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駒場学部講義2015 総合情報学特論III 「意識の神経科学:「気づき」と「サリエンシー」を手がかりに」

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駒場学部講義2015 総合情報学特論III 「意識の神経科学:「気づき」と「サリエンシー」を手がかりに」

  1. 1. 東京大学学際科学科 2015年度 総合情報学特論III 意識の神経科学:「気づき」と「サリエンシー」を手がかりに 6月10日(水) 3-4限 13:00-16:40, 駒場キャンパス15号館1階104講義室 吉田 正俊 (自然科学研究機構・生理学研究所・認知行動発達研究部門 助教) 要旨 Part 1: • Neural correlates of awareness can be studied using bistable percepts. • The dorsal and ventral visual pathways may have different roles on vision for action and vision for perception, respectively. • Feeling-of-something-happening in blindsight may arise from saliency computation. 要旨 Part 2: • Feeling-of-something-happening in blindsight can be explained by Sensorimotor contingency. • The positive symptom of schizophrenia can be explained by prediction error, which is equivalent to aberrant saliency hypothesis. • Presence can be the key to the sentient self, which is the integration of salience of extero- and interoceptive signals. PART 1 0. What is awareness, conscious experience and attention? -- Let’s start from examples • We cannot be aware of a big change in a scene when we do not pay attention to it. • At the same time, we have a conscious experience even before we notice it. (No hole in our picture!) 1. How to study consciousness? 1-1 Definition of consciousness • ?? “Consciousness is not a subject of science because we cannot define them.” • Let’s start from a common-sense definition, not from an analytic definition. 1 • A common-sense definition of consciousness: "consciousness refers to those states of sentience or awareness that typically begin when we wake from a dreamless sleep and continue through the day …” 1-2 Hard problem of consciousness • Philosophical zombie • The inverted spectrum • In his (Ned’s) class, ~2/3 of the students usually say, ‘Oh yeah, I see what you’re talking about’ and some of them even say, ‘Oh yeah, I’ve wondered about that since I was a kid.’ ~1/3 of people say, ‘I don’t know what you’re talking about.’ • The hard problem of consciousness: • the problem of explaining how and why we have qualia or phenomenal experiences — how sensations acquire characteristics, such as colors and tastes. • Awareness: “a state wherein we have access to some information, and can use that information in the control of behavior.” the psychological concept of mind” 2. Neural correlates of awareness • What are neural correlates? • Halle Berry の顔、名前を見たときだけ活動するニュ ーロン 2 =「Halle Berry という視覚刺激を提示した こと」の neural correlate 「Halle Berry を見たと いう経験」の neural correlate • An experimental manipulation is required by which a visual input is constant but perception of that visual stimulus varies. • Binocular rivalry (両眼視野闘争) • In binocular rivalry, the stimulus is stable but the content of awareness switches. Then, we can find the neural correlate of visual awareness by comparing two different perceptual reports. • Activity of IT neurons reflects the monkeys’ perceptual report 3 . This is strong evidence that IT neurons represent content of subjective experience. • 腹側視覚経路では、解剖学的により高次な領域でよ り主観的な情報が反映している • IT neurons represent a stage of processing beyond ambiguities and their activity reflects the brain's internal view of objects. • fMRI during binocular rivalry 4 • Neural correlates of awareness can be studied using bistable percepts. 3. Vision for perception and vision for action
