ISSN 1671-3710
CN 11-4766/R
主办:中国科学院心理研究所
出版:科学出版社

心理科学进展, 2018, 26(10): 1787-1793 doi: 10.3724/SP.J.1042.2018.01787

研究前沿

主动控制感的测量及认知神经机制

张淼1,2, 吴迪1,2, 李明3, 凌懿白1,2, 张明4,5, 赵科,1,4

1 中国科学院心理研究所, 脑与认知科学国家重点实验室, 北京 100101

2 中国科学院大学心理学系, 北京 100049

3 吉首大学, 湖南 吉首 416000

4 中国科学院心理研究所心理健康重点实验室, 北京 100101

5 大连医科大学心理学系, 大连 116044

The measurement and neural mechanism of sense of agency

ZHANG Miao1,2, WU Di1,2, LI Ming3, LING Yibai1,2, ZHANG Ming4,5, ZHAO Ke,1,4

1 State Key Laboratory of Brain and Cognitive Science, Institute of Psychology, Chinese Academy of Sciences, Beijing 100101, China

2 Department of Psychology, University of Chinese Academy of Sciences, Beijing 100049, China

3 Jishou University, Jishou 416000, China

4 Key Laboratory of Mental Health, Institute of Psychology, Chinese Academy of Sciences, Beijing 100101, China

5 Department of Psychology, Dalian Medical University, Dalian, China, Beijing 116044, China

通讯作者: 赵科, E-mail:zhaok@psych.ac.cn

收稿日期: 2017-11-16   网络出版日期: 2018-10-15

基金资助: *国家自然科学基金(31400876)
国家自然科学基金(61632004)
国家社科基金青年项目(16CMZ035)
中国科学院心理研究所心理健康重点实验室课题(KLMH2014ZG11)
中国科学院心理研究所心理健康重点实验室课题(KLMH2015G07)

Received: 2017-11-16   Online: 2018-10-15

摘要

主动控制感指预计动作和实际感觉反馈匹配会产生一种控制自己动作、作用于环境的主观体验, 是人类心理活动的基本特征之一。本文系统介绍主动控制感的测量方式, 尤其是主动动作的时间压缩效应这一内隐手段; 并从主观意识、动作的发生方式和动作结果特征三个方面探讨主动控制感的影响因素。基于动作的比较器模型, 解释主动控制感的产生原因; 并提供了主动控制感的认知神经脑机制的证据, 强调额叶和顶叶在主动控制感中的作用。未来研究应该更加注重外显与内隐测量的一致性, 特殊群体以及主动控制感的脑网络研究。

关键词: 主动控制感; 时间压缩效应; 反馈; 意识

Abstract

“Sense of agency” refers to the sense of controlling one’s actions and the course of events in the outside world. It is based on the match between the expected action and the real sensory feedback. The first part of this article introduces a method for implicit measurement of sense of agency. The second part deals with the factors that influence the sense of agency from the aspects of intentionality and sensory feedback. Part three provides the conclusion. With an emphasis on the functions of the prefrontal and parietal lobes, this article provides evidence to explain the mechanism of sense of agency through the theoretical model and cognitive neuroscience.

Keywords: sense of agency; binding effect; feedback; intentionality

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本文引用格式

张淼, 吴迪, 李明, 凌懿白, 张明, 赵科. 主动控制感的测量及认知神经机制. 心理科学进展, 2018, 26(10): 1787-1793 doi:10.3724/SP.J.1042.2018.01787

ZHANG Miao, WU Di, LI Ming, LING Yibai, ZHANG Ming, ZHAO Ke. The measurement and neural mechanism of sense of agency. Advances in Psychological Science, 2018, 26(10): 1787-1793 doi:10.3724/SP.J.1042.2018.01787

1 引言

主动动作是可预计、并伴随一定结果的动作(James, 1890; Ziessler, Nattkemper, & Frensch, 2004)。在被动接受外界的信息时, 人类可通过主动的动作与外界的环境发生交互, 从而适应环境和改变环境(Elsner & Hommel, 2004)。主动控制感是在主动动作过程中产生的控制自身动作, 进而控制外界事物和环境的一种主观感觉。本文讨论的主动控制感是基于动作的准备, 动作的发生和动作后效而产生的。根据动作的比较理论, 在动作前有一个副本预测动作的结果, 当预测结果和实际动作结果匹配时, 就会产生一种主动控制感, 反之则削弱这种主动控制感(Haggard, 2017)。主动控制感是人类心理活动的一个基本特征(Frith, 2014), 因其在生活中的普遍性, 主动控制感得到越来越多研究者的青睐。本文将从测量方式、影响因素和脑机制角度对主动控制感相关研究进行系统归纳和总结。

