Event-related potential

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A waveform showing several ERP components, including the N100 (labeled N1) and P300 (labeled P3). The ERP is plotted with negative voltages upward, a common, but not universal, practice in ERP research ComponentsofERP.svg
A waveform showing several ERP components, including the N100 (labeled N1) and P300 (labeled P3). The ERP is plotted with negative voltages upward, a common, but not universal, practice in ERP research

An event-related potential (ERP) is the measured brain response that is the direct result of a specific sensory, cognitive, or motor event. [1] 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.

Contents

ERPs are measured by means of electroencephalography (EEG). The magnetoencephalography (MEG) equivalent of ERP is the ERF, or event-related field. [2] Evoked potentials and induced potentials are subtypes of ERPs.

History

With the discovery of the electroencephalogram (EEG) in 1924, Hans Berger revealed that one could measure the electrical activity of the human brain by placing electrodes on the scalp and amplifying the signal. Changes in voltage can then be plotted over a period of time. He observed that the voltages could be influenced by external events that stimulated the senses. The EEG proved to be a useful source in recording brain activity over the ensuing decades. However, it tended to be very difficult to assess the highly specific neural process that are the focus of cognitive neuroscience because using pure EEG data made it difficult to isolate individual neurocognitive processes. Event-related potentials (ERPs) offered a more sophisticated method of extracting more specific sensory, cognitive, and motor events by using simple averaging techniques. In 1935–1936, Pauline and Hallowell Davis recorded the first known ERPs on awake humans and their findings were published a few years later, in 1939. Due to World War II not much research was conducted in the 1940s, but research focusing on sensory issues picked back up again in the 1950s. In 1964, research by Grey Walter and colleagues began the modern era of ERP component discoveries when they reported the first cognitive ERP component, called the contingent negative variation (CNV). [3] Sutton, Braren, and Zubin (1965) made another advancement with the discovery of the P3 component. [4] Over the next fifteen years, ERP component research became increasingly popular. The 1980s, with the introduction of inexpensive computers, opened up a new door for cognitive neuroscience research. Currently, ERP is one of the most widely used methods in cognitive neuroscience research to study the physiological correlates of sensory, perceptual and cognitive activity associated with processing information. [5]

Calculation

ERPs can be reliably measured using electroencephalography (EEG), a procedure that measures electrical activity of the brain over time using electrodes placed on the scalp. The EEG reflects thousands of simultaneously ongoing brain processes. This means that the brain response to a single stimulus or event of interest is not usually visible in the EEG recording of a single trial. To see the brain's response to a stimulus, the experimenter must conduct many trials and average the results together, causing random brain activity to be averaged out and the relevant waveform to remain, called the ERP. [6] [7]

The random (background) brain activity together with other bio-signals (e.g., EOG, EMG, EKG) and electromagnetic interference (e.g., line noise, fluorescent lamps) constitute the noise contribution to the recorded ERP. This noise obscures the signal of interest, which is the sequence of underlying ERPs under study. From an engineering point of view it is possible to define the signal-to-noise ratio (SNR) of the recorded ERPs. Averaging increases the SNR of the recorded ERPs making them discernible and allowing for their interpretation. This has a simple mathematical explanation provided that some simplifying assumptions are made. These assumptions are:

  1. The signal of interest is made of a sequence of event-locked ERPs with invariable latency and shape
  2. The noise can be approximated by a zero-mean Gaussian random process of variance which is uncorrelated between trials and not time-locked to the event (this assumption can be easily violated, for example in the case of a subject doing little tongue movements while mentally counting the targets in an experiment).

Having defined , the trial number, and , the time elapsed after the th event, each recorded trial can be written as where is the signal and is the noise (Under the assumptions above, the signal does not depend on the specific trial while the noise does).

The average of trials is

.

The expected value of is (as hoped) the signal itself, .

Its variance is

.

For this reason the noise amplitude of the average of trials is expected to deviate from the mean (which is ) by less or equal than in 68% of the cases. In particular, the deviation wherein 68% of the noise amplitudes lie is times that of a single trial. A larger deviation of can already be expected to encompass 95% of all noise amplitudes.

