N100

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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 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 (see visual N1, including an illustration), [1] olfactory, [2] heat, [3] pain, [3] balance, [4] respiration blocking, [5] and somatosensory stimuli. [6]

Contents

The auditory N100 is generated by a network of neural populations in the primary and association auditory cortices in the superior temporal gyrus in Heschl's gyrus [7] and planum temporale. [8] It also could be generated in the frontal and motor areas. [9] The area generating it is larger in the right hemisphere than the left. [7]

The N100 is preattentive and involved in perception because its amplitude is strongly dependent upon such things as the rise time of the onset of a sound, [10] its loudness, [11] interstimulus interval with other sounds, [12] and the comparative frequency of a sound as its amplitude increases in proportion to how much a sound differs in frequency from a preceding one. [13] Neuromagnetic research has linked it further to perception by finding that the auditory cortex has a tonotopic organization to N100. [14] However, it also shows a link to a person's arousal [15] and selective attention. [16] N100 is decreased when a person controls the creation of auditory stimuli, [17] such as their own voice. [18]

Types

There are three subtypes of adult auditory N100. [9]

Elicitation

The N100 is often known as the "auditory N100" because it is elicited by perception of auditory stimuli. Specifically, it has been found to be sensitive to things such as the predictability of an auditory stimulus, and special features of speech sounds such as voice onset time.

During sleep

It occurs during both REM and NREM stages of sleep though its time is slightly delayed. [19] During stage 2 NREM it seems responsible for the production of K-complexes. [20] N100 is reduced following total sleep deprivation and this associates with an impaired ability to consolidate memories. [21]

Stimulus repetition

The N100 depends upon unpredictability of stimulus: it is weaker when stimuli are repetitive, and stronger when they are random. When subjects are allowed to control stimuli, using a switch, the N100 may decrease. [17] This effect has been linked to intelligence, as the N100 attenuation for self-controlled stimuli occurs the most strongly (i.e., the N100 shrinks the most) in individuals who are also evaluated as having high intelligence. Indeed, researchers have found that in those with Down syndrome "the amplitude of the self-evoked response actually exceeded that of the machine-evoked potential". [17] Being warned about an upcoming stimulus also reduces its N100. [22]

The amplitude of N100 shows refractoriness upon repetition of a stimulus; in other words, it decreases at first upon repeated presentations of the stimulus, but after a short period of silence it returns to its previous level. [9] Paradoxically, at short repetition the second N100 is enhanced both for sound [23] and somatosensory stimuli. [6]

With paired clicks, the second N100 is reduced due to sensory gating. [24]

Voice onset time

The difference between many consonants is their voice onset time (VOT), the interval between consonant release (onset) and the start of rhythmic vocal cord vibrations in the vowel. The voiced stop consonants /b/, /d/ and /g/ have a short VOT, and unvoiced stop consonants /p/, /t/ and /k/ long VOTs. The N100 plays a role in recognizing the difference and categorizing these sounds: speech stimuli with a short 0 to +30 ms voice onset time evoke a single N100 response but those with a longer (+30 ms and longer) evoked two N100 peaks and these are linked to the consonant release and vocal cord vibration onset. [25] [26]

Top-down influences

Traditionally, 50 to 150 ms evoked potentials were considered too short to be influenced by top-down influences from the prefrontal cortex. However, it is now known that sensory input is processed by the occipital cortex by 56 ms and this is communicated to the dorsolateral frontal cortex where it arrives by 80 ms. [27] Research also finds that the modulation effects upon N100 are affected by prefrontal cortex lesions. [28] These higher-level areas create the attentive, repetition, and arousal modulations upon the sensory area processing reflected in N100. [29]

Another top-down influence upon N100 has been suggested to be efference copies from a person's intended movements so that the stimulation that results from them are not processed. [30] A person's own voice produces a reduced N100 [18] as does the effect of a self-initiated compared to externally created perturbation upon balance. [31]