2 主动控制感的测量方式

基于动作的主动控制感的度量有两种方式:一种是外显测量, 通过询问被试“这个动作是否就是你做的?” “你对你的动作控制的程度如何?”等问题来实现, 被试做这些判断需要把外在的事件归结于自己的意识动作而非其他原因; 另外一种外显测量的做法则是采用问卷调查, 最近有研究者编制了主动控制感的量表, 从正负性两个维度进行主动控制感的测量(Tapal, Oren, Dar, & Eitam, 2017)。这种外显测量可能有一个主观的认知偏差, 被试会片面夸大自己的主动控制感, 甚至把无关的事件也归结为自己的行动或动作结果, 尤其是当结果是正面或正性时。例如, 领导人总是倾向于把经济的增长归结为自己的政治行动(Haggard, 2017)。

因此, 一种更加简单、隐蔽且有效的方式-内隐测量备受研究者推崇。主动动作的时间压缩效应就是最有代表性的内隐测量方式。Haggard等人最早采用Libet时钟研究了该效应(图1a, Haggard, Clark, & Kalogeras, 2002; Libet, Gleason, Wright, & Pearl, 1983)。在他们的研究中, 实验分为三种条件:第一种为主动按键条件, 主动按键后间隔250 ms后有一听觉刺激, 要求被试报告按键时刻时钟指针的位置或声音刺激出现时的指针位置; 第二种是被动按键条件, 经颅性刺激(TMS)作用于被试大脑皮层的运动区造成被动按键, 间隔250 ms后同样有一个听觉刺激, 要求被试报告按键时的指针位置或声音刺激出现时的指针位置; 第三种是控制条件, 指针转动和声音刺激会在不同的区组随机呈现, 被试任务是报告动作发生时指针位置或者声音刺激出现时指针位置。结果表明, 在主动按键条件下, 被试报告自己动作的时间指针点相比控制条件靠后, 报告听觉刺激的呈现时间指针点比控制条件下提前。简言之, 在主动按键条件下, 动作发生到延迟声音刺激出现之间的间隔时间产生了时间维度上的主观压缩; 而被动按键条件没有发现该效应。主动动作的时间压缩效应测量主动控制感得到大量研究的支持(Buehner, 2012; Ebert & Wegner, 2010; Engbert & Wohlschläger, 2007; Hascalovitz & Obhi, 2015; Zhao, Chen, Yan, & Fu, 2013)。该范式能准确获得指针时间点信息, 并且能够分离动作压缩和动作结果两个方向的压缩。但也受到一些质疑(Gomes, 2002):时钟位置需要视觉加工而主动动作需体感运动信息加工, 存在跨通道的同步性问题; 另外, 动作发生时间点没有严格控制, 个体间在完成主动动作时存在差异。

图1

图1   主动控制感的内隐测量方式示意图


间隔估计范式 在这种范式中, 被试直接报告动作发生点与动作结果(声音刺激)之间的间隔时间(如Buehner & Humphreys, 2010; Engbert & Wohlschläger, 2007, 2008; Humphreys & Buehner, 2009)。其一般过程为:被试有意识的按键, 主动按键后间隔一段空白时间间隔后出现声音或视觉刺激。被试的任务是需要报告按键动作到动作结果(声音或视觉刺激)之间的间隔时间(图1b)。该方法简明易懂、操作性强, 但对于动作发生的起始计时点依然没有严格控制。在此基础上, 赵科等人(2013)改进了间隔估计方法, 并消除了估计时间间隔起始点不一致的问题(Zhao et al., 2013; 图1c)。其方法简述如下, 在屏幕上呈现刺激1(如方块), 被试要求主动按键的同时方块消失, 间隔一定的时间间隔出现刺激2(如声音), 被试要求估计刺激1消失到刺激2出现之间的间隔时间。

3 主动控制感的影响因素

影响动作的主动控制感的因素很多, 本文从主动动作的发生前的主观意识、动作的发生方式及动作的结果特征三方面进行归纳。首先是主观意识, 主观意识在研究领域内称“free will”。主观意识的重要性在最早的时间压缩研究中就得以体现, 被试有意识按键会产生主动动作的时间压缩效应, 经颅性刺激运动皮层产生的被动动作却产生相反的时间效应, 被试报告被动动作发生的时间估计点提前, 报告声音刺激出现的时间估计点靠后, 即被动动作到声音刺激的间隔估计时间反而延长了(Haggard, Clark, & Kalogeras, 2002)。对于被动的按键动作, 尽管也属于自己, 存在向我属性(feeling of “mineness”), 但都基于外周皮层而不是大脑的主观意识。赵科等人(2016)的研究也发现, 有意识的主动按键动作产生了时间压缩效应, 而通过自主/非自主动作装置实现的被动按键动作却没有产生时间压缩效应(Zhao et al., 2016)。其次, 如果主观意愿降低, 时间压缩效应也会被削弱。最近有研究要求被试在实验助手的胁迫下或者主动意识条件下给别人施加疼痛刺激。结果表明被试在胁迫而不是主动条件下, 产生更小的时间压缩效应, 并且这种效应在疼痛刺激的ERPs成分上表现为N1脑电成分的下降。研究者认为, 被试在胁迫条件下施加疼痛刺激比主动施加其主观意识水平较低(Caspar, Christensen, Cleeremans, & Haggard, 2016)。准备电位(readiness potential)是发生在动作前负性的脑电波成分, 也被认为主观意识的指标(Shibasaki & Hallett, 2006)。最近有研究发现该电位幅度越大则动作-结果之间的时间压缩效应更强, 主动控制感也增强(Jo, Wittmann, Hinterberger, & Schmidt, 2014)。

