The N400 response is one of the best-studied ERP components associated with language processing. Manipulations such as semantic priming ('coffee' → 'tea') and contextual fit ('I like my coffee with cream and sugar/socks') modulate the amplitude of the N400 response to words; this is referred to as the 'N400 effect'.
Two contrasting hypotheses address the functional interpretation of the N400 effect. The effect may reflect differences in the amount of effort needed to semantically integrate the incoming word into the previous sentential context; alternatively, it may reflect facilitated lexical access due to pre-activation by supportive contexts.
Previous imaging and patient data implicate the posterior middle temporal cortex in storing and activating information associated with lexical representations. Some evidence suggests that anterior temporal and inferior parietal cortices are involved in constructing new inputs and/or integrating them into temporary representations of the larger syntactic and semantic structure, and that the inferior frontal cortex subserves control processes of retrieval and selection of lexical representations that make the input available to integrative processes.
The functional anatomic model outlined leads to predictions about the locus of the N400 effect: if the effect reflects integration difficulty, it should localize to inferior frontal, anterior temporal or inferior parietal regions; if the effect reflects facilitated lexical access, it should localize to the posterior middle temporal cortex.
Functional MRI studies show semantic priming effects in the inferior frontal cortex only when there is a long interval between primes and targets. Priming effects are observed in the posterior middle temporal gyrus across all prime–target intervals, as is the N400 effect. Magnetoencephalography and event-related optical signal (EROS) studies of sentence context show semantic-priming and semantic-anomaly effects in the posterior temporal cortex at the same latency as the N400 effect.
The data from across these techniques provide strong evidence that the posterior middle temporal cortex is involved in generating the N400 effect, which supports the claim that the N400 effect is at least to some degree due to facilitated lexical access, and argues against an account in which the effect is purely integrative.
Measuring event-related potentials (ERPs) has been fundamental to our understanding of how language is encoded in the brain. One particular ERP response, the N400 response, has been especially influential as an index of lexical and semantic processing. However, there remains a lack of consensus on the interpretation of this component. Resolving this issue has important consequences for neural models of language comprehension. Here we show that evidence bearing on where the N400 response is generated provides key insights into what it reflects. A neuroanatomical model of semantic processing is used as a guide to interpret the pattern of activated regions in functional MRI, magnetoencephalography and intracranial recordings that are associated with contextual semantic manipulations that lead to N400 effects.
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We thank H. Kolk for extensive feedback on the manuscript. Preparation of this manuscript was supported by a National Science Foundation (NSF) Graduate Research Fellowship to E.F.L., 2R01DC05660 to D.P. and NSF DGE-0801465 to the University of Maryland.
- Event-related potentials
(ERPs). Electrical potentials that are generated in the brain as a consequence of the synchronized activation of neuronal networks by external stimuli. These evoked potentials are recorded at the scalp and consist of precisely timed sequences of waves or 'components'.
The smallest linguistic unit that has a specific meaning or syntactic function. For example, the word 'outplayed' consists of three morphemes: 'out', 'play' and 'ed'.
- Semantic structure
The mental representation of how the meanings of individual words or concepts are combined to yield the meaning of entire phrases or sentences.
- Syntactic structure
The mental representation of how the words and phrases within a sentence are organized to form a hierarchically nested structure.
- Discourse structure
The mental representation of the information in a conversation or speech act, specifying which information is common knowledge between the speakers, which events or individuals are the current topic of discussion and what are the speakers' communicative goals.
Auditory or visual strings that sound or look like words but are not actually words (for example, 'blick' but not 'lbikc').
A language impairment that is acquired as a result of stroke or other brain injury.
- Syntactic priming
The tendency in language production to formulate sentences with recently used syntactic constructions. For example, when one speaker in a conversation uses a passive construction, other speakers become more likely to use passives.
Individual units of speech sound used to store the sounds of words in memory.
- Split brain
A brain in which the two hemispheres have been separated by severing the commissures that connected them.
- Phase coherence
The degree to which two oscillating signals (for example, as measured with electrodes from the scalp) are synchronized.
- Stimulus-onset asynchrony
(SOA). The time between the onsets of sequentially presented stimuli.
- Equivalent current dipole (ECD) models
Estimates of the focal source of neuronal activation derived from multi-sensor data obtained by electroencephalography or magnetoencephalography. The activation is modelled as a point source with a position, direction and intensity.
- Distributed-source models
Estimates of the neuronal sources that underlie the brain activity that can be represented with magnetoencephalography or electroencephalography multi-channel data. Activity is not assumed to be focal, permitting a spatially distributed and perhaps more realistic visualization.
- Mental models
Internal psychological representations of actual or hypothetical situations that can guide perception, action and reasoning.
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Lau, E., Phillips, C. & Poeppel, D. A cortical network for semantics: (de)constructing the N400. Nat Rev Neurosci 9, 920–933 (2008). https://doi.org/10.1038/nrn2532
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