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A unifying(聯(lián)合的) view of the basis(基礎) of social cognitionVittorio Gallese1, Christian Keysers1,2 and Giacomo Rizzolatti1Department系 of Neuroscience, Section章節(jié) of Physiology生理學, University of Parma, ItalyBCN Neuroimaging神經影像 Center, University of Groningen, Antonius Deusinglaan 2, Groningen, The NetherlandsIn this article we provide a unifying neural hypothesis on how individuals understand the actions and emotions of others. 在這篇文章中我們對個體是如何理解別人的行動和感情的這一現(xiàn)象提供了一種聯(lián)合的神經學的假設。Our main claim is that the fundamental基本的 mechanism機械裝置 at the basis of the experiential經驗的 understanding of others actions is the activation激活 of the mirror 鏡子neuron system. 我們的主要觀點是,用來理解別人的行為基本裝置是“mirror neuron system”(鏡像神經系統(tǒng))的激活.A similar mechanism, but involving the activation of viscero發(fā)動機-motor centers, underlies在。之下 the experiential understanding of the emotions of others.理解別人的情感的基本裝置是一種相似的裝置,但它包含一個“viscera-motor centers”( “內臟發(fā)動機中心”)。Humans are an exquisitely精致 social species物種. Our survival and success depends crucially至關重要的 on our ability to thrive茁壯成長 in complex social situations. One of the most striking突出的features of our experience of others is its intuitive憑直覺感知的 nature.This implicit含蓄的,不言說的 grasp抓住,領會 of what other people do or feel will be the focus of our review回顧. We will posit假定 that, in our brain,there are neural mechanisms (mirror mechanisms) that allow us to directly understand the meaning of the actions and emotions of others by internally replicating復制 (simulat-simulating) them without any explicit明確的 reective mediation調解. 我們假定,在我們的腦中,有神經裝置允許我們通過在沒有任何明確調解的前提下直接理解別人的行為的和情感的含義。Conceptual reasoning is not necessary for this understanding. 概念推理對于這種理解并不是必要的。As human beings, of course, we are able to reason about others and to use this capacity能力 to understand other peoples minds at the conceptual,declarative?聲明的 level. 作為人類,我們能夠推出別人(的行為或情感),我們當然可以用這種能力在概念的、陳述的水平上來理解別人。Here we will argue, however, that the fundamental mechanism that allows us a direct experiential grasp of the mind of others is not conceptual reasoning but direct simulation模仿 of the observed events through the mirror mechanism. 然而,這里我們要提出的是,允許我們根據經驗的直接領會別人的心理活動的重要裝置不是概念的推理,而是通過鏡像裝置對被觀察到的事件的直接模仿。 The novelty新穎性 of our approach(。的方法) consists in providing for the rst time a neurophysiological account解釋 of the experiential(經驗的0) dimension尺寸,度量 of both action and emotion understanding.我們的研究方法的新穎性包括:對依據經驗的對于行為和情感的理解首次提供了神經科學上的解釋。What makes social interactions so different from our perception觀念 of the inanimate(使。沒有生命) world is that we witness目擊 the actions and emotions of others, but we also carry out拿出去similar actions and we experience similar emotions. 使得我們的社會互動如此不同于我們對于無機世界的觀念的是,我們看到別人的行為和情感,但我們也采取相似的行為并且經歷相似的情感。There is something shared分享的 between our rst- and third-person experience經驗 of these phenomena: the observer and the observed are both individuals endowed資助的 with a similar brainbody腦-身 system. 在第一人稱視角的經驗與第三人稱視角的經驗之間有共通的部分:觀察者與被觀察者都在應用一個相似的身心系統(tǒng)(mirror neuron system?)A crucial重要的 element元素 of social cognition is the brains capacity容量 to directly link the rst- 第一人稱and third-第三人稱person experiences體驗 of these phenomena (i.e. link I do and I feel with he does and he feels). 社會認知的一個重要因素是這些現(xiàn)象中大腦將第一、第三人稱視角經驗聯(lián)系起來的能力。(例如,將我做和我想與他做和他想聯(lián)系起來)We will dene this mechanism裝置 simulation.模仿我們將這種裝置定義為模仿In the rst part of this review we will show that in the case of actions, simulation is based on a neural system responding回應 both when we execute處決 a particular goal-directed目標導向的 action and when we observe someone else performing表現(xiàn)出 a similar action (mirror neuron system). 