动作的发生方式也是影响主动控制感的重要因素。主观意识在主动控制感中作用很大, 相比被动动作, 主动动作产生了更大的时间压缩效应。但该类研究大都采用主动按键动作, 忽略了动作方式的多样性。赵科等人(Zhao et al., 2013)采用间隔估计范式, 被试要求估计两个相继呈现的视觉刺激(S1和S2)之间的间隔时间。实验设置了三种条件, 分别为主动按键, 主动抬键和控制条件。结果表明主动按键条件下的时间压缩效应在估计时间间隔为150~1050 ms时都存在, 而主动抬键的时间压缩效应只有在估计时间间隔为150 ms和250 ms时存在。当估计的间隔时间很短时, 主动按键和抬键都产生了时间压缩效应; 当估计时间间隔变长时, 主动按键的时间压缩依然存在甚至可以达到数秒, 而主动抬键的时间压缩效应则消失了。主动按键和抬键动作时间压缩效应的异同可能原因是按键动作后有一个及时的触觉反馈。在赵科等人的另外一个研究里, 为排除按键动作的熟悉度影响, 研究者改变按键和抬键的方向, 分为向上按键、向下按键、向上抬键和向下抬键。结果表明不论动作发生方向如何, 自主按键都能产生更大的时间压缩效应。该研究的实验3则通过自主研发装置实现无触觉反馈的按键和抬键动作, 结果表明两种动作产生了相同的时间压缩效应(Zhao et al., 2016)。由此可见动作的发生方式和主动意识一样对于时间压缩效应非常重要。

第三个与动作有关的因素是动作结果的特征, 动作结果的特性(如效价, 结果发生的可能性与接近性)也会影响主动控制感。首先是动作结果的效价, 动作的结果可能是正性或负性的, 这些都能直接作用于主动控制感而造成时间的压缩或延长。在Haggard等人最近研究中采用不同效价的声音刺激作为动作结果, 当声音刺激为负性时相比中性产生了更小的时间压缩效应, 而正性和中性结果却无差异。研究者认为负性刺激降低了主动控制感从而削弱了时间压缩效应(Yoshie & Haggard, 2013; Yoshie & Haggard, 2017)。除了结果的效价, 结果发生的可能性与时间上的接近性也是影响主动控制感的因素。Buehner和Humphreys采用了一种新的范式探讨了结果发生的可能性的影响。在控制条件下, 被试面对连续呈现的两个声音(T1和T2), 反复呈现15次, T2发生的可能性与动作无关; 在因果关系条件下, 被试需要通过按键动作, 使得T2发生, 此条件下T2发生的可能性与动作建立了关联。在正式实验中, 被试被要求把自己的按键和T1,T2同步, 结果发现在因果关系的条件下产生了更大的时间压缩效应(Buehner & Humphreys, 2009)。Engbert等采用Libet时钟, 设置了两种动作结果(声音刺激)的概率条件, 一种条件下结果发生的概率是另外一种条件的两倍。结果发现被试报告的按键延迟的时间在高概率条件下比低概率条件往后延迟了27 ms, 产生了更大的时间压缩效应(Engbert & Wohlschlager, 2007)。动作-结果间的时间接近性也影响主动控制感, Haggard等人比较了不同的动作与听觉刺激间隔(250, 450, 或650 ms)情况下的时间压缩效应, 结果表明该效应在长时间间隔时会削弱(Haggard, Clark, & Kalogeras, 2002)。此外, 赵科等人(2013)在研究中也设置了不同的动作-结果时间间隔(150, 250, 350, 450, 550, 650, 750, 850, 950, 1050 ms), 同样发现随着时间间隔的不断增长, 时间压缩效应会逐渐减弱(Zhao et al., 2013)。Stetson等人(2006)发现当被试适应了按键动作到动作结果(彩色闪光)的固定间隔(135 ms)后,比之前距离按键动作时间间隔(44 ms)短的彩色闪光将会让被试产生一种动作和结果呈现顺序反转的错觉, 这也从另外一个角度证实了时间间隔的长短即时间上的接近性对主动控制感的影响。