  2. 2. • Two visual system hypothesis 5 • Dorsal pathway: Vision for action • Ventral pathway: Vision for perception • Optic ataxia (視覚性運動失調) • 右脳損傷あり。左周辺視野でのみ腕の傾きを間違 える。右手でも左手でも同様。 視覚障害はない。 6 • 原因部位は SPL-後頭皮質接合部 • 視覚背側経路への損傷が「行動のための視覚」に 影響を及ぼす。 • Visual form agnosia (視覚失認): • Subject DF: Bilateral damage in ventral visual pathway (Lateral occipital area: LO) • Very good performance in ‘posting’ task 7 • 「知覚のための視覚」は傷害されているのに「行 動のための視覚」が保存されている • DF matched her card orientation to the slot during the course of the movement, well before contacting the target. 8 • 行動のうちに知覚が現れてくる。 • Functional double dissociation • "Visual phenomenology ... can arise only from processing in the ventral stream ... visual-processing modules in the dorsal stream ... are not normally available to awareness." ("The visual brain in action" p.200-201) 9 • Dorsal and ventral visual pathways may have different roles on vision for action and vision for perception. 4. Blindsight (盲視) in human 4-1. What is blindsight? • “The visually evoked voluntary responses of patients with striate cortical destruction that are demonstrated despite a phenomenal blindness” 10 • Phenomenal consciousness can be dissociated from visual information processing. • 大脳皮質損傷によって「カエル脳」が える。 • 4-2. Case reports • G.Y. became blind in his right visual field due to traffic accident in eight years old. He was diagnosed as homonymous hemianopia. • Above-chance performance in forced-choice => blindsight 11 • 患者 G.Y.氏の「なにかあるかんじ」 • He has a ‘feeling’ of something happening in his blind field and, given the right conditions, that he is absolutely sure of the occurrence. • He described his experience as that of ‘a black shadow moving on a black background’, adding that ‘shadow is the nearest I can get to putting it into words so that people can understand’. • 被験者の内観報告:「縞模様が見えているわけでは ない」「上下どちらにあるかは雰囲気で分かる」 • 意識経験の「内容 content」はないけれども「現前 性 presence」はある? • 盲視における implicit perception • 1) Direct measure (左右方向の弁別)では左右方向 の情報処理の証拠はない • 2) Indirect measure (サッカードの終止位置)では 左右方向の情報処理をした証拠がある • 盲視における vision for action「運動中に視覚が現 れてくる」 5. Blindsight in monkey 5-1. Animal model of blindsight • Recovery after 2-3 months training 12 • Are the monkeys really ‘blind’ to the visual stimuli? • The monkeys behaved as if it is a No-target trial. • The monkeys are ‘not able to see’, as in human blindsight 13 . 5-2. Saliency in blindsight • Monkey without V1: Visually guided reaching => Visual saliency? • What is saliency? - The distinct subjective perceptual quality which makes some items in the world stand out from their neighbors and immediately grab our attention 14 • Saliency computational model 15 • Iterative calculation of center-surround differentiation and normalization • Intracortical lateral inhibition is mathematically equivalent to second derivative (=laplacian). It is used for edge detection. • Salient stimuli attract gazes of blindsight monkeys 16 • Color saliency or color identification? • The lesioned monkey responded to color saliency. • 盲視であるとはどのようなかんじ? • Feeling ‘atmosphere’ = saliency without visual consciousness • Dual system for conscious vision and saliency. PART 2 1. Active vision 1-1. Bayesian surprise • Saliency map: spatial outlier • Saliency model evaluates what is salient in a image in term of spatial configuration. • How to evaluate temporal outlier? • => Bayesian surprise • Why are you surprised when you see the sandstorm? • The sandstorm itself is not surprising. • The transition from the CNN news to the sandstorm is the surprise. • Bayesian surprise measures how much your belief
