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, 2019, 27(3): 508-521 doi: 10.3724/SP.J.1042.2019.00508

研究前沿

双通路理论视角下孤独症谱系障碍者的视线加工障碍

荆伟, 王庭照,

陕西师范大学教育学院, 西安 710062

The gaze processing impairment in individuals with autism spectrum disorder: A perspective from the two-process theory

JING Wei, WANG Tingzhao,

College of Education, Shaanxi Normal University, Xi’an 710062, China

通讯作者: 王庭照, E-mail:wangtingzhao@snnu.edu.cn

收稿日期: 2018-03-5   网络出版日期: 2019-03-15

基金资助: 2014年度教育部人文社会科学研究青年项目:“情境影响自闭症儿童运用眼睛注视线索习得词语的研究” 资助(14YJC880024)

Received: 2018-03-5   Online: 2019-03-15

摘要

临床行为观察发现, 孤独症谱系障碍个体普遍存在不能与他人建立视线接触, 不能追随他人视线看向目标物体等视线加工(gaze processing)障碍。然而已有实验研究发现该群体在实验情境中普遍存在视线接触(eye contact)异常, 但其视线追随行为(gaze following)则存在正常与异常并存的现象。基于视线加工双通路理论的启示, 该障碍可能是由于视线加工皮下通路先天功能异常而皮层通路后天发展异常所致。然而, 该理论尚缺乏皮下通路先天功能异常是视线接触障碍潜在神经机制的直接证据, 还需进一步考察皮下通路先天功能异常对视线追随障碍的影响作用, 以及皮层通路后天发展异常即其补偿机制的神经回路及早期形成过程。

关键词: 孤独症谱系障碍; 视线接触障碍; 视线追随障碍; 双通路理论

Abstract

Eye gaze is important social information in social interaction. However, clinical naturalistic observation found that autism spectrum disorder (ASD) individuals are impaired in gaze processing in the first year of life. In particular, individuals with ASD are unable to make eye contact with others and follow others' gaze to look at the target object. Furthermore, experimental studies found that while individuals with ASD presented impaired eye contact, there was contradictory phenomenon of a normal and abnormal coexistence in experimental contexts in gaze following of individuals with ASD. Based on the two- process theory of gaze processing, the group's subcortical pathway is born with functional abnormality, while the cortical pathway develops atypically, which is assumed as the neural mechanism of gaze processing impairment in individuals with ASD. However, the theoretical hypothesis still lacks the direct evidence supporting that the congenital functional abnormality of the subcortical pathway is the neural mechanism of eye contact impairment. Further studies are needed to investigate the effect of the congenital functional abnormality of the subcortical pathway on gaze following impairment and the neural circuit and early formation process of the compensation mechanism of the cortical pathway.

Keywords: autism spectrum disorder; eye contact impairment; gaze following impairment; two-process theory

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本文引用格式

荆伟, 王庭照. 双通路理论视角下孤独症谱系障碍者的视线加工障碍. 心理科学进展, 2019, 27(3): 508-521 doi:10.3724/SP.J.1042.2019.00508

JING Wei, WANG Tingzhao. The gaze processing impairment in individuals with autism spectrum disorder: A perspective from the two-process theory. Advances in Psychological Science, 2019, 27(3): 508-521 doi:10.3724/SP.J.1042.2019.00508

视线加工在社会性认知中起着核心作用(Itier & Batty, 2009)。临床行为观察研究发现, 孤独症谱系障碍(Autism Spectrum Disorder, ASD)个体普遍存在不能与他人建立视线接触、不能追随他人视线看向目标物体等视线加工障碍(Leekam, Lo Pez, & Moore, 2000; Tanaka & Sung, 2016)。因此, 由美国精神病学会制定的《精神疾病诊断和统计手册》第四版(Diagnostic and Statistical Manual of Mental Disorders, DSM-IV)将此列为ASD诊断依据之一(Psychiatric American Association, 2000)。然而, 通过对已有实验研究的梳理, 本文发现ASD个体在实验情境中普遍存在视线接触异常, 但其视线追随行为则存在正常与异常并存的现象。为了解释临床观察和实验情境之间、以及不同实验情境之间的矛盾, 本文将借鉴由面孔加工双通路理论发展而来的视线加工双通路理论, 初步探讨ASD个体视线加工障碍的潜在神经机制, 并在此基础之上提出未来的研究方向。

1 ASD个体实验情境中视线接触行为普遍存在异常

1.1 ASD个体视线接触异常的行为证据

行为研究结果证实, ASD个体在各类面孔加工任务中不存在ECE效应。例如, 在奇异球(oddball)探测任务中, TD个体探测直视面孔比斜视面孔更快更准, 表现出典型的ECE效应, 而ASD个体无此差异(Senju, Yaguchi, Tojo, & Hasegawa, 2003); 在性别识别(Pellicano & Macrae, 2009)和面孔记忆(Zaki & Johnson, 2013)任务中, TD儿童对直视面孔的反应时显著少于斜视面孔, 而直视面孔未促进ASD儿童的反应速度。与上述研究考察阈上面孔加工任务不同, 采用连续闪烁抑制范式(Continuous Flash Suppression, CFS)的研究发现, 在无意识条件下, TD被试对直视面孔的探测速度显著快于斜视面孔, 而ASD个体无此差异(Akechi et al., 2014)。另一以眼跳偏好指数为因变量的眼动研究亦发现, 在无意识条件下, TD成人更偏好于首先注视直视面孔, 而ASD成人则更偏好于首先注视斜视面孔(Madipakkam, Rothkirch, Dziobek, & Sterzer, 2017)。上述研究结果一致表明, ASD儿童或成人在各类阈上或阈下面孔加工任务中均不存在ECE效应。

近年来以高危ASD婴儿(兄弟姐妹被确诊为ASD的婴儿)为被试的前瞻研究也证实, ASD个体在生命早期, 在未确诊之前, 就缺乏对直视视线的敏感性和偏好性(Nele, Ellen, Petra, & Herbert, 2015; Nyström, Bölte, Falck-Ytter, & EASE, 2017)。例如, Nele等人(2015)发现, 相对于斜视面孔, 5个月的TD婴儿就表现出对直视面孔的注意偏好性, 而高危婴儿则未表现出此偏好性。Nyström等人(2017)以10个月的高危婴儿为被试, 呈现社会互动面孔视频, 采用以100 ms为单位的时间进程分析法, 进一步考察了该群体对视频中人物所呈现的直视视线的视觉注意行为。他们发现, 当视频中的人物注视被试时, TD婴儿的面孔注意偏好指数显著大于后期确诊的高危婴儿; 而当视频中的人物注视玩具时, 两者的面孔注意偏好指数无差异。这说明, TD婴儿对直视视线极为敏感。当他人直视自己时, 他们会立即增加对他人面孔的关注程度, 而后期确诊的高危婴儿无此敏感性。

1.2 ASD个体视线接触异常的生理证据

以生理指标为因变量的研究亦发现, ASD个体对直视视线和斜视视线的反应无差异(Helminen et al., 2017; Nuske, Vivanti, & Dissanayake, 2015)。例如, 心率减速(Heart Rate Deceleration, HRD)一般被认为是个体对刺激的注意集中程度和认知加工深度的有效测量指标。心率减速越大, 则表明个体对刺激的注意集中程度和认知加工深度越大。在Helminen等人(2017)的研究中, 研究者以2至5岁的低功能ASD儿童为被试, 分别呈现直视和斜视面孔, 测量两种视线所诱发的HRD信号。结果显示, 相对于斜视视线, 直视视线诱发了TD和智障儿童更大的HRD反应; 而ASD儿童与此相反。这说明, 对照组对直视视线的注意集中程度和认知加工深度显著高于斜视视线, 而ASD儿童与此相反。研究者认为这是由于ASD个体缺乏对直视视线所蕴含的社会性交流意图的理解。对于ASD个体而言, 直视视线不是一种更能够吸引个体注意的社会性突显信号。

1.3 ASD个体视线接触异常的脑电证据

脑电研究结果亦显示, ASD儿童或成人不存在ECE效应。相对于斜视视线, 直视视线在TD个体的右岛叶、颞顶联合区、额下回和上颞叶等社会脑皮层网络区域诱发更高的激活水平, 而ASD个体则恰好相反(Elsabbagh et al., 2015; Georgescu et al., 2013; Kylliäinen et al., 2012; Pitskel et al., 2011; von Dem Hagen, Stoyanova, Rowe, Baron- Cohen, & Calder, 2014)。例如, 一项运用Oddball视线探测任务的ERP研究发现, 直视视线在TD儿童的右半球枕颞区产生了较斜视视线更大的N2波, 而ASD儿童则不受视线方向的影响(Senju, Tojo, Yaguchi, & Hasegawa, 2005)。一项ASD成人的fMRI研究发现, 直视视线在TD成人右侧颞顶联合区、右前岛叶、左枕叶外侧皮层等脑区产生了较斜视视线更高的激活水平, 而ASD成人则与此相反(Pitskel et al., 2011)。另一项fMRI研究结果显示, 相对于斜视视线, 直视视线诱发了TD组杏仁核、后颞上沟、颞顶联合区以及前额叶中部等脑区更高的激活水平, 而ASD成人在相关脑区的激活水平与此相反(von Dem Hagen et al., 2014)。这说明, 相对于斜视视线, 直视视线未能诱发ASD成人或儿童社会脑神经网络更高的激活水平。

以高危婴儿为被试的前瞻神经生理研究进一步发现, 后期确诊的高危婴儿在确诊之前就未表现出ECE效应(Elsabbagh et al., 2015, 2012, 2009)。Elsabbagh等人(2009)首先发现对于10个月的高危婴儿而言, 直视面孔所诱发的P400波的潜伏期显著长于斜视面孔, 而TD婴儿与此相反。Elsabbagh等人(2012)进一步考察了3岁时确诊的高危婴儿在6个月时对动态视线转移的ERP信号反应与后期症状之间的关系。该研究发现, 较之于斜视视线, TD婴儿和未确诊的高危婴儿对直视视线在枕颞区产生了更大的P1、N260和P400波, 而确诊的高危婴儿未表现出此差异。在上述两个研究的基础之上, Elsabbagh等人(2015)进一步发现, 对高危婴儿而言, 不同视线朝向在枕颞区诱发的P400的潜伏期差异与亲子互动中的积极情感反应成正相关, 即表现出更多积极情感的高危婴儿对直视视线的反应速度显著快于斜视视线。上述结果说明, ASD个体在生命早期确诊之前就存在ECE障碍且与后期社会性障碍相关。

2 ASD个体实验情境中视线追随行为正常与异常并存

早期研究呈现孤立面孔, 采用Posner空间视线线索化范式, 考察ASD个体的视线追随行为。然而, 在真实生活中, 个体要成功追随他人视线, 首先要能够在复杂环境中捕获视线线索; 其次在追随他人视线对目标物体进行注意定向的基础之上, 还需要对目标物体进行注意加工。然而, 由于Posner范式呈现孤立面孔且以目标探测反应时为因变量, 因而个体无需捕获视线线索也无法测量目标物体的注意程度。上述因素导致Posner范式所得到的研究结果很难解释临床自然观察结果。因此, 研究者转而呈现具有生态效度的场景刺激, 同时搜集眼动数据, 考察ASD个体在场景刺激中的视线追随行为。通过对上述两类实验研究的梳理, 本文发现ASD个体的视线追随行为存在正常与异常并存的矛盾现象。

2.1 ASD个体面孔刺激中视线追随行为正常与异常并存

来自TD个体的研究显示, 在呈现面孔刺激的Posner范式中, 当视线方向与随后出现目标的位置一致时, 目标探测速度显著提高; 反之则显著降低(Schuller & Rossion, 2001, 2004)。即使视线线索对目标刺激没有预测作用, 仍诱导注意快速转移。研究者们将此现象称之为视线线索效应(Gaze Cuing Effect, GCE) (Driver et al., 1999)。该效应说明, 视线方向是一种非常有力的社会性线索, 能够诱导观察者将注意快速转移到视线所指示的方向上。然而, 来自ASD个体的研究发现, 该群体的视线追随行为在经典Posner范式中正常而在Posner范式变式中异常。

2.1.1 ASD个体经典Posner范式中视线追随行为正常

除极少数研究外(Gillespie-Lynch, Elias, Escudero, Hutman, & Johnson, 2013; Goldberg et al., 2008; Ristic et al., 2005), 绝大部分采用经典Posner范式的行为研究结果表明, ASD儿童、青少年或成人能够表现出正常的GCE效应(Ashwin, Hietanen, & Baron-Cohen, 2015; Böckler, Timmermans, Sebanz, Vogeley, & Schilbach, 2014; Chawarska, Klin, & Volkmar, 2003; Kirchgessner, Chuang, Patel, & Sereno, 2015; Kuhn et al., 2010; Nation & Penny, 2008)。例如,一项早期研究结果显示高功能ASD儿童与控制组一样, 即使视线线索对目标位置没有任何预测作用, 它仍能诱导该群体快速转移注意(Senju, Yoshikuni, Dairoku, & Hasegawa, 2004)。有研究证实, 虽然ASD幼童在自然情境中不能追随主试视线甚至头向或手势看向目标物体, 但在经典的Posner范式中却存在GCE效应(Chawarska et al., 2003)。与前述研究均以反应时为指标不同, 一项眼动研究也证实ASD个体在眼跳指标上亦存在GCE效应, 即一致条件下眼跳潜伏时间显著短于不一致条件(Kuhn et al., 2010)。这可能是因为在经典Posner范式中, 个体无需对视线线索进行初始探测且实验情境仅涉及视线方向属性, 因而ASD个体表现出正常的GCE效应。

2.1.2 ASD个体Posner范式变式中视线追随行为异常

近年来, 有研究表明在涉及视线社会意义的Posner范式的变式中, ASD个体的GCE效应异常(Ashwin et al., 2015; Gillespie-Lynch et al., 2013; Lassalle & Itier, 2014; Rombough & Iarocci, 2013; Zhao, Uono, Yoshimura, Kubota, & Toichi, 2017)。例如, Ashwin等人(2015)发现TD个体在面孔与视线朝向目标物体的条件下线索效应最大, 而ASD个体则在身体、面孔与视线均朝向目标物体的条件下线索效应最大。这可能是因为在面孔和视线朝向目标物体的条件下, TD个体认为对方注意到值得关注的事情而将注意力自动转向他人视线方向, 而在身体、面孔和视线均指向目标物体的条件下TD个体可能会推断对方只是身体方位转换而并不是注意到有趣事物, 而ASD个体只注意到身体方向这一显著特征。另一项研究还发现, TD个体的GCE效应存在视线线索与目标刺激之间社会背景关联性的促进效应, 即当线索为视线时, 目标刺激为人声的GCE效应显著大于音调; 当线索为箭头时, 两者之间无差异, 而ASD个体无此促进效应(Zhao et al., 2017)。上述研究说明, ASD个体可能将视线视为空间方向的物理性指示信号而非心理状态的社会性表征符号。因而, 该群体在涉及视线社会性意义的Posner范式的变式中表现异常。

2.2 ASD个体场景刺激中视线追随行为正常与异常并存

随着眼动追踪技术的普及, 研究者们还通过呈现场景刺激来考察ASD个体的视线追随行为。此类场景刺激一般在左右两边各呈现一个物体, 背景为空白墙面, 一位模特位于中央, 模特首先正视前方, 随后将视线和面孔朝向目标物体。研究者一般选取眼部至目标物体的眼跳次数和目标物体注视时间两类眼动指标, 分别考察被试对目标物体的注意定向准确性和注意加工偏好性。研究者们普遍认为两者分别反映了视线追随行为的准确性和视线指称意义的理解力。研究者们最初选取由视线转移至视频结束之间兴趣区内总注视点计算得到的宏观指标为测量指标。然而, 鉴于宏观指标不能灵敏地反映视线转移那一刻被试对目标物体的认知加工偏向, 研究者选取视线转移时兴趣区内首个注视点计算得到的微观指标为测量指标。通过对已有研究的梳理, 本文发现ASD个体的视线追随行为在简单场景宏观指标上正常, 而在简单场景微观指标上或复杂场景宏观指标上异常。

2.2.1 简单场景宏观指标上ASD个体的视线追随行为正常

在视线追随准确性方面, 以眼跳差值(视线转移至视频结束之间眼部至目标和非目标物体的眼跳次数之差)或眼跳差值比值(两者之差除以两者之和)等宏观指标为因变量的绝大部分研究结果显示, 在自由观察或词语习得任务中, ASD个体与对照组不存在差异(Billeci et al., 2016; Congiu, Fadda, Doneddu, & Striano, 2016; Falck-Ytter, Thorup, & Bölte, 2015; Thorup, Nyström, Gredebäck, Bölte, & Falck-Ytter, 2016; 荆伟, 方俊明, 赵微, 2014)。然而, 与上述研究以相对比值为因变量不同, 以绝对次数为因变量的研究发现ASD个体眼部至目标物体的眼跳次数显著少于TD组(Norbury, Griffiths, & Nation, 2010; 荆伟 等, 2014)。由于有研究证实ASD个体整个视频的总眼跳次数显著少于TD个体(Wang et al., 2015), 因而绝对值上的差异可能仅反映该群体在注意转移而非视线追随上存在障碍。上述研究结果表明ASD个体在简单场景刺激中能够追随他人视线对目标物体进行注意定向。这与ASD个体在经典Posner范式中表现出正常的视线线索注意定向能力相契合。

在视线意义理解力方面, 以注视时间比值(视线转移至视频结束之间目标物体注视时间除以整个视频注视时间)或差值比值(目标和非目标物体注视时间之差除以两者之和)等宏观指标为因变量的绝大部分研究结果亦显示, ASD儿童、青少年或成人与TD对照组不存在差异(Billeci et al., 2016; Congiu et al., 2016; Falck-Ytter, Fernell, Hedvall, von Hofsten, & Gillberg, 2012; Freeth, Ropar, Mitchell, Chapman, & Loher, 2011; Gliga, Elsabbagh, Hudry, Charman, & Johnson, 2012; Norbury et al., 2010; Swanson & Siller, 2013; 荆伟 et al., 2014)。然而, 以高危ASD婴儿为被试的前瞻研究发现36个月时确诊的高危婴儿在13个月时注视时间比值显著少于对照组(Bedford et al., 2012)。这可能是由于ASD个体的视线追随能力在生命早期滞后, 但随后发展起来的补偿机制可能使其存在一定发展。

2.2.2 简单场景微观指标上ASD个体的视线追随行为异常

同样在上述简单场景中, 少数选取微观指标为因变量的研究发现, 在准确性方面, 低功能而非高功能ASD个体与对照组存在差异。例如, 高功能ASD儿童的目标物体首次眼跳比值(视线转移后首次眼跳发生在眼部至目标物体之间的次数占有效试次的比值)与TD儿童无差异(Thorup, Kleberg, & Falck-Ytter, 2017); 后期确诊的高危婴儿在7和13个月时与未确诊的高危婴儿无差异(Bedford et al., 2012); 3岁左右ASD高危幼儿与TD幼儿亦无差异(Gliga et al., 2012)。然而, 低功能ASD幼童则显著低于生理年龄匹配的TD幼童(Gillespie-Lynch et al., 2013); 低功能ASD青少年亦显著低于智力水平匹配的TD儿童(荆伟 et al., 2014)。在理解力方面, 以目标物体首次注视时间(视线转移后目标物体首个注视点持续时间)为因变量的研究发现, 高功能ASD儿童显著低于TD儿童(Thorup et al., 2017), 低功能ASD儿童亦显著低于生理年龄匹配的TD儿童(Falck-Ytter et al., 2015)。由此可见, 在微观指标上, 高功能ASD个体可能具有正常的视线线索定向能力, 但视线意义解读能力存在障碍; 而低功能ASD个体可能在上述两方面均存在障碍。