4 主动控制感的理论模型与认知神经机制

主动控制感的产生机制被认为是基于动作控制系统的内在比较器模型(Blakemore, Wolpert, & Frith, 2000; Kawato, 1999; Lindner, Haarmeier, & Erb, Grodd, & Their, 2006; Wolpert, Ghahramani, & Jordan, 1995)。把动作系统看作一个内在比较过程这一思想可以追溯到赫尔姆霍兹(Helmholtz, 1866)。内在比较器模型把动作比作一个过程, 过程的两种状态分别是输入的“期望状态” (desired state)和输出的“估计的实际状态” (estimated actual state)。在这两种状态表征的基础上, 动作系统可以细分为3种内在比较过程(图2)。在反馈控制回路中, 比较器1对比“期望状态”和“估计的实际状态”的差异, 计算运动误差并将错误反馈给动作控制系统; 在动作发生前, 预测器接收动作指令传输的动作副本并产生一个预计的状态。比较器2则是对比预计的状态与期望的状态并将错误反馈给动作控制系统; 同时, 动作副本中包含一个预测的感知反馈信息, 这种预测是先于实际反馈的。比较器3则对预测的感知反馈信息与感知运动系统获取的实际感知进行对比, 将结果反馈给预测器并对其以后预测提供优化。一般认为, 预测的感知反馈信息和实际的感知反馈信息在匹配时产生主动控制感(Sense of Agency)并造成时间压缩效应(Moore & Obhi, 2012)。

图2

图2   动作的内在比较器模型(Synofzik, Vosgerau, & Newen, 2008)


目前关于主动控制感脑机制的研究还比较少, 但是这些研究都表明背外侧前额叶(dorsolateral prefrontal cortex)、辅助运动皮层和前辅助运动皮层(SMA and pre-SMA), 顶叶皮层(parietal cortex)等脑区在主动控制感产生中起了很重要的作用(Crivelli & Balconi, 2017)。其中额-顶网络的作用尤为突出。

早期的研究采用失用症患者(Apraxic patients), 他们的顶叶皮层存在损伤。在实验中, 他们被要求做按键动作并且可以通过屏幕看到自己手的动作或者实验者的动作。在每个试次中, 他们都需要报告屏幕上的手是自己还是别人的。病人和正常人都能正确识别自己的手势, 当实验者手势和自己的手势不一样时也能准确识别。但是当实验者的手势和自己的手势一样时, 病人的识别正确率明显下降, 即使病人自己的手势错误, 他们依然不能认为实验者的手势是别人的手势。这些结果可能表明, 如果顶叶受损, 被试不能比较和评估内在的和外在的动作反馈, 说明顶叶皮层在主动控制感中发挥着重要作用(Sirigu, Daprati, Pradat- Diehl, Franck, & Jeannerod, 1999)。Fink等人(1999)在研究顶叶控制主动动作中的作用时, 考察了正确的视觉反馈和错误的视觉反馈(镜子的反向成像)条件下手做动作时后侧顶叶的情况, 结果表明后侧顶叶在手的同向和反向动作中都有激活, 尤其是错误视觉反馈的条件下(Fink et al., 1999)。这些结果都显示了后侧顶叶在比较动作和动作结果中的作用。MacDonald和Paus (2003)在此基础上采用TMS分别刺激被试的后侧顶叶皮层和颞叶皮层, 在主动动作条件下, 被刺激后侧顶叶皮层的被试判断动作和视觉反馈之间的延迟成绩有明显降低。研究者认为后顶叶皮层在动作副本和视觉结果反馈的匹配中即主动控制感上发挥了重要作用。

另外一些研究则表明位于顶下小叶的角回是主动控制感的负责脑区。Blanke, Ortigue, Landis和Seeck (2002)的研究发现, 如果给开颅后的癫痫病人的右侧角回区域施加电刺激, 病人会产生一种动作已经不属于自己的体验。另外有研究发现外显的控制感判断任务中, 如果被试把视觉的反馈归结为非自己动作造成时角回会显著激活(Farrer & Frith, 2002)。元分析的结果也发现颞顶交接处含角回是无主动控制感的基础(Sperduti, Delaveau, Fossati, & Nadel, 2011)。在另外一项fMRI研究中, 被试被要求根据呈现的线索提示按左右键, 结果显示线索提示与左右按键一致时比不一致产生了更快的按键反应。相应的, 被试在一致的情况下产生了更大的主动控制感。磁共振结果表明角回在主动控制感评分低的情况下会产生更强激活(Chambon, Wenke, Fleming, Prinz, & Haggard, 2013)。采用TMS刺激角回, 会让被试产生更低的主动控制感评分(Chambon, Moore, & Haggard, 2015)。同时, 精神分裂症患者的主动控制感比正常人强, 会产生更大的主动动作的时间压缩效应, 功能磁共振结果也表明精神分裂症患者在线索和动作不匹配的情况下, 前额叶和角回的连接减弱(Voss et al., 2017)。

额叶中的辅助运动区或前辅助运动区也被认为是与主动控制感密切相连的一个脑区。如前所述, 有研究发现前辅助运动区产生脑电的准备电位, 准备电位越大产生的时间压缩效应就越大。Moore, Ruge, Wenke, Rothwell和Haggard (2010)发现TMS抑制pre-SMA也造成了主动动作时间压缩效应的消失和主动控制感降低。背外侧前额叶是与动作选择有关的一个脑区。施加微弱电流刺激的经颅直流电刺激(tDCS)技术作用于大脑该区域, 发现时间压缩效应降低(Khalighinejad & Haggard, 2015)。Khalighinejad等人(2015)综述并做了几个实验, 在实验中要求被试主动选择一种动作, 动作后有一个结果, 通过荟萃分析, 研究者发现阳性的刺激会促进主动动作的时间压缩效应产生(Khalighinejad, Di Costa, & Haggard, 2016)。

