When on board H.M.S. “Beagle,” as naturalist, I was much struck with certain facts in the distribution of the inhabitants of South America, and in the geological relations of the present to the past inhabitants of that continent. These facts seemed to me to throw some light on the origin of species—that mystery of mysteries, as it has been called by one of our greatest philosophers. On my return home, it occurred to me, in 1837, that something might perhaps be made out on this question by patiently accumulating and reflecting on all sorts of facts which could possibly have any bearing on it. After five years' work I allowed myself to speculate on the subject, and drew up some short notes; these I enlarged in 1844 into a sketch of the conclusions, which then seemed to me probable: from that period to the present day I have steadily pursued the same object. I hope that I may be excused for entering on these personal details, as I give them to show that I have not been hasty in coming to a decision.
My work is now nearly finished; but as it will take me two or three more years to complete it, and as my health is far from strong, I have been urged to publish this Abstract. I have more especially been induced to do this, as Mr. Wallace, who is now studying the natural history of the Malay archipelago, has arrived at almost exactly the same general conclusions that I have on the origin of species. Last year he sent to me a memoir on this subject, with a request that I would forward it to Sir Charles Lyell, who sent it to the Linnean Society, and it is published in the third volume of the Journal of that Society. Sir C. Lyell and Dr. Hooker, who both knew of my work—the latter having read my sketch of 1844—honoured me by thinking it advisable to publish, with Mr. Wallace's excellent memoir, some brief extracts from my manuscripts.
This Abstract, which I now publish, must necessarily be imperfect. I cannot here give references and authorities for my several statements; and I must trust to the reader reposing some confidence in my accuracy. No doubt errors will have crept in, though I hope I have always been cautious in trusting to good authorities alone. I can here give only the general conclusions at which I have arrived, with a few facts in illustration, but which, I hope, in most cases will suffice. No one can feel more sensible than I do of the necessity of hereafter publishing in detail all the facts, with references, on which my conclusions have been grounded; and I hope in a future work to do this. For I am well aware that scarcely a single point is discussed in this volume on which facts cannot be adduced, often apparently leading to conclusions directly opposite to those at which I have arrived. A fair result can be obtained only by fully stating and balancing the facts and arguments on both sides of each question; and this cannot possibly be here done.
I much regret that want of space prevents my having the satisfaction of acknowledging the generous assistance which I have received from very many naturalists, some of them personally unknown to me. I cannot, however, let this opportunity pass without expressing my deep obligations to Dr. Hooker, who for the last fifteen years has aided me in every possible way by his large stores of knowledge and his excellent judgment.
In considering the Origin of Species, it is quite conceivable that a naturalist, reflecting on the mutual affinities of organic beings, on their embryological relations, their geographical distribution, geological succession, and other such facts, might come to the conclusion that each species had not been independently created, but had descended, like varieties, from other species. Nevertheless, such a conclusion, even if well founded, would be unsatisfactory, until it could be shown how the innumerable species inhabiting this world have been modified, so as to acquire that perfection of structure and coadaptation which most justly excites our admiration. Naturalists continually refer to external conditions, such as climate, food, etc., as the only possible cause of variation. In one very limited sense, as we shall hereafter see, this may be true; but it is preposterous to attribute to mere external conditions, the structure, for instance, of the woodpecker, with its feet, tail, beak, and tongue, so admirably adapted to catch insects under the bark of trees. In the case of the misseltoe, which draws its nourishment from certain trees, which has seeds that must be transported by certain birds, and which has flowers with separate sexes absolutely requiring the agency of certain insects to bring pollen from one flower to the other, it is equally preposterous to account for the structure of this parasite, with its relations to several distinct organic beings, by the effects of external conditions, or of habit, or of the volition of the plant itself.
The author of the “Vestiges of Creation” would, I presume, say that, after a certain unknown number of generations, some bird had given birth to a woodpecker, and some plant to the misseltoe, and that these had been produced perfect as we now see them; but this assumption seems to me to be no explanation, for it leaves the case of the coadaptations of organic beings to each other and to their physical conditions of life, untouched and unexplained.
It is, therefore, of the highest importance to gain a clear insight into the means of modification and coadaptation. At the commencement of my observations it seemed to me probable that a careful study of domesticated animals and of cultivated plants would offer the best chance of making out this obscure problem. Nor have I been disappointed; in this and in all other perplexing cases I have invariably found that our knowledge, imperfect though it be, of variation under domestication, afforded the best and safest clue. I may venture to express my conviction of the high value of such studies, although they have been very commonly neglected by naturalists.
