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导读
ERP是Event-related Potentials的简称。ERP波形通常用正峰和负峰来描述,并用P1、P2和N1、N2等这样的符号来表示波形的极性与所在时间段或达到峰值的位置,例如,N1表示波形中的第一个负峰,P2表示第二个正峰。但要注意的是相同的符号不等同于任何功能上的相关。以下简要介绍了经典的ERP成分,旨在为对认知神经心理学研究感兴趣的小伙伴提供参考。
视觉感官反应
C1:通常起始于刺激后40-60ms,在80-100ms达到峰值。最大幅值出现在头皮后部的中线电极位置,具有极性变化,来自V1区(初级视觉皮层),在距状裂周围。可通过上视野的刺激诱发负向C1波。
推荐阅读:
Source locations of pattern-specific components of human visual evoked potentials. I. Component of striate cortical origin.
Localization of neural generators during visual selective attention.
Cortical sources of the early components of the visual evoked potential.
P1:通常起始于刺激后60-90ms,在100-130ms达到峰值。位于两侧枕区,可能会与C1波重叠。fMRI确定P1波早期起源于背侧纹外皮层(枕中回),晚期起源主要在腹侧梭状回的腹侧部分。
推荐阅读:
Sensory gain control (amplification) as a mechanism of selective attention: electrophysiological and neuroimaging evidence.
Event-related potential studies of attention.
Flow of activation from V1 to frontal cortex in humans.
N1:视觉N1包含多个子成分。子成分1在刺激后100-150ms达到峰值,位于头前部电极位置;子成分2在刺激后150-200ms达到峰值,来自顶叶皮层;子成分3在刺激后150-200ms达到峰值,来自外侧枕叶皮层,这个子成分可能体现了某种与辨别相关的处理过程。
推荐阅读:
The visual N1 component as an index of a discrimination process.
Neural systems mediating selective attention.
Localizing Visual Discrimination Processes in Time and Space.
A brain event related to the making of a sensory discrimination.
N170与顶正波:顶正波(vertex positive potential,VPP)是面孔刺激与非面孔刺激的差异波,在150-200ms间达到峰值,位于中央区头皮的中线部位。N170是指在右半球外侧枕叶电极位置发现的,面孔刺激比非面孔刺激诱发更负的电位,约在170ms左右达到峰值。倒置面孔比正向面孔诱发更大的N170,称为倒置效应或面孔特异性。
推荐阅读:
A face-responsive potential recorded from the human scalp.
Electrophysiological Studies of Face Perception in Humans.
Spatio-temporal localization of the face inversion effect: an event-related potentials study.
The N170: Understanding the Time Course of Face Perception in the Human Brain.
Does physical interstimulus variance account for early electrophysiological face sensitive responses in the human brain? Ten lessons on the N170.
Meaningful processing of meaningless stimuli: The influence of perceptual experience on early visual processing of faces.
A Neural Basis for Expert Object Recognition.
Long-term expertise with artificial objects increases visual competition with early face categorization processes.
P2:位于头前部和中央区,靶刺激相对罕见时反应会增强。例如,在视觉Oddball范式中,奇异刺激会诱发出比标准刺激更大的P2和P3波幅,但区别是P2效应仅在刺激是基于相对简单的刺激特征进行定义时才会出现,而P3是在任意复杂刺激下都会出现。
推荐阅读:
Electrophysiological correlates of feature analysis during visual search.
听觉感官反应
早潜伏期反应:听觉刺激开始后的10ms内会观察到一系列ERP峰。脑干诱发反应(brainstemevoked responses,BERs)或称听觉脑干反应(auditory brainstem responses,ABRs),用于评价听觉通道的完整性。中潜伏期反应是指10-50ms之间的反应,源于内侧膝状体和初级听觉皮层。长潜伏期反应通常的顺序是P50(又叫P1)、N100(又叫N1)、P160(又叫P2),受高级认知因素的调控。N1:包含几种不同的子成分。(1)额-中央部分,约75ms达到峰值,源于背侧颞叶听觉皮层;(2)顶区,约100ms达到峰值,源位置尚未知;(3)约150ms达到峰值,可能源于额上回,N1波对注意力敏感。
推荐阅读:
Sensory ERP Components.