额叶在主动控制感中的作用也体现在其它手段的研究中, 如赵科等人最近的ERPs研究, 通过比较时间感知阶段主动按键和主动抬键条件下的差异, 结果显示前额叶在主动控制感中起了很大作用。在时间感知阶段的空白时间间隔中, 主动按键比主动抬键产生了更大的脑电P1成分, 两种条件下都产生了相同的P2脑电成分。根据脑地形图, 这两成分都主要头皮分布为前额叶。根据动作比较器模型, 作者认为第一个脑电成分P1可能反映了预测结果与触觉反馈的比较。第二个成分P2则反映了动作与视觉反馈结果的比较(Zhao et al., 2014)。最近有研究发现青少年和老年人的主动控制感明显削弱, 可能也与其额叶功能降低有关(Cavazzana, Begliomini, & Bisiacchi, 2017)。

5 问题与展望

总之, 主动控制感是人类心理活动的一个基本特征, 主动控制感的相关研究也越来越引起广大研究者的兴趣, 但是揭示主动控制感的认知机制及其脑机制仍然是研究者面临的巨大挑战, 存在诸多问题和难点。

第一, 主动控制感的外显和内隐测量之间的对应关系亟待研究。现有的研究分别从两个独立的维度去测量主动控制感, 外显测量和内隐测量的一致性需要大量的实证研究来验证。

第二, 不同人群的主动控制感的测量需要进一步研究。尽管目前研究发现精神分裂症患者的主动控制感高于常人, 但是对于其他特殊人群例如帕金森患者、成瘾患者、抑郁症患者等的主动控制感缺乏相关研究。研究这些特定群体的主动控制感, 有助于揭示主动控制感的特征, 并有助于提高其心理健康的水平, 提升其生活的质量。

第三, 主动控制感的脑机制需要更深入的研究。现在对主动控制感脑机制方面的研究较少, 对于主动控制感产生的脑区存在很大的争议。额叶和顶叶在主动控制感中起到何种作用, 额叶和顶叶在主动控制感产生的过程中如何进行信息交流等问题还需要在大量的实证研究中做进一步的探讨。未来可以采用颅内记录、脑磁成像等方法进一步构建主动控制感的脑网络图谱。

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When a voluntary action is causally linked with a sensory outcome, the action and its consequent effect are perceived as being closer together in time. This effect is called intentional binding. Although many experiments were conducted on this phenomenon, the underlying neural mechanisms are not well understood. While intentional binding is specific to voluntary action, we presumed that preconscious brain activity (the readiness potential, RP), which occurs before an action is made, might play an important role in this binding effect. In this study, the brain dynamics were recorded with electroencephalography (EEG) and analyzed in single-trials in order to estimate whether intentional binding is correlated with the early neural processes. Moreover, we were interested in different behavioral performance between meditators and non-meditators since meditators are expected to be able to keep attention more consistently on a task. Thus, we performed the intentional binding paradigm with twenty mindfulness meditators and compared them to matched controls. Although, we did not observe a group effect on either behavioral data or EEG recordings, we found that self-initiated movements following ongoing negative deflections of slow cortical potentials (SCPs) result in a stronger binding effect compared to positive potentials, especially regarding the perceived time of the consequent effect. Our results provide the first direct evidence that the early neural activity within the range of SCPs affects perceived time of a sensory outcome that is caused by intentional action.

Kawato M., ( 1999).

Internal models for motor control and trajectory planning

Current Opinion in Neurobiology, 9( 6), 718-727.

DOI:10.1016/S0959-4388(99)00028-8      URL     PMID:10607637      [本文引用: 1]

Abstract A number of internal model concepts are now widespread in neuroscience and cognitive science. These concepts are supported by behavioral, neurophysiological, and imaging data; furthermore, these models have had their structures and functions revealed by such data. In particular, a specific theory on inverse dynamics model learning is directly supported by unit recordings from cerebellar Purkinje cells. Multiple paired forward inverse models describing how diverse objects and environments can be controlled and learned separately have recently been proposed. The 'minimum variance model' is another major recent advance in the computational theory of motor control. This model integrates two furiously disputed approaches on trajectory planning, strongly suggesting that both kinematic and dynamic internal models are utilized in movement planning and control.

Khalighinejad N., & Haggard, P. ( 2015).

Modulating human sense of agency with non-invasive brain stimulation

Cortex, 69, 93-103.