From these considerations, I shall devote the first chapter of this Abstract to Variation under Domestication. We shall thus see that a large amount of hereditary modification is at least possible; and, what is equally or more important, we shall see how great is the power of man in accumulating by his Selection successive slight variations. I will then pass on to the variability of species in a state of nature; but I shall, unfortunately, be compelled to treat this subject far too briefly, as it can be treated properly only by giving long catalogues of facts. We shall, however, be enabled to discuss what circumstances are most favourable to variation. In the next chapter the Struggle for Existence amongst all organic beings throughout the world, which inevitably follows from their high geometrical powers of increase, will be treated of. This is the doctrine of Malthus, applied to the whole animal and vegetable kingdoms. As many more individuals of each species are born than can possibly survive; and as, consequently, there is a frequently recurring struggle for existence, it follows that any being, if it vary however slightly in any manner profitable to itself, under the complex and sometimes varying conditions of life, will have a better chance of surviving, and thus be naturally selected. From the strong principle of inheritance, any selected variety will tend to propagate its new and modified form.
This fundamental subject of Natural Selection will be treated at some length in the fourth chapter; and we shall then see how Natural Selection almost inevitably causes much Extinction of the less improved forms of life, and induces what I have called Divergence of Character. In the next chapter I shall discuss the complex and little known laws of variation and of correlation of growth. In the four succeeding chapters, the most apparent and gravest difficulties on the theory will be given: namely, first, the difficulties of transitions, or in understanding how a simple being or a simple organ can be changed and perfected into a highly developed being or elaborately constructed organ; secondly, the subject of Instinct, or the mental powers of animals; thirdly, Hybridism, or the infertility of species and the fertility of varieties when intercrossed; and fourthly, the imperfection of the Geological Record. In the next chapter I shall consider the geological succession of organic beings throughout time; in the eleventh and twelfth, their geographical distribution throughout space; in the thirteenth, their classification or mutual affinities, both when mature and in an embryonic condition. In the last chapter I shall give a brief recapitulation of the whole work, and a few concluding remarks.