  3. 3. changed by the data. • Bayesian surprise is defined as the difference between prior and posterior. =>KL divergence • Surprise is better predictor than saliency for gaze patterns during free-viewing 1-2. Predictive coding • ヘルムホルツ的視覚観 17 • ニューロンは特徴検出器(フィルタ,template)であ るという考え • でもニューロンの応答はすぐ adapt する。=> サプ ライズ検出器なんじゃないか? • V1 response can be modeled by surprise • A neuron has a belief (=model = prior) • Feed-forward prediction error = Bayesian Surprise = Attention • Backward prediction = Internal model = Conscious perception • Binocular rivalry can be explained by predictive coding 18 1-3 Friston’s free energy principle • An organism is able to minimize the free energy • by reducing bayesian surprise (internal state) => predictive coding • by changing sensation (action) => active inference 19 . • Active inference = interplay between action and sensation = sensorimotor contingency 1-4. Enactive view: Sensorimotor contingency theory • Standard view: Seeing is making an internal representation • New view: Seeing is knowing about things to do 20 • Alva Noe のエナクション説 21 :対象に向かって近 づくと対象の姿が大きくなる。私たちはこのような sensorimotor dependence に精通している。私たち の知覚能力は、この種の感覚-運動的知識を持って いるということによってはじめて成り立つ。 • Hurley and Noë’s argument 22 : • Based on Enactive view, sensorimotor contingency, rather than brain region, is the determinant of conscious experience. • This is empirically testable. sensory input ? (=> Externalism, enactivism) or brain activity? (=> Internalism) • Phantom limb: the case for brain activity => Internalist view • Inverted glass: the case for sensory input 23 =>Externalist view • Which occurs in blindsight? • Internalist view: 「なにかあるかんじ」は上丘の 活動に固有の経験であるが、V1 の活動に固有の 経験である「ビビッドな赤」にマスクされている。 V1 損傷によって隠れていた「なにかあるかんじ」 が現れる。 • Enactive view: 意識経験の感覚運動ループ説:「な にかあるかんじ」は機能回復に伴って感覚運動ル ープの形成による環境の操作可能性が拡大した ことによって生成した、ある種の視覚意識ではな いか。 • 盲視の例は externalist 説を支持しているのではな いか? • 1) Blindsight is not available just after the lesion. • 2) さらに empirical にテストできる。Normal subject で V1 を一時的に suppress したら feeling-of-something は起こるか? 盲視で SC を 一時的に抑えたら何が起こるか? • 盲視を現象学的に理解する • 現象学者の Zahavi は現象的意識の構成的特徴とし て「前反省的自己意識」が不可欠であると書く 24 。 • 前反省的自己意識:経験的現象が現象学的な意味 での反省を経る前から直接的一人称的に与えら れているということ (first-person perspective を 持っている) • <=>反省的自己意識:(いわゆる「自己意識」) • 「なにかあるかんじ」とは意識経験である • 盲視の研究から示唆された視覚の二つのレイヤ ー(「視覚的意識」と「なにかあるかんじ」)はど ちらとも前反省的自己意識を持っているといえ る。 • 「なにかあるかんじ」(=「視覚の現前性」)とは無 意識の過程ではなくて、「視覚的意識」(=「意識 の内容」)を図とするならば、それに対する地の関 係として意識経験を構成するものなのかも。 1-5. Let’s combine everything • 視覚の腹側経路 = 知覚のための視覚 • Conscious experience = prediction in predictive coding • Bottom up attention (or bayesian surprise) = prediction error • 視覚の背側経路 = 行動のための視覚 • なにかがあるかんじ = presence = サリエンシ ー: shaped by sensorimotor contingency • こちらもある種の conscious experience 2. Schizophrenia 2-1. What is schizophrenia? • A chronic mental illness characterized by persistent psychotic and negative symptoms and relatively subtle cognitive impairment. 25 • Positive symptoms (陽性症状) = Psychosis (精神症 状): • Delusions (妄想): A fixed implausible, preoccupying belief (例:「私の耳の後ろにマイク ロチップが埋め込まれていて、それが私の考えを 支配している」 • Hallucinations (幻覚): A voice, vision, or other percept in the absence of a stimulus. 典型的には