DOI:10.1016/j.cortex.2015.04.015      URL     PMID:26004997      [本文引用: 2]

Human voluntary actions are accompanied by a distinctive subjective experience termed “sense of agency”. We performed three experiments using transcranial direct current stimulation (tDCS) to modulate brain circuits involved in control of action, while measuring stimulation-induced changes in one implicit measure of sense of agency, namely the perceived temporal relationship between a voluntary action and tone triggered by the action. Participants perceived such tones as shifted towards the action that caused them, relative to baseline conditions with tones but no actions. Actions that caused tones were perceived as shifted towards the tone, relative to baseline actions without tones. This ‘intentional binding’ was diminished by anodal stimulation of the left parietal cortex [targeting the angular gyrus (AG)], and, to a lesser extent, by stimulation targeting the left dorsolateral prefrontal cortex (DLPFC), (Experiment 1). Cathodal AG stimulation had no effect (Experiment 2). Experiment 3 replicated the effect of left anodal AG stimulation for actions made with either the left or the right hand, and showed no effect of right anodal AG stimulation. The angular gyrus has been identified as a key area for explicit agency judgements in previous neuroimaging and lesion studies. Our study provides new causal evidence that the left angular gyrus plays a key role in the perceptual experience of agency.

Khalighinejad N., Di C S., & Haggard P . ( 2016).

Endogenous action selection processes in dorsolateral prefrontal cortex contribute to sense of agency: A meta- analysis of tDCS studies of ‘intentional binding’

Brain Stimulation, 9( 3), 372-379.

[本文引用: 1]

Libet B., Gleason C. A., Wright E. W., & Pearl D. K . ( 1983).

Time of conscious intention to act in relation to onset of cerebral activity (readiness-potential) the unconscious initiation of a freely voluntary act

Brain, 106( 3), 623-642.

[本文引用: 1]

Lindner A., Haarmeier T., Erb M., Grodd W., & Thier P . ( 2006).

Cerebrocerebellar circuits for the perceptual cancellation of eye-movement-induced retinal image motion

Journal of Cognitive Neuroscience, 18( 11), 1899-1912.

[本文引用: 1]

MacDonald P. A., & Paus, T. ( 2003).

The role of parietal cortex in awareness of self-generated movements: A transcranial magnetic stimulation study

Cerebral Cortex, 13( 9), 962-967.

[本文引用: 1]

Moore J. W., & Obhi, S. S . ( 2012).

Intentional binding and the sense of agency: A review

Consciousness and Cognition, 21( 1), 546-561.

DOI:10.1016/j.concog.2011.12.002      URL     PMID:22240158      [本文引用: 1]

It is nearly 1002years since Patrick Haggard and colleagues first reported the ‘intentional binding’ effect ( Haggard, Clark, & Kalogeras, 2002 ). The intentional binding effect refers to the subjective compression of the temporal interval between a voluntary action and its external sensory consequence. Since the first report, considerable interest has been generated and a fascinating array of studies has accumulated. Much of the interest in intentional binding comes from the promise to shed light on human agency. In this review we survey studies on intentional binding, focusing, in particular, on the link between intentional binding and the sense of agency (the experience of controlling action to influence events in the environment). We suggest that, whilst it is yet to be fully explicated, the link between intentional binding and the sense of agency is compelling. We conclude by considering outstanding questions and future directions for research on intentional binding.

Moore J. W., Ruge D., Wenke D., Rothwell J., & Haggard P . ( 2010).

Disrupting the experience of control in the human brain: Pre-supplementary motor area contributes to the sense of agency

Proceedings of the Royal Society of London B: Biological Sciences, 277, 2503-2509.

[本文引用: 1]

Shibasaki H., & Hallett, M. ( 2006).

What is the Bereitschaftspotential?

Clinical Neurophysiology, 117( 11), 2341-2356.

DOI:10.1016/j.clinph.2006.04.025      URL     PMID:16876476      [本文引用: 1]

Abstract Since discovery of the slow negative electroencephalographic (EEG) activity preceding self-initiated movement by Kornhuber and Deecke [Kornhuber HH, Deecke L. Hirnpotential nderungen bei Willkurbewegungen und passiven Bewegungen des Menschen: Bereitschaftspotential und reafferente Potentiale. Pflugers Archiv 1965;284:1-17], various source localization techniques in normal subjects and epicortical recording in epilepsy patients have disclosed the generator mechanisms of each identifiable component of movement-related cortical potentials (MRCPs) to some extent. The initial slow segment of BP, called 'early BP' in this article, begins about 2 s before the movement onset in the pre-supplementary motor area (pre-SMA) with no site-specificity and in the SMA proper according to the somatotopic organization, and shortly thereafter in the lateral premotor cortex bilaterally with relatively clear somatotopy. About 400 ms before the movement onset, the steeper negative slope, called 'late BP' in this article (also referred to as NS'), occurs in the contralateral primary motor cortex (M1) and lateral premotor cortex with precise somatotopy. These two phases of BP are differentially influenced by various factors, especially by complexity of the movement which enhances only the late BP. Event-related desynchronization (ERD) of beta frequency EEG band before self-initiated movements shows a different temporospatial pattern from that of the BP, suggesting different neuronal mechanisms for the two. BP has been applied for investigating pathophysiology of various movement disorders. Volitional motor inhibition or muscle relaxation is preceded by BP quite similar to that preceding voluntary muscle contraction. Since BP of typical waveforms and temporospatial pattern does not occur before organic involuntary movements, BP is used for detecting the participation of the 'voluntary motor system' in the generation of apparently involuntary movements in patients with psychogenic movement disorders. In view of Libet et al.'s report [Libet B, Gleason CA, Wright EW, Pearl DK. Time of conscious intention to act in relation to onset of cerebral activity (readiness-potential). The unconscious initiation of a freely voluntary act. Brain 1983;106:623-642] that the awareness of intention to move occurred much later than the onset of BP, the early BP might reflect, physiologically, slowly increasing cortical excitability and, behaviorally, subconscious readiness for the forthcoming movement. Whether the late BP reflects conscious preparation for intended movement or not remains to be clarified.

