N400 (neuroscience)

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The N400 is a component of time-locked EEG signals known as event-related potentials (ERP). It is a negative-going deflection that peaks around 400 milliseconds post-stimulus onset, although it can extend from 250-500 ms, and is typically maximal over centro-parietal electrode sites. The N400 is part of the normal brain response to words and other meaningful (or potentially meaningful) stimuli, including visual and auditory words, sign language signs, pictures, faces, environmental sounds, and smells. [1] [2] [3]

Contents

History

The N400 was first discovered by Marta Kutas and Steven Hillyard in 1980. [4] They conducted the first experiment looking at the response to unexpected words in read sentences, expecting to elicit a P300 component. The P300 had previously been shown to be elicited by unexpected stimuli. Kutas and Hillyard therefore used sentences with anomalous endings (i.e.I take coffee with cream and dog), expecting to see a P300 to the unexpected sentence-final words. However, instead of eliciting a large positivity, these anomalous endings elicited a large negativity, relative to the sentences with expected endings (i.e. He returned the book to the library) In the same paper, they confirmed that the negativity was not just caused by any unexpected event at the end of a sentence, since a semantically expected but physically unexpected word (i.e. She put on her high-heeled SHOES) elicited a P300 instead of negativity in the N400 window. This finding showed that the N400 is related to semantic processing, and is not just a response to unexpected words.

Component characteristics

The N400 is characterized by a distinct pattern of electrical activity that can be observed at the scalp. As its name indicates, this waveform peaks around 400 ms post-stimulus onset, with negativity that can be observed in the time window ranging from 250-500 ms. This latency (delay between stimulus and response) is very stable across tasks—little else besides age affects the timing of its peak. [2] The N400 is a negative component, relative to reference electrodes placed on the mastoid processes (the bony ridge behind the ear), and relative to a 100 ms pre-stimulus baseline. Its amplitude can range from -5 to 5 microvolts. However, it is important to note that in studies using the N400 as a dependent measure, the relative amplitude of the waveform compared to another experimental condition (the "N400 effect") is more important than its absolute amplitude. The N400 itself is not always negative—it is just a more negative-going deflection than that seen to other conditions. Its distribution is maximal over centro-parietal electrode sites, and is slightly larger over the left side of the head for visual words, although the distribution can change slightly depending on the eliciting stimulus. [2]

Main paradigms

A typical experiment designed to study the N400 will usually involve the visual presentation of words, either in sentence or list contexts. In a typical visual N400 experiment, for example, subjects will be seated in front of a computer monitor while words are presented one-by-one at a central screen location. Stimuli must be presented centrally because eye movements will generate large amounts of electrical noise that will mask the relatively small N400 component. Subjects will often be given a behavioral task (e.g., making a word/nonword decision, answering a comprehension question, responding to a memory probe), either after each stimulus or at longer intervals, to ensure that subjects are paying attention. Note, however, that overt responses by the subject are not required to elicit the N400—passively viewing stimuli will still evoke this response.

An example of an experimental task used to study the N400 is a priming paradigm. Subjects are shown a list of words in which a prime word is either associatively related to a target word (e.g. bee and honey), semantically related (e.g. sugar and honey) or a direct repetition (e.g. honey and honey). The N400 amplitude seen to the target word (honey) will be reduced upon repetition due to semantic priming. [1] The amount of reduction in amplitude can be used to measure the degree of relatedness between the words.

Another widely used experimental task used to study the N400 is sentence reading. In this kind of study, sentences are presented to subjects centrally, one word at a time, until the sentence is completed. Alternatively, subjects could listen to a sentence as natural auditory speech. Again, subjects may be asked to respond to comprehension questions periodically throughout the experiment, although this is not necessary. Experimenters can choose to manipulate various linguistic characteristics of the sentences, including contextual constraint or the cloze probability of the sentence-final word (see below for a definition of cloze probability) to observe how these changes affect the waveform's amplitude.

As previously mentioned, the N400 response is seen to all meaningful, or potentially meaningful, stimuli. As such, a wide range of paradigms can be used to study it. Experiments involving the presentation of spoken words, [5] acronyms, [6] pictures embedded at the end of sentences, [7] music, [8] words related to current context or orientation [9] and videos of real-word events, [10] have all been used to study the N400, just to name a few.

Functional sensitivity

Extensive research has been carried out to better understand what kinds of experimental manipulations do and do not affect the N400. General findings are discussed below.

Factors that affect N400 amplitude

The frequency of a word's usage is known to affect the amplitude of the N400. With all else being constant, highly frequent words elicit reduced N400s relative to infrequent words. [11] As previously mentioned, N400 amplitude is also reduced by repetition, such that a word's second presentation exhibits a more positive response when repeated in context. [12] These findings suggest that when a word is highly frequent or has recently appeared in context, it eases the semantic processing thought to be indexed by the N400, reducing its amplitude.