No one ought to feel surprise at much remaining as yet unexplained in regard to the origin of species and varieties, if he makes due allowance for our profound ignorance in regard to the mutual relations of all the beings which live around us. Who can explain why one species ranges widely and is very numerous, and why another allied species has a narrow range and is rare? Yet these relations are of the highest importance, for they determine the present welfare, and, as I believe, the future success and modification of every inhabitant of this world. Still less do we know of the mutual relations of the innumerable inhabitants of the world during the many past geological epochs in its history. Although much remains obscure, and will long remain obscure, I can entertain no doubt, after the most deliberate study and dispassionate judgment of which I am capable, that the view which most naturalists entertain, and which I formerly entertained—namely, that each species has been independently created—is erroneous. I am fully convinced that species are not immutable; but that those belonging to what are called the same genera are lineal descendants of some other and generally extinct species, in the same manner as the acknowledged varieties of any one species are the descendants of that species. Furthermore, I am convinced that Natural Selection has been the main but not exclusive means of modification.
搭乘皇家“貝格爾”號(hào)周游世界時(shí),身為博物學(xué)者,我對(duì)南美洲的生物分布以及現(xiàn)存生物和古生物的地質(zhì)關(guān)系頗為留意。某些情況似乎讓我對(duì)物種起源略有所悟——這是個(gè)謎中謎,正如一位極偉大的哲學(xué)家說(shuō)過(guò)的?;貒?guó)后的1837年,我靈機(jī)一動(dòng),耐心深入搜集有關(guān)的資料,加以融會(huì)貫通,這個(gè)問(wèn)題也許可以有所心得的嘛。經(jīng)過(guò)五年努力之后,我斗膽對(duì)該主題進(jìn)行了思辨,并記了些簡(jiǎn)短的筆記;1844年又把它擴(kuò)充為一份結(jié)論提綱,當(dāng)時(shí)在我看來(lái)算是有點(diǎn)眉目了。從那以后,我始終不渝,孜孜以求。希望讀者原諒我扯個(gè)人的瑣事,說(shuō)出來(lái)是為了表明,我的決定并非草率做出。
如今,我的研究工作即將告一段落,但離徹底完成還需投入兩三年時(shí)間,而且我現(xiàn)在身體不算強(qiáng)壯,便有人勸我先發(fā)表了這份摘要再說(shuō)。特別促使我這樣做的,是研究馬來(lái)群島博物學(xué)的華萊士先生對(duì)于物種起源所做的一般結(jié)論,竟幾乎和我不謀而合。去年,他把一份有關(guān)本主題的研究報(bào)告寄給了我,要求我轉(zhuǎn)交查爾斯·賴爾(Charles Lyell)爵士,爵士把它交給了林奈學(xué)會(huì),刊登在該學(xué)會(huì)第三卷學(xué)報(bào)上。賴爾爵士和胡克博士都了解我的研究,胡克還讀過(guò)我1844年寫(xiě)的綱要,他們認(rèn)為最好把我原稿的若干章節(jié)和華萊士先生的優(yōu)秀論文同時(shí)發(fā)表。我不勝榮幸。
現(xiàn)在發(fā)表的這個(gè)摘要必定不夠完善。這里無(wú)法為我的若干論述提出參考文獻(xiàn)和權(quán)威典籍,有必要拜托讀者對(duì)我的論述精確性有所信任。錯(cuò)誤在所難免,雖然自認(rèn)一貫小心謹(jǐn)慎,只信賴可靠的典籍。本書(shū)僅僅能給出我所得到的一般結(jié)論,用少量事實(shí)來(lái)做實(shí)例,希望在大多數(shù)情況下這就足夠了。今后有必要把我做結(jié)論所依據(jù)的全部事實(shí)以及參考資料一五一十發(fā)表出來(lái),這一點(diǎn)我比誰(shuí)都念念不忘的;希望將來(lái)的著作中能做到。我很清楚,本書(shū)所討論的,幾乎沒(méi)有一點(diǎn)不能用事實(shí)來(lái)舉證,而絕不會(huì)引出同我的結(jié)論直接背道而馳的東西。只有對(duì)每一個(gè)問(wèn)題的正反兩方面事實(shí)和論點(diǎn)加以充分論述,反復(fù)權(quán)衡,才能得出公平的結(jié)果,但這里做不到。
我得到了許多學(xué)者的慷慨相助,其中有些素不相識(shí);很遺憾,由于篇幅的限制,無(wú)法一一鳴謝。然而,機(jī)會(huì)難得,一定要對(duì)胡克博士深切致謝,最近十五年來(lái),他以豐富的學(xué)識(shí)和卓越的判斷力,千方百計(jì)鼎力相助。