Sirigu A., Daprati E., Pradat-Diehl P., Franck N., & Jeannerod M . ( 1999).

Perception of self-generated movement following left parietal lesion

Brain, 122( 10), 1867-1874.

DOI:10.1093/brain/122.10.1867      URL     PMID:10506089      [本文引用: 1]

Abstract Three apraxic patients with lesions in the left parietal cortex were required to execute finger movements with either hand, while the visual feedback they received about the movement was manipulated systematically. We used a device which allowed us to present on a video monitor either the patient's hand or the examiner's hand simultaneously performing an identical or a different movement. In each trial, patients were required to decide whether the hand shown on the screen was their own or not. Hand movements produced in response to verbal command included simple (single-finger extension) and complex gestures (multi-finger extension). Ownership judgements were analysed and compared with those produced by six normal controls and two non-apraxic neurological patients. Apraxic patients and controls accurately recognized their own hand on the screen (own movement condition) and correctly identified the viewed hand as the examiner's when it performed a movement different from their own movement (incongruent movement condition). However, when the viewed hand was the examiner's hand executing their own movement (congruent movement condition), apraxic patients were significantly more impaired than controls. When the results were analysed as a function of gesture type, the number of correct responses was significantly lower for apraxic patients with respect to controls only for complex gestures. Interestingly, when patients executed the finger gestures inaccurately, they still failed to recognize the examiner's hand as alien, and claimed that the correct movement presented on the screen was their own. These results confirm that parietal lesions alter the representational aspects of gestures, and suggest a failure in evaluating and comparing internal and external feedback about movement. We conclude that the parietal cortex plays an important role in generating and maintaining a kinaesthetic model of ongoing movements.

Sperduti M., Delaveau P., Fossati P., & Nadel J . ( 2011).

Different brain structures related to self-and external- agency attribution: A brief review and meta-analysis

Brain Structure and Function, 216( 2), 151-157.

[本文引用: 1]

Stetson C., Cui X., Montague P. R., & Eagleman D. M . ( 2006).

Motor-sensory recalibration leads to an illusory reversal of action and sensation

Neuron, 51( 5), 651-659.

[本文引用: 1]

Synofzik M., Vosgerau G., & Newen A . ( 2008).

Beyond the comparator model: a multifactorial two-step account of agency

Consciousness and Cognition, 17( 1), 219-239.

DOI:10.1016/j.concog.2007.03.010      URL     PMID:17482480      [本文引用: 1]

There is an increasing amount of empirical work investigating the sense of agency, i.e. the registration that we are the initiators of our own actions. Many studies try to relate the sense of agency to an internal feed-forward mechanism, called the omparator model . In this paper, we draw a sharp distinction between a non-conceptual level of feeling of agency and a conceptual level of judgement of agency. By analyzing recent empirical studies, we show that the comparator model is not able to explain either. Rather, we argue for a two-step account: a multifactorial weighting process of different agency indicators accounts for the feeling of agency, which is, in a second step, further processed by conceptual modules to form an attribution judgement. This new framework is then applied to disruptions of agency in schizophrenia, for which the comparator model also fails. Two further extensions are discussed: We show that the comparator model can neither be extended to account for the sense of ownership (which also has to be differentiated into a feeling and a judgement of ownership) nor for the sense of agency for thoughts. Our framework, however, is able to provide a unified account for the sense of agency for both actions and thoughts.

Tapal A., Oren E., Dar R., & Eitam B . ( 2017).

The Sense of Agency Scale: A Measure of Consciously Perceived Control over One's Mind, Body, and the Immediate Environment

Frontiers in Psychology, 8, 1552.

[本文引用: 1]

Voss M., Chambon V., Wenke D., Kühn S., & Haggard P . ( 2017).

In and out of control: brain mechanisms linking fluency of action selection to self-agency in patients with schizophrenia

Brain, 140( 8), 2226-2239.

[本文引用: 1]

Wolpert D. M., Ghahramani Z., & Jordan M. I . ( 1995).

