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遺傳學 1910

果蠅中的限性遺傳

湯瑪斯·亨特·摩根

一隻白眼果蠅,把基因釘在了染色體上——遺傳從此成了能畫出地圖的東西。

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In depth · the introduction

一隻生來眼色不對的果蠅,把遺傳從一本抽象的規則書,變成了一張你真能畫出來的地圖。

核心想法

孟德爾已經表明,性狀是作為一個個離散的「因子」傳下去的,可沒人知道這些因子住在哪裡。摩爾根找到了答案——靠的是一次偶然。在成千上萬隻紅眼果蠅之中,冒出了一隻白眼的雄蠅;而那白眼的遺傳方式,唯有當牠的因子搭乘在某一條特定染色體——X,也就是幫著決定果蠅是雄是雌的那條——之上時,才說得通。

由於 X 的傳遞方式,兒子那唯一的一份拷貝只能來自母親。於是,藏在 X 上的隱性性狀可以跳過女兒,到孫輩的雄性身上現身——這正是摩爾根看到的那個古怪模式。基因不再是一個記帳符號;牠有了一個實在的家。

它是如何誕生的

故事的舞台,是哥倫比亞大學裡一間狹小、飄著香蕉味的實驗室,人人稱它「蠅室」。摩爾根其實是個懷疑者——他既懷疑孟德爾的因子,也懷疑「染色體攜帶遺傳」這個想法,養果蠅,多少是為了檢驗這些說法。然後,在 1910 年,那隻孤零零的白眼雄蠅出現了。摩爾根和他的合作者們——包括他的妻子、生物學家莉蓮·沃恩·摩爾根,以及一群出色的學生——把牠繁育、把牠的後代數清、把那個模式讀了出來。懷疑者,成了染色體理論最有力的擁護者。

它為何重要

這就是那個給了基因一個地址的實驗。它把兩個一直彼此繞圈的想法焊在了一起——孟德爾那些看不見的因子,與生物學家在顯微鏡下真能看見的染色體——並把遺傳學變成了一門關於位置與地圖的科學。它還頭一回解釋了,為什麼色盲、血友病這類人類病症會從母親傳給兒子,卻放過女兒。

一個可以想像的畫面

把每條染色體想成圖書館裡某一層特定的書架,把基因想成架上的一本書。白眼這本書,放在 X 這層架子上。女兒繼承兩層 X 架子,父母各給一層,所以單單一本白眼書,很容易被一本正常的書蓋過。可兒子只繼承一層 X 架子——而且只來自母親。要是她那層架子上擺著白眼書,又沒有別的來推翻牠,他讀到的就是白眼。用下面的雜交工具,換一換哪位親本是白眼,看看後代的模式怎樣翻轉。

一張雜交圖,兩個按鈕用來選擇親本方向。三行——親本、子代、孫代——畫出雌、雄果蠅的紅眼或白眼,讓你看到白眼如何隨著哪位親本攜帶牠而消失、又重現。

它的位置

摩爾根的果蠅,居於一條偉大鏈條的中段。孟德爾(1866)找到了遺傳的規則,卻不知道載體是誰;摩爾根把那些規則安放到了染色體上;又過了四十年,埃弗里(1944),而後沃森與克里克(1953),才揭示出那載體究竟由什麼構成——DNA。從蠅室裡起步的那些遺傳圖譜,一路不斷地延伸到今天的人類基因組計畫。

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