of Cognitive Neuroscience.

[39]  I Rosén,et al.  Semantic processing without conscious identification: evidence from event-related potentials. , 2000, Journal of experimental psychology. Learning, memory, and cognition.

[40]  T. Curran.  Brain potentials of recollection and familiarity , 2000, Memory & cognition.

[41]  E Hennighausen,et al.  Missed prime words within the attentional blink evoke an N400 semantic priming effect. , 2001, Psychophysiology.

[42]  Kara D. Federmeier,et al.  Effects of transient, mild mood states on semantic memory organization and use: an event-related potential investigation in humans , 2001, Neuroscience Letters.

[43]  S. Kotz,et al.  ERP Evidence for a Sex-Specific Stroop Effect in Emotional Speech , 2003, Journal of Cognitive Neuroscience.

[44]  Marta Kutas,et al.  Potato not Pope: human brain potentials to gender expectation and agreement in Spanish spoken sentences , 2003, Neuroscience Letters.

[45]  P. Hagoort.  Interplay between Syntax and Semantics during Sentence Comprehension: ERP Effects of Combining Syntactic and Semantic Violations , 2003, Journal of Cognitive Neuroscience.

[46]  Stefan R. Schweinberger,et al.  Covert Recognition and the Neural System for Face Processing , 2003, Cortex.

[47]  Erik D. Reichle,et al.  The E-Z Reader model of eye-movement control in reading: Comparisons to other models , 2003, Behavioral and Brain Sciences.

[48]  P. Hagoort,et al.  Integration of Word Meaning and World Knowledge in Language Comprehension , 2004, Science.

[49]  S. Kelly,et al.  Neural correlates of bimodal speech and gesture comprehension , 2004, Brain and Language.

[50]  L. Osterhout,et al.  Neural correlates of second-language word learning: minimal instruction produces rapid change , 2004, Nature Neuroscience.

[51]  W. Witzke,et al.  How vegetative is the vegetative state? Preserved semantic processing in VS patients--evidence from N 400 event-related potentials. , 2005, NeuroRehabilitation.

[52]  C. Umilta,et al.  Electrophysiological correlates of stimulus-driven multiplication facts retrieval , 2004, Neuropsychologia.

[53]  Walter Ritter,et al.  Repetition and semantic priming of nonwords: implications for theories of N400 and word recognition. , 2004, Psychophysiology.

[54]  M. Kutas,et al.  Association of schizotypy with semantic processing differences: An event-related brain potential study , 2005, Schizophrenia Research.

[55]  Ying Choon Wu,et al.  Meaningful gestures: electrophysiological indices of iconic gesture comprehension. , 2005, Psychophysiology.

[56]  Kara D. Federmeier,et al.  Right hemisphere sensitivity to word- and sentence-level context: evidence from event-related brain potentials. , 2005, Journal of Experimental Psychology. Learning, Memory and Cognition.

[57]  H. Kolk,et al.  Accessing world knowledge: evidence from N400 and reaction time priming. , 2005, Brain research. Cognitive brain research.

[58]  Katherine A. DeLong,et al.  Probabilistic word pre-activation during language comprehension inferred from electrical brain activity , 2005, Nature Neuroscience.

[59]  Colin M. Brown,et al.  Anticipating upcoming words in discourse: evidence from ERPs and reading times. , 2005, Journal of experimental psychology. Learning, memory, and cognition.

[60]  Joel L. Voss,et al.  Fluent Conceptual Processing and Explicit Memory for Faces Are Electrophysiologically Distinct , 2006, The Journal of Neuroscience.

[61]  Y. Sakai.  An ERP study of the integration process between a noun and a numeral classifier , 2006 .

[62]  C. Petten,et al.  Neural localization of semantic context effects in electromagnetic and hemodynamic studies , 2006, Brain and Language.

[63]  Manuela Friedrich,et al.  Early N400 development and later language acquisition. , 2006, Psychophysiology.

[64]  Sotaro Kita,et al.  On-line Integration of Semantic Information from Speech and Gesture: Insights from Event-related Brain Potentials , 2007, Journal of Cognitive Neuroscience.

[65]  Joel L. Voss,et al.  Validating neural correlates of familiarity , 2007, Trends in Cognitive Sciences.

[66]  J. Debruille,et al.  The N400 potential could index a semantic inhibition , 2007, Brain Research Reviews.

[67]  H. Kolk,et al.  Late positivities in unusual situations , 2007, Brain and Language.

[68]  Monica Fabiani,et al.  Imaging cortical dynamics of language processing with the event-related optical signal , 2007, Proceedings of the National Academy of Sciences.