Early ERP components to gaps in white noise: onset and offset effects.
Human Auditory Evoked Potentials.
Auditory event-related potentials studies of information processing during human sleep.
Two separate frontal components in the N1 wave of the human auditory evoked response.
The effects of channel-selective attention on the mismatch negativity wave elicited by deviant tones.
体感、嗅觉和味觉反应
N10:来自外周神经,体感刺激诱发的是反应动作电位而不是突触后电位;N10反应后跟随一系列皮下成分,约10-20ms;短潜伏期和中潜伏期的皮层成分,约20-100ms;随后便是N1约150ms;P2约 200ms。
推荐阅读:
Differential effects of active attention and age on event-related potentials to visual and olfactory stimuli.
Segregation of gustatory cortex in response to salt and umami taste studied through event-related potentials.
N2家族
N2a或称MMN:N2a由听觉失匹配条件下自动诱发的效应,刺激与任务无关时也会被诱发,这一效应通常被称为失匹配负波。约100-200ms附近的负向波。
N2b或称前部N2成分:异常刺激是与任务相关的,对于听觉刺激,主要在中央区位置,视觉刺激在某些情况下也会诱发N2b成分。
N2c或称后部N2成分:主要分布于头皮后部区域,由任务相关的靶刺激诱发,很像P3波形,而且不常出现的靶刺激会诱发更大的N2c成分。
N2pc或称对侧负波:出现时间与N2c成分大致相同,出现在被注意物体的对侧后部头皮,反应了注意力集中的某个方面。一般在完成工作记忆任务时出现。
推荐阅读:
Psychophysiology of N200/N400: A review and classification scheme.
A brain event related to the making of a sensory discrimination.
Memory-based detection of task-irrelevant visual changes.
A dedicated low-level mechanism for visual-deviance detection.
The scalp topography of potentials in auditory and visual discrimination tasks.
Novelty and conflict in the categorization of complex stimuli.
Influence of cognitive control and mismatch on the N2 component of the ERP: A review.
Inhibition, response mode, and stimulus probability: a comparative event-related potential study.
Predictive value of novel stimuli modifies visual event-related potentials and behavior.
An ERP study of the temporal course of the Stroop color-word interference effect.
The feedback-related negativity refects the binary evaluation of good versus bad outcomes.
Correct and Incorrect Responses in a Choice Reaction Time Task and the Endogenous Components of the Evoked Potential.
Time-course of hemispheric preference for processing contralateral relevant shapes: P1pc, N1pc, N2pc, N3pc.
P3家族
P3a:额区最大,由意外而不经常的,即刺激中不可预测且出现概率较小的变化所诱发。
P3b:顶区最大,由意外而不经常的,即刺激中不可预测且出现概率较小的变化所诱发,但是该成分仅出现在这些刺激的变化与任务相关时,绝大多数研究者通常将P3b成分就用P3来指代。
这里,给大家分享一篇文章经典再读 | P3a与P3b的整合理论,以加深对P3a与P3b的理解。
类P3反应:在被试的刺激序列中,出现的不可预期、令人意外、不寻常或使人惊讶的任务无关的刺激会在额区诱发一个类似P3的反应。
注:P3波的标志性特点是其对靶概率的敏感性,靶概率越低,振幅就越大;被试对任务付出更多的努力时,振幅也越大,P3振幅=不确定性(U)×(概率P+资源分配R);P3的潜伏期必须依赖分类刺激所需的时间,但对分类后的过程却没有逻辑上的依赖关系。可以用P3的潜伏期来确定一个给定的实验操作是影响分类的过程,还是影响反应选择与执行的过程。
推荐阅读:
Neuropsychology of P300.
Two varieties of long-latency positive waves evoked by unpredictable auditory stimuli in man.
Stimulus novelty, task relevance and the visual evoked potential in man.