2.2.3 复杂场景宏观指标上ASD个体的视线追随行为异常

在上述简单场景中, 无关干扰信息极少, 视线线索易于捕获, 因而ASD个体视线追随行为仅在微观指标上异常而在宏观指标上正常。近年来, 研究者们发现, 当干扰信息增多视线线索微弱时, ASD个体在宏观指标上也将异常。例如, 一项呈现三个物体的研究发现, ASD个体的眼跳差值显著低于对照组(Falck-Ytter et al., 2012)。Thorup等人(2016)发现, ASD高危婴儿在视线和面孔同时朝向目标物体时眼跳比值显著高于仅视线朝向目标物体, 而对照组无此差异。另一项研究发现, 当人物看向目标物体而非目标物体同时开始振动时, TD成人能够克服物理性特征的干扰作用稳定跟随他人视线看向并注视目标物体, 而ASD成人则不能克服此干扰(Aldaqre, Paulus, & Sodian, 2015)。此外, Congiu等人(2016)还发现, 在目标物体的位置对被试而言是可知的知觉条件下, ASD儿童视线追随行为正常, 而在目标物体位置不可知的表征条件下, ASD儿童则表现出障碍。他们认为视线对于ASD个体而言可能更多是一种空间方向的物理性指示信号而非心理状态的社会性表征符号。该结果与前述涉及视线社会性意义的Posner范式变式中的结果相印证。

与上述研究呈现人为控制的场景不同, 还有一部分研究呈现具有复杂背景的真实生活场景。他们发现, 在自由观察任务中, ASD个体对目标物体的注视时间比值显著少于对照组(Fletcher- Watson, Leekam, Benson, Frank, & Findlay, 2009; Riby, Hancock, Jones, & Hanley, 2013; Wang et al., 2015)。Ridy等人(2013)还发现, 即使要求被试对目标物体进行命名, ASD个体也未能像对照组那样, 增加目标物体的注视时间。不过, 一项引入时间变量的研究发现, 当视线呈现2秒时ASD个体对目标物体的注视时间显著低于对照组, 而呈现5秒时则无差异。通过以0.5秒为间隔的时间进程分析发现, 视线呈现1.5秒后TD组对目标物体的注视时间就显著提高, 而ASD组直到4秒后才显著提高(Freeth, Chapman, Ropar, & Mitchell, 2010)。此研究结果说明, 在真实的复杂场景中, 由于干扰信息众多, 因而ASD个体可能需要更多时间才能将注意资源分配至目标物体。

3 TD个体视线加工双通路理论的启示

面孔加工双通路理论提出, 除后天经验驱动形成的皮层通路外, 面孔加工还涉及一条包括杏仁核、上丘和枕核在内的作为人类先天机制而存在的皮下通路。与皮层通路对高空间频率面孔信息进行慢速有意识加工不同, 皮下通路对低空间频率面孔信息进行快速无意识的初始探测和定向, 先行于并促进皮层通路加工过程, 进而在皮层通路乃至社会脑的后天形成发展中发挥着重要作用。近年来, 为了解释ECE效应, 研究者们将皮下通路探测的社会性信息由面孔、恐惧表情或生物运动等扩展至直视视线, 进一步提出了视线加工双通路理论(Johnson, Senju, & Tomalski, 2015; Senju & Johnson, 2009)。

3.1 TD个体皮下通路对直视视线的快速无意识探测

Morris, Öhman和Dolan (1999)提出上丘-丘脑枕核-杏仁核皮下快速通路之后, 大量以“皮层盲视”病人以及正常个体为被试的脑成像和脑损伤研究为该通路的存在提供了证据。研究者们发现, 与中性表情相比, 在无意识知觉条件下, 负性表情能够引起皮下通路中各成分的特异性激活以及各成分之间功能连接的增强(见综述:陈珊珊, 蔡厚德, 2015)。近年来, 运用CFS范式呈现阈下刺激的研究证据显示, 与负性表情一样, 阈下直视视线亦能快速激活该皮下通路。例如, 以反应时为因变量的行为研究发现, 相比斜视面孔, TD个体能更快克服CFS抑制作用探测到直视面孔(Akechi et al., 2014; Stein, Senju, Peelen, & Sterzer, 2011)。以眼跳偏好指数为因变量的眼动研究亦发现, 在完全不能知觉到面孔的CFS条件下, 相对于斜视面孔, TD成人更偏好于首先注视直视面孔(Madipakkam et al., 2017; Rothkirch, Madipakkam, Rehn, & Sterzer, 2015)。在一项呈现阈上表情面孔考察视线方向对表情识别影响作用的ERP研究中, 研究者在刺激呈现后85 ms观测到直视视线诱发的神经反应信号, 而情绪表情诱发的神经反应信号则出现在115 ms (Klucharev & Sams, 2004)。这说明皮下通路对直视视线的反应十分迅速, 发生在皮层通路对情绪表情进行完整地细节性分析之前。上述研究为皮下通路对直视视线的快速无意识探测能力提供了行为神经水平上的间接证据。

近年来, 脑功能成像研究则为该理论提供了直接证据。他们发现阈下直视能够诱发皮下通路尤其是杏仁核的激活反应。例如, 来自于皮层盲视患者的研究发现, 较之于斜视视线, 当观察到直视视线时, 患者皮下通路中的杏仁核激活水平显著提高(Burra et al., 2013)。一项TD个体的研究也发现, 在无意识条件下, 直视视线在包括上丘和杏仁核在内的双侧皮下网络区域的激活水平显著高于斜视视线(Sato, Kochiyama, Uono, & Toichi, 2016)。另一来自于灵长类动物的研究甚至在猴子的杏仁核中直接探测到一类对直视视线做出选择性反应的神经元(Mosher, Zimmerman, & Gothard, 2014)。当观察者注视被观察者眼部而被观察者注视别处时, 杏仁核中的部分神经元产生激活反应且反应极为迅速, 平均激活潜伏时间仅为118 ms。而当观察者与被观察者建立视线接触时, 已处于激活状态的“眼部注视”神经元中的一部分神经元的激活水平进一步增强。这表明视线接触能够诱发皮下通路中杏仁核更强的神经反应信号。

3.2 TD个体皮下通路对皮层通路加工过程的促进作用

除上述研究证实直视视线能够快速激活皮下通路外, 还有大量来自脑功能成像的研究证实, 直视视线还能够促进面孔加工皮层通路的神经活动。例如, 当个体观察直视视线时, 社会脑皮层网络, 如梭状回(George, Driver, & Dolan, 2001; Grossmann & Johnson, 2007)、颞上沟(Conty, N'Diaye, Tijus, & George, 2007; Grossmann & Johnson, 2007)、前额内侧(Grossmann & Johnson, 2007; Schilbach et al., 2006)、眶额皮层(Conty et al., 2007; Grossmann & Johnson, 2007)等脑区的激活水平显著高于斜视视线。来自神经生理研究的证据亦显示, 阈上直视视线在刺激呈现之后150~220 ms之间在前额内侧、颞上沟和眶额皮层等社会脑皮层网络所诱发的ERP信号显著大于斜视视线(Conty et al., 2007), 而阈下直视视线在刺激呈现200 ms和250 ms时在额顶皮层也诱发了较斜视视线更大的负波(Yokoyama, Noguchi, & Kita, 2013)。上述研究结果说明, 较之于斜视视线, 直视视线能够促进社会脑皮层网络的激活水平。同时, 还有大量研究证实, 较之于斜视视线, 当被试感知直视视线时, 面孔加工皮下与皮层通路之间的功能连接亦显著提高。例如, 在一项考察视线方向对性别识别任务影响作用的研究中, 研究者发现相较于斜视面孔, 在直视面孔加工中梭状回面孔区与杏仁核之间的神经活动显示出更高的耦合水平(George et al., 2001)。另一项考察视线接触对表情识别任务中社会脑神经网络激活水平影响作用的研究发现, 较之于自由观看, 当建立视线接触时, TD个体的社会脑神经网络激活水平在各类表情面孔中均显著提高, 且与杏仁核之间的功能连接亦显著提高(Hadjikhani et al., 2017)。依据双通路理论, 皮下通路不仅能对面孔信息进行快速无意识的初始探测和定向, 而且还能够调节皮层通路的加工过程。因此, 研究者们推测直视视线对皮层通路的促进作用可能是源于直视视线能够快速激活皮下通路进而促进皮层通路的加工过程(Johnson et al., 2015; Senju & Johnson, 2009)。

皮下通路对皮层通路的调节作用主要涉及以下两个神经回路。由上丘-丘脑枕核-杏仁核组成的皮下快速通路通过上丘接收视网膜神经节大细胞传递的低空间频率视觉信息, 经由丘脑枕核快速传递至杏仁核, 再由杏仁核向前额叶和顶叶等注意神经皮层网络进行投射, 从而调控皮层通路的社会性信息加工过程(Pourtois, Schettino, & Vuilleumier, 2013)。该通路在没有视觉皮层参与的情况下不仅能够对社会性刺激进行快速无意识初始探测(Diano, Celeghin, Bagnis, & Tamietto, 2017; Gainotti, 2012), 而且还涉及以社会性行为动机为基础的注意选择过程(Troiani, Price, & Schultz, 2012; Troiani & Schultz, 2013)。在Troiani等人(2012, 2013)的研究中, 与房屋刺激相比, 在阈下恐惧面孔的早期加工过程中, 虽然腹侧视觉流中面孔敏感区域(梭状回或海马旁回)和初级视觉皮层均未激活, 但伴随皮下通路中上丘、丘脑枕核和杏仁核的激活, 部分背侧视觉流(顶下叶皮层)激活, 且杏仁核与右丘脑枕核和左侧顶下叶之间的功能连接增强。这说明在无意识条件下杏仁核依然能够将信息传递至注意神经皮层网络, 引导个体将注意转移至社会性突显刺激。另一丘脑枕核-皮层环路则强调丘脑枕核基于注意调节和选择在皮层网络之间的同步化作用(Malekmohammadi, Elias, & Pouratian, 2015; Saalmann, Pinsk, Wang, Li, & Kastner, 2012)。Saalmann等人(2012)在猴子进行视觉空间注意任务的同时, 记录了丘脑枕核、V4和枕颞皮层等脑区神经元的电活动。研究结果显示, 上述脑区神经元的兴奋水平在低频波段α波(8~15 Hz)有很强的一致性。Malekmohammadi等人(2015)甚至在两名癫痫症病人处于清醒状态进行自发行为时, 同时记录了丘脑和皮层的电生理活动。他们的研究结果进一步证实, 丘脑是基于一种丘脑低频振荡相位调节皮层高频振荡振幅的“相位振幅耦合”机制对皮层活动进行调节的。上述研究结果表明, 丘脑枕核通过同步相邻皮层神经元的振荡水平, 根据注意分配调节皮层细胞激活水平的同步化, 由此提高信息在相邻皮层间传递的效率, 使信息以一种自动化的前注意方式在相邻皮层网络之间进行快速传递(Benarroch, 2015; Villeneuve, Thompson, Hess, & Casanova, 2012; Ward, Calder, Parker, & Arend, 2007)。

3.3 TD个体皮下通路对社会脑发展形成的重要作用

来自新生婴儿的研究证明, TD个体在生命初始即对社会性信息具有先天的注意加工倾向。例如, 较之于斜视面孔, 新生儿更倾向于注视直视面孔(Farroni, Menon, & Johnson, 2006)。与“非面孔”刺激或随机物理运动等非社会性刺激比较而言, 新生儿更偏好于注视“类面孔”和生物运动等社会性刺激(Di Giorgio et al., 2016)。由于上述研究有效地控制了后天社会性经验的影响作用, 因而他们认为TD新生儿的先天社会性注意倾向可能正是源于皮下通路对社会性信息的先天快速注意定向机制。据此, 从发展的视角, 双通路理论进一步提出, TD个体的先天社会性注意倾向使皮层网络获得了更多的后天社会性经验, 从而使个体逐渐形成一个由腹内侧前额叶、颞下回、颞上沟、梭状回、扣带回等皮层组织以及上丘、枕核、杏仁核等皮下组织组成的神经网络结构, 即专门加工社会性信息的社会脑(Johnson et al., 2015; Senju & Johnson, 2009)。有研究证实, 社会脑的形成是一个受经验驱动起始于婴儿期早期直到青春期后期才逐渐成熟的漫长发展过程(Amestoy, Guillaud, Bouvard, & Cazalets, 2015; Meaux et al., 2014)。例如, 梭状回面孔特异性(Aylward et al., 2005), 与眼睛视线加工和社会意图解读(Pelphrey, Viola, & McCarthy, 2004)相关的颞下回(Shaw et al., 2008), 以及皮下与皮层通路之间的功能联结均直到青春期才发展成熟(Thomas et al., 2001)。来自杏仁核损伤病人的研究还证实, 童年期杏仁核损伤病人在心理理论推理任务中表现出障碍, 而成年期杏仁核损伤病人则表现正常。这说明皮下通路中的杏仁核在社会脑心理理论神经系统的发展过程中起到了重要作用(Shaw et al., 2004)。

4 ASD个体视线加工障碍的潜在神经机制

由于大量研究证实ASD个体的杏仁核存在形态(Ecker, Suckling, Deoni, & Et, 2012; Schumann & Amaral, 2006)和功能异常(Gotts et al., 2012; Philip et al., 2012), Senju和Johnson等人(2009)首先提出ASD个体的ECE障碍可能是来源于杏仁核受损而导致的皮下通路功能异常。同时, 鉴于皮下通路对皮层通路后天发展的调节作用(Schultz, 2005), 本文在Senju和Johnson等人的基础之上进一步提出, ASD个体可能因缺乏皮下通路的调节作用而导致皮层通路后天发展异常。因而, 笔者推测ASD个体视线加工障碍可能既源于皮下通路先天功能异常也源于皮层通路后天发展异常。

4.1 ASD个体的视线加工皮下通路或存在先天功能异常

依据面孔加工双通路理论, TD个体的皮下通路对包括直视视线在内的面孔、恐惧表情或生物运动等社会性刺激具有快速无意识探测加工能力。然而, 来自ASD个体的研究发现, 在性别识别和情绪效价判断任务中, 当刺激呈现时间为33 ms时, ASD个体未表现出与TD个体一样的阈下面孔加工能力(Vanmarcke & Wagemans, 2017)。Guillon等人(2016)发现, 与不会被知觉为面孔的倒立“类面孔”刺激比较而言, ASD幼童指向会被知觉为面孔的正立“类面孔”刺激的首次眼跳比值更低, 而对照组与此相反(Guillon et al., 2016)。另一来自ASD个体的fMRI研究还直接观测到在阈下恐惧面孔加工中, 控制组在杏仁核、枕核和上丘等皮下结构有显著激活反应, 而ASD在上述脑区则无激活反应(Kleinhans et al., 2011)。上述结果为ASD个体皮下通路存在功能异常提供给了有力证据。另一项以高危新生儿为被试的前瞻研究还发现, 与TD新生儿相反, 相对于正立“类面孔”和生物运动等社会性刺激, 后期确诊的高危新生儿更偏好于注视倒立“类面孔”和随机物理运动等非社会性刺激。这说明ASD个体的皮下通路功能异常很可能是先天性的(Di Giorgio et al., 2016)。

4.2 ASD个体的视线加工皮层通路或存在后天发展异常

来自脑功能成像的前瞻研究发现, 后期确诊的高危ASD婴儿6个月时, 枕中回、右契叶和右舌回等初级视觉信息加工皮层的表面积显著大于TD婴儿(Hazlett et al., 2017), 而额下回和颞上沟等社会脑皮层网络由社会性刺激所诱发的激活水平显著低于TD婴儿(Lloyd Fox et al., 2017)。由此可见, ASD个体的大脑皮层早在生命之初, 确诊之前, 在行为障碍出现之前, 就表现出初级感知觉皮层过度增长而高级社会脑皮层激活不足的异常发展模式。另一来自父母为盲人的新生婴儿的研究证实, 当缺乏视线加工的早期后天社会性经验时, 此类婴儿在12个月时表现出视线加工行为障碍。这说明, 早期后天社会性经验在大脑社会性功能发展中具有重要作用(Senju et al., 2015)。结合该结果, 本文认为面孔加工双通路理论能够为上述异常发展模式提供一定的理论解释:皮下通路先天功能异常导致ASD个体在生命之初即缺乏先天社会性注意倾向, 进而剥夺了皮层通路获得早期后天社会性经验的机会, 导致皮层通路的社会性信息输入减少而非社会性信息输入增多, 因而面孔加工皮层通路呈现出异常发展模式, 进而导致社会脑的发展异常(Grice et al., 2005; Kleinhans et al., 2011; Schultz 2005)。

脑成像研究已经证实, 作为社会脑的重要组成部分, ASD个体的视线加工皮层通路也存在此异常发展模式(Greene et al., 2011; Lajiness-O Neill et al., 2014; Pelphrey, Morris, & McCarthy, 2005; Vaidya et al., 2011)。例如, 一项研究结果显示, 在视线方向与目标物体不一致条件下, TD个体颞上沟的激活水平显著高于一致条件, 而ASD个体颞上沟的激活水平在两种条件下无差异。他们认为这是由于ASD个体的颞上沟能够识别视线方向但不能解读视线意图(Pelphrey et al., 2005)。另一项研究报告显示, 相较于中性视线, 视线线索在TD个体的左额下回、扣带回等社会脑皮层网络诱发了更高激活水平, 而ASD个体则仅在契叶等初级视觉皮层网络有更高激活水平(Greene et al., 2011)。这种初级感知觉皮层过度激活而高级社会脑皮层激活不足的异常模式, 使ASD个体不能理解视线所蕴含的社会性意义, 因而只能基于低水平的物理性的方向属性而非高水平的社会性的指称意义, 以一种加工非社会性信息的补偿机制对视线线索进行加工。

4.3 基于上述潜在机制对ASD个体视线加工障碍的解释

为何ASD个体的视线接触行为普遍存在异常, 而视线追随行为却正常与异常并存?基于上述潜在机制, 我们认为这可能是因为, 一方面, 视线接触效应主要涉及皮下通路对直视视线的快速无意识反应能力, 而ASD个体由于皮下通路功能异常, 因而直视视线不能有效激活皮下通路。那么, 在实验情境中, ASD个体在各类面孔认知加工任务中均不能表现出直视视线的促进作用; 而在临床观察中, ASD个体可能因为缺乏对直视视线的敏感性进而不能与他人建立视线接触。另一方面, 视线追随行为则不仅涉及皮下通路对视线线索的快速无意识探测能力, 还涉及皮层通路对视线线索的空间方向和指称意图的加工能力。当实验情境要求被试从背景中捕获视线线索时, ASD个体可能因皮下通路异常而缺乏此视线线索捕获能力, 进而导致视线追随行为异常; 当实验情境涉及视线的社会背景信息时, ASD个体可能因皮层通路异常而缺乏视线意义解读能力, 进而导致视线追随行为异常。只有当实验情境既不涉及视线线索探测也不涉及视线意义解读时, ASD个体可能凭借皮层通路的补偿机制表现出正常的视线追随行为。