Development in children

The N100 is a slow-developing evoked potential. From one to four years of age, a positive evoked potential, P100, is the predominant peak. [32] Older children start to develop a negative evoked potential at 200 ms that dominates evoked potentials until adolescence; [33] this potential is identical to the adult N100 in scalp topography and elicitation, but with a much later onset. The magnetic M100 (measured by MEG rather than EEG) is, likewise, less robust in children than in adults. [34] An adult-like N100-P200 complex only develops after 10 years of age. [35]

The various types of N100 mature at different times. Their maturation also varies with the side of the brain: N100a in the left hemisphere is mature before three years of age but this does not happen in the right hemisphere until seven or eight years of age. [33]

Clinical use

The N100 may be used to test for abnormalities in the auditory system where verbal or behavioral responses cannot be used, [36] such with individuals in coma; in such cases, it can help predict the probability of recovery. [37] [38] Another application is in assessing the optimal level of sedation in intensive critical care. [39]

High density mapping of the location of the generators of M100 is being researched as a means of presurgical neuromapping needed for neurosurgery. [40]

Many cognitive or other mental impairments are associated with changes in the N100 response, including the following:

The N100 is 10 to 20% larger than normal when the auditory stimulus is synchronized with the diastolic phase of the cardiac blood pressure pulse. [45]

Relationship to mismatch negativity

The Mismatch negativity (MMN) is an evoked potential that occurs at roughly the same time as N100 in response to rare auditory events. It differs from the N100 in that:

Though this suggests that they are separate processes, arguments have been made that this is not necessarily so and that they are created by the "relative activation of multiple cortical areas contributing to both of these 'components'". [49]

History

Pauline A. Davis at Harvard University first recorded the wave peak now identified with N100. [50] The present use of the N1 to describe this peak originates in 1966 [51] and N100 later in the mid 1970s. [52] The origin of the wave for a long time was unknown and only linked to the auditory cortex in 1970. [9] [53]

Due to magnetoencephalography, research is increasingly done upon M100, the magnetic counterpart of the electroencephalographic N100. Unlike electrical fields which face the high resistance of the skull and generate secondary or volume currents, magnetic fields which are orthogonal to them have a homogeneous permeability through the skull. This enables the location of sources generating fields that are tangent to the head surface with an accuracy of a few millimeters. [54] New techniques, such as event-related beam-forming with magnetoencephalography, allow sufficiently accurate location of M100 sources to be clinically useful for preparing surgery upon the brain. [40]

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.

<span class="mw-page-title-main">Sensory nervous system</span> Part of the nervous system responsible for processing sensory information

The sensory nervous system is a part of the nervous system responsible for processing sensory information. A sensory system consists of sensory neurons, neural pathways, and parts of the brain involved in sensory perception and interoception. Commonly recognized sensory systems are those for vision, hearing, touch, taste, smell, balance and visceral sensation. Sense organs are transducers that convert data from the outer physical world to the realm of the mind where people interpret the information, creating their perception of the world around them.

<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.

Evoked fields are part of the magnetoencephalogram. They are brain signals evoked by sensory stimulation, but usually buried by the ongoing brain activity. Repeating the stimulus multiple times and averaging the signals reduces the uncorrelated ongoing activity and reveals the evoked field. Evoked fields are the magnetoencephalographic equivalent to evoked potentials, which are part of the electroencephalogram.

Brainwave entrainment, also referred to as brainwave synchronization or neural entrainment, refers to the observation that brainwaves will naturally synchronize to the rhythm of periodic external stimuli, such as flickering lights, speech, music, or tactile stimuli.