N400 amplitude is also sensitive to a word's orthographic neighborhood size, or how many other words differ from it by only one letter (e.g. boot and boat). Words with large neighborhoods (that have many other physically similar items) elicit larger N400 amplitudes than do words with small neighborhoods. [13] This finding also holds true for pseudowords, or pronounceable letter strings that are not real words (e.g. flom), which are not themselves meaningful but look like words. This has been taken as evidence that the N400 reflects general activation in the comprehension network, such that an item that looks like many words (regardless of whether it itself is a word) partially activates the representations of similar-looking words, producing a more negative N400.

The N400 is sensitive to priming: in other words, its amplitude is reduced when a target word is preceded by a word that is semantically, morphologically, or orthographically related to it. [1]

In a sentence context, an important determinant of N400 amplitude elicited by a word is its cloze probability. Cloze probability is defined as the probability of the target word completing that particular sentence frame. Kutas and Hillyard (1984) found that a word's N400 amplitude has a nearly inverse linear relationship with its cloze probability. [14] That is, as a word becomes less expected in context, its N400 amplitude is increased relative to more expected words. Words that are incongruent with a context (and thus have a cloze probability of 0) elicit large N400 amplitudes as well (although the amplitude of the N400 for incongruent words is also modulated by the cloze probability of the congruent word that would have been expected in its place [15] Relatedly, the N400 amplitude elicited by open-class words (i.e. nouns, verbs, adjectives, and adverbs) is reduced for words appearing later in a sentence compared to earlier words. [11] Taken together, these findings suggest that when the prior context builds up meaning, it makes the processing of upcoming words that fit with that context easier, reducing the N400 amplitude they elicit.

Factors that do not affect N400 amplitude

While the N400 is larger to unexpected items at the end of a sentence, its amplitude is generally unaffected by negation that causes the last word to be untrue and thus anomalous. [16] For example, in the sentence A sparrow is a building, the N400 response to building is more negative than the N400 response to bird in the sentence A sparrow is a bird. In this case, building has a lower cloze probability, and so it is less expected than bird. However, if negation is added to both sentences in the form of the word not (i.e. A sparrow is not a building and A sparrow is not a bird), the N400 amplitude to building will still be more negative than that seen to bird. This suggests that the N400 responds to the relationship between words in context, but is not necessarily sensitive to the sentence's truth value. More recent research, however, has demonstrated that the N400 can sometimes be modulated by quantifiers or adjectives that serve negation-like purposes, [17] or by pragmatically licensed negation. [18]

Additionally, grammatical violations do not elicit a large N400 response. Rather, these types of violations show a large positivity from about 500-1000 ms after stimulus onset, known as the P600. [2]

Factors that affect N400 latency

A striking feature of the N400 is the general invariance of its peak latency. Although many different experimental manipulations affect the amplitude of the N400, few factors (aging and disease states and language proficiency being rare examples) alter the time it takes for the N400 component to reach a peak amplitude. [19]

Sources

Although localization of the neural generators of an ERP signal is difficult due to the spreading of current from the source to the sensors, multiple techniques can be used to provide converging evidence about possible neural sources. [20] Using methods such as recordings directly off the surface of the brain or from electrodes implanted in the brain, evidence from brain damaged patients, and magnetoencephalographic (MEG) recordings (which measure magnetic activity at the scalp associated with the electrical signal measured by ERPs), the left temporal lobe has been highlighted as an important source for the N400, [21] with additional contributions from the right temporal lobe. [22] More generally, however, activity in a wide network of brain areas is elicited in the N400 time window, suggesting a highly distributed neural source. [2]

Theories

There is still much debate as to exactly what kind of neural and comprehension processes the N400 indexes. Some researchers believe that the underlying processes reflected in the N400 occur after a stimulus has been recognized. For example, Brown and Hagoort (1993) believe that the N400 occurs late in the processing stream, and reflects the integration of a word's meaning into the preceding context (see Kutas & Federmeier, 2011, [2] for a discussion). However, this account has not explained why items that themselves have no meaning (e.g. pseudowords without defined associations) also elicit the N400. Other researchers believe that the N400 occurs much earlier, before words are recognized, and represents neurolinguistics, orthographic or phonological analysis. [23]

More recent accounts posit that the N400 represents a broader range of processes indexing access to semantic memory. According to this account, it represents the binding of information obtained from stimulus input with representations from short- and long-term memory (such as recent context, and accessing a word's meaning in long term memory) that work together to create meaning from the information available in the current context (Federmeier & Laszlo, 2009; see Kutas & Federmeier, 2011 [2] ).