因而, 在简单场景宏观指标上或经典Posner范式中, 视线线索一般位于视域中央, 干扰信息极少, 且仅涉及视线方向加工, ASD个体无需探测视线线索也无需解读视线意义, 因而表现出了正常的视线追随行为; 而在复杂场景中视线线索一般位于外周视域, 干扰信息极多, 需要被试首先启动皮下通路捕获视线线索, 因而ASD个体便表现出了异常的视线追随行为; 而在Posner范式的变式中又因涉及视线线索的社会背景信息, 需要被试解读视线线索意义, 因而ASD个体也表现出了异常的视线追随行为; 而在临床观察中, 真实社会生活情境既涉及视线线索捕获也涉及视线意义解读, 所以ASD便普遍表现出了异常的视线追随行为。综上, 皮下通路功能异常可能不仅使ASD个体对直视视线缺乏敏感性进而表现出视线接触障碍, 而且还使ASD个体因缺乏在复杂情境中捕获视线线索的能力而表现出视线追随障碍; 而皮层通路异常发展使ASD个体在涉及社会背景信息的情境中表现出视线追随障碍。由此可见, 该机制能够较好地解释ASD个体在实验情境和临床观察之间、不同视线加工行为之间, 以及视线追随行为在不同实验情境之间存在的矛盾。

5 总结与展望

通过对已有研究的梳理, 本文发现在实验情境中, ASD个体在各类面孔认知加工任务中普遍不存在ECE效应, 而视线追随行为则存在异常与正常并存的矛盾现象。借鉴视线加工双通路理论, 本文推测其潜在神经机制可能是视线加工的皮下通路先天功能异常而皮层通路后天发展异常。皮下通路先天功能异常使ASD个体缺乏对直视视线的敏感性和偏好性以及在复杂情境中捕获视线线索的能力, 而皮下通路后天发展异常使ASD个体以一种基于物理性方向而非社会性意义的补偿机制对视线线索进行加工。因而, 在无关干扰信息众多而视线线索尤为微弱的真实社会生活情境中, ASD个体便普遍表现出不能与他人建立视线接触、不能追随他人视线看向目标物体等视线加工障碍。双通路理论聚焦于早期先天皮下通路, 同时兼顾皮层通路后天发展及其与皮下通路先天机制之间的关系, 能够将不同视线加工障碍纳入统一的理论框架中, 以一种动态发展联系的视角进行解释。然而, 该理论虽颇具前景, 但尚不完备, 今后研究者还需从以下三个方面进一步深入探讨:

5.1 验证皮下通路先天功能异常是视线接触障碍的神经机制

我们已有大量不同水平的研究证据显示TD个体普遍存在ECE效应, 且其潜在神经机制是皮下通路对直视视线的快速无意识探测能力。而ASD个体普遍不存在ECE效应, 且该群体所普遍表现出的阈下面孔加工障碍表明该群体的皮下通路或存在功能异常。据此, 研究者们推测ASD个体的视线接触障碍可能源于皮下通路功能异常导致该群体对直视视线的加工缺乏敏感性和优势性。然而, 由于目前我们尚未直接观察到ASD个体皮下通路对直视视线的神经反应模式, 因而上述理论假设还缺乏来自ASD个体生命早期的神经生理水平上的直接有力证据。今后研究者可对上述假设进行验证。

5.2 考察皮下通路先天功能异常对视线追随障碍的影响作用

鉴于微观指标的灵敏性, 笔者推测ASD个体在简单场景微观指标上表现出来的障碍很可能也是源于皮下快速通路先天异常所导致的视线线索快速无意识探测障碍。依据双通路理论提出的皮下通路主要对低空间频率社会性信息进行快速无意识加工, 笔者预测ASD个体的皮下通路功能异常将导致该群体在面孔刺激中, 当视线呈现在阈下或低空间频率条件下ASD个体将表现出视线加工障碍, 反之则正常; 在场景刺激中, 当视线呈现在外周视域或快速呈现时障碍明显表现在宏观指标上, 反之则微弱仅表现在微观指标上。因此, 为了验证前述假设并澄清前述矛盾, 笔者认为今后的研究可尝试在面孔刺激中引入视线呈现的意识状态和空间频率、在场景刺激中引入视线呈现位置和时间考察皮下通路先天功能异常对ASD个体视线追随障碍的影响作用。

5.3 揭示皮层通路后天发展异常的神经回路及早期形成过程

已有脑成像研究证实ASD个体的视线加工皮层通路在Posner范式下激活模式异常, 但由于相关研究证据积累不够, 关于此异常模式, 研究者们目前仅仅初步证实ASD个体的初级感知觉皮层过度激活而高级社会性皮层激活不足, 而该补偿机制具体涉及哪些大脑皮层结构还不得而知。因而, 今后研究者可考虑引入视线线索的意识状态、空间频率或视线指称意义等变量, 比较阈上和阈下、高空间频率和低空间频率, 或方向性和指称性条件下ASD个体视线线索注意定向行为的神经激活反应模式, 以期逐步揭示该补偿机制的神经回路。同时, 鉴于双通路理论所提出的皮下通路在皮层通路以及社会脑后天发展中的重要作用, 今后的研究还应考虑选取高危ASD婴儿为被试, 聚焦于视线加工皮层通路异常发展的早期形成过程, 着重考察皮下通路与皮层通路之间的功能连接。目前, 仅有Hazlett等人(2017)和Lloyd Fox等人(2017)的脑功能成像前瞻研究初步证实在生命之初确诊之前, 6个月的ASD婴儿的皮层通路就表现出异常发展模式。如果有更多此类脑功能成像前瞻研究进一步探究皮层通路异常发展模式的早期形成过程, 那么ASD的诊断和干预将能够凭借此类早期神经征兆大大提前。

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Journal of Cognitive Neuroscience, 17( 2), 308-319.

DOI:10.1162/0898929053124884      URL     PMID:15811242      [本文引用: 1]

Behavioral studies suggest that children under age 10 process faces using a piecemeal strategy based on individual distinctive facial features, whereas older children use a configural strategy based on the spatial relations among the face's features. The purpose of this study was to determine whether activation of the fusiform gyrus, which is involved in face processing in adults, is greater during face processing in older children (12 14 years) than in younger children (8 10 years). Functional MRI scans were obtained while children viewed faces and houses. A developmental change was observed: Older children, but not younger children, showed significantly more activation in bilateral fusiform gyri for faces than for houses. Activation in the fusiform gyrus correlated significantly with age and with a behavioral measure of configural face processing. Regions believed to be involved in processing basic facial features were activated in both younger and older children. Some evidence was also observed for greater activation for houses versus faces for the older children than for the younger children, suggesting that processing of these two stimulus types becomes more differentiated as children age. The current results provide biological insight into changes in visual processing of faces that occur with normal development.

Bedford R., Elsabbagh M., Gliga T., Pickles A., Senju A., Charman T., & Johnson M H. .,( 2012).

Precursors to social and communication difficulties in infants at-risk for autism: Gaze following and attentional engagement

Journal of Autism and Developmental Disorders, 42( 10), 2208-2218. doi: 10.1007/s10803-012-1450-y.

URL     PMID:22278030      [本文引用: 2]

Whilst joint attention (JA) impairments in autism have been widely studied, little is known about the early development of gaze following, a precursor to establishing JA. We employed eye-tracking to record gaze following longitudinally in infants with and without a family history of autism spectrum disorder (ASD) at 7 and 13months. No group difference was found between at-risk and low-risk infants in gaze following behaviour at either age. However, despite following gaze successfully at 13months, at-risk infants with later emerging socio-communication difficulties (both those with ASD and atypical development at 36months of age) allocated less attention to the congruent object compared to typically developing at-risk siblings and low-risk controls. The findings suggest that the subtle emergence of difficulties in JA in infancy may be related to ASD and other atypical outcomes.

Benarroch E.E . ( 2015).

Pulvinar: Associative role in cortical function and clinical correlations

Neurology, 84( 7), 738-748.

DOI:10.1212/WNL.0000000000001276      URL     PMID:25609762      [本文引用: 1]

Abstract The pulvinar is the largest nucleus of the thalamus and has strong connectivity with the visual cortex. The pulvinar is a prototypic association nucleus that participates in reciprocal cortico-cortical interactions and promotes synchronized oscillatory activity in functionally related areas of the cortex. Via its connections with the superior colliculus (SC) and areas of the dorsal visual stream projecting to the posterior parietal cortex (PPC), the pulvinar is an important component of the visual attention network. Visual salience is likely an important function of the pulvinar. The pulvinar also participates in mechanisms of social cognition and blindsight. Unilateral lesions of the pulvinar result in a contralateral neglect syndrome resembling that resulting from lesions of the posterior parietal cortex. The pulvinar is susceptible to damage in epilepsy and prion disorders, and may contribute to cognitive and visual manifestations of Lewy body disease. The anatomy, physiology, and putative function of the pulvinar and its involvement in neurologic disorders have been the subject of comprehensive reviews.(1-3</SUP)

Billeci L., Narzisi A., Campatelli G., Crifaci G., Calderoni S., Gagliano A., .. ALERT group ( 2016).

Disentangling the initiation from the response in joint attention: An eye-tracking study in toddlers with autism spectrum disorders

Translational Psychiatry, 6, e808.

DOI:10.1038/tp.2016.75      URL     PMID:27187230      [本文引用: 2]

Joint attention (JA), whose deficit is an early risk marker for autism spectrum disorder (ASD), has two dimensions: (1) responding to JA and (2) initiating JA. Eye-tracking technology has largely been used to investigate responding JA, but rarely to study initiating JA especially in young children with ASD. The aim of this study was to describe the differences in the visual patterns of toddlers with ASD and those with typical development (TD) during both responding JA and initiating JA tasks. Eye-tracking technology was used to monitor the gaze of 17 children with ASD and 15 age-matched children with TD during the presentation of short video sequences involving one responding JA and two initiating JA tasks (initiating JA-1 and initiating JA-2). Gaze accuracy, transitions and fixations were analyzed. No differences were found in the responding JA task between children with ASD and those with TD, whereas, in the initiating JA tasks, different patterns of fixation and transitions were shown between the groups. These results suggest that children with ASD and those with TD show different visual patterns when they are expected to initiate joint attention but not when they respond to joint attention. We hypothesized that differences in transitions and fixations are linked to ASD impairments in visual disengagement from face, in global scanning of the scene and in the ability to anticipate object's action.

Böckler A., Timmermans B., Sebanz N., Vogeley K., & Schilbach L . ( 2014).

Effects of observing eye contact on gaze following in High-Functioning autism

Journal of Autism and Developmental Disorders, 44( 7), 1651-1658. doi: 10.1007/s10803-014-2038-5.

URL     PMID:24442835      [本文引用: 1]

Abstract Observing eye contact between others enhances the tendency to subsequently follow their gaze and has been suggested to function as a social signal that adds meaning to an upcoming action or event. The present study investigated effects of observed eye contact in high-functioning autism (HFA). Two faces on a screen either looked at or away from each other before providing congruent or incongruent gaze cues to one of two target locations. In contrast to control participants, HFA participants did not depict enhanced gaze following after observing eye contact. Individuals with autism, hence, do not seem to process observed mutual gaze as a social signal indicating the relevance of upcoming (gaze) behaviour. This may be based on the reduced tendency of individuals with HFA to engage in social gaze behavior themselves, and might underlie some of the characteristic deficiencies in social communicative behaviour in autism.

Burra N., Hervais-Adelman A., Kerzel D., Tamietto M., de Gelder B., & Pegna A. J . ( 2013).

Amygdala activation for eye contact despite complete cortical blindness

The Journal of Neuroscience, 33( 25), 10483-10489.

DOI:10.1523/JNEUROSCI.3994-12.2013      URL     PMID:23785160      [本文引用: 1]

Cortical blindness refers to the loss of vision that occurs after destruction of the primary visual cortex. Although there is no sensory cortex and hence no conscious vision, some cortically blind patients show amygdala activation in response to facial or bodily expressions of emotion. Here we investigated whether direction of gaze could also be processed in the absence of any functional visual cortex. A well-known patient with bilateral destruction of his visual cortex and subsequent cortical blindness was investigated in an fMRI paradigm during which blocks of faces were presented either with their gaze directed toward or away from the viewer. Increased right amygdala activation was found in response to directed compared with averted gaze. Activity in this region was further found to be functionally connected to a larger network associated with face and gaze processing. The present study demonstrates that, in human subjects, the amygdala response to eye contact does not require an intact primary visual cortex.

Chawarska K., Klin A., & Volkmar F . ( 2003).

Automatic attention cueing through eye movement in 2-year-old children with autism

Child Development, 74( 4), 1108-1122. doi: 10.1111/1467-8624.00595.

URL     PMID:12938707      [本文引用: 2]

http://www.jstor.org/stable/3696211

Congiu S., Fadda R., Doneddu G., & Striano T . ( 2016).

Impaired representational gaze following in children with autism spectrum disorder

Research in Developmental Disabilities, 57, 11-17. doi: 10.1016/j.ridd.2016.06.008.

URL     PMID:27348855      [本文引用: 3]

Using eye-tracking methodology, we compared spontaneous gaze following in young children with Autism Spectrum Disorder (mean age 5.8 years) to that of typically developing children (mean age 5.7 years). Participants saw videos in which the position of a hidden object was either perceptually visible or was only represented in another person mind. The findings indicate that children with Autism Spectrum Disorder were significantly less accurate in gaze following and observed the attended object for less time than typically developing children only in the Representational Condition. These results show that children with Autism Spectrum Disorder are responsive to gaze as a perceptual cue although they ignore its representational meaning.

Conty L., N'Diaye K., Tijus C., & George N . ( 2007).

When eye creates the contact! ERP evidence for early dissociation between direct and averted gaze motion processing

Neuropsychologia, 45( 13), 3024-3037. doi: 10.1016/j.neuropsychologia.2007.05.017.

URL     PMID:17644145      [本文引用: 3]

Direct gaze or eye contact is a strong social signal, which triggers distinct processes as compared to other gaze directions. Thus, direct gaze could be early dissociated from averted gaze during the processing of gaze direction. In order to examine the processing of gaze contact relative to averted gaze, we studied evoked potentials in human adults in response to the apparent motion of gaze. Deviated and frontal faces were presented with a fixed gaze direction, followed by an apparent movement of the eyes either toward the subject or away from him/her. The results showed that the perception of direct relative to averted gaze evoked a greater, later and longer lasting N170, suggesting that gaze contact recruited more resources than averted gaze in the early stage of gaze direction processing. Furthermore, direct and averted motion of gaze elicited distinct ERP components between 160 and 210 ms, initiated over centro-parietal electrodes. Source reconstruction revealed the involvement of the Theory-of-Mind network, including the regions of the superior temporal sulcus, the medial prefrontal and the orbitofrontal cortices, in this early dissociation. In addition, the perception of gaze contact relative to averted gaze yielded increased fronto-central P3a and parieto-occipital P3b. All the results were significant whatever the head orientation. Our findings show that gaze contact, as compared to other gaze directions, is an essential social cue which recruits early specific processes. The dissociation between direct and averted gaze processing occurs as soon as 160 ms, involving the social brain network.

Di Giorgio E., Frasnelli E., Salva O. R., Luisa S. M., Puopolo M., Tosoni D., … Vallortigara G . ( 2016).

Difference in visual social predispositions between newborns at low-and high-risk for autism

Scientific Reports, 6, 26395.

DOI:10.1038/srep26395      URL     PMID:27198160      [本文引用: 2]

Abstract Some key behavioural traits of Autism Spectrum Disorders (ASD) have been hypothesized to be due to impairments in the early activation of subcortical orienting mechanisms, which in typical development bias newborns to orient to relevant social visual stimuli. A challenge to testing this hypothesis is that autism is usually not diagnosed until a child is at least 3 years old. Here, we circumvented this difficulty by studying for the very first time, the predispositions to pay attention to social stimuli in newborns with a high familial risk of autism. Results showed that visual preferences to social stimuli strikingly differed between high-risk and low-risk newborns. Significant predictors for high-risk newborns were obtained and an accurate biomarker was identified. The results revealed early behavioural characteristics of newborns with familial risk for ASD, allowing for a prospective approach to the emergence of autism in early infancy.

Diano M., Celeghin A., Bagnis A., & Tamietto M . ( 2017).

Amygdala response to emotional stimuli without awareness: Facts and interpretations

Frontiers in Psychology, 7, 2029.

DOI:10.3389/fpsyg.2016.02029      URL     PMID:5222876      [本文引用: 1]

Over the past two decades, evidence has accumulated that the human amygdala exerts some of its functions also when the observer is not aware of the content, or even presence, of the triggering emotional stimulus. Nevertheless, there is as of yet no consensus on the limits and conditions that affect the extent of amygdala response without focused attention or awareness. Here we review past and recent studies on this subject, examining neuroimaging literature on healthy participants as well as brain-damaged patients, and we comment on their strengths and limits. We propose a theoretical distinction between processes involved inattentional unawareness, wherein the stimulus is potentially accessible to enter visual awareness but fails to do so because attention is diverted, and insensory unawareness, wherein the stimulus fails to enter awareness because its normal processing in the visual cortex is suppressed. We argue this distinction, along with data sampling amygdala responses with high temporal resolution, helps to appreciate the multiplicity of functional and anatomical mechanisms centered on the amygdala and supporting its role in non-conscious emotion processing. Separate, but interacting, networks relay visual information to the amygdala exploiting different computational properties of subcortical and cortical routes, thereby supporting amygdala functions at different stages of emotion processing. This view reconciles some apparent contradictions in the literature, as well as seemingly contrasting proposals, such as the dual stage and the dual route model. We conclude that evidence in favor of the amygdala response without awareness is solid, albeit this response originates from different functional mechanisms and is driven by more complex neural networks than commonly assumed. Acknowledging the complexity of such mechanisms can foster new insights on the varieties of amygdala functions without awareness and their impact on human behavior.

Doi H. & Shinohara K., ( 2015).

Unconscious presentation of fearful face modulates electrophysiological responses to emotional prosody

Cerebral Cortex, 25(3), 817-832.

DOI:10.1093/cercor/bht282      URL     PMID:24108801     

Abstract Cross-modal integration of visual and auditory emotional cues is supposed to be advantageous in the accurate recognition of emotional signals. However, the neural locus of cross-modal integration between affective prosody and unconsciously presented facial expression in the neurologically intact population is still elusive at this point. The present study examined the influences of unconsciously presented facial expressions on the event-related potentials (ERPs) in emotional prosody recognition. In the experiment, fearful, happy, and neutral faces were presented without awareness by continuous flash suppression simultaneously with voices containing laughter and a fearful shout. The conventional peak analysis revealed that the ERPs were modulated interactively by emotional prosody and facial expression at multiple latency ranges, indicating that audio-visual integration of emotional signals takes place automatically without conscious awareness. In addition, the global field power during the late-latency range was larger for shout than for laughter only when a fearful face was presented unconsciously. The neural locus of this effect was localized to the left posterior fusiform gyrus, giving support to the view that the cortical region, traditionally considered to be unisensory region for visual processing, functions as the locus of audiovisual integration of emotional signals. The Author 2013. Published by Oxford University Press. All rights reserved. For Permissions, please e-mail: journals.permissions@oup.com.