Wide amplitude noise (such as eye blinks or movement artifacts) are often several orders of magnitude larger than the underlying ERPs. Therefore, trials containing such artifacts should be removed before averaging. Artifact rejection can be performed manually by visual inspection or using an automated procedure based on predefined fixed thresholds (limiting the maximum EEG amplitude or slope) or on time-varying thresholds derived from the statistics of the set of trials.[ citation needed ]

Nomenclature

ERP waveforms consist of a series of positive and negative voltage deflections, which are related to a set of underlying components. [8] Though some ERP components are referred to with acronyms (e.g., contingent negative variation  – CNV, error-related negativity  – ERN), most components are referred to by a letter (N/P) indicating polarity (negative/positive), followed by a number indicating either the latency in milliseconds or the component's ordinal position in the waveform. For instance, a negative-going peak that is the first substantial peak in the waveform and often occurs about 100 milliseconds after a stimulus is presented is often called the N100 (indicating its latency is 100 ms after the stimulus and that it is negative) or N1 (indicating that it is the first peak and is negative); it is often followed by a positive peak, usually called the P200 or P2. The stated latencies for ERP components are often quite variable, particularly so for the later components that are related to the cognitive processing of the stimulus. For example, the P300 component may exhibit a peak anywhere between 250 ms – 700 ms.

Advantages and disadvantages

Relative to behavioral measures

Compared with behavioral procedures, ERPs provide a continuous measure of processing between a stimulus and a response, making it possible to determine which stage(s) are being affected by a specific experimental manipulation. Another advantage over behavioral measures is that they can provide a measure of processing of stimuli even when there is no behavioral change. However, because of the significantly small size of an ERP, it usually takes a large number of trials to accurately measure it correctly. [9]

Relative to other neurophysiological measures

Invasiveness

Unlike microelectrodes, which require an electrode to be inserted into the brain, and PET scans that expose humans to radiation, ERPs use EEG, a non-invasive procedure.

Spatial and temporal resolution

ERPs provide excellent temporal resolution—as the speed of ERP recording is only constrained by the sampling rate that the recording equipment can feasibly support, whereas hemodynamic measures (such as fMRI, PET, and fNIRS) are inherently limited by the slow speed of the BOLD response. The spatial resolution of an ERP, however, is much poorer than that of hemodynamic methods—in fact, the location of ERP sources is an inverse problem that cannot be exactly solved, only estimated. Thus, ERPs are well suited to research questions about the speed of neural activity, and are less well suited to research questions about the location of such activity. [1]

Cost

ERP research is much cheaper to do than other imaging techniques such as fMRI, PET, and MEG. This is because purchasing and maintaining an EEG system is less expensive than the other systems.

Clinical

Physicians and neurologists will sometimes use a flashing visual checkerboard stimulus to test for any damage or trauma in the visual system. In a healthy person, this stimulus will elicit a strong response over the primary visual cortex located in the occipital lobe, in the back of the brain.

ERP component abnormalities in clinical research have been shown in neurological conditions such as:

Research

ERPs are used extensively in neuroscience, cognitive psychology, cognitive science, and psycho-physiological research. Experimental psychologists and neuroscientists have discovered many different stimuli that elicit reliable ERPs from participants. The timing of these responses is thought to provide a measure of the timing of the brain's communication or timing of information processing. For example, in the checkerboard paradigm described above, healthy participants' first response of the visual cortex is around 50–70 ms. This would seem to indicate that this is the amount of time it takes for the transduced visual stimulus to reach the cortex after light first enters the eye. Alternatively, the P300 response occurs at around 300ms in the oddball paradigm, for example, regardless of the type of stimulus presented: visual, tactile, auditory, olfactory, gustatory, etc. Because of this general invariance with regard to stimulus type, the P300 component is understood to reflect a higher cognitive response to unexpected and/or cognitively salient stimuli. The P300 response has also been studied in the context of information and memory detection. [23] In addition, there are studies on abnormalities of P300 in depression. Depressed patients tend to have a reduced P200 and P300 amplitude and a prolonged P300 latency. [20]

Due to the consistency of the P300 response to novel stimuli, a brain–computer interface can be constructed which relies on it. By arranging many signals in a grid, randomly flashing the rows of the grid as in the previous paradigm, and observing the P300 responses of a subject staring at the grid, the subject may communicate which stimulus he is looking at, and thus slowly "type" words. [24]

Another area of research in the field of ERP lies in the efference copy. This predictive mechanism plays a central role in for example human verbalization. [25] [26] Efference copies, however, do not only occur with spoken words, but also with inner language - i.e. the quiet production of words - which has also been proven by event-related potentials. [27]

Other ERPs used frequently in research, especially neurolinguistics research, include the ELAN, the N400, and the P600/SPS. The analysis of ERP data is also increasingly supported by machine learning algorithms. [28] [29]

Number of trials

A common issue in ERP studies is whether the observed data have a sufficient number of trials to support statistical analysis. [30] The background noise in any ERP for any individual can vary. Therefore simply characterizing the number of ERP trials needed for a robust component response is inadequate. ERP researchers can use metrics like the standardized measurement error (SME) to justify the examination of between-condition or between-group differences [31] or estimates of internal consistency to justify the examination of individual differences. [32] [33] [30]