An internal model for sensorimotor integration

Science, 269( 5232), 1880-1882.

[本文引用: 1]

Yoshie M., & Haggard, P. ( 2013).

Negative emotional outcomes attenuate sense of agency over voluntary actions

Current Biology, 23( 20), 2028-2032.

DOI:10.1016/j.cub.2013.08.034      URL     PMID:24094850      [本文引用: 1]

Abstract Sense of agency (SoA) refers to the feeling that one's voluntary actions produce external sensory events [1, 2]. Several psychological theories hypothesized links between SoA and affective evaluation [3-6]. For example, people tend to attribute positive outcomes to their own actions, perhaps reflecting high-level narrative processes that enhance self-esteem [3]. Here we provide the first evidence that such emotional modulations also involve changes in the low-level sensorimotor basis of agency. The intentional binding paradigm [1] was used to quantify the subjective temporal compression between a voluntary action and its sensory consequences, providing an implicit measure of SoA. Emotional valence of action outcomes was manipulated by following participants' key-press actions with negative or positive emotional vocalizations [7], or neutral sounds. We found that intentional binding was reduced for negative compared to positive or neutral outcomes. Discriminant analyses identified a change in time perception of both actions and their negative outcomes, demonstrating that the experience of action itself is subject to affective modulation. A small binding benefit was also found for positive action outcomes. Emotional modulation of SoA may contribute to regulating social behavior. Correctly tracking the valenced effects of one's voluntary actions on other people could underlie successful social interactions. Copyright 2013 The Authors. Published by Elsevier Inc. All rights reserved.

Yoshie M., & Haggard, P. ( 2017).

Effects of emotional valence on sense of agency require a predictive model

Scientific Reports, 7( 1), 8733.

DOI:10.1038/s41598-017-08803-3      URL     PMID:5562802      [本文引用: 1]

Sense of agency (SoA), a feeling that one’s voluntary actions produce events in the external world, is a key factor behind every goal-directed human behaviour. Recent studies have demonstrated that SoA is reduced when one’s voluntary action causes negative outcomes, compared to when it causes positive outcomes. It is yet unclear whether this emotional modulation of SoA is caused by predicting the outcome valence (predictionhypothesis) or by retrospectively interpreting the outcome (postdictionhypothesis). To address this, we emulated a social situation where one’s voluntary action was followed by either another’s negative emotional vocalisation or positive emotional vocalisation. Crucially, the relation between an action and the emotional valence of its outcome was predictable in some blocks of trials, but unpredictable in other blocks. Quantitative, implicit measures of SoA based on the intentional binding effect supported the prediction hypothesis. Our findings imply that the social-emotional modulation of SoA is based on predicting the emotional valence of action outcomes.

Zhao K., Chen Y. H., Yan W. J., & Fu X. L . ( 2013).

To bind or not to bind? Different temporal binding effects from voluntary pressing and releasing actions

PloS One, 8( 5), e64819.

[本文引用: 4]

Zhao K., Gu R. L., Wang L., Xiao P., Chen Y. H., Liang J ., .. & Fu, X. L. ( 2014).

Voluntary pressing and releasing actions induce different senses of time: Evidence from event-related brain responses

Scientific Reports, 4, 6047.

[本文引用: 1]

Zhao K., Hu L., Qu F. B., Cui Q., Piao Q. H., Xu H ., .. & Fu, X. L. ( 2016).

Voluntary action and tactile sensory feedback in the intentional binding effect

Experimental Brain Research, 234( 8), 2283-2292.

DOI:10.1007/s00221-016-4633-5      URL     PMID:27038203      [本文引用: 2]

The intentional binding effect refers to a subjective compression over a temporal interval between the start point initialized by a voluntary action and the endpoint signaled by an external sensory...

Ziessler M., Nattkemper D., & Frensch P. A . ( 2004).

The role of anticipation and intention in the learning of effects of self-performed actions

Psychological Research, 68( 2-3), 163-175.

DOI:10.1007/s00426-003-0153-6      URL     PMID:14634810      [本文引用: 1]

Abstract The anticipative learning model for acquiring action-effect relations states that the acquisition of action-effect relations depends on processes that are part of action planning, in particular the anticipation of possible effects. Experiment 1 shows that response planning is indeed crucial for the learning of response effects. In this experiment distractors (tones) were presented either during response preparation or in the time interval between response execution and the presentation of a response effect. Response-effect learning was impaired when the distractors were presented during response preparation. The finding is consistent with the assumption that the distractors impaired the anticipation of potential effects and therefore reduced effect learning. In Experiment 2 all responses had two effects. Participants were instructed to produce one of the effects. Under this condition, response-effect learning was only found for the instructed effect, not for the non-instructed effect. The two experiments thus support the view that response-effect learning is selective and depends on the anticipation of potential effects during response planning. The results are discussed in terms of a model that explains both the learning of response-effect relations and the use of these effects for action control within the same theoretical framework. Copyright 2003 Springer-Verlag

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