Driver J., Davis G., Ricciardelli P., Kidd P., Maxwell E., & Baron-Cohen S . ( 1999).

Gaze perception triggers reflexive visuospatial orienting

Visual Cognition, 6( 5), 509-540.

DOI:10.1080/135062899394920      URL     [本文引用: 1]

This paper seeks to bring together two previously separate research traditions: research on spatial orienting within the visual cueing paradigm and research into social cognition, addressing our tendency to attend in the direction that another person looks. Cueing methodologies from mainstream attention research were adapted to test the automaticity of orienting in the direction of seen gaze. Three studies manipulated the direction of gaze in a computerized face, which appeared centrally in a frontal view during a peripheral letter-discrimination task. Experiments 1 and 2 found faster discrimination of peripheral target letters on the side the computerized face gazed towards, even though the seen gaze did not predict target side, and despite participants being asked to ignore the face. This suggests reflexive covert and/or overt orienting in the direction of seen gaze, arising even when the observer has no motivation to orient in this way. Experiment 3 found faster letter discrimination on the side the computerized face gazed towards even when participants knew that target letters were four times as likely on the opposite side. This suggests that orienting can arise in the direction of seen gaze even when counter to intentions. The experiments illustrate that methods from mainstream attention research can be usefully applied to social cognition, and that studies of spatial attention may profit from considering its social function.

Ecker C., Suckling J., Deoni S. C., Lombardo M. V., Bullmore E. T., Baroncohen S., .. Murphy D. G . ( 2012).

Brain anatomy and its relationship to behavior in adults with autism spectrum disorder: A multicenter magnetic resonance imaging study

Archives of General Psychiatry, 69( 2), 195-209.

DOI:10.1001/archgenpsychiatry.2011.1251      URL     PMID:22310506      [本文引用: 1]

ContextThere is consensus that autism spectrum disorder (ASD) is accompanied by differences in neuroanatomy. However, the neural substrates of ASD during adulthood, as well as how these relate to behavioral variation, remain poorly understood.ObjectiveTo identify brain regions and systems associated with ASD in a large, well-characterized sample of adults.DesignMulticenter case-control design using quantitative magnetic resonance imaging.SettingMedical Research Council UK Autism Imaging Multicentre Study (MRC AIMS), with sites comprising the Institute of Psychiatry, Kings College London; the Autism Research Centre, University of Cambridge; and the Autism Research Group, University of Oxford.ParticipantsEighty-nine men with ASD and 89 male control participants who did not differ significantly in mean age (26 and 28 years, respectively) and full-scale IQ (110 and 113, respectively).Main Outcome Measures(1) Between-group differences in regional neuroanatomy assessed by voxel-based morphometry and (2) distributed neural systems maximally correlated with ASD, as identified by partial least-squares analysis.ResultsAdults with ASD did not differ significantly from the controls in overall brain volume, confirming the results of smaller studies of individuals in this age group without intellectual disability. However, voxelwise comparison between groups revealed that individuals with ASD had significantly increased gray matter volume in the anterior temporal and dorsolateral prefrontal regions and significant reductions in the occipital and medial parietal regions compared with controls. These regional differences in neuroanatomy were significantly correlated with the severity of specific autistic symptoms. The large-scale neuroanatomic networks maximally correlated with ASD identified by partial least-squares analysis included the regions identified by voxel-based analysis, as well as the cerebellum, basal ganglia, amygdala, inferior parietal lobe, cingulate cortex, and various medial, orbital, and lateral prefrontal regions. We also observed spatially distributed reductions in white matter volume in participants with ASD.ConclusionsAdults with ASD have distributed differences in brain anatomy and connectivity that are associated with specific autistic features and traits. These results are compatible with the concept of autism as a syndrome characterized by atypical neural onnectivity.

Elsabbagh M., Bruno R., Wan M. W., Charman T., Johnson M. H., Green J., & the BASIS Team ( 2015).

Infant neural sensitivity to dynamic eye gaze relates to quality of parent-infant interaction at 7-months in infants at risk for autism

Journal of Autism and Developmental Disorders, 45( 2), 283-291. doi: 10.1007/s10803-014-2192-9.

URL     PMID:4309912      [本文引用: 3]

Links between brain function measures and quality of parent–child interactions within the early developmental period have been investigated in typical and atypical development. We examined such links in a group of 104 infants with and without a family history for autism in the first year of life. Our findings suggest robust associations between event related potential responses to eye gaze and observed parent–infant interaction measures. In both groups, infants with more positive affect exhibit stronger differentiation to gaze stimuli. This association was observed with the earlier P100 waveform component in the control group but with the later P400 component in infants at-risk. These exploratory findings are critical in paving the way for a better understanding of how infant laboratory measures may relate to overt behavior and how both can be combined in the context of predicting risk or clinical diagnosis in toddlerhood.

Elsabbagh M., Mercure E., Hudry K., Chandler S., Pasco G., Charman T., … the BASIS Team ( 2012).

Infant neural sensitivity to dynamic eye gaze is associated with later emerging autism

Current Biology, 22( 4), 338-342. doi: 10.1016/j.cub.2011.12.056.

[本文引用: 2]

Elsabbagh M., Volein A., Csibra G., Holmboe K., Garwood H., Tucker L., … Johnson M. H . ( 2009).

Neural correlates of eye gaze processing in the infant broader autism phenotype

Biological Psychiatry, 65( 1), 31-38. doi: 10.1016/j.biopsych.2008.09.034.

URL     PMID:19064038      [本文引用: 2]

Studies of infant siblings of children diagnosed with autism have allowed for a prospective approach to study the emergence of autism in infancy and revealed early behavioral characteristics of the broader autism phenotype. In view of previous findings of atypical eye gaze processing in children and adults with autism, the aim of this study was to examine the early autism phenotype in infant siblings of children diagnosed with autism spectrum disorder (sib-ASD), focusing on the neural correlates of direct compared with averted gaze. A group of 19 sib-ASD was compared with 17 control infants with no family history of ASD (mean age = 10 months) on their response to direct versus averted gaze in static stimuli. Relative to the control group, the sib-ASD group showed prolonged latency of the occipital P400 event-related potentials component in response to direct gaze, but they did not differ in earlier components. Similarly, time-frequency analysis of high-frequency oscillatory activity in the gamma band showed group differences in response to direct gaze, where induced gamma activity was late and less persistent over the right temporal region in the sib-ASD group. This study suggests that a broader autism phenotype, which includes an atypical response to direct gaze, is manifest early in infancy.

Falck-Ytter T., Fernell E., Hedvall Å., von Hofsten C., & Gillberg C . ( 2012).

Gaze performance in children with autism spectrum disorder when observing communicative actions

Journal of Autism and Developmental Disorders, 42( 10), 2236-2245. doi: 10.1007/s10803-012-1471-6.

URL     PMID:22354708      [本文引用: 2]

The main purpose of this eye tracking study was to map the correlates of gaze performance in a brief test of spontaneous gaze and point-gesture following in young children with autistic disorder (AD), Pervasive developmental disorder—not otherwise specified (PDD-NOS), or typical development (TD). Gaze measures included the children’s spontaneous tendency to look at the correct (attended) toy, and the latency of their correct responses. In addition to group differences (AD vs. TD), we found that in AD, accuracy of performance was specifically related to adaptive communication skills. The study also indicated that the latency of correct gaze shifts is related to verbal intelligence. These results have direct implications for our understanding of (responsive) joint attention impairments in AD.

Falck-Ytter T., Thorup E., & Bölte S . ( 2015).

Brief report: Lack of processing bias for the objects other people attend to in 3-year-olds with autism

Journal of Autism and Developmental Disorders, 45( 6), 1897-1904. doi: 10.1007/s10803-014-2278-4.

URL     PMID:4441907      [本文引用: 2]

Whether gaze following—a key component of joint attention—is impaired in children with autism spectrum disorder (ASD) is currently debated. Functional gaze following involves saccading towards the attended rather than unattended targets (accuracy) as well as a subsequent processing bias for attended objects. Using non-invasive eye tracking technology, we show that gaze following accuracy is intact in intellectually low-functioning 3-year-olds with ASD. However, analyses of the duration of first fixations at the objects in the scene revealed markedly weaker initial processing bias for attended objects in children with ASD compared to children with typical development and non-autistic children with developmental delays. Limited processing bias for the objects other people attend to may negatively affect learning opportunities in ASD.

Farroni T., Menon E., & Johnson M. H . ( 2006).

Factors influencing newborns’ preference for faces with eye contact

Journal of Experimental Child Psychology, 95( 4), 298-308.

DOI:10.1016/j.jecp.2006.08.001      URL     PMID:17030037      [本文引用: 1]

We investigated newborns’ sensitivity to the direction of gaze of another’s face by using a preferential looking technique. This study extends earlier work on a preference for faces with direct gaze in newborns. In Experiment 1, we replicate the basic finding of Farroni and colleagues that newborns prefer to look at faces with direct gaze. In Experiments 2 and 3, we establish that a preference for faces with direct gaze in newborns is present only within the context of an upright face and a straight head, suggesting that relatively primitive configuration-sensitive mechanisms may be operating. Overall, these results further the view that relatively simple perceptual biases in newborns may be an essential foundation for later social-cognitive development.

Fletcher-Watson S., Leekam S. R., Benson V., Frank M. C., & Findlay J. M . ( 2009).

Eye-movements reveal attention to social information in autism spectrum disorder

Neuropsychologia, 47( 1), 248-257.

DOI:10.1016/j.neuropsychologia.2008.07.016      URL     PMID:18706434      [本文引用: 1]

Autism spectrum disorder (ASD) is a neurodevelopmental condition in which children show reduced attention to social aspects of the environment. However in adults with ASD, evidence for social attentional deficits is equivocal. One problem is that many paradigms present social information in an unrealistic, isolated way. This study presented adults and adolescents, with and without ASD, with a complex social scene alongside another, non-social scene, and measured eye-movements during a 3-s viewing period. Analyses first identified viewing time to different regions and then investigated some more complex issues. These were: the location of the very first fixation in a trial (indicating attentional priority); the effect of a task instruction on scan paths; the extent to which gaze-following was evident; and the degree to which participants scan paths were influenced by the low-level properties of a scene. Results indicate a superficially normal attentional preference for social information in adults with ASD. However, more sensitive measures show that ASD does entail social attention problems across the lifespan, supporting accounts of the disorder which emphasise lifelong neurodevelopmental atypicalities. These subtle abnormalities may be sufficient to produce serious difficulties in real-life scenarios.

Freeth M., Chapman P., Ropar D., & Mitchell P . ( 2010).

Do gaze cues in complex scenes capture and direct the attention of high functioning adolescents with ASD? Evidence from eye-tracking

Journal of Autism and Developmental Disorders, 40( 5), 534-547.

DOI:10.1007/s10803-009-0893-2      URL     PMID:19904597      [本文引用: 1]

Visual fixation patterns whilst viewing complex photographic scenes containing one person were studied in 24 high-functioning adolescents with Autism Spectrum Disorders (ASD) and 24 matched typically developing adolescents. Over two different scene presentation durations both groups spent a large, strikingly similar proportion of their viewing time fixating the person face. However, time-course analyses revealed differences between groups in priorities of attention to the region of the face containing the eyes. It was also noted that although individuals with ASD were rapidly cued by the gaze direction of the person in the scene, this was not followed by an immediate increase in total fixation duration at the location of gaze, which was the case for typically developing individuals.

Freeth M., Ropar D., Mitchell P., Chapman P., & Loher S . ( 2011).

Brief report: How adolescents with ASD process social information in complex scenes. Combining evidence from eye movements and verbal descriptions

Journal of Autism and Developmental Disorders, 41( 3), 364-371. doi: 10.1007/s10803-010-1053-4.

URL     PMID:20559702      [本文引用: 1]

We investigated attention, encoding and processing of social aspects of complex photographic scenes. Twenty-four high-functioning adolescents (aged 11–16) with ASD and 24 typically developing matched control participants viewed and then described a series of scenes, each containing a person. Analyses of eye movements and verbal descriptions provided converging evidence that both groups displayed general interest in the person in each scene but the salience of the person was reduced for the ASD participants. Nevertheless, the verbal descriptions revealed that participants with ASD frequently processed the observed person’s emotion or mental state without prompting. They also often mentioned eye-gaze direction, and there was evidence from eye movements and verbal descriptions that gaze was followed accurately. The combination of evidence from eye movements and verbal descriptions provides a rich insight into the way stimuli are processed overall. The merits of using these methods within the same paradigm are discussed.

Gainotti G. ( 2012).

Unconscious processing of emotions and the right hemisphere

Neuropsychologia, 50( 2), 205-218.

DOI:10.1016/j.neuropsychologia.2011.12.005      URL     PMID:22197572      [本文引用: 1]

This survey takes into account the unconscious aspects of emotions and the critical role played in them by the right hemisphere, considering different acceptations of the term ‘unconscious’. In a preliminary step, the nature of emotions, their componential and hierarchical organization and the relationships between emotions and hemispheric specialization are shortly discussed, then different aspects of emotions are surveyed: first are reviewed studies dealing with the unconscious processing of emotional information, taking separately into account various lines of research. All these studies suggest that unconscious processing of emotional information is mainly subsumed by a right hemisphere subcortical route, through which emotional stimuli quickly reach the amygdala. We afterwards inquire if a right hemisphere dominance can also be observed in automatic emotional action schemata and if ‘non-removed preverbal implicit memories’ also have a preferential link with the right hemisphere. Finally, we try to evaluate if the right hemisphere may also play a critical role in dynamic unconscious phenomena, such as anosognosia/denial of hemiplegia in patients with unilateral brain lesions. In the last part of the review, the reasons that could subsume the right hemisphere dominance for unconscious emotions are shortly discussed.

George N., Driver J., & Dolan R. J . ( 2001).

Seen gaze-direction modulates fusiform activity and its coupling with other brain areas during face processing

Neuroimage, 13( 1), 1102-1112.

DOI:10.1006/nimg.2001.0769      URL     PMID:11352615      [本文引用: 2]

Gaze-contact is often a preliminary to social interaction and so constitutes a signal for the allocation of processing resources to the gazing face. We investigated how gaze direction influences face processing in an fMRI study, where seen gaze and head direction could independently be direct or deviated. Direct relative to averted gaze elicited stronger activation for faces in ventral occipitotemporal cortices around the fusiform gyrus, regardless of head orientation. Moreover, direct gaze led to greater correlation between activity in the fusiform and the amygdala, a region associated with emotional responses and stimulus saliency. By contrast, faces with averted gaze (again, regardless of head orientation) yielded increased correlation between activity in the fusiform and the intraparietal sulcus, a region associated with shifting attention to the periphery.

Georgescu A. L., Kuzmanovic B., Schilbach L., Tepest R., Kulbida R., Bente G., & Vogeley K . ( 2013).

Neural correlates of “social gaze” processing in high-functioning autism under systematic variation of gaze duration

NeuroImage: Clinical, 3, 340-351.

DOI:10.1016/j.nicl.2013.08.014      URL     PMID:24273718      [本文引用: 1]

61We investigate the neural processing of gaze direction and duration in HFA.61We use a combined categorical and parametric approach to analyze fMRI data.61The social neural network is not modulated by direct gaze cues in HFA.61Persons with HFA are impaired in using subtle aspects of gaze to understand others.

Gillespie-Lynch K., Elias R., Escudero P., Hutman T., & Johnson S. P . ( 2013).

Atypical gaze following in autism: A comparison of three potential mechanisms

Journal of Autism and Developmental Disorders, 43( 12), 2779-2792. doi: 10.1007/s10803-013-1818-7.

URL     PMID:23619947      [本文引用: 3]

In order to evaluate the following potential mechanisms underlying atypical gaze following in autism, impaired reflexive gaze following, difficulty integrating gaze and affect, or reduced understanding of the referential significance of gaze, we administered three paradigms to young children with autism (N = 21) and chronological (N = 21) and nonverbal mental age (N = 21) matched controls. Children with autism exhibited impaired reflexive gaze following. The absence of evidence of integration of gaze and affect, regardless of diagnosis, indicates ineffective measurement of this construct. Reduced gaze following was apparent among children with autism during eye-tracking and in-person assessments. Word learning from gaze cues was better explained by developmental level than autism. Thus, gaze following may traverse an atypical, rather than just delayed, trajectory in autism.

Gliga T., Elsabbagh M., Hudry K., Charman T., & Johnson M. H . ( 2012).

Gaze following, gaze reading, and word learning in children at risk for autism

Child Development, 83( 3), 926-938.

DOI:10.1111/j.1467-8624.2012.01750.x      URL     PMID:22462503      [本文引用: 2]

This study investigated gaze-following abilities as a prerequisite for word learning, in a population expected to manifest a wide range of social and communicative skills—children with a family history of autism. Fifty-three 3-year-olds with or without a family history of autism took part in a televised word-learning task. Using an eye-tracker to monitor children’s gaze behavior, it was shown that the ability to follow gaze was necessary but not sufficient for successful word learning. Those children who had poor social and communicative skills followed gaze to the labeled object but did not then learn the associated word. These findings shed light on the conditions that lead to successful word learning in typical and atypical populations.

Goldberg M. C., Mostow A. J., Vecera S. P., Larson J. G., Mostofsky S. H., Mahone E. M., & Denckla M. B . ( 2008).

Evidence for impairments in using static line drawings of eye gaze cues to orient visual-spatial attention in children with high functioning autism

Journal of Autism and Developmental Disorders, 38( 8), 1405-1413. doi: 10.1007/s10803-007-0506-x.

URL     PMID:18074212      [本文引用: 1]

We examined the ability to use static line drawings of eye gaze cues to orient visual-spatial attention in children with high functioning autism (HFA) compared to typically developing children (TD). The task was organized such that on valid trials , gaze cues were directed toward the same spatial location as the appearance of an upcoming target, while on invalid trials gaze cues were directed to an opposite location. Unlike TD children, children with HFA showed no advantage in reaction time (RT) on valid trials compared to invalid trials (i.e., no significant validity effect). The two stimulus onset asynchronies (200 ms, 700 ms) did not differentially affect these findings. The results suggest that children with HFA show impairments in utilizing static line drawings of gaze cues to orient visual-spatial attention.

Gotts S. J., Simmons W. K., Milbury L. A., Wallace G. L., Cox R. W., & Martin A . ( 2012).

Fractionation of social brain circuits in autism spectrum disorders

Brain, 135( 9), 2711-2725.

[本文引用: 1]

Greene D. J., Colich N., Iacoboni M., Zaidel E., Bookheimer S. Y., & Dapretto M . ( 2011).

Atypical neural networks for social orienting in autism spectrum disorders

NeuroImage, 56( 1), 354-362.

DOI:10.1016/j.neuroimage.2011.02.031      URL     PMID:21334443      [本文引用: 2]

78Greater activation for social vs. nonsocial orienting in typical development than ASD. 78No differences in behavioral performance in typical development and ASD. 78Group X cue condition interactions confirm group differences in brain activity.

Grice S. J., Halit H., Farroni T., Baron-Cohen S., Bolton P., & Johnson M. H . ( 2005).

Neural correlates of eye-gaze detection in young children with autism

Cortex, 41( 3), 342-353. doi: 10.1016/S0010-9452(08)70271-5.