在第一部分我們將展示行為方面的案例,模仿基于神經系統(tǒng),在我們實際執(zhí)行目標導向的行為以及我們觀察某人表現(xiàn)相似的行為時,它都會有回應。In the second part, we will show that a similar mirroring mechanism, bridging?橋,連接 rst- and third-person experiences,also exists for emotions.第二部分,我們將展示,在情感方面也存在一個相似的裝置將第一第三視角經驗連接起來。Action understanding: cognitive and motor mechanisms 行為理解:認知和發(fā)動裝置The conventional依照傳統(tǒng)的 conceptual觀念的,概念的approach for understanding actions performed by others is to consider seen actions in a similar way to all other visual stimuli.傳統(tǒng)觀念中理解別人表現(xiàn)出的行為的方法就是按照與看待其他視覺刺激的方式相似的方式.Let us imagine a boy throwing stones into a river: the visual system describes the boy, the stone, the river, the movement of his arm, and the ying of the stones. The integration of all these separate elements produces the neural input to a central conceptual system that will interpret and assign meaning to the visual representation (e.g. see Fodor 1,2).讓我們想象In recent years, a different proposal提議 has been advanced提前了on how others actions can be understood. 近些年來,一種就“別人的行為何以被理解”的新的提議日趨高昂。This proposal is mostly based on the discovery of a set of neurons calledmirror neurons. 這種提議建立在對于鏡像神經元的發(fā)現(xiàn)之上These neurons, originally found in the Ventral腹側的 Premotor(運動前區(qū))cortex 皮層(area F5) of the macaque monkey(獼猴),respond both when the monkey performs a particular goal-directed action, and when it observes another individual performing a similar action 3,4.這些神經元,最初是在獼猴的前運動皮層腹側(F5區(qū))發(fā)現(xiàn)的,這些神經元在猴子執(zhí)行一種目標導向的行動以及當它觀察里一個個體執(zhí)行相似的行動時都會有所反應。%Mirror neurons are a particular class of visuo-motor neurons, originally discovered in a sec-tor of the ventral premotor cortex of monkeys,called area F5 (REFS 10,11).Rizzolatti, G., Fadiga, L., Fogassi, L. & Gallese, V.%直方圖單位 縱坐標 spikes/s 尖峰放電 閃電強波 橫坐標 s(左至右增大)Figure 1.Example of a monkey F5 mirror neuron responding to action observation in full vision (a) and hidden (b) conditions. 例:一只猴子的f5鏡像神經元在完全可以看到的條件下和部分內容被隱藏的條件下對行動觀察都有所反應。The lower part of each panel illustrates the experimenters action as observed from the monkeys vantage point: 每個儀表板的較低部分暗示從猴子的優(yōu)勢眼觀察到的實驗者的行動。the experimenters hand starting from a fixed position, moving toward an object and grasping it (a,b), or mimicking grasping, also either in full vision(c) or hidden conditions (d). 實驗者的手從一個固定位置開始,向一個物體移動并且抓住它,或者模仿抓的動作。這兩種都是在完全可以看到的條件下和部分內容被隱藏的條件下進行的。In these last two conditions the monkey observed the same movements as in (a) and (b), but without the target object. 在這些后面的兩種條件下,猴子觀察和a和b中同樣的運動,但是沒有目標導向。The gray square on the right in (b) and (d) represents the opaque sliding screen that prevented the monkey from seeing the experimenters action performed behind it. B與d中的右側灰色區(qū)域表示阻止猴子看到其后面的實驗者的行動的不透明的滑動屏幕。A metallic frame was interposed between experimenter and monkey in all conditions. 在所有條件下一個金屬框架都被設置于實驗者和猴子之間。The asterisk indicates the location of a marker on this frame. 星號表示這個框架上的標記位置。In hidden conditions (b,d), the experimenters hand started disappearing from the monkeys vision when crossing the marker position. 在隱藏的條件下(b.d),在穿過標記位置時,實驗者的手開始從猴子的視線中消失。The upper part of each panel shows raster displays and histograms of ten consecutive(連續(xù)的) trials recorded during the experimenters hand movement. 儀表板的偏上的部分表示光柵顯示器和在實驗者的手運動過程中記錄的十次連續(xù)的實驗的直方圖。Above each raster, kinematic recordings (colored traces) of the experimenters hand are shown. 在每個直方圖上方,實驗者的手的運動學記錄(彩色的痕跡)被展示出來。The illustrated neuron responded to the observation of grasping and holding in full vision (a) and in the hidden condition (b). 有注釋的神經元對全視覺及部分視覺條件下的抓握動作的觀察都有反應。