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神經科學 1906

神經系統的整合作用

查爾斯·斯科特·謝靈頓

神經系統不是一塊配電板,而是一台整合機——反射是它的單元,突觸是它的閥門。

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In depth · the introduction

你的身體由數十億個各自獨立的細胞構成,卻能作為一個整體行動。謝靈頓追問這是如何做到的——答案,竟藏在一個卑微如反射的東西裡。

核心想法

膝跳反射,手從燙爐上猛地縮回——這些反射看起來簡單得無足輕重。查爾斯·謝靈頓卻把它們看作理解整個神經系統的鑰匙。他意識到,大腦與脊髓並不只是像電線那樣傳遞信號;它們在整合信號——把一些相加,把另一些抵消,將一股相互競爭的信息洪流,化為一個被協調起來的動作。

通過研究反射,他推算出了一個神經細胞與下一個神經細胞「對話」之處的法則——那就是突觸,一個他幫忙創造的詞。他表明:要彎曲一個關節,身體會在興奮一塊肌肉的同時,主動關掉它相對的那塊(交互支配);而許多信號會匯集到通往肌肉的同一條「最終公路」上,它一次卻只能聽從其中一個。

它是如何誕生的

大約在 1900 年,科學家剛剛弄清,神經系統是由一個個獨立的細胞——神經元——構成的,而非一張連續的網。但沒人知道,這些細胞是如何協同產生協調動作的。謝靈頓,一位英國生理學家,自下而上地攻克這個難題,對脊髓的反射做了一絲不苟的實驗。

他把成果匯成 1904 年在耶魯所做的十場講座,並於 1906 年以《神經系統的整合作用》出版。這本書純粹從反射在時間上的表現,一路推理到突觸的種種性質,而那時還沒人能看見一個突觸。它重塑了生理學;1932 年,謝靈頓因這一脈工作分享了諾貝爾獎。

它為何重要

謝靈頓把神經系統從一塊配電板,變成了某種會在微觀層面「思考」的東西——一台權衡相互競爭的訴求、再做出決定的機器。他那個「神經細胞把興奮與抑制相加、然後才動作」的想法,是每一個神經元如何工作的基本圖景,甚至也是現代人工智慧裡那些人工「神經元」被設想出來的樣子。交互支配與最終公路,至今仍是每一位醫生、每一位研究運動的學生必學的內容。

一個可以想像的畫面

想像一間擠滿人的屋子,只有一道門,許多人同時朝守門人下達命令——有人使勁要把門推開,有人拼命要把門拉上。守門人沒法聽從所有人;他把「推」相加、把「拉」相減,只做那個由這筆總帳決定的動作。運動神經元就是這位守門人,就是那條「最終公路」;每一塊肌肉,都由同時抵達的全部興奮與全部抑制的「流水總帳」來驅動。

可互動的反射:兩個滑桿設定有多少興奮與抑制抵達同一個運動神經元;只有當興奮勝出足夠多,神經元才發放,使肢體屈曲、相對的肌肉放鬆。

它的位置

謝靈頓,立在現代腦科學的源頭。在他之前不久,聖地亞哥·拉蒙·伊·卡哈爾證明神經系統由一個個獨立的細胞構成;謝靈頓則表明這些細胞如何協同行動。在他之後,艾倫·霍奇金與安德魯·赫胥黎——也在這座圖書館裡——破解了沿單個神經元飛馳的電脈衝,而他自己的學生約翰·埃克爾斯,揭示了他當年只能推斷的那個突觸的化學。他那個興奮與抑制的平衡,是這座圖書館裡每一個「把眾多輸入整合為單一輸出」的系統的近親,一直延伸到今天人工智慧背後的那個人工神經元。

The original document
Original source text
Charles S. Sherrington · The Integrative Action of the Nervous System · New York: Charles Scribner's Sons, 1906 · the ten Silliman Lectures delivered at Yale University, 1904
The lectures set out from a single question: how does a body made of countless separate cells act as one individual? Sherrington's answer is that the nervous system integrates — it binds the parts into a coordinated whole — and that the elementary act of integration can be studied in its simplest form, the reflex. The book builds, lecture by lecture, from that unit up to the unified animal.
The simple reflex as the unit of integration
He treats the reflex — a stimulus at a receptor producing a coordinated response — as the indivisible unit of nervous action, while warning that a truly 'simple' reflex is a convenient abstraction: in the living animal no reaction stands alone, each is shaped by all the others competing for the same machinery.
The synapse: a one-way valve between neurons
From the timing of reflexes — their delay, their one-way conduction, their fatigue and summation — Sherrington infers that the junction between one neuron and the next has special properties, unlike conduction along a nerve fibre. For this junction he had earlier established the name synapse. He could not see it; he reasoned it into existence from how reflexes behave.
Reciprocal innervation of antagonistic muscles
A limb's flexor and extensor pull against each other, so to bend a joint the body must both excite one muscle and relax its antagonist. Sherrington shows that the same reflex that excites a muscle simultaneously inhibits its opponent — reciprocal innervation — making inhibition as active and central a process as excitation.
The final common path
Many separate reflex arcs converge on the same motor neuron running out to a muscle. That motoneuron is the final common path: it must serve all the arcs that claim it, and at any instant it can obey only one, so the convergence forces the nervous system to choose — to integrate competing demands into a single coordinated act.
[ … ]
From these pieces the later lectures assemble the coordinated reflexes — the scratch-reflex, the flexion and crossed-extension reflexes of standing and stepping, postural tone — and close on the nervous system as the organ that compounds reflexes into the behaviour of the whole individual.
Liverpool · 1906