URL     PMID:15871599      [本文引用: 1]

Various reports have demonstrated difficulties in eye-gaze processing in older children and adults with autism. However, little is known about the neural or developmental origin of such difficulties. In the present study, we used high-density Event-Related Potentials (HD-ERPs) to record the neural correlates of gaze processing in young children with autism, and their age-matched controls. In addition, to determine normal gaze processing development we also tested a non-autism adult group. The data obtained from the children with autism resembled that previously observed in typical 4-month old infants. In contrast, the control group showed the same pattern as typical adults. These findings suggest that the neural correlates of gaze direction processing may be delayed in young children with autism.

Grossmann T., & Johnson M.H . ( 2007).

The development of the social brain in human infancy

European Journal of Neuroscience, 25( 4), 909-919.

DOI:10.1111/j.1460-9568.2007.05379.x      URL     PMID:17331189      [本文引用: 4]

Much research has focused on how the adult human brain processes the social world, yet until recently little was known about the early development of these abilities. Developmental studies inform debates about the specificity of social functions in the adult cortex. This review highlights recent work, mainly based on electroencephalography/event-related potential methods, examining the precursors of the human social brain network during infancy in several domains such as face and eye gaze processing, the perception of emotions, decoding biological motion, perceiving human actions and joint attention. The findings illustrate that the human brain is fundamentally adapted to develop within a social context, and that this context contributes to many of the specializations seen in the adult cortex.

Guillon Q., Rogé B., Afzali M. H., Baduel S., Kruck J., & Hadjikhani N . ( 2016).

Intact perception but abnormal orientation towards face-like objects in young children with ASD

Scientific Reports, 6, 22119.

DOI:10.1038/srep22119      URL     PMID:26912096      [本文引用: 2]

There is ample behavioral evidence of diminished orientation towards faces as well as the presence of face perception impairments in autism spectrum disorder (ASD), but the underlying mechanisms of these deficits are still unclear. We used face-like object stimuli that have been shown to evoke pareidolia in typically developing (TD) individuals to test the effect of a global face-like configuration on orientation and perceptual processes in young children with ASD and age-matched TD controls. We show that TD children were more likely to look first towards upright face-like objects than children with ASD, showing that a global face-like configuration elicit a stronger orientation bias in TD children as compared to children with ASD. However, once they were looking at the stimuli, both groups spent more time exploring the upright face-like object, suggesting that they both perceived it as a face. Our results are in agreement with abnormal social orienting in ASD, possibly due to an abnormal tuning of the subcortical pathway, leading to poor orienting and attention towards faces. Our results also indicate that young children with ASD can perceive a generic face holistically, such as face-like objects, further demonstrating holistic processing of faces in ASD.

Hadjikhani N., Zurcher N. R., Lassalle A., Hippolyte L., Ward N., & Johnels J. Å . ( 2017).

The effect of constraining eye-contact during dynamic emotional face perception—An fMRI study

Social Cognitive and Affective Neuroscience, 12( 7), 1197-1207.

DOI:10.1093/scan/nsx046      URL     PMID:28402536      [本文引用: 1]

Eye-contact modifies how we perceive emotions and modulates activity in the social brain network. Here, using fMRI, we demonstrate that adding a fixation cross in the eye region of dynamic facial emotional stimuli significantly increases activation in the social brain of healthy, neurotypical participants when compared with activation for the exact same stimuli observed in a free-viewing mode. In addition, using PPI analysis, we show that the degree of amygdala connectivity with the rest of the brain is enhanced for the constrained view for all emotions tested except for fear, and that anxiety and alexithymia modulate the strength of amygdala connectivity for each emotion differently. Finally, we show that autistic traits have opposite effects on amygdala connectivity for fearful and angry emotional expressions, suggesting that these emotions should be treated separately in studies investigating facial emotion processing.

Hazlett H. C., Gu H., Munsell B. C., Kim S. H., Styner M., Wolff J. J., … Piven J . ( 2017).

Early brain development in infants at high risk for autism spectrum disorder

Nature, 542( 7641), 348-351.

DOI:10.1038/nature21369      URL     PMID:5336143      [本文引用: 2]

Brain enlargement has been observed in children with Autism Spectrum Disorder (ASD), but the timing of this phenomenon and its relationship to the appearance of behavioral symptoms is unknown. Retrospective head circumference and longitudinal brain volume studies of 2 year olds followed up at age 4 years, have provided evidence that increased brain volume may emerge early in development.1,2Studies of infants at high familial risk for autism can provide insight into the early development of autism and have found that characteristic social deficits in ASD emerge during the latter part of the first and in the second year of life3,4. These observations suggest that prospective brain imaging studies of infants at high familial risk for ASD might identify early post-natal changes in brain volume occurring before the emergence of an ASD diagnosis. In this prospective neuroimaging study of 106 infants at high familial risk of ASD and 42 low-risk infants, we show that cortical surface area hyper-expansion between 6-12 months of age precedes brain volume overgrowth observed between 12-24 months in the 15 high-risk infants diagnosed with autism at 24 months. Brain volume overgrowth was linked to the emergence and severity of autistic social deficits. A deep learning algorithm primarily using surface area information from brain MRI at 6 and 12 months of age predicted the diagnosis of autism in individual high-risk children at 24 months (with a positive predictive value of 81%, sensitivity of 88%). These findings demonstrate that early brain changes unfold during the period in which autistic behaviors are first emerging.

Helminen T. M., Leppänen J. M., Eriksson K., Luoma A., Hietanen J. K., & Kylliäinen A . ( 2017).

Atypical physiological orienting to direct gaze in low‐functioning children with autism spectrum disorder

Autism Research, 10( 5), 810-820.

DOI:10.1002/aur.1738      URL     [本文引用: 2]

Reduced use of eye contact is a prominent feature in individuals with autism spectrum disorder (ASD). It has been proposed that direct gaze does not capture the attention of individuals with ASD. Experimental evidence is, however, mainly restricted to relatively high-functioning school-aged children or adults with ASD. This study investigated whether 2–5-year-old low-functioning children with severe ASD differ from control children in orienting to gaze stimuli, as measured with the heart rate deceleration response. Responses were measured to computerized presentations of dynamic shifts of gaze direction either toward (direct) or away (averted) from the observing child. The results showed a significant group by gaze direction interaction effect on heart rate responses (permuted P65=65.004), reflecting a stronger orienting response to direct versus averted gaze in typically developing (N65=6517) and developmentally delayed (N65=6516) children but not in children with ASD (N65=6512). The lack of enhanced orienting response to direct gaze in the ASD group was not caused by a lack of looking at the eye region, as confirmed by eye tracking. The results suggest that direct gaze is not a socially salient, attention-grabbing signal for low-functioning children with ASD. Autism Res 2017, 10: 810–820.

Hood B. M., Macrae C. N., Cole-Davies V., & Dias M . ( 2003).

Eye remember you: The effects of gaze direction on face recognition in children and adults

Developmental Science, 6( 1), 67-71.

DOI:10.1111/1467-7687.00256      URL    

Abstract Children and adults were tested on a forced-choice face recognition task in which the direction of eye gaze was manipulated over the course of the initial presentation and subsequent test phase of the experiment. To establish the effects of gaze direction on the encoding process, participants were presented with to-be-studied faces displaying either direct or deviated gaze (i.e. encoding manipulation). At test, all the faces depicted persons with their eyes closed. To investigate the effects of gaze direction on the efficiency of the retrieval process, a second condition (i.e. retrieval manipulation) was run in which target faces were presented initially with eyes closed and tested with either direct or deviated gaze. The results revealed the encoding advantages enjoyed by faces with direct gaze was present for both children and adults. Faces with direct gaze were also recognized better than faces with deviated gaze at retrieval, although this effect was most pronounced for adults. Finally, the advantage for direct gaze over deviated gaze at encoding was greater than the advantage for direct gaze over deviated gaze at retrieval. We consider the theoretical implications of these findings.

Itier R.J., & Batty M. ,( 2009).

Neural bases of eye and gaze processing: The core of social cognition

Neuroscience & Biobehavioral Reviews, 33( 6), 843-863. doi: 10.1016/ j.neubiorev.2009.02.004.

URL     PMID:19428496      [本文引用: 1]

Eyes and gaze are very important stimuli for human social interactions. Recent studies suggest that impairments in recognizing face identity, facial emotions or in inferring attention and intentions of others could be linked to difficulties in extracting the relevant information from the eye region including gaze direction. In this review, we address the central role of eyes and gaze in social cognition. We start with behavioral data demonstrating the importance of the eye region and the impact of gaze on the most significant aspects of face processing. We review neuropsychological cases and data from various imaging techniques such as fMRI/PET and ERP/MEG, in an attempt to best describe the spatio-temporal networks underlying these processes. The existence of a neuronal eye detector mechanism is discussed as well as the links between eye gaze and social cognition impairments in autism. We suggest impairments in processing eyes and gaze may represent a core deficiency in several other brain pathologies and may be central to abnormal social cognition.

Johnson M. H., Senju A., & Tomalski P . ( 2015).

The two-process theory of face processing: Modifications based on two decades of data from infants and adults

Neuroscience & Biobehavioral Reviews, 50, 169-179. doi: 10.1016/j.neubiorev.2014.10.009.

URL     PMID:25454353      [本文引用: 3]

Johnson and Morton (1991. Biology and Cognitive Development: The Case of Face Recognition. Blackwell, Oxford) used Gabriel Horn's work on the filial imprinting model to inspire a two-process theory of the development of face processing in humans. In this paper we review evidence accrued over the past two decades from infants and adults, and from other primates, that informs this two-process model. While work with newborns and infants has been broadly consistent with predictions from the model, further refinements and questions have been raised. With regard to adults, we discuss more recent evidence on the extension of the model to eye contact detection, and to subcortical face processing, reviewing functional imaging and patient studies. We conclude with discussion of outstanding caveats and future directions of research in this field.

Kirchgessner M. A., Chuang A. Z., Patel S. S., & Sereno A. B . ( 2015).

Intact reflexive but deficient voluntary social orienting in autism spectrum disorder

Frontiers in Neuroscience, 9, 453. doi: 10.3389/fnins.2015.00453.

URL     PMID:4665260      [本文引用: 1]

Impairment in social interactions is a primary characteristic of people diagnosed with autism spectrum disorder (ASD). Although these individuals tend to orient less to naturalistic social cues than do typically developing (TD) individuals, laboratory experiments testing social orienting in ASD have been inconclusive, possibly because of a failure to fully isolate reflexive (stimulus-driven) and voluntary (goal-directed) social orienting processes. The purpose of the present study was to separately examine potential reflexive and/or voluntary social orienting differences in individuals with ASD relative to TD controls. Subjects (ages 7 14) with high-functioning ASD and a matched control group completed three gaze cueing tasks on an iPad in which individuals briefly saw a face with averted gaze followed by a target after a variable delay. Two tasks were 100% predictive with either all congruent (target appears in gaze direction) or all incongruent (target appears opposite from gaze direction) trials, respectively. Another task was non-predictive with these same trials (half congruent and half incongruent) intermixed randomly. Response times (RTs) to the target were used to calculate reflexive (incongruent condition RT ongruent condition RT) and voluntary (non-predictive condition RT redictive condition RT) gaze cueing effects. Subjects also completed two additional non-social orienting tasks (ProPoint and AntiPoint). Subjects with ASD demonstrate intact reflexive but deficient voluntary gaze following. Similar results were found in a separate test of non-social orienting. This suggests problems with using social cues, but only in a goal-directed fashion, in our sample of high-functioning individuals with ASD. Such findings may not only explain inconclusive previous findings but more importantly be critical for understanding social dysfunctions in ASD and for developing future interventions.

Kleinhans N. M., Richards T., Johnson L. C., Weaver K. E., Greenson J., Dawson G., & Aylward E . ( 2011).

FMRI evidence of neural abnormalities in the subcortical face processing system in ASD

NeuroImage, 54( 1), 697-704.

DOI:10.1016/j.neuroimage.2010.07.037      URL     PMID:3426450      [本文引用: 2]

Recent evidence suggests that a rapid, automatic face detection system is supported by subcortical structures including the amygdala, pulvinar, and superior colliculus. Early-emerging abnormalities in these structures may be related to reduced social orienting in children with autism, and subsequently, to aberrant development of cortical circuits involved in face processing. Our objective was to determine whether functional abnormalities in the subcortical face processing system are present in adults with autism spectrum disorders (ASD) during supraliminal fearful face processing. Participants included twenty-eight individuals with ASD and 25 controls group-matched on age, IQ, and behavioral performance. The ASD group met diagnostic criteria on the ADI-R, ADOS-G, and DSM-IV. Both the ASD and control groups showed significant activation in bilateral fusiform gyri. The control group exhibited additional significant responses in the right amygdala, right pulvinar, and bilateral superior colliculi. In the direct group comparison, the controls showed significantly greater activation in the left amygdala, bilateral fusiform gyrus, right pulvinar, and bilateral superior colliculi. No brain region showed significantly greater activation in the ASD group compared to the controls. Thus, basic rapid face identification mechanisms appear to be functional in ASD. However, individuals with ASD failed to engage the subcortical brain regions involved in face detection and automatic emotional face processing, suggesting a core mechanism for impaired socioemotional processing in ASD. Neural abnormalities in this system may contribute to early-emerging deficits in social orienting and attention, the putative precursors to abnormalities in social cognition and cortical face processing specialization.Research Highlights?Rapid face detection is supported by the amygdala, pulvinar, and superior colliculus. ?Autism involves reduced activation in these subcortical face processing regions. ?Social orienting deficits may stem from neural abnormalities in subcortical structures. ?Social orienting deficits precede abnormalities in social cognition in autism.

Klucharev V. & Sams M., ( 2004).

Interaction of gaze direction and facial expressions processing: ERP study

Neuroreport, 15( 4), 621-625.

[本文引用: 1]

Kuhn G., Benson V., Fletcher-Watson S., Kovshoff H., McCormick C. A., Kirkby J., & Leekam S. R . ( 2010).

Eye movements affirm: Automatic overt gaze and arrow cueing for typical adults and adults with autism spectrum disorder

Experimental Brain Research, 201( 2), 155-165. doi: 10.1007/s00221-009-2019-7.

URL     PMID:19798489      [本文引用: 2]

People with autism spectrum disorder (ASD) show reduced interest towards social aspects of the environment and a lesser tendency to follow other people gaze in the real world. However, most studies have shown that people with ASD do respond to eye-gaze cues in experimental paradigms, though it is possible that this behaviour is based on an atypical strategy. We tested this possibility in adults with ASD using a cueing task combined with eye-movement recording. Both eye gaze and arrow pointing distractors resulted in overt cueing effects, both in terms of increased saccadic reaction times, and in proportions of saccades executed to the cued direction instead of to the target, for both participant groups. Our results confirm previous reports that eye gaze cues as well as arrow cues result in automatic orienting of overt attention. Moreover, since there were no group differences between arrow and eye gaze cues, we conclude that overt attentional orienting in ASD, at least in response to centrally presented schematic directional distractors, is typical.

Kylliäinen A., Wallace S., Coutanche M. N., Leppänen J. M., Cusack J., Bailey A. J., & Hietanen J. K . ( 2012).

Affective-motivational brain responses to direct gaze in children with autism spectrum disorder

Journal of Child Psychology and Psychiatry, 53( 7), 790-797.

DOI:10.1111/j.1469-7610.2011.02522.x      URL     PMID:22276654      [本文引用: 1]

Background: It is unclear why children with autism spectrum disorders (ASD) tend to be inattentive to, or even avoid eye contact. The goal of this study was to investigate affective otivational brain responses to direct gaze in children with ASD. To this end, we combined two measurements: skin conductance responses (SCR), a robust arousal measure, and asymmetry in frontal electroencephalography (EEG) activity which is associated with motivational approach and avoidance tendencies. We also explored whether degree of eye openness and face familiarity modulated these responses.Methods: Skin conductance responses and frontal EEG activity were recorded from 14 children with ASD and 15 typically developing children whilst they looked at familiar and unfamiliar faces with eyes shut, normally open or wide-open. Stimuli were presented in such a way that they appeared to be looming towards the children.Results: In typically developing children, there were no significant differences in SCRs between the different eye conditions, whereas in the ASD group the SCRs were attenuated to faces with closed eyes and increased as a function of the degree of eye openness. In both groups, familiar faces elicited marginally greater SCRs than unfamiliar faces. In typically developing children, normally open eyes elicited greater relative left-sided frontal EEG activity (associated with motivational approach) than shut eyes and wide-open eyes. In the ASD group, there were no significant differences between the gaze conditions in frontal EEG activity.Conclusions: Collectively, the results replicate previous finding in showing atypical modulation of arousal in response to direct gaze in children with ASD but do not support the assumption that this response is associated with an avoidant motivational tendency. Instead, children with ASD may lack normative approach-related motivational response to eye contact.

Lajiness-O Neill R., Richard A. E., Moran J. E., Olszewski A., Pawluk L., Jacobson D., .. Bowyer S. M . ( 2014).

Neural synchrony examined with magnetoencephalography (MEG) during eye gaze processing in autism spectrum disorders: Preliminary findings

Journal of Neurodevelopmental Disorders, 6, 15 doi: 10.1186/1866-1955-6-15.

URL     PMID:40728455      [本文引用: 1]

Background Gaze processing deficits are a seminal, early, and enduring behavioral deficit in autism spectrum disorder (ASD); however, a comprehensive characterization of the neural processes mediating abnormal gaze processing in ASD has yet to be conducted. Methods This study investigated whole-brain patterns of neural synchrony during passive viewing of direct and averted eye gaze in ASD adolescents and young adults (MAge???=???16.6) compared to neurotypicals (NT) (MAge???=???17.5) while undergoing magnetoencephalography. Coherence between each pair of 54 brain regions within each of three frequency bands (low frequency (0 to 15 Hz), beta (15 to 30 Hz), and low gamma (30 to 45 Hz)) was calculated. Results Significantly higher coherence and synchronization in posterior brain regions (temporo-parietal-occipital) across all frequencies was evident in ASD, particularly within the low 0 to 15 Hz frequency range. Higher coherence in fronto-temporo-parietal regions was noted in NT. A significantly higher number of low frequency cross-hemispheric synchronous connections and a near absence of right intra-hemispheric coherence in the beta frequency band were noted in ASD. Significantly higher low frequency coherent activity in bilateral temporo-parieto-occipital cortical regions and higher gamma band coherence in right temporo-parieto-occipital brain regions during averted gaze was related to more severe symptomology as reported on the Autism Diagnostic Interview-Revised (ADI-R). Conclusions The preliminary results suggest a pattern of aberrant connectivity that includes higher low frequency synchronization in posterior cortical regions, lack of long-range right hemispheric beta and gamma coherence, and decreased coherence in fronto-temporo-parietal regions necessary for orienting to shifts in eye gaze in ASD; a critical behavior essential for social communication.

Lassalle A., & Itier R.J . ( 2014).

Autistic traits influence gaze-oriented attention to happy but not fearful faces

Social Neuroscience, 10( 1), 70-88. doi: 10.1080/ 17470919.2014.958616.