<span class="mw-page-title-main">Neural oscillation</span> Brainwaves, repetitive patterns of neural activity in the central nervous system

Neural oscillations, or brainwaves, are rhythmic or repetitive patterns of neural activity in the central nervous system. Neural tissue can generate oscillatory activity in many ways, driven either by mechanisms within individual neurons or by interactions between neurons. In individual neurons, oscillations can appear either as oscillations in membrane potential or as rhythmic patterns of action potentials, which then produce oscillatory activation of post-synaptic neurons. At the level of neural ensembles, synchronized activity of large numbers of neurons can give rise to macroscopic oscillations, which can be observed in an electroencephalogram. Oscillatory activity in groups of neurons generally arises from feedback connections between the neurons that result in the synchronization of their firing patterns. The interaction between neurons can give rise to oscillations at a different frequency than the firing frequency of individual neurons. A well-known example of macroscopic neural oscillations is alpha activity.

<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.

Sensory gating describes neural processes of filtering out redundant or irrelevant stimuli from all possible environmental stimuli reaching the brain. Also referred to as gating or filtering, sensory gating prevents an overload of information in the higher cortical centers of the brain. Sensory gating can also occur in different forms through changes in both perception and sensation, affected by various factors such as "arousal, recent stimulus exposure, and selective attention."

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.

Echoic memory is the sensory memory that registers specific to auditory information (sounds). Once an auditory stimulus is heard, it is stored in memory so that it can be processed and understood. Unlike most visual memory, where a person can choose how long to view the stimulus and can reassess it repeatedly, auditory stimuli are usually transient and cannot be reassessed. Since echoic memories are heard once, they are stored for slightly longer periods of time than iconic memories. Auditory stimuli are received by the ear one at a time before they can be processed and understood.

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 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.

Somatosensory evoked potential 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 combining SEP recordings at different levels of the somatosensory pathways, it is possible to assess the transmission of the afferent volley from the periphery up to the cortex. SEP components include a series of positive and negative deflections that can be elicited by virtually any sensory stimuli. For example, SEPs can be obtained in response to a brief mechanical impact on the fingertip or to air puffs. However, SEPs are most commonly elicited by bipolar transcutaneous electrical stimulation applied on the skin over the trajectory of peripheral nerves of the upper limb or lower limb, and then recorded from the scalp. In general, somatosensory stimuli evoke early cortical components, generated in the contralateral primary somatosensory cortex (S1), related to the processing of the physical stimulus attributes. About 100 ms after stimulus application, additional cortical regions are activated, such as the secondary somatosensory cortex (S2), and the posterior parietal and frontal cortices, marked by a parietal P100 and bilateral frontal N140. SEPs are routinely used in neurology today to confirm and localize sensory abnormalities, to identify silent lesions and to monitor changes during surgical procedures.

<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.

The oddball paradigm is an experimental design used within psychology research. Presentations of sequences of repetitive stimuli are infrequently interrupted by a deviant stimulus. The reaction of the participant to this "oddball" stimulus is recorded.

The frequency following response (FFR), also referred to as frequency following potential (FFP) or envelope following response (EFR), is an evoked potential generated by periodic or nearly-periodic auditory stimuli. Part of the auditory brainstem response (ABR), the FFR reflects sustained neural activity integrated over a population of neural elements: "the brainstem response...can be divided into transient and sustained portions, namely the onset response and the frequency-following response (FFR)". It is often phase-locked to the individual cycles of the stimulus waveform and/or the envelope of the periodic stimuli. It has not been well studied with respect to its clinical utility, although it can be used as part of a test battery for helping to diagnose auditory neuropathy. This may be in conjunction with, or as a replacement for, otoacoustic emissions.

Musicogenic seizure, also known as music-induced seizure, is a rare type of seizure, with an estimated prevalence of 1 in 10,000,000 individuals, that arises from disorganized or abnormal brain electrical activity when a person hears or is exposed to a specific type of sound or musical stimuli. There are challenges when diagnosing a music-induced seizure due to the broad scope of triggers, and time delay between a stimulus and seizure. In addition, the causes of musicogenic seizures are not well-established as solely limited cases and research have been discovered and conducted respectively. Nevertheless, the current understanding of the mechanism behind musicogenic seizure is that music triggers the part of the brain that is responsible for evoking an emotion associated with that music. Dysfunction in this system leads to an abnormal release of dopamine, eventually inducing seizure.

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