  4. 4. 「患者の行動にコメントしたり、患者を侮 する 声」 • Negative symptoms (陰性症状): • Apathy, reduced social interactions, poor self-care • Socio-developmental-cognitive モデル 26 • 統合失調症は主に思春期から青年期くらいの時 期に発症する。それ以前に神経発達の障害が見ら れる。これが社会的困難と重なることによって発 症する。 • 統合失調症研究が難しい理由 • Schizophrenia is not a single entity: • Paranoid schizophrenia (妄想型: delusions or hallucinations) • Hebephrenic schizophrenia (破瓜型: thought disorder) • Catatonic schizophrenia (緊張型: abnormal motor behavior) • No single cause: • No single gene (RGS4, DISC1, DTNBP1, NRG1, HTR2A,...) • No single environmental factor • No single brain area (STG, Str, HC, AI, ACC, DLPFC,...) • No single neurotransmitter (DA, Glu, 5HT, ACh,...) • No animal model but endophenotype (中間表現型): • Pre-pulse inhibition, latent inhibition • Anti-saccade, inhibition-of-return • Working memory task • We need a working hypothesis which explains how the symptoms develop. 2-2. Predictive coding (Cris Frith) • Frith のコンパレーター仮説 = predictive coding • In Type I (acute) schizophrenic patients, intentions of will lead to actions, but these willed intentions are not monitored correctly. This apparent discrepancy between will and action gives rise to experiential (1st rank) positive symptoms (e.g. delusions of control and passivity) 27 . • どうして自分をくすぐることはできないの? 28 • 自分の行動による感覚への影響を予測してしま っているから。 • 時間差によって予測不可能にすると自分をくす ぐることができる! • 統合失調症患者は自分をくすぐることができる 29 • 自分の行動による感覚への影響をキャンセルア ウトすることに失敗している=> Agency (主体感) の失調 • 自己運動の結果をキャンセルしている脳部位: SII, ACC 30 2-3. Bayesian brain (Karl Friston) • 統合失調症での幻覚、妄想のベイズ脳的説明 • What do you see in this scene? (Schizophrenia Block Course 2012 TNU Zurich 05_Computational Models Studying Schizophrenia - an Overview.pdf) • Normal vision: sensory evidence に対して prior が適切な prediction を作ることによって prediction error=0 にする。 • Delusion(妄想): 適切な prediction を作ったのに prediction error=0 にならない。そこで prior を変 えて prediction error=0 にした。 • Hallucination(幻覚)の場合: sensory evidence な しで prior が不適切な prediction を作るが prediction error=0 になっている。 • Hollow-mask illusion • 統合失調症患者は hollow-mask にだまされない (Schizophrenia Block Course 2012 TNU Zurich 11_Modeling perceptual abnormalities.pdf) • 統合失調症患者は depth-inversion にだまされない 31 • Normal: prior > evidence • Schizophrenia: prior < evidence • 機能的結合:ERP を DCM によって解析 32 • 健常群のデータ:=> IPS->LOC のトップダウンの 情報伝達 • 統合失調症患者のデータ:=> V1->LOC のボトム アップの情報伝達 2-4. Aberrant salience (Shitij Kapur) • 発症初期に注目する • At-risk mental state (ARMS) • Ultra-high risk - transition rate: 35% and 40% within 12 months 33 • Prodromal (100%) - retrospective • First episode • 中安による「初期分裂病」の概念 • 1.自生思考: 「自分で意識して考えていることと 無関係な考えが,急に発作的にどんどん押し寄せ てくる」 • 2.気付き亢進: 「他人の声や不意の音,たとえば 戸を開閉する音や近くを走る電車の音などを聞 くとビクッとして落ち着かなくなる」 • 3.緊迫困惑気分: 「何かが差し迫っているようで 緊張を要するものの,なぜそんな気持ちになるの か分からなくて戸惑っている」 • 4.即時的認知の障害 - 即時理解の障害: 「他人 の話の内容,テレビの内容などが理解しにくくて, なかなか頭に入らない」 • 統合失調症から回復した当事者が前駆期の神経症 状について回顧的に記述した例 • 「いちばん最初に起きたのは、私の脳の眠ってい た部分の一部が目覚めて、さまざなま人、出来事、 場所、考えに対して興味が惹かれるようになった ことです。それらは普段だったらなんの印象も憶 えないようなものでした。」 • 「全てのことになにか圧倒的なまでに意味深い ものがあるように思えるのです…知らない人が