URL     PMID:25222883      [本文引用: 1]

The relationship between autistic traits and gaze-oriented attention to fearful and happy faces was investigated at the behavioral and neuronal levels. Upright and inverted dynamic face stimuli were used in a gaze-cueing paradigm while event related potentials (ERPs) were recorded. Participants responded faster to gazed-at than to non-gazed-at targets, and this gaze orienting effect (GOE) diminished with inversion, suggesting it relies on facial configuration. It was also larger for fearful than happy faces but only in participants with high autism-spectrum quotient (AQ) scores. While the GOE to fearful faces was of similar magnitude regardless of AQ scores, a diminished GOE to happy faces was found in participants with high AQ scores. At the ERP level, a congruency effect on target-elicited P1 component reflected enhanced visual processing of gazed-at targets. In addition, cue-triggered early directing attention negativity and anterior directing attention negativity reflected, respectively, attention orienting and attention holding at gazed-at locations. These neural markers of spatial attention orienting were not modulated by emotion and were not found in participants with high AQ scores. Together, these findings suggest that autistic traits influence attention orienting to gaze and its modulation by social emotions such as happiness.

Leekam S. R., LóPez B., & Moore C . ( 2000).

Attention and joint attention in preschool children with autism

Developmental Psychology, 36( 2), 261-273.

DOI:10.1037/0012-1649.36.2.261      URL     PMID:10749083      [本文引用: 2]

Three experiments examined the role of attention in explaining dyadic (child-adult) and triadic (child-adult-object) joint attention difficulties in autism. Experiments 1 and 2 investigated children's ability to orient to an adult's attention bid and to follow the direction of a human or nonhuman cue. Experiment 3 tested ability to disengage and shift attention to objects. Results showed autism-specific difficulties at both dyadic and triadic levels. Children with autism were less responsive than developmentally delayed controls in orienting to attention bids and in following a human head-turn cue yet had no difficulty in shifting attention and were faster overall in orienting to targets. Results suggest a specific developmental delay in which children with autism rely on the presence of objects in the visual field to guide action. The relation between this problem and autistic children's difficulties with human communicative signals is discussed.

Lloyd Fox S., Blasi A., Pasco G., Gliga T., Jones E. J. H., Murphy D. G. M., .. Johnson M. H . ( 2017).

Cortical responses before 6 months of life associate with later autism

European Journal of Neuroscience, 47( 6), 736-749.

DOI:10.1111/ejn.13757      URL     PMID:29057543      [本文引用: 2]

Autism spectrum disorder (ASD) is a common, highly heritable, developmental disorder and later‐born siblings of diagnosed children are at higher risk of developingASDthan the general population. Although the emergence of behavioural symptoms ofASDin toddlerhood is well characterized, far less is known about development during the first months of life of infants at familial risk. In a prospective longitudinal study of infants at familial risk followed to 3602months, we measured functional near‐infrared spectroscopy (fNIRS) brain responses to social videos of people (i.e. peek‐a‐boo) compared to non‐social images (vehicles) and human vocalizations compared to non‐vocal sounds. At 4–602months, infants who went on to developASDat 302years (N=025) evidenced‐reduced activation to visual social stimuli relative to low‐risk infants (N=0216) across inferior frontal (IFG) and posterior temporal (pSTS‐TPJ) regions of the cortex. Furthermore, these infants also showed reduced activation to vocal sounds and enhanced activation to non‐vocal sounds within left lateralized temporal (aMTG‐STG/pSTS‐TPJ) regions compared with low‐risk infants and high‐risk infants who did not developASD(N=0215). The degree of activation to both the visual and auditory stimuli correlated with parent‐reportedASDsymptomology in toddlerhood. These preliminary findings are consistent with later atypical social brain responses seen in children and adults withASD, and highlight the need for further work interrogating atypical processing in early infancy and how it may relate to later social interaction and communication difficulties characteristic of ASD.

Macrae C. N., Hood B. M., Milne A. B., Rowe A. C., & Mason M. F . ( 2002).

Are you looking at me? Eye gaze and person perception

Psychological Science, 13( 5), 460-464.

DOI:10.1111/1467-9280.00481      URL     PMID:12219814     

Previous research has highlighted the pivotal role played by gaze detection and interpretation in the development of social cognition. Extending work of this kind, the present research investigated the effects of eye gaze on basic aspects of the person090009perception process, namely, person construal and the extraction of category090009related knowledge from semantic memory. It was anticipated that gaze direction would moderate the efficiency of the mental operations through which these social090009cognitive products are generated. Specifically, eye gaze was expected to influence both the speed with which targets could be categorized as men and women and the rate at which associated stereotypic material could be accessed from semantic memory. The results of two experiments supported these predictions: Targets with nondeviated (i.e., direct) eye gaze elicited facilitated categorical responses. The implications of these findings for recent treatments of person perception are considered.

Madipakkam A. R., Rothkirch M., Dziobek I., & Sterzer P . ( 2017).

Unconscious avoidance of eye contact in autism spectrum disorder

Scientific Reports, 7( 1), 13378.

DOI:10.1038/s41598-017-13945-5      URL     PMID:29042641      [本文引用: 2]

Abstract Atypical responses to direct gaze are one of the most characteristic hallmarks of autism spectrum disorder (ASD). The cause and mechanism underlying this phenomenon, however, have remained unknown. Here we investigated whether the atypical responses to eye gaze in autism spectrum disorder is dependent on the conscious perception of others' faces. Face stimuli with direct and averted gaze were rendered invisible by interocular suppression and eye movements were recorded from participants with ASD and an age and sex matched control group. Despite complete unawareness of the stimuli, the two groups differed significantly in their eye movements to the face stimuli. In contrast to the significant positive saccadic index observed in the TD group, indicating an unconscious preference to the face with direct gaze, the ASD group had no such preference towards direct gaze and instead showed a tendency to prefer the face with averted gaze, suggesting an unconscious avoidance of eye contact. These results provide the first evidence that the atypical response to eye contact in ASD is an unconscious and involuntary response. They provide a better understanding of the mechanism of gaze avoidance in autism and might lead to new diagnostic and therapeutic interventions.

Malekmohammadi M., Elias W. J., & Pouratian N . ( 2015).

Human thalamus regulates cortical activity via spatially specific and structurally constrained Phase-Amplitude coupling

Cerebral Cortex, 25( 6), 1618-1628.

DOI:10.1093/cercor/bht358      URL     PMID:24408958      [本文引用: 2]

Although the thalamus is believed to regulate and coordinate cortical activity both within and across functional regions, such as motor and visual cortices, direct evidence for such regulation and the mechanism of regulation remains poorly described. Using simultaneous invasive recordings of cortical and thalamic electrophysiological activity in 2 awake and spontaneously behaving human subjects, we provide direct evidence of thalamic regulation of cortical activity through a mechanism of phase-amplitude coupling (PAC), in which the phase of low frequency oscillations regulates the amplitude of higher frequency oscillations. Specifically, we show that cortical PAC between the theta phase and beta amplitude is spatially dependent on and time variant with the magnitude of thalamocortical theta coherence. Moreover, using causality analysis and MR diffusion tractography, we provide evidence that thalamic theta activity drives cortical theta oscillations and PAC across structures and that these thalamocortical relationships are structurally constrained by anatomic pathways. This relationship allows for new evidence of thalamocortical PAC. Given the diffuse connectivity of the thalamus with the cerebral cortex, thalamocortical PAC may play an important role in addressing the binding problem, including both integration and segregation of information within and across cortical areas.

Morris J. S., Öhman A., & Dolan R. J . ( 1999).

A subcortical pathway to the right amygdala mediating “unseen” fear

Proceedings of the National Academy of Sciences, 96( 4), 1680-1685.

DOI:10.1073/pnas.96.4.1680      URL     PMID:9990084      [本文引用: 1]

Neuroimaging studies have shown differential amygdala responses to masked ("unseen") emotional stimuli. How visual signals related to such unseen stimuli access the amygdala is unknown. A possible pathway, involving the superior colliculus and pulvinar, is suggested by observations of patients with striate cortex lesions who show preserved abilities to localize and discriminate visual stimuli that are not consciously perceived ("blindsight"). We used measures of right amygdala neural activity acquired from volunteer subjects viewing masked fear-conditioned faces to determine whether a colliculo-pulvinar pathway was engaged during processing of these unseen target stimuli. Increased connectivity between right amygdala, pulvinar, and superior colliculus was evident when fear-conditioned faces were unseen rather than seen. Right amygdala connectivity with fusiform and orbitofrontal cortices decreased in the same condition. By contrast, the left amygdala, whose activity did not discriminate seen and unseen fear-conditioned targets, showed no masking-dependent changes in connectivity with superior colliculus or pulvinar. These results suggest that a subcortical pathway to the right amygdala, via midbrain and thalamus, provides a route for processing behaviorally relevant unseen visual events in parallel to a cortical route necessary for conscious identification.

Mosher C. P., Zimmerman P. E., & Gothard K. M . ( 2014).

Neurons in the monkey amygdala detect eye contact during naturalistic social interactions

Current Biology, 24( 20), 2459-2464. doi: 10.1016/j.cub.2014.08.063.

URL     PMID:4253056      [本文引用: 1]

During natural exploration of the visual world, social primates gaze at the eyes of their peers. Mosher et al. report the existence of neurons in the amygdala, the social-emotional center of the brain, that signal eye contact. These cells may be implicated in mental disorders marked by abnormal eye contact and impaired social behaviors.

Nation K. & Penny S., ( 2008).

Sensitivity to eye gaze in autism: Is it normal? Is it automatic? Is it social?

Development and Psychopathology, 20( 1), 79-97.

DOI:10.1017/S0954579408000047      URL     PMID:18211729      [本文引用: 1]

Children with autism are developmentally delayed in following the direction of another person's gaze in social situations. A number of studies have measured reflexive orienting to eye gaze cues using Posner-style laboratory tasks in children with autism. Some studies observe normal patterns of cueing, suggesting that children with autism are alert to the significance of the eyes, whereas other studies reveal an atypical pattern of cueing. We review this contradictive evidence to consider the extent to which sensitivity to gaze is normal, and ask whether apparently normal performance may be a consequence of atypical (nonsocial) mechanisms. Our review concludes by highlighting the importance of adopting a developmental perspective if we are to understand the reasons why people with autism process eye gaze information atypically.

Nele D., Ellen D., Petra W., & Herbert R . ( 2015).

Social information processing in infants at risk for ASD at 5 months of age: The influence of a familiar face and direct gaze on attention allocation

Research in Autism Spectrum Disorders, 17, 95-105. doi: 10.1016/j.rasd.2015.06.006.

URL     PMID:4885968      [本文引用: 2]

A visual paired comparison eye-tracking paradigm was used to measure preferences for socially relevant faces and subsequent objects in a 5-month-old low- and high-risk group for autism spectrum disorder (ASD). In a first condition the mother's face was presented next to a stranger's face, in the second condition two stranger's faces were presented with direct versus averted gaze. Afterwards two objects were presented at the same places where previously the faces appeared. Both groups showed a similar preference for their mother's face over a stranger's face, while only the low-risk group tended to show a preference for the face with direct gaze over the face with averted gaze. No difference between groups was found for looking at the objects. These findings show that differences in social attention between both groups are subtle and depend on the manipulations used.

Norbury C. F., Griffiths H., & Nation K . ( 2010).

Sound before meaning: Word learning in autistic disorders

Neuropsychologia, 48( 14), 4012-4019.

DOI:10.1016/j.neuropsychologia.2010.10.015      URL     PMID:20951710      [本文引用: 2]

Successful word learning depends on the integration of phonological and semantic information with social cues provided by interlocutors. How then, do children with autism spectrum disorders (ASDs) learn new words when social impairments pervade? We recorded the eye-movements of verbally-able children with ASD and their typical peers while completing a word learning task in a social context. We assessed learning of semantic and phonological features immediately after learning and again four weeks later. Eye-movement data revealed that both groups could follow social cues, but that typically developing children were more sensitive to the social informativeness of gaze cues. In contrast, children with ASD were more successful than peers at mapping phonological forms to novel referents; however, this advantage was not maintained over time. Typical children showed clear consolidation of learning both semantic and phonological information, children with ASD did not. These results provide unique evidence of qualitative differences in word learning and consolidation and elucidate the different mechanisms underlying the unusual nature of autistic language.

Nuske H. J., Vivanti G., & Dissanayake C . ( 2015).

No evidence of emotional dysregulation or aversion to mutual gaze in preschoolers with autism spectrum disorder: An eye-tracking pupillometry study

Journal of Autism and Developmental Disorders, 45( 11), 3433-3445. doi: 10.1007/s10803-015-2479-5.

URL     PMID:26031923      [本文引用: 1]

The 'gaze aversion hypothesis', suggests that people with Autism Spectrum Disorder (ASD) avoid mutual gaze because they experience it as hyper-arousing. To test this hypothesis we showed mutual and averted gaze stimuli to 23 mixed-ability preschoolers with ASD (M Mullen DQ = 68) and 21 typically-developing preschoolers, aged 2-5 years, using eye-tracking technology to measure visual attention and emotional arousal (i.e., pupil dilation). There were no group differences in attention to the eye region or pupil dilation. Both groups dilated their pupils more to mutual compared to averted gaze. More internalizing symptoms in the children with ASD related to less emotional arousal to mutual gaze. The pattern of results suggests that preschoolers with ASD are not dysregulated in their responses to mutual gaze.

Nyström P., Bölte S., Falck-Ytter T., & EASE T . ( 2017).

Responding to other people`s direct gaze: Alterations in gaze behavior in infants at risk for autism occur on very short timescales

Journal of Autism and Developmental Disorders, 47( 11), 3498-3509.

DOI:10.1007/s10803-017-3253-7      URL     PMID:5633639      [本文引用: 2]

Atypical gaze processing has been reported in children with autism spectrum disorders (ASD). Here we explored how infants at risk for ASD respond behaviorally to others’ direct gaze. We assessed 10-month-olds with a sibling with ASD (high risk group; n65=6561) and a control group (n65=6518) during interaction with an adult. Eye-tracking revealed less looking at the adult in the high risk group during 300–100002ms after the adult initiated direct gaze: a short alteration that is likely to go unnoticed by the naked eye. Data aggregated over longer segments (the traditional eye-tracking approach) showed no group differences. Although findings are limited by lack of outcome data, they are in line with theories linking atypical eye processing to the emergence of ASD. The online version of this article (doi:10.1007/s10803-017-3253-7) contains supplementary material, which is available to authorized users.

Pellicano E., & Macrae C.N . ( 2009).

Mutual eye gaze facilitates person categorization for typically developing children, but not for children with autism

Psychonomic Bulletin & Review, 16( 6), 1094-1099. doi: 10.3758/ PBR.16.6.1094.

[本文引用: 1]

Pelphrey K. A., Morris J. P., & McCarthy G . ( 2005).

Neural basis of eye gaze processing deficits in autism

Brain, 128( 5), 1038-1048.

DOI:10.1093/brain/awh404      URL     PMID:15758039      [本文引用: 2]

Impairments in using eye gaze to establish joint attention and to comprehend the mental states and intentions of other people are striking features of autism. Here, using event-related functional MRI (fMRI), we show that in autism, brain regions involved in gaze processing, i ncluding the superior temporal sulcus (STS) region, are not sensitive to intenti ons conveyed by observed gaze shifts. On congruent trials, subjects watched as a virtual actor looked towards a checkerboard that appeared in her visual field, confirming the subject's expectation regarding what the actor ‘ought to do’in this context. On incongruent trials, she looked towards empty space, violating the subject's expectation. Consistent with a prior report from our laboratory t hat used this task in neurologically normal subjects, ‘errors’(incongruent tri als) evoked more activity in the STS and other brain regions linked to social co gnition, indicating a strong effect of intention in typically developing subject s (n=9). The same brain regions were activated during observation of gaze shifts in subjects with autism (n=10), but did not differentiate congruent and incongr uent trials, indicating that activity in these regions was not modulated by the context of the perceived gaze shift. These results demonstrate a difference in t he response of brain regions underlying eye gaze processing in autism. We conclu de that lack of modulation of the STS region by gaze shifts that convey differen t intentions contributes to the eye gaze processing deficits associated with aut ism.

Pelphrey K. A., Viola R. J., & McCarthy G . ( 2004).

When strangers pass: Processing of mutual and averted social gaze in the superior temporal sulcus

Psychological Science, 15( 9), 598-603.

DOI:10.1111/j.0956-7976.2004.00726.x      URL     PMID:15327630      [本文引用: 1]

Abstract Using functional magnetic resonance imaging (fMRI), we investigated brain activity evoked by mutual and averted gaze in a compelling and commonly experienced social encounter. Through virtual-reality goggles, subjects viewed a man who walked toward them and shifted his neutral gaze either toward (mutual gaze) or away (averted gaze) from them. Robust activity was evoked in the superior temporal sulcus (STS) and fusiform gyrus (FFG). For both conditions, STS activity was strongly right lateralized. Mutual gaze evoked greater activity in the STS than did averted gaze, whereas the FFG responded equivalently to mutual and averted gaze. Thus, we show that the STS is involved in processing social information conveyed by shifts in gaze within an overtly social context. This study extends understanding of the role of the STS in social cognition and social perception by demonstrating that it is highly sensitive to the context in which a human action occurs.

Philip R. C. M., Dauvermann M. R., Whalley H. C., Baynham K., Lawrie S. M., & Stanfield A. C . ( 2012).

A systematic review and meta-analysis of the fMRI investigation of autism spectrum disorders

Neuroscience & Biobehavioral Reviews, 36( 2), 901-942. doi: https://doi.org/10.1016/j.neubiorev.2011.10.008.

[本文引用: 1]

Pitskel N. B., Bolling D. Z., Hudac C. M., Lantz S. D., Minshew N. J., Vander Wyk B. C., & Pelphrey K. A . ( 2011).

Brain mechanisms for processing direct and averted gaze in individuals with autism

Journal of Autism and Developmental Disorders, 41( 12), 1686-1693. doi: 10.1007/s10803-011-1197-x.

URL     PMID:21484518      [本文引用: 2]

Prior studies have indicated brain abnormalities underlying social processing in autism, but no fMRI study has specifically addressed the differential processing of direct and averted gaze, a critical social cue. Fifteen adolescents and adults with autism and 14 typically developing comparison participants viewed dynamic virtual-reality videos depicting a simple but realistic social scenario, in which an approaching male figure maintained either direct or averted gaze. Significant group by condition interactions reflecting differential responses to direct versus averted gaze in people with autism relative to typically developing individuals were identified in the right temporoparietal junction, right anterior insula, left lateral occipital cortex, and left dorsolateral prefrontal cortex. Our results provide initial evidence regarding brain mechanisms underlying the processing of gaze direction during simple social encounters, providing new insight into the social deficits in individuals with autism.

Pourtois G., Schettino A., & Vuilleumier P . ( 2013).

Brain mechanisms for emotional influences on perception and attention: What is magic and what is not

Biological Psychology, 92( 3), 492-512.

[本文引用: 1]

Psychiatric American Association . ( 2000).

Diagnostic and statistical manual of mental disorders

DSM-IV-TR fourth edition (text revision).

DOI:10.1007/978-0-387-78665-0_5485      URL     [本文引用: 1]

DefinitionDefinitionThere are up to four editions of the DSM. This is a clinical guide, with practical aims, that contains a brief description of the different mental disorders with their diagnostic c

Riby D. M., Hancock P., Jones N., & Hanley M . ( 2013).

