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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。

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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。

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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。

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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(RPreadiness 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(LRPlateralized 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 Neuropsychology27(2), 183-215.

Helfrich, R. F. , &  Knight, R. T. . (2019). Handbook of Clinical Neurology[M]. Vol.160 (Cognitive neurophysiology: Event-related potentials, Chapter 36).

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