關(guān)于物種起源,學(xué)者們?nèi)绻麑?duì)生物的相互親緣關(guān)系、胚胎關(guān)系、地理分布、地質(zhì)演替等等加以思考,那就可以想見(jiàn)會(huì)得出如下結(jié)論:物種不是獨(dú)立創(chuàng)造出來(lái)的,而是與變種一樣,是從其他物種傳承下來(lái)的。然而,這一結(jié)論即使有根有據(jù),也不能令人滿意,除非我們能夠證明,這個(gè)世界的無(wú)數(shù)物種如何變異才獲得了令人贊不絕口的完善構(gòu)造和相互適應(yīng)性。學(xué)者們始終把可能的變異僅僅歸因于外界條件,如氣候、食物等。從某一狹義來(lái)說(shuō),正如后文即將看到的,這可能是正確的;但是,例如把啄木鳥(niǎo)的構(gòu)造,它的腳、尾、喙、舌,如此絕妙地適應(yīng)于在樹(shù)皮下捉昆蟲(chóng),也僅僅歸因于外界條件,則是十分荒謬的。再如槲寄生,它從某些樹(shù)木吸取營(yíng)養(yǎng),種子必須由某些鳥(niǎo)傳播,而且是雌雄異花,絕對(duì)需要借助某些昆蟲(chóng)來(lái)完成異花授粉。用外界條件、習(xí)性或植株本身的意志作用來(lái)解釋這種寄生生物的構(gòu)造以及它和若干種不同生物的關(guān)系,也同樣荒謬絕倫。
我想,“創(chuàng)世遺跡”論者會(huì)斷言,經(jīng)過(guò)不計(jì)其數(shù)的世代,某鳥(niǎo)生下了啄木鳥(niǎo),某植物生下了槲寄生,且創(chuàng)造得如我們所見(jiàn)的一樣完美;但依我看,這種假設(shè)無(wú)法自圓其說(shuō),未觸及和解釋生物的相互適應(yīng)性,以及對(duì)其生活條件的適應(yīng)性。
因此,弄清變異和適應(yīng)的途徑至關(guān)重要。剛開(kāi)始觀察時(shí),我就覺(jué)得仔細(xì)研究馴養(yǎng)動(dòng)物和栽培植物,對(duì)于解決這個(gè)難題也許會(huì)提供最好的機(jī)會(huì)。果然功夫不負(fù)有心人,通過(guò)這種和所有其他的復(fù)雜個(gè)例,我一個(gè)勁兒地發(fā)現(xiàn),有關(guān)馴養(yǎng)變異的知識(shí)即使不完善,也能提供最好、最可靠的線索。我在此斗膽聲明,我堅(jiān)信這種研究?jī)r(jià)值很高,雖然學(xué)者們往往加以忽視。
有鑒于此,本書(shū)第一章討論馴養(yǎng)變異。我們將看到,大量的遺傳變異至少是可能的,更有甚者,我們將看到,人類通過(guò)選擇積累連續(xù)的微小變異,能耐是何等巨大。然后將討論物種在自然狀況下的變異性;不幸的是,討論這個(gè)問(wèn)題不得不簡(jiǎn)而又簡(jiǎn),因?yàn)橹挥辛_列長(zhǎng)篇的事實(shí)才能加以妥當(dāng)處理。然而,我們還能得以討論什么環(huán)境條件最有利于變異。第三章討論全世界所有生物之間的生存斗爭(zhēng),這是以幾何級(jí)數(shù)高度增殖的必由之路。這就是馬爾薩斯(Malthus)學(xué)說(shuō)在整個(gè)動(dòng)物界和植物界的應(yīng)用。每一物種所出生的個(gè)體,大大超過(guò)其可能生存的數(shù)量,于是生存斗爭(zhēng)反復(fù)出現(xiàn),結(jié)果任何生物所發(fā)生的變異,無(wú)論多么微小,只要在復(fù)雜多變的生活條件下以任何方式有利于自身,就會(huì)有較好的生存機(jī)會(huì),這樣便被自然選擇了。根據(jù)強(qiáng)有力的遺傳原理,任何被選中的變種都傾向于繁殖其變異了的新形態(tài)。
自然選擇的基本問(wèn)題在第四章詳述;我們將看到,自然選擇幾乎不可避免地導(dǎo)致較少改進(jìn)的生物類型大量絕滅,并且引發(fā)我所謂的“性狀分歧”(Divergence of Character)。第五章討論復(fù)雜的、不為人知的變異法則和相關(guān)生長(zhǎng)法則。接下來(lái)的四章將對(duì)本學(xué)說(shuō)所存在的最明顯最重大的難點(diǎn)加以討論:第一,過(guò)渡的難點(diǎn),也就是難以了解簡(jiǎn)單生物或簡(jiǎn)單器官如何變化和改善成高度發(fā)展的生物或構(gòu)造精密的器官;第二,本能的問(wèn)題,即動(dòng)物的精神力;第三,雜交現(xiàn)象,即物種雜交的不育性和變種雜交的能育性;第四,地質(zhì)記錄不完全。第十章考察生物在整個(gè)時(shí)間上的地質(zhì)演替。第十一章和第十二章討論生物在整個(gè)空間上的地理分布。第十三章論述生物的分類或相互的親緣關(guān)系,包括成熟期和胚胎期。最后一章對(duì)全書(shū)做一扼要的復(fù)述,加上簡(jiǎn)短的結(jié)束語(yǔ)。
只要承認(rèn)對(duì)周圍全部生物的相互關(guān)系是多么無(wú)知,關(guān)于物種和變種的起源至今還不甚了了,就不足為奇了。誰(shuí)能解釋某一個(gè)物種為什么分布范圍廣而且為數(shù)眾多,而另一個(gè)近緣物種為什么分布范圍狹而為數(shù)稀少?這種關(guān)系至關(guān)重要,決定著世界一切生物現(xiàn)在的繁盛,并且我相信也決定著它們未來(lái)的成功和變異。至于世界上無(wú)數(shù)生物在史上諸多既往地質(zhì)時(shí)代里的相互關(guān)系,我們就所知甚少了。雖然諸多問(wèn)題至今模糊不清,而且還會(huì)長(zhǎng)期如此,但經(jīng)過(guò)盡可能從容的斟酌研究和冷靜判斷,我毫不懷疑,許多學(xué)者還保持著的和我以前所持的觀點(diǎn)——即每一物種都是獨(dú)立創(chuàng)造出來(lái)的——是錯(cuò)誤的。我完全相信,物種不是一成不變的。那些所謂同屬的物種都是其單元屬已然滅絕的另一物種的直系后裔,正如任何一個(gè)物種的公認(rèn)變種乃是那個(gè)物種的后裔一樣。另外,我還相信自然選擇是變異的主要而非唯一的途徑。
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