However, the neuron response was virtually absent in the two conditions in which the observed action was mimed (c,d). 但是,這種神經元的反應在被觀察到的行為是模擬的這種條件下事實上消失了。Histograms bin width 1/420 ms. 直方圖寬度1/420 ms,Ordinates: spikes.s K1 ; Abscissae: time. Reproduced with permission from 5.%【相似實驗】%The core核 of the proposal提議 is that the observation of an action leads to the activation of parts of the same cortical皮層的 neural network網絡that is active活躍的during its execution執(zhí)行. 這個提議的關鍵是,對于行為的觀察這一活動導致了執(zhí)行相同行為時會被激活的同一皮層的神經網絡的激活。The observer understands the action because he know its outcomes輸出 when he does it. 觀察者理解行為因為他知道在他做這件事的時候它的結果。Action understanding does not depend, according to this view, on the activation of visual representations表征 (an activation obviously present) followed by their Interpretation解釋 by the central conceptual system, but by the penetration 滲透of visual information into the experiential 根據經驗的(rst person) motor發(fā)動 knowledge of the observer.根據這種觀點,行為理解并不依賴于他們經由中心概念系統(tǒng)解釋后的視覺表征的激活(顯著表現(xiàn)出的激活),而是通過視覺信息對于觀察者的根據經驗的(第一人稱視角)發(fā)動知識的“滲透”。Evidence for motor involvement牽連 in action understanding: monkey data 行為理解中的發(fā)動牽連的證據:猴子數據In addition to previous evidence (see above), the proposed link between action understanding and observers motor activation激活 was recently supported by two studies on mirror neuron properties性能. 除了先前的證據,被提出的在行為理解與觀察者的發(fā)動激活之間的關系最近得到兩項鏡像神經元性能研究的支持。In both of them, 在這兩項研究中the activity of F5 mirror neurons was studied in conditions in which the monkey could understand the meaning of the occurring action, but had no access to the visual features that typically顯著性的 trigger引發(fā),觸發(fā) mirror neurons. F5區(qū)的鏡像神經元在猴子可以理解出現(xiàn)的行為的含義但沒有機會接觸可以顯著的引發(fā)f5區(qū)的視覺特征。在這兩項研究中,f5區(qū)鏡像神經元是在猴子可以理解呈現(xiàn)的行為的含義的條件下進行的,但沒有機會接觸可以顯著的激活f5區(qū)的視覺特征。If mirror neurons mediate調解 action understanding, one should also expect期待 their activation激活 in these conditions, reecting反映 the meaning含義of the observed action.如果鏡像神經元調解行為理解,那么,就也應該期待在這些條件下的他們的激活,這種激活反映了被觀察到的行為的含義。In the rst study, mirror neurons were tested in two conditions: in one, the monkey could see the entire action (e.g. a hand grasping action); in the other, the same action was presented but its nal critical part, that is the handobject interaction, was hidden by a screen (Figure 1).在第一項研究中,鏡像神經元是在兩種條件下被測試的:其中一項中,猴子可以看見全部的行為(例如:一只手的“抓”動作);在另一項研究中,同一種行為被表現(xiàn)出來但是其最后的關鍵部分,是手物互動被屏幕所隱藏了。Thus, in the hidden condition the monkey only knew that the object was present behind the screen. 所以,在隱藏條件下,猴子只是知道物體在屏幕后面。The results showed that more than half of the recorded neurons also responded in the hidden condition 5.結果顯示超過一半的被記錄的神經元對于隱藏條件下的實驗也有所反應。In the second study, F5 mirror neurons were recorded when the monkey saw and heard executed執(zhí)行的 noisy actions (e.g. breaking peanuts, tearing sheets of paper apart), only saw, or only heard the same actions performed by another individual (Figure 2).Figure 2. Responses of two neurons selectively activated by action-related sounds (e.g. breaking peanuts, ripping paper).對與行為相聯(lián)系的聲音存在選擇性激活的兩種神經元的反應。(例如,砸花生,撕紙)Responses to two different actions for each neuron are shown, with the action triggering the most activation in blue. 所示的每種神經元對不同行為的反應,行為觸發(fā)了絕大部分藍色表示的激活。VCS, vision plus sound condition; V, vision-only condition; S, sound-only condition; M, motor condition, in which the monkey touched the object. VCS,視覺加上聲音條件;V,只有視覺條件;S,只有聲音條件;M,發(fā)動條件,猴子碰到了物體。The vertical black lines indicate either the time when the sound would have occurred (V) or the moment the monkey touched the object (M). 垂直的黑線表示要么是聲音會出現(xiàn)的時刻(V),要么是猴子接觸了物體的時刻。