[69]  Gina R. Kuperberg,et al.  Neural mechanisms of language comprehension: Challenges to syntax , 2007, Brain Research.

[70]  Ken A Paller,et al.  Neural correlates of conceptual implicit memory and their contamination of putative neural correlates of explicit memory. , 2007, Learning & memory.

[71]  Kara D. Federmeier.  Thinking ahead: the role and roots of prediction in language comprehension. , 2007, Psychophysiology.

[72]  Megan A. Boudewyn,et al.  Processing new and repeated names: Effects of coreference on repetition priming with speech and fast RSVP , 2007, Brain Research.

[73]  M. Rugg,et al.  Event-related potentials and recognition memory , 2007, Trends in Cognitive Sciences.

[74]  Marta Kutas,et al.  Interplay between computational models and cognitive electrophysiology in visual word recognition , 2007, Brain Research Reviews.

[75]  Roel M. Willems,et al.  Seeing and Hearing Meaning: ERP and fMRI Evidence of Word versus Picture Integration into a Sentence Context , 2008, Journal of Cognitive Neuroscience.

[76]  Kara D. Federmeier,et al.  What's 'Right' in Language Comprehension: Event-Related Potentials Reveal Right Hemisphere Language Capabilities , 2008, Lang. Linguistics Compass.

[77]  Peter,et al.  Semantic Unification , 2008 .

[78]  Ellen F. Lau,et al.  A cortical network for semantics: (de)constructing the N400 , 2008, Nature Reviews Neuroscience.

[79]  M. van Elk,et al.  Semantics in action: An electrophysiological study on the use of semantic knowledge for action , 2008, Journal of Physiology-Paris.

[80]  Phillip J. Holcomb,et al.  Two Neurocognitive Mechanisms of Semantic Integration during the Comprehension of Visual Real-world Events , 2008, Journal of Cognitive Neuroscience.

[81]  Matthias Schlesewsky,et al.  An alternative perspective on “semantic P600” effects in language comprehension , 2008, Brain Research Reviews.

[82]  Methodological considerations related to sleep paradigm using event related potentials. , 2008, Biological research.

[83]  Kara D. Federmeier,et al.  Chapter 1 Time for Meaning: Electrophysiology Provides Insights into the Dynamics of Representation and Processing in Semantic Memory , 2009 .

[84]  W. Prinz,et al.  N400-like negativities in action perception reflect the activation of two components of an action representation , 2009, Social neuroscience.

[85]  Ellen F. Lau,et al.  A lexical basis for N400 context effects: Evidence from MEG , 2009, Brain and Language.

[86]  M. Kutas,et al.  The Cambridge Handbook of Linguistic Code-switching: Code-switching and the brain , 2009 .

[87]  Jiro Gyoba,et al.  Event-related potentials during the evaluation of the appropriateness of cooperative actions , 2009, Neuroscience Letters.

[88]  Alice Mado Proverbio,et al.  RP and N400 ERP components reflect semantic violations in visual processing of human actions , 2009, Neuroscience Letters.

[89]  Maren Grigutsch,et al.  The neural correlates of infant and adult goal prediction: evidence for semantic processing systems. , 2009, Developmental psychology.

[90]  Kara D. Federmeier,et al.  A Beautiful Day in the Neighborhood: An Event-Related Potential Study of Lexical Relationships and Prediction in Context. , 2009, Journal of memory and language.

[91]  Karen M. Evans,et al.  See what I mean? An ERP study of the effect of background knowledge on novel object processing , 2009, Memory & cognition.

[92]  Van Berkum,et al.  The neuropragmatics of 'simple' utterance comprehension: An ERP review , 2009 .

[93]  Kristina Küper,et al.  Electrophysiology reveals semantic priming at a short SOA irrespective of depth of prime processing , 2009, Neuroscience Letters.

[94]  Jonathan Grainger,et al.  ERP effects of short interval masked associative and repetition priming , 2009, Journal of Neurolinguistics.

[95]  Gina R. Kuperberg,et al.  Language in Schizophrenia Part 1: An Introduction , 2010, Lang. Linguistics Compass.

[96]  Kara D. Federmeier,et al.  Language of the Aging Brain: Event-Related Potential Studies of Comprehension in Older Adults , 2010, Lang. Linguistics Compass.

[97]  Angela D. Friederici,et al.  Maturing brain mechanisms and developing behavioral language skills , 2010, Brain and Language.

[98]  Kara D. Federmeier,et al.  The N400 as a snapshot of interactive processing: Evidence from regression analyses of orthographic neighbor and lexical associate effects. , 2011, Psychophysiology.

[99]  Seana Coulson.  The Cambridge Handbook of Psycholinguistics: Cognitive Neuroscience of Figurative Language , 2012 .