Another account is that the N400 reflects prediction error or surprisal. Word-based surprisal was a strong predictor of N400 amplitude in an ERP corpus. [24] In addition, connectionist models make use of prediction error for learning and linguistic adaptation, and these models can explain several N400/P600 results in terms of prediction error propagation for learning. [25]

It may also be that the N400 reflects a combination of these or other factors. Nieuwland et al. (2019) argue that the N400 is actually made up of two sub-components, with predictability affecting the early part of the N400 (200-500 ms after stimulus onset) and plausibility affecting it later (350-650 ms after stimulus onset). [26] This suggests that the N400 reflects both access to semantic memory (which is sensitive to prediction), and semantic integration (sensitive to plausibility).

As research in the field of electrophysiology continues to progress, these theories will likely be refined to include a complete account of just what the N400 represents.

See also

Related Research Articles

<span class="mw-page-title-main">Neurolinguistics</span> Neuroscience and linguistics-related studies

Neurolinguistics is the study of neural mechanisms in the human brain that control the comprehension, production, and acquisition of language. As an interdisciplinary field, neurolinguistics draws methods and theories from fields such as neuroscience, linguistics, cognitive science, communication disorders and neuropsychology. Researchers are drawn to the field from a variety of backgrounds, bringing along a variety of experimental techniques as well as widely varying theoretical perspectives. Much work in neurolinguistics is informed by models in psycholinguistics and theoretical linguistics, and is focused on investigating how the brain can implement the processes that theoretical and psycholinguistics propose are necessary in producing and comprehending language. Neurolinguists study the physiological mechanisms by which the brain processes information related to language, and evaluate linguistic and psycholinguistic theories, using aphasiology, brain imaging, electrophysiology, and computer modeling.

<span class="mw-page-title-main">Event-related potential</span> Brain response that is the direct result of a specific sensory, cognitive, or motor event

An event-related potential (ERP) is the measured brain response that is the direct result of a specific sensory, cognitive, or motor event. More formally, it is any stereotyped electrophysiological response to a stimulus. The study of the brain in this way provides a noninvasive means of evaluating brain functioning.

<span class="mw-page-title-main">P300 (neuroscience)</span> Event-related potential

The P300 (P3) wave is an event-related potential (ERP) component elicited in the process of decision making. It is considered to be an endogenous potential, as its occurrence links not to the physical attributes of a stimulus, but to a person's reaction to it. More specifically, the P300 is thought to reflect processes involved in stimulus evaluation or categorization.

The mismatch negativity (MMN) or mismatch field (MMF) is a component of the event-related potential (ERP) to an odd stimulus in a sequence of stimuli. It arises from electrical activity in the brain and is studied within the field of cognitive neuroscience and psychology. It can occur in any sensory system, but has most frequently been studied for hearing and for vision, in which case it is abbreviated to vMMN. The (v)MMN occurs after an infrequent change in a repetitive sequence of stimuli For example, a rare deviant (d) stimulus can be interspersed among a series of frequent standard (s) stimuli. In hearing, a deviant sound can differ from the standards in one or more perceptual features such as pitch, duration, loudness, or location. The MMN can be elicited regardless of whether someone is paying attention to the sequence. During auditory sequences, a person can be reading or watching a silent subtitled movie, yet still show a clear MMN. In the case of visual stimuli, the MMN occurs after an infrequent change in a repetitive sequence of images.

The contingent negative variation (CNV) is a negative slow surface potential, as measured by electroencephalography (EEG), that occurs during the period between a warning stimulus or signal and an imperative ("go") stimulus. The CNV was one of the first event-related potential (ERP) components to be described. The CNV component was first described by W. Grey Walter and colleagues in an article published in Nature in 1964. The importance of this finding was that it was one of the first studies which showed that consistent patterns of the amplitude of electric responses could be obtained from the large background noise which occurs in EEG recordings and that this activity could be related to a cognitive process such as expectancy.

The P600 is an event-related potential (ERP) component, or peak in electrical brain activity measured by electroencephalography (EEG). It is a language-relevant ERP component and is thought to be elicited by hearing or reading grammatical errors and other syntactic anomalies. Therefore, it is a common topic of study in neurolinguistic experiments investigating sentence processing in the human brain.

The early left anterior negativity is an event-related potential in electroencephalography (EEG), or component of brain activity that occurs in response to a certain kind of stimulus. It is characterized by a negative-going wave that peaks around 200 milliseconds or less after the onset of a stimulus, and most often occurs in response to linguistic stimuli that violate word-category or phrase structure rules. As such, it is frequently a topic of study in neurolinguistics experiments, specifically in areas such as sentence processing. While it is frequently used in language research, there is no evidence yet that it is necessarily a language-specific phenomenon.