Spontaneous and cued gaze-following in autism and Williams syndrome

Journal of Neurodevelopmental Disorders, 5( 1), 1-12. doi: 10.1186/1866-1955-5-13.

URL     PMID:3766200      [本文引用: 2]

AbstractBackgroundFrom a young age the typical development of social functioning relies upon the allocation of attention to socially relevant information, which in turn allows experience at processing such information and thus enhances social cognition. As such, research has attempted to identify the developmental processes that are derailed in some neuro-developmental disorders that impact upon social functioning. Williams syndrome (WS) and autism are disorders of development that are characterized by atypical yet divergent social phenotypes and atypicalities of attention to people.MethodsWe used eye tracking to explore how individuals with WS and autism attended to, and subsequently interpreted, an actor eye gaze cue within a social scene. Images were presented for 3 seconds, initially with an instruction simply to look at the picture. The images were then shown again, with the participant asked to identify the object being looked at. Allocation of eye gaze in each condition was analyzed by analysis of variance and accuracy of identification was compared with tests.ResultsParticipants with WS allocated more gaze time to face and eyes than their matched controls, both with and without being asked to identify the item being looked at; while participants with autism spent less time on face and eyes in both conditions. When cued to follow gaze, participants with WS increased gaze to the correct targets; those with autism looked more at the face and eyes but did not increase gaze to the correct targets, while continuing to look much more than their controls at implausible targets. Both groups identified fewer objects than their controls.ConclusionsThe atypicalities found are likely to be entwined with the deficits shown in interpreting social cognitive cues from the images. WS and autism are characterized by atypicalities of social attention that impact upon socio-cognitive expertise, but, importantly, the type of atypicality is syndrome specific.

Ristic J., Mottron L., Friesen C. K., Iarocci G., Burack J. A., & Kingstone A . ( 2005).

Eyes are special but not for everyone: The case of autism

Cognitive Brain Research, 24( 3), 715-718. doi: 10.1016/j.cogbrainres.2005.02.007.

URL     PMID:16099372      [本文引用: 1]

Current research indicates that human gaze direction is a special cue for shifting attention for one of two reasons: (1) it reflects social desires and intentions and (2) its basic perceptual features usually correspond to important events in the environment. This study, conducted with individuals with autism and with age- and IQ-matched typically developing individuals, dissociates these two often-confounded explanations and demonstrates that eyes appear to be special for typically developing individuals because of their social power, whereas gaze effects are mediated by feature correspondence among persons with autism.

Rombough A. & Iarocci G., ( 2013).

Orienting in response to gaze and the social use of gaze among children with autism spectrum disorder

Journal of Autism and Developmental Disorders, 43( 7), 1584-1596. doi: 10.1007/s10803-012-1704-8.

URL     PMID:23124360      [本文引用: 1]

Potential relations between gaze cueing, social use of gaze, and ability to follow line of sight were examined in children with autism and typically developing peers. Children with autism (mean age = 10 years) demonstrated intact gaze cueing. However, they preferred to follow arrows instead of eyes to infer mental state, and showed decreased accuracy in following line of sight when several visual distracters were present. Performance across tasks was not correlated for either group. Findings suggest that children with autism are less inclined to prioritize and select eyes, particularly in visually-rich environments. Gaze-following deficits may lie at the level of selective attention, rather than cueing-a possibility that can be explored with more complex and ecologically valid tasks.

Rothkirch M., Madipakkam A. R., Rehn E., & Sterzer P . ( 2015).

Making eye contact without awareness

Cognition, 143, 108-114.

DOI:10.1016/j.cognition.2015.06.012      URL     PMID:26133642      [本文引用: 1]

Direct gaze is a potent non-verbal signal that establishes a communicative connection between two individuals, setting the course for further interactions. Although consciously perceived faces with direct gaze have been shown to capture attention, it is unknown whether an attentional preference for these socially meaningful stimuli exists even in the absence of awareness. In two experiments, we recorded participants’ eye movements while they were exposed to faces with direct and averted gaze rendered invisible by interocular suppression. Participants’ inability to correctly guess the occurrence of the faces in a manual forced-choice task demonstrated complete unawareness of the faces. However, eye movements were preferentially directed towards faces with direct compared to averted gaze, indicating a specific sensitivity to others’ gaze directions even without awareness. This oculomotor preference suggests that a rapid and automatic establishment of mutual eye contact constitutes a biological advantage, which could be mediated by fast subcortical pathways in the human brain.

Saalmann Y. B., Pinsk M. A., Wang L., Li X., & Kastner S . ( 2012).

Pulvinar regulates information transmission between cortical areas based on attention demands

Science, 337( 6095), 753-756.

[本文引用: 2]

Sato W., Kochiyama T., Uono S., & Toichi M . ( 2016).

Neural mechanisms underlying conscious and unconscious attentional shifts triggered by eye gaze

NeuroImage, 124, Part A, 118-126. doi: 10.1016/j.neuroimage.2015.08.061.

URL     PMID:26343316      [本文引用: 1]

Behavioral studies have shown that eye gaze triggers attentional shifts both with and without conscious awareness. However, the neural substrates of conscious and unconscious attentional shifts triggered by eye gaze remain unclear. To investigate this issue, we measured brain activity using event-related functional magnetic resonance imaging while participants observed averted or straight eye-gaze cues presented supraliminally or subliminally in the central visual field and then localized a subsequent target in the peripheral visual field. Reaction times for localizing the targets were shorter under both supraliminal and subliminal conditions when eye-gaze cues were directionally congruent with the target locations than when they were directionally neutral. Conjunction analyses revealed that a bilateral cortical network, including the middle temporal gyri, inferior parietal lobules, anterior cingulate cortices, and superior and middle frontal gyri, was activated more in response to averted eyes than to straight eyes under both supraliminal and subliminal conditions. Interaction analyses revealed that the right inferior parietal lobule was specifically active when participants viewed averted eyes relative to straight eyes under the supraliminal condition; the bilateral subcortical regions, including the superior colliculus and amygdala, and the middle temporal and inferior frontal gyri in the right hemisphere were activated in response to averted versus straight eyes under the subliminal condition. These results suggest commonalities and differences in the neural mechanisms underlying conscious and unconscious attentional shifts triggered by eye gaze.

Schilbach L., Wohlschlaeger A. M., Kraemer N. C., Newen A., Shah N. J., Fink G. R., & Vogeleyabc K . ( 2006).

Being with virtual others: Neural correlates of social interaction

Neuropsychologia, 44( 5), 718-730.

DOI:10.1016/j.neuropsychologia.2005.07.017      URL     PMID:16171833      [本文引用: 1]

To characterize the neural correlates of being personally involved in social interaction as opposed to being a passive observer of social interaction between others we performed an fMRI study in which participants were gazed at by virtual characters (ME) or observed them looking at someone else (OTHER). In dynamic animations virtual characters then showed socially relevant facial expressions as they would appear in greeting and approach situations (SOC) or arbitrary facial movements (ARB).Differential neural activity associated with ME &gt; OTHER was located in anterior medial prefrontal cortex in contrast to the precuneus for OTHER &gt; ME. Perception of socially relevant facial expressions (SOC &gt; ARB) led to differentially increased neural activity in ventral medial prefrontal cortex. Perception of arbitrary facial movements (ARB &gt; SOC) differentially activated the middle temporal gyrus.The results, thus, show that activation of medial prefrontal cortex underlies both the perception of social communication indicated by facial expressions and the feeling of personal involvement indicated by eye gaze. Our data also demonstrate that distinct regions of medial prefrontal cortex contribute differentially to social cognition: whereas the ventral medial prefrontal cortex is recruited during the analysis of social content as accessible in interactionally relevant mimic gestures, differential activation of a more dorsal part of medial prefrontal cortex subserves the detection of self-relevance and may thus establish an intersubjective context in which communicative signals are evaluated.

Schuller A. & Rossion B., ( 2001).

Spatial attention triggered by eye gaze increases and speeds up early visual activity

Neuroreport, 12( 11), 2381-2386.

DOI:10.1097/00001756-200108080-00019      URL     PMID:11496114      [本文引用: 1]

What are the neuronal correlates of reflexive shifts of attention triggered by eye gaze direction? Event related potentials (ERPs) were measured on 14 subjects performing a spatial attention task where eye gaze direction of a face cued the location of a forthcoming target. Subjects were faster in detecting a validly cued target, i.e. one appearing at the location the eye was gazing at, compared to invalidly cued targets, despite the non-predictive value of the eye cues. ERP results showed an enhanced and earlier occipito-parietal P1 and N1 for valid trials, demonstrating the early modulation of visual input by attentional allocation. These findings provide the first evidence that social attention can rapidly modify the processing of visual information in extrastriate cortex.

Schuller A. & Rossion B., ( 2004).

Perception of static eye gaze direction facilitates subsequent early visual processing

Clinical Neurophysiology, 115( 5), 1161-1168.

DOI:10.1016/j.clinph.2003.12.022      URL     PMID:15066541      [本文引用: 1]

Objective: Using event-related potentials (ERPs), it has been recently shown that a reflexive shift of attention following the observation of a dynamic eye gaze cue enhances and speeds up early visual processing of a target presented at the gazed-at location. Here we investigate whether similar early sensory modulations are also elicited by static gaze cues, or if previously described attentional effects were caused mainly by visual motion cues and not by eye gaze direction per se. Furthermore, we explore if these possible attentional orienting effects reflect facilitation of the processing of cued stimuli, inhibition of the unattended stimuli, or both. Methods: Subjects were presented with a face looking to the right or left visual field (VF), or straight away, before the occurrence of a lateralized target to detect. There were 3 conditions in this nonpredictive cueing task: (1) target presented in the VF indicated by the eye gaze direction (congruent); (2) opposite to the eye gaze direction (incongruent); or (3) preceded by a straight gazing face (neutral). Results: Subjects were faster at detecting congruently than incongruently and neutrally cued targets. Facilitation effects were observed on early ERP components: the occipital P1 and occipito-temporal N1 components were speeded up as early as 100 ms following stimulus onset (P1), and enhanced (P1 and N1) in response to congruent trials, particularly in the right hemisphere. Conclusions: Spatial attention triggered by static eye gaze direction produces response facilitations predominantly lateralized to the right hemisphere 鈥 from the early sensory stages of visual processing. Significance: This study provides the first evidence of a speeding up and amplification of early visual processing following attention triggered by static eye gaze perception.

Schultz R.T . ( 2005).

Developmental deficits in social perception in autism: The role of the amygdala and fusiform face area

International Journal of Developmental Neuroscience, 23( 2-3), 125-141. doi: 10.1016/j.ijdevneu.2004.12.012.

[本文引用: 2]

Schumann C.M., & Amaral D.G . ( 2006).

Stereological analysis of amygdala neuron number in autism

The Journal of Neuroscience, 26( 29), 7674-7679.

[本文引用: 1]

Senju A., & Johnson M.H . ( 2009).

Atypical eye contact in autism: Models, mechanisms and development

Neuroscience & Biobehavioral Reviews, 33( 8), 1204-1214. doi: 10.1016/j.neubiorev.2009.06.001.

URL     PMID:19538990      [本文引用: 3]

An atypical pattern of eye contact behaviour is one of the most significant symptoms of Autism Spectrum Disorder (ASD). Recent empirical advances have revealed the developmental, cognitive and neural basis of atypical eye contact behaviour in ASD. We review different models and advance a new ‘fast-track modulator model’. Specifically, we propose that atypical eye contact processing in ASD originates in the lack of influence from a subcortical face and eye contact detection route, which is hypothesized to modulate eye contact processing and guide its emergent specialization during development.

Senju A., Tojo Y., Yaguchi K., & Hasegawa T . ( 2005).

Deviant gaze processing in children with autism: An ERP study

Neuropsychologia, 43( 9), 1297-1306. doi: 10.1016/j.neuropsychologia.2004.12.002.

URL     PMID:15949514     

http://linkinghub.elsevier.com/retrieve/pii/S0028393205000035

Senju A. & Hasegawa T., ( 2005).

Direct gaze captures visuospatial attention

Visual Cognition, 12( 1), 127-144.

[本文引用: 1]

Senju A., Vernetti A., Ganea N., Hudry K., Tucker L., & Charman T & Johnson M. H ., et al.( 2015).

Early social experience affects the development of eye gaze processing

Current Biology, 25( 23), 3086-3091. doi: 10.1016/j.cub. 2015.10.019.

URL     PMID:4683081      [本文引用: 1]

Eye gaze is a key channel of non-verbal communication in humans [1,2,3]. Eye contact with others is present from birth [4], and eye gaze processing is crucial for social learning and adult-infant communication [5,6,7]. However, little is known about the effect of selectively different experience of eye contact and gaze communication on early social and communicative development. To directly address this question, we assessed 14 sighted infants of blind parents (SIBPs) longitudinally at 6–10 and 12–1602months. Face scanning [8] and gaze following [7,9] were assessed using eye tracking. In addition, naturalistic observations were made when the infants were interacting with their blind parent and with an unfamiliar sighted adult. Established measures of emergent autistic-like behaviors [10] and standardized tests of cognitive, motor, and linguistic development [11] were also collected. These data were then compared with those obtained from a group of infants of sighted parents. Despite showing typical social skills development overall, infants of blind parents allocated less attention to adult eye movements and gaze direction, an effect that increased between 6–10 and 12–1602months of age. The results suggest that infants adjust their use of adults’ eye gaze depending on gaze communication experience from early in life. The results highlight that human functional brain development shows selective experience-dependent plasticity adaptive to the individual’s specific social environment. 61Infants of blind parents allocate less attention to adults’ eyes and gaze direction61Differences in gaze processing in these infants become greater after 1202months of age61Overall social communication skills development of them were within the typical range Infants of blind parents allocate less attention to adults’ eyes and gaze direction Differences in gaze processing in these infants become greater after 1202months of age Overall social communication skills development of them were within the typical range Senju et02al. show that infants of blind parents allocate less attention to adults’ eyes and to the object adults are looking at, despite showing typical overall social communication. The results highlight that human functional brain development shows selective experience-dependent plasticity attuned to the individual’s specific environment.

Senju A., Yaguchi K., Tojo Y., & Hasegawa T . ( 2003).

Eye contact does not facilitate detection in children with autism

Cognition, 89( 1), B43-B51. doi: 10.1016/S0010- 0277(03)00081-7.

URL     PMID:12893128      [本文引用: 1]

Eye contact is crucial in achieving social communication. Deviant patterns of eye contact behavior are found in individuals with autism, who suffer from severe social and communicative deficits. This study used a visual oddball paradigm to investigate whether children with high functioning autism have difficulty in detecting mutual gaze under experimental conditions. The results revealed that children with autism were no better at detecting direct gaze than at detecting averted gaze, which is unlike normal children. This suggests that whereas typically developing children have the ability to detect direct gaze, children with autism do not. This might result in altered eye-contact behavior, which hampers subsequent development of social and communicative skills.

Senju A., Yoshikuni T., Dairoku H., & Hasegawa T. K . ( 2004).

Reflexive orienting in response to eye gaze and an arrow in children with and without autism

Journal of Child Psychology and Psychiatry, 45( 3), 445-458.

DOI:10.1111/j.1469-7610.2004.00236.x      URL     PMID:15055365      [本文引用: 1]

Background: This study investigated whether another person's social attention, specifically the direction of their eye gaze, and a non-social directional cue, an arrow, triggered reflexive orienting in children with and without autism in an experimental situation.

Shaw P., Kabani N. J., Lerch J. P., Eckstrand K., Lenroot R., Gogtay N., .. Wise S. P . ( 2008).

Neurodevelopmental trajectories of the human cerebral cortex

Journal of Neuroscience, 28( 14), 3586-3594.

DOI:10.1523/JNEUROSCI.5309-07.2008      URL     PMID:18385317      [本文引用: 1]

Abstract Understanding the organization of the cerebral cortex remains a central focus of neuroscience. Cortical maps have relied almost exclusively on the examination of postmortem tissue to construct structural, architectonic maps. These maps have invariably distinguished between areas with fewer discernable layers, which have a less complex overall pattern of lamination and lack an internal granular layer, and those with more complex laminar architecture. The former includes several agranular limbic areas, and the latter includes the homotypical and granular areas of association and sensory cortex. Here, we relate these traditional maps to developmental data from noninvasive neuroimaging. Changes in cortical thickness were determined in vivo from 764 neuroanatomic magnetic resonance images acquired longitudinally from 375 typically developing children and young adults. We find differing levels of complexity of cortical growth across the cerebrum, which align closely with established architectonic maps. Cortical regions with simple laminar architecture, including most limbic areas, predominantly show simpler growth trajectories. These areas have clearly identified homologues in all mammalian brains and thus likely evolved in early mammals. In contrast, polysensory and high-order association areas of cortex, the most complex areas in terms of their laminar architecture, also have the most complex developmental trajectories. Some of these areas are unique to, or dramatically expanded in primates, lending an evolutionary significance to the findings. Furthermore, by mapping a key characteristic of these development trajectories (the age of attaining peak cortical thickness) we document the dynamic, heterochronous maturation of the cerebral cortex through time lapse sequences ("movies").

Shaw P., Lawrence E. J., Radbourne C., Bramham J., Polkey C. E., & David A. S . ( 2004).

The impact of early and late damage to the human amygdala on 'theory of mind' reasoning

Brain A Journal of Neurology, 127( 7), 1535-1548.

DOI:10.1093/brain/awh168      URL     PMID:15155523      [本文引用: 1]

There is a burgeoning interest in the neural basis of the ability to attribute mental states to others; a capacity referred to as 'theory of mind' (ToM). We examined the effects of lesions of the amygdala which arise at different stages of development on this key aspect of social cognition. Tests of ToM, executive and general neuropsychological function were given to subjects with lesions of the amygdala arising congenitally or in early childhood ('early damage', n = 15), subjects who acquired damage to the amygdala in adulthood ('late damage' n = 11) and matched clinical (n = 14) and healthy comparison groups (n = 38). Subjects with early damage to the amygdala, particularly if the lesion was associated with childhood onset of seizures, were impaired relative to all other groups on more advanced tests of ToM reasoning, such as detecting tactless or ironic comments or interpreting non-literal utterances. These deficits held for subjects with either left or right early amygdala damage and encompassed the understanding of both the beliefs and emotional states of others. In contrast, subjects who acquired damage to the amygdala in adulthood (usually as part of an anterior temporal lobectomy) were not impaired in ToM reasoning relative to both clinical and healthy controls, supporting the position that the amygdala is not part of the neural circuitry mediating the 'on-line' performance of ToM reasoning. In line with theories which claim that ToM is an independent faculty of cognition, we found that the pattern of results held after co-varying for measures of executive function, memory and general intellectual functioning. We discuss the results in the light of recent theories which link early developmental insults to the amygdala with the ToM impairments which are thought to be a core neurocognitive deficit found in disorders such as autism. We conclude that the amygdala may play an important role in the neural systems supporting the normal development of ToM reasoning.

Stein T., Senju A., Peelen M. V., & Sterzer P . ( 2011).

Eye contact facilitates awareness of faces during interocular suppression

Cognition, 119( 2), 307-311. doi: 10.1016/ j.cognition.2011.01.008.