Traces(痕跡) under the spike(釘)-density(密度) functions(功能) in the sound-only conditions are oscillograms(波形圖) of the sounds(聲音) played back. spike-density函數下面的痕跡是在只有聲音條件下的播放出來的聲音的波形圖。Modied(修良的) with permission許可 from 6. 在第二項研究中,f5區(qū)鏡像神經元在猴子看到或聽到行動被執(zhí)行時被記錄。Like all F5 mirror neurons, the mirror neurons in this study had motor properties and red during execution執(zhí)行eksikju:t of those actions, the observation of which also triggered their activation. 正如f5區(qū)所有的鏡像神經元,這項研究中的鏡像神經元具有發(fā)動性能,并在執(zhí)行這些行為時被激活,對于這些行為的觀察也觸發(fā)了他們的激活。The results showed that about 15% of mirror neurons responsive to presentation of actions accompanied by sounds also responded to the presentation of the sound alone. 結果顯示,在對行為和聲音共同出現(xiàn)反應的鏡像神經元中有15%的神經元會對單獨出現(xiàn)的聲音起反應。These audiovisual視聽的mirror neurons therefore represent actions independently獨立的 of whether these actions are performed, heard or seen 6.所以不論這些行為是表現(xiàn)出來的,聽到的,還是看到的,這些視聽鏡像神經元都是單獨表征行為的。These studies suggest that the activity of mirror neurons correlates with action understanding. 這些研究說明鏡像神經元的活動與行為理解是相關的。The sensory features of the perceived actions (partially seen or just heard) are fundamental to the activation of mirror neurons only in as much as they trigger the motor representation表現(xiàn)表征 of the same相同 actions within the observer/listener brain.被感知到的行為的感覺特征(部分的被看見或只是被聽到)對于鏡像神經元的活動只有在他們觸發(fā)觀察者大腦中已有的行為的表征時是很重要的。Evidence for motor involvement in action understanding: human data行為理解中發(fā)動牽連的證據:人類數據Several studies using different methodologies have shown that, in humans too, the observation of actions performed by others activates cortical motor representations. 幾項運用了不同方法的研究表明,在人類身上也表現(xiàn)出相同的東西,對于被試行為的觀察會激活皮層的發(fā)動表征。The human mirror neuron system is formed(成形) by a cortical network composed(組成) of the rostral(有喙的) part of the inferior(下級) parietal lobule and by the caudal sector (pars opercularis) of the inferior(下級) frontal(額) gyrus (IFG), plus the adjacent (鄰近的)part of the premotor cortex (see 7).The human mirror neuron system resonates(共鳴) in response to a wider range of actions than the monkey system. 相對于猴子的鏡像神經元系統(tǒng)而言,人類鏡像神經元系統(tǒng)能對更廣泛的行為有反應。First, whereas the presence of an object the target of the action appears to be necessary to activate the mirror neuron system in the monkey 3,4, the observation of intransitive and mimed(模擬的) actions is able to activate the human system 810. 首先,不論物體是否出現(xiàn)行為的目標對于激活猴子的鏡像神經元系統(tǒng)是必要的,對于不及物的和模擬的行為的觀察都會激活人類的鏡像神經元系統(tǒng)。Second, TMS() experiments have shown that, in humans, motor evoked(引起) potentials(電位) (MEPs) recorded from the muscles of an observer, are facilitated(得到易化) when an individual observes intransitive(不及物的), meaningless hand/arm gestures, as well as when an individual observes a transitive(及物的) action 11,12. 第二,經顱磁刺激技術實驗表明,在人類身上,從觀察者的肌肉記錄到的運動誘發(fā)電位,在個體觀察不及物行為、無意義的手的或胳膊的姿勢時得到易化,就像個體觀察一項及物行為時一樣。Taken together, these data show that the human motor system codes both the goal of an observed action and the way in which the observed action is performed. 綜合考慮,這些數據表明人類的發(fā)動系統(tǒng)對所觀察到的行為的目標和行為所執(zhí)行的方式都會進行編碼。In summary, these data indicate that when we see someone performing an action, besides the activation of various visual areas, there is a concurrent(同時的) activation of part of the same motor circuits(環(huán)路) that are recruited(復原,得到補充) when we ourselves perform that action. 總而言之,這些數據暗示,當我們看到某人執(zhí)行某一行為時,除了激活各種視覺區(qū)域外,同時,一個相同的發(fā)動環(huán)路也被激活,并且這個環(huán)路會在我們執(zhí)行那個行為時激活。Although we do not overtly reproduce the observed action, part of our motor system becomes active as if we were executing that very same action that we are observing. 