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The P600 is an
event-related potential An event-related potential (ERP) is the measured brain response that is the direct result of a specific sense, sensory, cognition, cognitive, or motor system, motor event. More formally, it is any stereotyped electrophysiology, electrophysiologi ...
(ERP) component, or peak in electrical brain activity measured by
electroencephalography Electroencephalography (EEG) is a method to record an electrogram of the spontaneous electrical activity of the brain. The biosignals detected by EEG have been shown to represent the postsynaptic potentials of pyramidal neurons in the neocortex ...
(EEG). It is a language-relevant ERP component and is thought to be elicited by hearing or reading
grammatical In linguistics, grammaticality is determined by the conformity to language usage as derived by the grammar of a particular variety (linguistics), speech variety. The notion of grammaticality rose alongside the theory of generative grammar, the go ...
errors and other
syntactic In linguistics, syntax () is the study of how words and morphemes combine to form larger units such as phrases and sentences. Central concerns of syntax include word order, grammatical relations, hierarchical sentence structure (constituency), ...
anomalies. Therefore, it is a common topic of study in neurolinguistic experiments investigating
sentence processing Sentence processing takes place whenever a reader or listener processes a language utterance, either in isolation or in the context of a conversation or a text. Many studies of the human language comprehension process have focused on reading of s ...
in the human brain. The P600 can be elicited in both visual (reading) and auditory (listening) experiments, and is characterized as a positive-going deflection with an onset around 500 milliseconds after the stimulus that elicits it; it often reaches its peak around 600 milliseconds after presentation of the stimulus (hence its name), and lasts several hundred milliseconds.Different authors give slightly different time periods for the P600; for example, claim it may start as early as 400 milliseconds, and describes it as occurring "between 600–1000 ms. In other words, in the EEG waveform it is a large peak in the positive direction, which starts around 500 milliseconds after the subject sees or hears a stimulus. It is typically thought of as appearing mostly on centro- parietal electrodes (i.e., over the posterior part of the center of the scalp), but frontal P600s have also been observed in several studies. In EEG, however, this distribution at the scalp does not mean the P600 is coming from that part of the brain; a 2007 study using
magnetoencephalography Magnetoencephalography (MEG) is a functional neuroimaging technique for mapping brain activity by recording magnetic fields produced by electrical currents occurring naturally in the brain, using very sensitive magnetometers. Arrays of SQUIDs (s ...
(MEG) speculates that the generators of the P600 are in the posterior
temporal lobe The temporal lobe is one of the four Lobes of the brain, major lobes of the cerebral cortex in the brain of mammals. The temporal lobe is located beneath the lateral fissure on both cerebral hemispheres of the mammalian brain. The temporal lobe ...
, behind
Wernicke's area Wernicke's area (; ), also called Wernicke's speech area, is one of the two parts of the cerebral cortex that are linked to speech, the other being Broca's area. It is involved in the comprehension of written and spoken language, in contrast to B ...
. The P600 was first reported by Lee Osterhout and Phillip Holcomb in 1992. It is also sometimes called the syntactic positive shift (SPS), since it has a positive polarity and is usually elicited by syntactic phenomena.