URL     PMID:21316650      [本文引用: 1]

Eye contact captures attention and receives prioritized visual processing. Here we asked whether eye contact might be processed outside conscious awareness. Faces with direct and averted gaze were rendered invisible using interocular suppression. In two experiments we found that faces with direct gaze overcame such suppression more rapidly than faces with averted gaze. Control experiments ruled out the influence of low-level stimulus differences and differential response criteria. These results indicate an enhanced unconscious representation of direct gaze, enabling the automatic and rapid detection of other individuals making eye contact with the observer.

Swanson M.R., & Siller M. , ( 2013).

Patterns of gaze behavior during an eye-tracking measure of joint attention in typically developing children and children with autism spectrum disorder

Research in Autism Spectrum Disorders, 7( 9), 1087-1096. doi: 10.1016/j.rasd.2013.05.007.

URL     [本文引用: 1]

This study evaluated whether diagnostic classifications or features of ASD were associated with individual differences in children's gaze pattern during an eye-tracking measure of joint attention. The sample included 21 children with ASD (mean age, 7.3卤1.5 years) and 24 typically developing children (mean age, 6.8卤1.6 years), matched on receptive language abilities. Results revealed no significant group differences on global measures of gaze allocation (total gaze time allocation). However, significant group differences emerged using a measure evaluating a microstructure of children's gaze (duration of first fixation). In addition, individual differences in children's gaze pattern were reliably predicted by parent report measures of children's social abilities. The majority of children in this sample (including all typically developing children and those children with ASD who scored lowest on the SRS Social Awareness subscale) showed significant modulation in eye-gaze between the two experimental conditions. In contrast, children with ASD who also scored the highest on the SRS Social Awareness subscale consistently failed to modulate their eye gaze in accordance with the experimental condition. This failure to flexibly modulate gaze in the context of a joint attention eye-tracking paradigm may reveal children's limited awareness of social cues that may further limit social learning.

Tanaka J.W., & Sung A. , ( 2016).

The “eye avoidance” hypothesis of autism face processing

Journal of Autism and Developmental Disorders, 46( 5), 1538-1552.

DOI:10.1007/s10803-013-1976-7      URL     PMID:24150885      [本文引用: 1]

Although a growing body of research indicates that children with autism spectrum disorder (ASD) exhibit selective deficits in their ability to recognize facial identities and expressions, the source of their face impairment is, as yet, undetermined. In this paper, we consider three possible accounts of the autism face deficit: (1) the holistic hypothesis, (2) the local perceptual bias hypothesis and (3) the eye avoidance hypothesis. A review of the literature indicates that contrary to the holistic hypothesis, there is little evidence to suggest that individuals with autism do perceive faces holistically. The local perceptual bias account also fails to explain the selective advantage that ASD individuals demonstrate for objects and their selective disadvantage for faces. The eye avoidance hypothesis provides a plausible explanation of face recognition deficits where individuals with ASD avoid the eye region because it is perceived as socially threatening. Direct eye contact elicits a increased physiological response as indicated by heightened skin conductance and amygdala activity. For individuals with autism, avoiding the eyes is an adaptive strategy, however, this approach interferes with the ability to process facial cues of identity, expressions and intentions, exacerbating the social challenges for persons with ASD.

Thomas K. M., Drevets W. C., Whalen P. J., Eccard C. H., Dahl R. E., Ryan N. D., & Caseya B. J . ( 2001).

Amygdala response to facial expressions in children and adults

Biological Psychiatry, 49( 4), 309-316.

DOI:10.1016/S0006-3223(00)01066-0      URL     PMID:11239901      [本文引用: 1]

Background: The amygdala plays a central role in the human response to affective or emotionally charged stimuli, particularly fear-producing stimuli. We examined the specificity of the amygdala response to facial expressions in adults and children. Methods: Six adults and 12 children were scanned in a 1.5-T scanner during passive viewing of fearful and neutral faces using an EPI BOLD sequence. All scans were registered to a reference brain, and analyses of variance were conducted on the pooled data to examine interactions with age and gender. Results: Overall, we observed predominantly left amygdala and substantia innominata activity during the presentation of nonmasked fearful faces relative to fixation, and a decrease in activation in these regions with repeated exposure to the faces. Adults showed increased left amygdala activity for fearful faces relative to neutral faces. This pattern was not observed in the children who showed greater amygdala activity with neutral faces than with fearful faces. For the children, there was an interaction of gender and condition whereby boys but not girls showed less activity with repeated exposure to the fearful faces. Conclusions: This is the first study to examine developmental differences in the amygdala response to facial expressions using functional magnetic resonance imaging.

Thorup E., Kleberg J. L., & Falck-Ytter T . ( 2017).

Gaze following in children with autism: Do high interest objects boost performance?

Journal of Autism and Developmental Disorders, 47( 3), 626-635. doi: 10.1007/s10803-016-2955-6.

URL     PMID:27987062      [本文引用: 2]

This study tested whether including objects perceived as highly interesting by children with autism during a gaze following task would result in increased first fixation durations on the target objects. It has previously been found that autistic children differentiate less between an object another person attends to and unattended objects in terms of this measure. Less differentiation between attended and unattended objects in ASD as compared to control children was found in a baseline condition, but not in the high interest condition. However, typically developing children differentiated less between attended and unattended objects in the high interest condition than in the baseline condition, possibly reflecting reduced influence of gaze cues on object processing when objects themselves are highly interesting.

Thorup E., Nyström P., Gredebäck G., Bölte S., & Falck-Ytter T . ( 2016).

Altered gaze following during live interaction in infants at risk for autism: An eye tracking study

Molecular Autism, 7, 12. doi: 10.1186/s13229- 016-0069-9.

URL     PMID:4729153      [本文引用: 2]

The ability to follow gaze is an important prerequisite for joint attention, which is often compromised in children with autism spectrum disorder (ASD). The direction of both the head and eyes provides cues to other people attention direction, but previous studies have not separated these factors and their relation to ASD susceptibility. Development of gaze following typically occurs before ASD diagnosis is possible, and studies of high-risk populations are therefore important. Eye tracking was used to assess gaze following during interaction in a group of 10-month-old infants at high familial risk for ASD (high-risk group) as well as a group of infants with no family history of ASD (low-risk group). The infants watched an experimenter gaze at objects in the periphery. Performance was compared across two conditions: one in which the experimenter moved both the eyes and head toward the objects (Eyes and Head condition) and one that involved movement of the eyes only (Eyes Only condition). A group by condition interaction effect was found. Specifically, whereas gaze following accuracy was comparable across the two conditions in the low-risk group, infants in the high-risk group were more likely to follow gaze in the Eyes and Head condition than in the Eyes Only condition. In an ecologically valid social situation, responses to basic non-verbal orienting cues were found to be altered in infants at risk for ASD. The results indicate that infants at risk for ASD may rely disproportionally on information from the head when following gaze and point to the importance of separating information from the eyes and the head when studying social perception in ASD. The online version of this article (doi:10.1186/s13229-016-0069-9) contains supplementary material, which is available to authorized users.

Troiani V., Price E. T., & Schultz R. T . ( 2012).

Unseen fearful faces promote amygdala guidance of attention

Social Cognitive and Affective Neuroscience, 9( 2), 133-140.

DOI:10.1093/scan/nss116      URL     PMID:23051897      [本文引用: 2]

Abstract Little is known about the network of brain regions activated prior to explicit awareness of emotionally salient social stimuli. We investigated this in a functional magnetic resonance imaging study using a technique that combined elements of binocular rivalry and motion flash suppression in order to prevent awareness of fearful faces and houses. We found increased left amygdala and fusiform gyrus activation for fearful faces compared to houses, despite suppression from awareness. Psychophysiological interaction analyses showed that amygdala activation was associated with task-specific (fearful faces greater than houses) modulation of an attention network, including bilateral pulvinar, bilateral insula, left frontal eye fields, left intraparietal sulcus and early visual cortex. Furthermore, we report an unexpected main effect of increased left parietal cortex activation associated with suppressed fearful faces compared to suppressed houses. This parietal finding is the first report of increased dorsal stream activation for a social object despite suppression, which suggests that information can reach parietal cortex for a class of emotionally salient social objects, even in the absence of awareness.

Troiani V., & Schultz R.T . ( 2013).

Amygdala, pulvinar, and inferior parietal cortex contribute to early processing of faces without awareness

Frontiers in Human Neuroscience, 7, 241.

DOI:10.3389/fnhum.2013.00241      PMID:3674317      [本文引用: 2]

The goals of the present study were 2-fold. First, we wished to investigate the neural correlates of stimulus-driven processing of stimuli strongly suppressed from awareness and in the absence of top-down influences. We accomplished this using a novel approach in which participants performed an orthogonal task atop a flash suppression noise image to prevent top-down search. Second, we wished to investigate the extent to which amygdala responses differentiate between suppressed stimuli (fearful faces and houses) based on their motivational relevance. Using continuous flash suppression (CFS) in conjunction with fMRI, we presented fearful faces, houses, and a no stimulus control to one eye while participants performed an orthogonal task that appeared atop the flashing Mondrian image presented to the opposite eye. In 29 adolescents, we show activation in subcortical regions, including the superior colliculus, amygdala, thalamus, and hippocampus for suppressed objects (fearful faces and houses) compared to a no stimulus control. Suppressed stimuli showed less activation compared to a no stimulus control in early visual cortex (EVC), indicating that object information was being suppressed from this region. Additionally, we find no activation in regions associated with conscious processing of these percepts (fusiform gyrus and/or parahippocampal cortex) as assessed by mean activations and multi-voxel patterns. A psychophysiological interaction analysis (PPI) that seeded the amygdala showed task-specific (fearful faces greater than houses) modulation of right pulvinar and left inferior parietal cortex. Taken together, our results support a role for the amygdala in stimulus-driven attentional guidance toward objects of relevance and a potential mechanism for successful suppression of rivalrous stimuli.

Vaidya C. J., Foss-Feig J., Shook D., Kaplan L., Kenworthy L., & Gaillard W. D . ( 2011).

Controlling attention to gaze and arrows in childhood: An fMRI study of typical development and Autism Spectrum Disorders

Developmental Science, 14( 4), 911-924. doi: 10.1111/ j.1467-7687.2011.01041.x.

URL     PMID:21676110      [本文引用: 1]

Functional magnetic resonance imaging was used to examine functional anatomy of attention to social (eye gaze) and nonsocial (arrow) communicative stimuli in late childhood and in a disorder defined by atypical processing of social stimuli, Autism Spectrum Disorders (ASD). Children responded to a target word (090004LEFT090005/090004RIGHT090005) in the context of a distracting arrow or averted gaze pointing in a direction that was congruent, incongruent, or neutral (bar without arrowheads, central gaze) relative to the target word. Despite being irrelevant to the target task, both arrow and averted gaze facilitated responses (Congruent vs. Neutral trials) to the same extent in the two groups and led to interference (Incongruent vs. Congruent trials), which was greater from arrows in ASD than control children. In the brain, interaction between group and distracter-domain was observed in frontal-temporal regions during facilitation and frontal-striatal regions during interference. During facilitation, regions associated with attention to gaze in control children (left superior temporal sulcus, premotor) were associated with attention to arrows in ASD children; gaze was associated with medial temporal involvement in ASD children. During interference, regions associated with arrows in control children (anterior cingulate, right caudate) were activated in response to gaze in ASD children; further, left dorsolateral prefrontal cortex, a region not observed in control children, was activated during gaze-interference in ASD children. Thus, functional anatomy was atypical in ASD children during spontaneous processing of social and nonsocial communicative cues.

Vanmarcke S. & Wagemans J., ( 2017).

Priming facial gender and emotional valence: The influence of spatial frequency on face perception in ASD

Journal of Autism and Developmental Disorders, 47( 4), 927-946.

DOI:10.1007/s10803-016-3017-9      URL     [本文引用: 1]

Adolescents with and without autism spectrum disorder (ASD) performed two priming experiments in which they implicitly processed a prime stimulus, containing high and/or low spatial frequency information, and then explicitly categorized a target face either as male/female (gender task) or as positive/negative (Valence task). Adolescents with ASD made more categorization errors than typically developing adolescents. They also showed an age-dependent improvement in categorization speed and had more difficulties with categorizing facial expressions than gender. However, in neither of the categorization tasks, we found group differences in the processing of coarse versus fine prime information. This contradicted our expectations, and indicated that the perceptual differences between adolescents with and without ASD critically depended on the processing time available for the primes.

Villeneuve M. Y., Thompson B., Hess R. F., & Casanova C . ( 2012).

Pattern-motion selective responses in MT, MST and the pulvinar of humans

European Journal of Neuroscience, 36( 6), 2849-2858.

DOI:10.1111/j.1460-9568.2012.08205.x      URL     PMID:22759086      [本文引用: 1]

Abstract Plaid stimuli are often used to investigate the mechanisms involved in the integration and segregation of motion information. Considering the perceptual importance of such mechanisms, only a very limited number of visual brain areas have been found to be specifically involved in motion integration. These are the human (h)MT+ complex, area V3 and the pulvinar. The hMT+ complex can be functionally subdivided into two separate areas, middle temporal area (MT) and medial superior temporal area (MST); however, it is currently unclear whether these distinct sub-regions have different responses to plaid stimuli. To address this issue we used functional magnetic resonance imaging to quantify the relative response of MT and MST to component and pattern motion. Participants viewed plaid stimuli that were constrained to result in the perception of either component motion (segregation of motion information) or pattern motion (integration of motion information). MT/MST segregation was achieved using a moving dot stimulus that allowed stimulation of each visual hemifield either in unison or separately. We found pattern motion selective responses in both MT and MST. Consistent with previous reports, activity indicative of pattern motion selectivity was also found in the pulvinar as well as in other extrastriate areas. These results demonstrate that MT, MST and the pulvinar are involved in the complex motion integration mechanisms that are triggered by plaid stimuli. This reinforces the concept that integrative computations take place in a distributed neuronal circuit both in cortical and sub-cortical networks.

von Dem Hagen E. A. H.., Stoyanova R. S., Rowe J. B., Baron-Cohen S., & Calder A. J . ( 2014).

Direct gaze elicits atypical activation of the theory-of-mind network in autism spectrum conditions

Cerebral Cortex, 24( 6), 1485-1492.

DOI:10.1093/cercor/bht003      URL     PMID:4014180      [本文引用: 2]

Eye contact plays a key role in social interaction and is frequently reported to be atypical in individuals with autism spectrum conditions (ASCs). Despite the importance of direct gaze, previous functional magnetic resonance imaging in ASC has generally focused on paradigms using averted gaze. The current study sought to determine the neural processing of faces displaying direct and averted gaze in 18 males with ASC and 23 matched controls. Controls showed an increased response to direct gaze in brain areas implicated in theory-of-mind and gaze perception, including medial prefrontal cortex, temporoparietal junction, posterior superior temporal sulcus region, and amygdala. In contrast, the same regions showed an increased response to averted gaze in individuals with an ASC. This difference was confirmed by a significant gaze direction group interaction. Relative to controls, participants with ASC also showed reduced functional connectivity between these regions. We suggest that, in the typical brain, perceiving another person gazing directly at you triggers spontaneous attributions of mental states (e.g. he is "interested" in me), and that such mental state attributions to direct gaze may be reduced or absent in the autistic brain.

Wang S., Jiang M., Duchesne X. M., Laugeson E. A., Kennedy D. P., Adolphs R., & Zhao Q . ( 2015).

Atypical visual saliency in autism spectrum disorder quantified through model-based eye tracking

Neuron, 88( 3), 604-616. doi: 10.1016/j.neuron.2015.09.042.

URL     PMID:26593094      [本文引用: 2]

Wang et02al. use a comprehensive saliency model and eye tracking to quantify the relative contributions of each image attribute to visual saliency. People with ASD demonstrate atypical visual attention across multiple levels and categories of objects.

Ward R., Calder A. J., Parker M., & Arend I . ( 2007).

Emotion recognition following human pulvinar damage

Neuropsychologia, 45( 8), 1973-1978. doi: https://doi.org/ 10.1016/j.neuropsychologia.2006.09.017.

URL     PMID:17250857      [本文引用: 1]

Pulvinar activation has been observed while viewing fearful expressions, but the necessity of this activation to their recognition has not been previously assessed. We measured the processing of emotional facial expressions in a rare patient with complete unilateral loss of the pulvinar. With brief presentations, patient CJ was incapable of recognizing fearful expressions in his contralesional field. Three other patients, with damage limited to the anterior and to the lateral pulvinar, showed no deficits in recognition. In conjunction with anatomical studies of the monkey pulvinar, these results suggest that fear recognition is mediated by the human medial pulvinar. We outline the possible role of the pulvinar in fear recognition, considering both the pulvinar's direct and indirect cortical connections with the amygdala, and we suggest that the integrative role of the pulvinar may be primary. Our results suggest that the cortex in isolation from the entire pulvinar is incapable of recognizing fearful expressions.

Yokoyama T., Noguchi Y., & Kita S . ( 2013).

Unconscious processing of direct gaze: Evidence from an ERP study

Neuropsychologia, 51( 7), 1161-1168.

DOI:10.1016/j.neuropsychologia.2013.04.002      URL     PMID:23603242      [本文引用: 1]

61It is unknown how invisible direct gaze is processed by the brain.61We recorded the ERPs of direct gaze during interocular suppression.61ERPs for direct gaze were larger than for averted gaze when faces were invisible.61This was seen at 200, 240, and 320ms after stimulus onset over the frontal areas.

Zaki S.R., & Johnson S.A . ( 2013).

The role of gaze direction in face memory in autism spectrum disorder

Autism Research, 6( 4), 280-287. doi: 10.1002/aur.1292.

URL     PMID:23512772      [本文引用: 1]

We tested the hypothesis that the direction of gaze of target faces may play a role in reported face recognition deficits in those with an autism spectrum disorder (ASD). In previous studies, typically developing children and adults better remembered faces in which the eyes were gazing directly at them compared with faces in which the eyes were averted. In the current study, high-functioning children and adolescents with an ASD and age- and IQ-matched typically developing controls were shown a series of pictures of faces in a study phase. These pictures were of individuals whose gaze was either directed straight ahead or whose gaze was averted to one side. We tested the memory for these study faces in a recognition task in which the faces were shown with their eyes closed. The typically developing group better remembered the direct-gaze faces, whereas the ASD participants did not show this effect. These results imply that there may be an important link between gaze direction and face recognition abilities in ASD. Autism Res 2013, 090305090305: 090305090305090009090305090305. 0008 2013 International Society for Autism Research, Wiley Periodicals, Inc.

Zhao S., Uono S., Yoshimura S., Kubota Y., & Toichi M . ( 2017).

Atypical gaze cueing pattern in a complex environment in individuals with ASD

Journal of Autism and Developmental Disorders, 47( 7), 1978-1986.

DOI:10.1007/s10803-017-3116-2      URL     [本文引用: 2]

Clinically, social interaction, including gaze-triggered attention, has been reported to be impaired in autism spectrum disorder (ASD), but psychological studies have generally shown intact gaze-triggered attention in ASD. These studies typically examined gaze-triggered attention under simple environmental conditions. In real life, however, the environment is complex. Previous studies have shown that an enhanced cueing effect was found when using eye gaze compared with arrow cues in unpredictably complex conditions in typically developing (TD) individuals. However, in the current study, compared with TD individuals, the cueing effect failed to enhance when using eye gaze compared with arrow cues under complex conditions in individuals with ASD. This may reflect the atypical style of gaze-triggered attention when individuals with ASD adapt to environmental complexity.

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