  5. 5. 道を歩いているのを見ると、そこに私が解釈しな ければならないなにか徴(sign)があるように思え ました」 • 「私が入院をした頃には、窓枠の光や空の青さが あまりに重要な意味を持ちすぎて叫びたくなる ような、そんな「覚醒状態」の段階にまで到達し ていました」 • => サリエンシーの亢進? • Aberrant salience hypothesis of Psychosis 34 • (2)->(3): During the prodromal period (前駆期) there is a context-independent firing of DA neurons and subsequent DA release. This produces a perplexing sense of novelty in patients. Patients continue to accumulate several experiences of altered novelty and salience without a clear explanation for them. • (3)->(4): The perplexing sense crystallizes into a delusion – and then it all ‘makes sense’ to the patient. A delusion is a ‘top-down’ cognitive explanation that the individual imposes on these aberrant novelty and salience experiences in an effort to make sense of them. • (4)->(5) : These delusions or hallucinations impact on the patient's behavior, and this is typically when patients are brought to care and antipsychotics are administered. • (6)->(8) : Antipsychotics, by blocking dopamine transmission, attenuate aberrant salience. Antipsychotics do not directly erase delusions but (with it) new aberrant salience is less likely to form. They do not immediately abandon the delusion or hallucination but instead report that it ‘doesn't bother me as much anymore’. • How to test aberrant salience? - “Salience attribution task” (SAT) 35 • The individual at risk assigns a meaning to irrelevant cues 2-5. Visual saliency への影響 (ヒト患者) • 統合失調症研究のストラテジー • 行動指標の選択- ヒト、動物の両方で評価可能, 神経回路解明につながる • フリービューイング課題 36 • 仮説:眼球運動計測によってサリエンシーの異常が 検出できるのでは? • 仮説:統合失調症であるとは?視覚サリエンシーが 亢進し、すべてが意味ありげに見える(動機サリエ ンシーが高い) • フリービューイング課題 • 統制群では、視線はサリエンシーの高いところに 向かう。それからよりサリエンシーの低いところ に向かう。 • 統合失調症患者では、視線がサリエンシーの高い ところに誘引されやすい。「異常サリエンス説」 と整合的。 2-6. Visual saliency への影響 (動物モデル) • トランスレータブル指標に向けて • 統合失調症のモデル動物で、患者で見いだされた 視線パターンは再現されるか? • どの脳部位がこのような視線パターンに寄与し ているか? • そもそもマーモセットで視線計測は可能か? 37 3. Presence (現前性、存在感) 3-1. Presence とは? • the concept of presence is used to refer to the subjective sense of reality of the world and of the self within the world 38 • Depersonalization disorder (DPD): • a subjective feeling of internal and/or external change, experienced as one of strangeness or unreality” • chronic expression of symptoms • Depersonalization: loss of subjective sense of reality of the self • Derealization: loss of subjective sense of reality of the world • 仮説:DPD であるとは? • サリエンシーの低下 =>自分が世界の外にあるよ うに思える • Presence and predictive coding 38 • agency - presence は互いに影響を与え合う。 • Presence is the result of successful suppression by top-down predictions of informative interoceptive signals evoked (directly) by autonomic control signals and (indirectly) by bodily responses to afferent sensory signals 3-2. サリエンスネットワーク • Insula and awareness 39 • the cortical basis for awareness is an ordered set of representations of all feelings at each immediate moment extending across a finite period of time. • The key to the cortical (that is, mental) representation of the sentient self is the integration of salience across all relevant conditions at each moment. • Salience network 40 • Composed of dosal anterior insula and ACC. • Salience network works as dynamic switching between central-executive network (for externally directed action) and default mode network (for internally directed action) • 統合失調症とサリエンスネットワーク 41 • 初期症状の段階でサリエンスネットワーク (AI-ACC)の灰白質の容量が小さい References
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