Elicitation

The P600 was originally considered a "syntactic" ERP component, as it is elicited by several types of syntactic phenomena, including ungrammatical stimuli,
garden-path sentence A garden-path sentence is a grammatically correct sentence that starts in such a way that a reader's most likely interpretation will be incorrect; the reader is lured into a parse that turns out to be a dead end or yields a clearly unintended me ...
s that require reanalysis, complex sentences with a large number of thematic roles, and the processing of filler-gap dependencies (such as
wh-word An interrogative word or question word is a function word used to ask a question, such as ''what, which'', ''when'', ''where'', ''who, whom, whose'', ''why'', ''whether'' and ''how''. They are sometimes called wh-words, because in English most ...
s that appear at the beginning of a sentence in
English English usually refers to: * English language * English people English may also refer to: Peoples, culture, and language * ''English'', an adjective for something of, from, or related to England ** English national ide ...
but are actually interpreted somewhere else). ;Grammatical errors A P600 may be elicited by several kinds of grammatical errors in sentences, such as problems in
agreement Agreement may refer to: Agreements between people and organizations * Gentlemen's agreement, not enforceable by law * Trade agreement, between countries * Consensus, a decision-making process * Contract, enforceable in a court of law ** Meeting o ...
, such as "the child *''throw'' the toy". In addition to this sort of subject-verb disagreement, P600s have also been elicited by disagreements in tense,
gender Gender is the range of characteristics pertaining to femininity and masculinity and differentiating between them. Depending on the context, this may include sex-based social structures (i.e. gender roles) and gender identity. Most cultures u ...
,
number A number is a mathematical object used to count, measure, and label. The original examples are the natural numbers 1, 2, 3, 4, and so forth. Numbers can be represented in language with number words. More universally, individual numbers c ...
, and case, as well as
phrase structure Phrase structure rules are a type of rewrite rule used to describe a given language's syntax and are closely associated with the early stages of transformational grammar, proposed by Noam Chomsky in 1957. They are used to break down a natural langu ...
violations. A 2009 study has suggested that these errors elicit stronger P600s than the other syntactic stimuli that have been implicated. ;Garden paths P600s are also known to occur when a sentence contains no outright grammatical error, but must be parsed in a different way than the reader originally expects. These sentences are known as "garden path" sentences, because the reader follows one interpretation of the sentence only to realize later that this interpretation was wrong and he must backtrack to understand the sentence. For example, found P600s elicited by the word ''to'' in sentences such as
The broker persuaded to sell the stock was tall.
In sentences such as this, the preferred reading is to interpret "persuaded" as the main verb of the sentence (i.e., "the broker persuaded me"), and upon seeing the word ''to'' the reader has to re-analyze the sentence to mean something more like "the broker that was persuaded to sell the stock, he was tall". ;Syntactic errors in music P600s are also elicited by errors in musical harmony, such as when a chord is played out of key with the rest of a musical phrase. This implies that P600s are not "language-specific," but "can be elicited in nonlinguistic (but rule-governed) sequences." ;Dependencies and complexity Some studies have found a P600 elicited by words where there is no grammatical error and no "garden path" (i.e., where the word is exactly what the reader would expect), but when the sentence is complex because there are a number of
noun phrase In linguistics, a noun phrase, or nominal (phrase), is a phrase that has a noun or pronoun as its head or performs the same grammatical function as a noun. Noun phrases are very common cross-linguistically, and they may be the most frequently oc ...
s active. This has most often been the case when the reader has to "re-activate" a word that appeared earlier in the sentence. For example, in a sentence like "Who did you imitate?", the word ''who'' appears in the beginning of the sentence but is actually the direct object of ''imitate'', and must be interpreted in that way (i.e., as "you imitated who?"); several studies have found that after the reader sees the word ''imitate'' he or she has a P600 response, possibly as a result of re-activating ''who''. These sorts of P600s get stronger as the number of noun phrases active in the sentences increases, suggesting that the P600 generator is sensitive to the level of complexity in a sentence. ;Semantic attraction demonstrated a so-called "semantic P600" in sentences that are grammatically correct but semantically anomalous, and in which syntactic reanalysis is more appealing than semantic reanalysis. For example, a P600 may be elicited in the following sentence:
The hearty meal was devouring the kids.
This suggests that the reader would rather interpret the sentence as containing a morphosyntactic error (saying "devour''ing''" instead of "devour''ed by''") rather than a semantic one (meals can't devour kids, but can be devoured by them). The interpretation of "semantic P600s" has attracted considerable attention and controversy in the literature.


Interpretations

There are several theories about what computational processes the P600 may be triggered by. Because it often happens in response to grammatical violations or garden path sentences, one theory is that the P600 reflects processes of ''revision'' (i.e., trying to "rescue" the interpretation of a sentence that can't be processed normally because of structural errors) and ''reanalysis'' (i.e., trying to rearrange the structure of a sentence that has been interpreted incorrectly because of a garden path). On the other hand, other models suggest that the P600 may not reflect these processes in particular, but just the amount of time and effort in general it takes to build up coherent structure in a sentence, or the general processes of creating or destroying syntactic structure (not specifically because of repair). Another proposal is that the P600 does not necessarily reflect any linguistic processes per se, but is similar to the P300 in that it is triggered when a subject encounters "improbable" stimuli—since ungrammatical sentences are relatively rare in natural speech, a P600 may not be a linguistic response but simply an effect of the subject's "surprise" upon encountering an unexpected stimulus. Another account is that the P600 reflects error/surprisal propagation due to learning processes that take place during linguistic adaptation and this account has been implemented in a connectionist model that explains several P600/N400 results.


See also

*
Bereitschaftspotential In neurology, the Bereitschaftspotential or BP (German for "readiness potential"), also called the pre-motor potential or readiness potential (RP), is a measure of activity in the motor cortex and supplementary motor area of the brain leading up t ...
* C1 and P1 *
Contingent negative variation 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 ...
*
Difference due to memory Difference due to memory (Dm) indexes differences in neural activity during the study phase of an experiment for items that subsequently are remembered compared to items that are later forgotten. It is mainly discussed as an event-related potential ...
*
Early left anterior negativity The early left anterior negativity (commonly referred to as ELAN) 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 negativ ...
* Error-related negativity * Late positive component * Lateralized readiness potential * Mismatch negativity * N2pc *
N100 In neuroscience, the N100 or N1 is a large, negative-going evoked potential measured by electroencephalography (its equivalent in magnetoencephalography is the M100); it peaks in adults between 80 and 120 milliseconds after the onset of a stimul ...
* N170 * N200 * N400 *
P3a 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 l ...
*
P3b 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 (usually relative to a reference behind the ear ...
*
P200 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-synapti ...
*
P300 (neuroscience) 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' ...
*
Somatosensory evoked potential Somatosensory evoked potential (SEP or SSEP) is the electrical activity of the brain that results from the stimulation of touch. SEP tests measure that activity and are a useful, noninvasive means of assessing somatosensory system functioning. By co ...
*
Visual N1 The visual N1 is a visual evoked potential, a type of event-related electrical potential (ERP), that is produced in the brain and recorded on the scalp. The N1 is so named to reflect the polarity and typical timing of the component. The "N" indi ...


Notes


References


Bibliography

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External links


Video of the N400 and P600 visualized as animated scalp topographies
{{Neuroscience Electroencephalography Evoked potentials Neurolinguistics