Recovery cycles of event-related potentials in multiple detection tasks.
Neural Origins of the P300.
Updating P300: an integrative theory of P3a and P3b.
Neuropsychology and neuropharmacology of P3a and P3b.
P3a from Visual Stimuli: Typicality, Task, and Topography.
P3a, perceptual distinctiveness, and stimulus modality.
P3a and P3b from typical auditory and visual stimuli.
Evidence for an Integrative Role of P3b in Linking Reaction to Perception.
The event-related brain potential as an index of display-monitoring workload.
The dynamics of P300 during dual-task performance.
语言相关的ERP成分
N400:负向波形,在中央和顶区电极位置的波幅最大,而且右半球的振幅比左半球稍大。但是N400主要产生于左侧颞叶,近期研究发现,左侧前额叶对N400也有贡献。典型的N400见于违反语义期待的反应。只有有意义的刺激可以诱发N400或者类似N400的活动。
P600:由违反语法规则诱发,约在300-500ms的左侧额叶负波。功能词会在左前部电极位置诱发一个叫N280的成分,而实义词没有这个成分;相反,实义词会诱发一个功能词没有的N400。
推荐阅读:
Language-Related ERP Components.
Reading senseless sentences: brain potentials reflect semantic incongruity.
Event-related potentials to grammatical errors and semantic anomalies.
Electrophysiological evidence for the effect of interactive imagery on episodic memory: Encouraging familiarity for non-unitized stimuli during associative recognition.
Event-related brain potentials reflect semantic priming in an object decision task.
Seeing and hearing meaning: ERP and fMRI evidence of word versus picture integration into a sentence context.
Rapid Distributed Fronto-parieto-occipital Processing Stages During Working Memory in Humans.
Event-related brain potentials elicited by syntactic anomaly.
错误相关的ERP成分
错误相关负波:(ERN,error-related negativty),由错误反应后的负反馈所诱发的。ERN通常被认为产生于前扣带回的背侧部分(dACC),但其实ERN的形成可能包含多个神经源的贡献,它反映了反应监测系统的活动,该系统对预期和实际反应之间的冲突比较敏感,或者会产生反应依赖的情绪反应。
推荐阅读:
A Neural System for Error Detection and Compensation.
Effects of errors in choice reaction tasks on the ERP under focused and devided attention.
ERP correlates of conscious error recognition: aware and unaware errors in an antisaccade task.
Is the ‘error negativity’ specific to errors?
Action-monitoring deficits in obsessive-compulsive disorder.
反应相关的ERP成分
准备电位(RP,readiness potential):或称BP(bereitschafts potential)。起始于一系列临时手动反应之前(约1秒左右)的一个负向慢波,位于额叶和中央区,准备电位的头部地形图分布依赖于使用哪一个效应器进行反应,因身体两侧的差别而异,以及因同侧身体使用不同的效应器而异。
偏侧化准备电位(LRP,lateralized readiness potential):一个在对侧半球比在同侧半球(相对于反应手而言)幅值更大的负波,与反应手之间具有侧向化关系,而其他成分没有,容易判断实验操作对LRP的时刻或振幅产生的影响。LRP至少有一部分产生于运动皮层。
推荐阅读:
Topography of the human motor potential.
Brain potential changes in voluntary and passive movements in humans: readiness potential and reafferent potentials.
The Lateralized Readiness Potential.
Effects of preliminary perceptual output on neuronal activity of the primary motor cortex.
Both primary motor cortex and supplementary motor area play an important role in complex finger movement.
参考资料:
Steven J. Luck. (2009). 事件相关电位基础 第二版[M]. 上海:华东师范大学出版社.
Key, A. P. F. , Dove, G. O. , & Maguire, M. J. . (2005). Linking Brainwaves to the Brain: An ERP Primer. Developmental Neuropsychology, 27(2), 183-215.
Helfrich, R. F. , & Knight, R. T. . (2019). Handbook of Clinical Neurology[M]. Vol.160 (Cognitive neurophysiology: Event-related potentials, Chapter 36).