In neuroscience, the N100 or N1 is a large, negative-going evoked potential measured by electroencephalography ; it peaks in adults between 80 and 120 milliseconds after the onset of a stimulus, and is distributed mostly over the fronto-central region of the scalp. It is elicited by any unpredictable stimulus in the absence of task demands. It is often referred to with the following P200 evoked potential as the "N100-P200" or "N1-P2" complex. While most research focuses on auditory stimuli, the N100 also occurs for visual, olfactory, heat, pain, balance, respiration blocking, and somatosensory stimuli.

Music semantics refers to the ability of music to convey semantic meaning. Semantics are a key feature of language, and whether music shares some of the same ability to prime and convey meaning has been the subject of recent study.

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The P3a, or novelty P3, is a component of time-locked (EEG) signals known as event-related potentials (ERP). The P3a is a positive-going scalp-recorded brain potential that has a maximum amplitude over frontal/central electrode sites with a peak latency falling in the range of 250–280 ms. The P3a has been associated with brain activity related to the engagement of attention and the processing of novelty.

In neuroscience, the visual P200 or P2 is a waveform component or feature of the event-related potential (ERP) measured at the human scalp. Like other potential changes measurable from the scalp, this effect is believed to reflect the post-synaptic activity of a specific neural process. The P2 component, also known as the P200, is so named because it is a positive going electrical potential that peaks at about 200 milliseconds after the onset of some external stimulus. This component is often distributed around the centro-frontal and the parieto-occipital areas of the scalp. It is generally found to be maximal around the vertex of the scalp, however there have been some topographical differences noted in ERP studies of the P2 in different experimental conditions.

The N200, or N2, is an event-related potential (ERP) component. An ERP can be monitored using a non-invasive electroencephalography (EEG) cap that is fitted over the scalp on human subjects. An EEG cap allows researchers and clinicians to monitor the minute electrical activity that reaches the surface of the scalp from post-synaptic potentials in neurons, which fluctuate in relation to cognitive processing. EEG provides millisecond-level temporal resolution and is therefore known as one of the most direct measures of covert mental operations in the brain. The N200 in particular is a negative-going wave that peaks 200-350ms post-stimulus and is found primarily over anterior scalp sites. Past research focused on the N200 as a mismatch detector, but it has also been found to reflect executive cognitive control functions, and has recently been used in the study of language.

The late positive component or late positive complex (LPC) is a positive-going event-related brain potential (ERP) component that has been important in studies of explicit recognition memory. It is generally found to be largest over parietal scalp sites, beginning around 400–500 ms after the onset of a stimulus and lasting for a few hundred milliseconds. It is an important part of the ERP "old/new" effect, which may also include modulations of an earlier component similar to an N400. Similar positivities have sometimes been referred to as the P3b, P300, and P600. Here, we use the term "LPC" in reference to this late positive component.

<span class="mw-page-title-main">P3b</span>

The P3b is a subcomponent of the P300, an event-related potential (ERP) component that can be observed in human scalp recordings of brain electrical activity. The P3b is a positive-going amplitude peaking at around 300 ms, though the peak will vary in latency from 250 to 500 ms or later depending upon the task and on the individual subject response. Amplitudes are typically highest on the scalp over parietal brain areas.

N2pc refers to an ERP component linked to selective attention. The N2pc appears over visual cortex contralateral to the location in space to which subjects are attending; if subjects pay attention to the left side of the visual field, the N2pc appears in the right hemisphere of the brain, and vice versa. This characteristic makes it a useful tool for directly measuring the general direction of a person's attention with fine-grained temporal resolution.

Linguistic prediction is a phenomenon in psycholinguistics occurring whenever information about a word or other linguistic unit is activated before that unit is actually encountered. Evidence from eyetracking, event-related potentials, and other experimental methods indicates that in addition to integrating each subsequent word into the context formed by previously encountered words, language users may, under certain conditions, try to predict upcoming words. In particular, prediction seems to occur regularly when the context of a sentence greatly limits the possible words that have not yet been revealed. For instance, a person listening to a sentence like, "In the summer it is hot, and in the winter it is..." would be highly likely to predict the sentence completion "cold" in advance of actually hearing it. A form of prediction is also thought to occur in some types of lexical priming, a phenomenon whereby a word becomes easier to process if it is preceded by a related word. Linguistic prediction is an active area of research in psycholinguistics and cognitive neuroscience.

<span class="mw-page-title-main">Jabberwocky sentence</span>

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Kara D. Federmeier is a professor in the Department of Psychology, Department of Kinesiology, and the Program in Neuroscience at the University of Illinois at Urbana-Champaign. She is known for her work using human electrophysiology to understand the neural basis of cognition, with a focus on language and memory in both younger and older adults.

Seana Coulson is a cognitive scientist known for her research on the neurobiology of language and studies of how meaning is constructed in human language, including experimental pragmatics, concepts, semantics, and metaphors. She is a professor in the Cognitive Science department at University of California, San Diego, where her Brain and Cognition Laboratory focuses on the cognitive neuroscience of language and reasoning.

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