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遗传学 1910

果蝇中的限性遗传

托马斯·亨特·摩尔根

一只白眼果蝇,把基因钉在了染色体上——遗传从此成了能画出地图的东西。

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In depth · the introduction

一只生来眼色不对的果蝇,把遗传从一本抽象的规则书,变成了一张你真能画出来的地图。

核心想法

孟德尔已经表明,性状是作为一个个离散的「因子」传下去的,可没人知道这些因子住在哪里。摩尔根找到了答案——靠的是一次偶然。在成千上万只红眼果蝇之中,冒出了一只白眼的雄蝇;而那白眼的遗传方式,唯有当它的因子搭乘在某一条特定染色体——X,也就是帮着决定果蝇是雄是雌的那条——之上时,才说得通。

由于 X 的传递方式,儿子那唯一的一份拷贝只能来自母亲。于是,藏在 X 上的隐性性状可以跳过女儿,到孙辈的雄性身上现身——这正是摩尔根看到的那个古怪模式。基因不再是一个记账符号;它有了一个实在的家。

它是如何诞生的

故事的舞台,是哥伦比亚大学里一间狭小、飘着香蕉味的实验室,人人称它「蝇室」。摩尔根其实是个怀疑者——他既怀疑孟德尔的因子,也怀疑「染色体携带遗传」这个想法,养果蝇,多少是为了检验这些说法。然后,在 1910 年,那只孤零零的白眼雄蝇出现了。摩尔根和他的合作者们——包括他的妻子、生物学家莉莲·沃恩·摩尔根,以及一群出色的学生——把它繁育、把它的后代数清、把那个模式读了出来。怀疑者,成了染色体理论最有力的拥护者。

它为何重要

这就是那个给了基因一个地址的实验。它把两个一直彼此绕圈的想法焊在了一起——孟德尔那些看不见的因子,与生物学家在显微镜下真能看见的染色体——并把遗传学变成了一门关于位置与地图的科学。它还头一回解释了,为什么色盲、血友病这类人类病症会从母亲传给儿子,却放过女儿。

一个可以想象的画面

把每条染色体想成图书馆里某一层特定的书架,把基因想成架上的一本书。白眼这本书,放在 X 这层架子上。女儿继承两层 X 架子,父母各给一层,所以单单一本白眼书,很容易被一本正常的书盖过。可儿子只继承一层 X 架子——而且只来自母亲。要是她那层架子上摆着白眼书,又没有别的来推翻它,他读到的就是白眼。用下面的杂交工具,换一换哪位亲本是白眼,看看后代的模式怎样翻转。

一张杂交图,两个按钮用来选择亲本方向。三行——亲本、子代、孙代——画出雌、雄果蝇的红眼或白眼,让你看到白眼如何随着哪位亲本携带它而消失、又重现。

它的位置

摩尔根的果蝇,居于一条伟大链条的中段。孟德尔(1866)找到了遗传的规则,却不知道载体是谁;摩尔根把那些规则安放到了染色体上;又过了四十年,埃弗里(1944),而后沃森与克里克(1953),才揭示出那载体究竟由什么构成——DNA。从蝇室里起步的那些遗传图谱,一路不断地延伸到今天的人类基因组计划。

The original document
Original source text
T. H. Morgan · Science, new series, 32 (812): 120–122 · July 22, 1910
The paper opens not with a theory but with an accident — a single fly that should not have looked the way it did:
In a pedigree culture of Drosophila which had been running for nearly a year through a considerable number of generations, a male appeared with white eyes.
The crosses
Morgan bred the white-eyed male to its red-eyed sisters. The entire first generation (F₁) had the ordinary brilliant red eyes — white had vanished, behaving as a simple recessive. He then let the red-eyed F₁ flies interbreed.
In the second generation (F₂) white reappeared — but with a twist no autosomal factor could produce. Morgan's tally was 2459 red-eyed females, 1011 red-eyed males, and 782 white-eyed males. Not one white-eyed female appeared. (The white flies fell short of a clean 3 : 1 ratio because, as Morgan noted, white-eyed individuals were the weaker and died off more often.)
[ … ]
The explanation
Morgan proposed that the factor for red eye-colour is carried together with the factor that determines sex — what he wrote as “X.” A daughter receives an X from each parent; a son receives his single X only from his mother, and a Y from his father. The white factor, riding on the X, therefore could not reach a son except through his mother — which is exactly why white eyes surfaced only in F₂ males.
The reciprocal cross sealed it. Mate a red-eyed male to a white-eyed female and the result inverts: every F₁ daughter is red and every F₁ son is white — the “criss-cross” of a trait passing from mother to son. Ordinary Mendelian characters give identical results whichever parent carries them; a character that does not is one tied to the sex chromosome.
T. H. Morgan · Columbia University, New York · 1910