盡管我們不會過度再生那個被觀察到的行為,我們的激活系統(tǒng)部分被激活了“就像”我們在執(zhí)行那個與我們所觀察的行為非常相似的行為。The mechanism of action understanding, based on the mirror neuron mechanism, is conceptually similar to the proposal for how action understanding takes place according to phenomenologists, and Merleau-Ponty in particular 13. 建立在鏡像神經元機制基礎上的行為理解的機制,與現(xiàn)象學家提出的行為理解是如何發(fā)生的這一提議在概念上是相似的。For example, the description(描述)by Merleau- Ponty of what it means to understand an action (The sense感覺 of gesture姿勢 is not given, but understood, that is recaptured再體驗 by an act of the spectators觀眾 part; 13, p. 185), expresses表達 nicely the direct experiential understanding of the observed actions mediated(調解的) by the mirror mechanism.舉個例子,Merleau- Ponty對于它對于行為理解的意義的描述,很好的表達了直接的根據經驗的對于所觀察行為的經由鏡像機制調解的理解。It is important to stress here that, in the present review, we discuss only how the meaning of action is understood and not how the intention of the actions agent is captured. 在這里強調一下是很重要的,在目前的回顧中,我們只是討論了行為的意義是如何被理解的,而不是行為的意圖是如何被識破的。Although we are inclined to believe that simulation probably underlies intention understanding(see 1421) too, this issue is outside the aim of the present review.盡管我們傾向于認為模擬大概也在對于意圖的理解之下,這個話題確實在目前的回顧的目標之外。Emotion understanding 情感理解So far, we have discussed the neural mechanism underlying action understanding. 至今為止,我們已經討論了行為理解之下的神經裝置。Does a similar mechanism mediate our understanding of the emotions of others?是否存在一個相似的裝置調節(jié)我們對于他人情感的理解呢? In the next sections we will show that a similar mechanism is also involved in our capacity to understand and experience the emotional states of others. 在下一部分,我們將會展示一個在我們的能力中也有相似的裝置來理解和體驗別人的情感情形。We will focus on the emotion of disgust, for which rich empirical(經驗的) evidence has recently been acquired(掌握了). 我們將會關注厭惡情感,因為針對這種情感的經驗證據已經被掌握了。 We will discuss in particular the role of the insula(腦島) that appears to play a fundamental role in the feeling and understanding of this basic emotion (see Box 1).我們將特別討論在感覺和理解基本情感過程中發(fā)揮重要作用的腦島。Box 1. Amygdala midl, insula, and selectivity for particular emotions表格1.杏仁核,腦島,對于特定情感的選擇性In this review we focus on the role played by the insula in the perception and experience of disgust. 在這片回顧里,我們將集中精力于腦島在厭惡情感的知覺與體驗中發(fā)揮的作用。Many authors have emphasized a major role played by the amygdala in the perception and experience of fear (e.g. 64). 許多作者都強調過杏仁核在恐懼情感的知覺與體驗中發(fā)揮的作用。Phillips et al. 47 found activations of amygdala as well as of the insula insjul to fearful versus neutral faces, and of the insula but not of the amygdala for disgusted versus neutral facial expressions. Phillips et al.發(fā)現(xiàn)杏仁核的激活就像腦島在相對于帶有中性表情的臉對帶有恐懼表情臉表現(xiàn)出激活一樣,而腦島的激活并不像杏仁核在相對于帶有中性表情的臉對于帶有厭惡表情的臉的激活一樣。Zald 46 reviewed the functional imaging literature on the amygdala and showed that although fearful facial expressions are the most robust rubst elicitors of amygdala activation,other facial expressions, including disgust, often also activate it. Zald 46回顧了對于杏仁核的功能性成像的文獻,表明,盡管“恐懼”面部表情是杏仁核激活的最強大的激發(fā)子,其他面部表情,包括“厭惡”表情,也能激活它。This last nding was not conrmed by our data 55. 最后一項發(fā)現(xiàn)尚未為我們的數據所證實。This controversy is probably due to the fact that the amygdala is a complex structure,containing medial nuclei that are involved in olfactory lfktri processing, and lateral ltrl nuclei that appear more selectively involved in the visual and auditory processing of threat 65. 這種爭論大概要歸咎于杏仁核是一個復雜的結構,包括中間的與嗅覺過程相關的核,側面的對視覺和聽覺威脅的過程表現(xiàn)出更大的選擇性的核。Given the susceptibility artifacts encountered in fMRI and the spatial limits of PET, it is difcult, if not hopeless,

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