See also

Related Research Articles

An evoked potential or evoked response is an electrical potential in a specific pattern recorded from a specific part of the nervous system, especially the brain, of a human or other animals following presentation of a stimulus such as a light flash or a pure tone. Different types of potentials result from stimuli of different modalities and types. Evoked potential is distinct from spontaneous potentials as detected by electroencephalography (EEG), electromyography (EMG), or other electrophysiologic recording method. Such potentials are useful for electrodiagnosis and monitoring that include detections of disease and drug-related sensory dysfunction and intraoperative monitoring of sensory pathway integrity.

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 stimuli, including visual and auditory words, sign language signs, pictures, faces, environmental sounds, and smells.

<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.

In cognitive psychology, the Eriksen flanker task is a set of response inhibition tests used to assess the ability to suppress responses that are inappropriate in a particular context. The target is flanked by non-target stimuli which correspond either to the same directional response as the target, to the opposite response, or to neither. The task is named for American psychologists Barbara. A. Eriksen & Charles W. Eriksen, who first published the task in 1974, and for the flanker stimuli that surround the target. In the tests, a directional response is assigned to a central target stimulus. Various forms of the task are used to measure information processing and selective attention.

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 lateralized readiness potential (LRP) is an event-related brain potential, or increase in electrical activity at the surface of the brain, that is thought to reflect the preparation of motor activity on a certain side of the body; in other words, it is a spike in the electrical activity of the brain that happens when a person gets ready to move one arm, leg, or foot. It is a special form of bereitschaftspotential. LRPs are recorded using electroencephalography (EEG) and have numerous applications in cognitive neuroscience.

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.

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 (ERP) effect that appears in studies employing a subsequent memory paradigm, in which ERPs are recorded when a participant is studying a list of materials and trials are sorted as a function of whether they go on to be remembered or not in the test phase. For meaningful study material, such as words or line drawings, items that are subsequently remembered typically elicit a more positive waveform during the study phase. This difference typically occurs in the range of 400–800 milliseconds (ms) and is generally greatest over centro-parietal recording sites, although these characteristics are modulated by many factors.

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.

<span class="mw-page-title-main">Visual N1</span>

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" indicates that the polarity of the component is negative with respect to an average mastoid reference. The "1" originally indicated that it was the first negative-going component, but it now better indexes the typical peak of this component, which is around 150 to 200 milliseconds post-stimulus. The N1 deflection may be detected at most recording sites, including the occipital, parietal, central, and frontal electrode sites. Although, the visual N1 is widely distributed over the entire scalp, it peaks earlier over frontal than posterior regions of the scalp, suggestive of distinct neural and/or cognitive correlates. The N1 is elicited by visual stimuli, and is part of the visual evoked potential – a series of voltage deflections observed in response to visual onsets, offsets, and changes. Both the right and left hemispheres generate an N1, but the laterality of the N1 depends on whether a stimulus is presented centrally, laterally, or bilaterally. When a stimulus is presented centrally, the N1 is bilateral. When presented laterally, the N1 is larger, earlier, and contralateral to the visual field of the stimulus. When two visual stimuli are presented, one in each visual field, the N1 is bilateral. In the latter case, the N1's asymmetrical skewedness is modulated by attention. Additionally, its amplitude is influenced by selective attention, and thus it has been used to study a variety of attentional processes.

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.

Error-related negativity (ERN), sometimes referred to as the Ne, is a component of an event-related potential (ERP). ERPs are electrical activity in the brain as measured through electroencephalography (EEG) and time-locked to an external event or a response. A robust ERN component is observed after errors are committed during various choice tasks, even when the participant is not explicitly aware of making the error; however, in the case of unconscious errors the ERN is reduced. An ERN is also observed when non-human primates commit errors.

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.

The N170 is a component of the event-related potential (ERP) that reflects the neural processing of faces, familiar objects or words. Furthermore, the N170 is modulated by prediction error processes.

<span class="mw-page-title-main">Oddball paradigm</span> Psychology research paradigm

The oddball paradigm is an experimental design used within psychology research. The oddball paradigm relies on the brain's sensitivity to rare deviant stimuli presented pseudo-randomly in a series of repeated standard stimuli. The oddball paradigm has a wide selection of stimulus types, including stimuli such as sound duration, frequency, intensity, phonetic features, complex music, or speech sequences. The reaction of the participant to this "oddball" stimulus is recorded.

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Further reading