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物种起源|The Origin of Species

第七章　本能|CHAPTER VII　INSTINCT

属类： 双语小说 【分类】其他读物 阅读：[7133]
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本能与习性可比，但起源不同——本能的分级——蚜虫和蚁——本能是变异的——家养的本能，其起源——杜鹃、鸵鸟以及寄生蜂的自然本能——蓄奴蚁——蜜蜂，营造蜂房的本能——自然选择学说应用于本能的难点——中性或不育的昆虫——提要

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Instincts comparable with habits, but different in their origin— Instincts graduated—Aphides and ants—Instincts variable— Domestic instincts, their origin—Natural instincts of the cuckoo, ostrich, and parasitic bees—Slave-making ants—Hive-bee, its cell- making instinct—Difficulties on the theory of the Natural Selection of instincts—Neuter or sterile insects—Summary

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本能问题原本可以纳入前面的章节。但我想，单独讨论比较方便，尤其是蜜蜂筑巢的本能是如此奇妙，在许多读者看来大概是一个足以推翻我的全部学说的难点。我先要声明一点，就是我不准备讨论智力的起源，就如我未曾讨论生命本身的起源一样。我们所要讨论的，只是同纲动物中本能的多样性，以及其他精神品质的多样性。

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The subject of instinct might have been worked into the previous chapters; but I have thought that it would be more convenient to treat the subject separately, especially as so wonderful an instinct as that of the hive-bee making its cells will probably have occurred to many readers, as a difficulty sufficient to overthrow my whole theory. I must premise, that I have nothing to do with the origin of the primary mental powers, any more than I have with that of life itself. We are concerned only with the diversities of instinct and of the other mental qualities of animals within the same class.

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我并不想给本能下任何定义。容易阐明，这一术语通常包含着若干不同的精神活动；但是，说本能促使杜鹃迁徙并把蛋下在别种鸟巢里，每一个人都知道这是什么意义。我们自己需要经验才能完成的一种活动，而被一种没有经验的动物，特别是被幼小动物所完成，并且许多个体并不知道为了什么目的却按照同一方式去完成时，一般就称为本能。但是我能阐明，这些性状没有一个是普遍的。如于贝尔（Pierre Huber）所说的，甚至在自然系统中低级的动物里，小量的判断或理性也常发生作用。

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I will not attempt any definition of instinct. It would be easy to show that several distinct mental actions are commonly embraced by this term; but every one understands what is meant, when it is said that instinct impels the cuckoo to migrate and to lay her eggs in other birds’ nests. An action, which we ourselves should require experience to enable us to perform, when performed by an animal, more especially by a very young one, without any experience, and when performed by many individuals in the same way, without their knowing for what purpose it is performed, is usually said to be instinctive. But I could show that none of these characters of instinct are universal. A little dose, as Pierre Huber expresses it, of judgment or reason, often comes into play, even in animals very low in the scale of nature.

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弗·居维叶（Frederick Cuvier）等老一辈玄学者曾把本能与习性加以比较。我想，这一比较对于完成本能活动时的心理状态，提供了精确的概念，但不一定涉及它的起源。许多习惯性活动是在无意识下进行的，甚至不少直接与我们有意识的意志相反！然而意志和理性可以改变它们。习性容易与其他习性，与一定的时期，与身体的状态相联系。习性一经获得，常常终生保持不变。还可以指出本能和习性之间的其他若干类似点。有如反复歌唱一首名曲，在本能里也是一种活动有节奏地随着另一活动；如果一个人歌唱时，或在反复背诵东西时被打断了，一般地他就被迫走回头路，恢复已经成为习惯的思路。于贝尔发现能制造很复杂茧床的毛毛虫（caterpillar）就是如此；如果在完成构造第六阶段时把它抓出，放在只完成构造第三阶段的茧床里，这个毛毛虫仅重筑第四、第五、第六个阶段的构造。然而，如果把完成构造第三阶段的毛毛虫，放在已完成构造第六阶段的茧床里，那么工作已大都完成了，可是它并没有从中感到受益，反而不知所措，并且为了完成茧床，它似乎不得不从构造第三阶段开始，它是在这里放下的，就这样试图去完成已经完成了的工作。

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Frederick Cuvier and several of the older metaphysicians have compared instinct with habit. This comparison gives, I think, a remarkably accurate notion of the frame of mind under which an instinctive action is performed, but not of its origin. How unconsciously many habitual actions are performed, indeed not rarely in direct opposition to our conscious will! yet they may be modified by the will or reason. Habits easily become associated with other habits, and with certain periods of time and states of the body. When once acquired, they often remain constant throughout life. Several other points of resemblance between instincts and habits could be pointed out. As in repeating a well-known song, so in instincts, one action follows another by a sort of rhythm; if a person be interrupted in a song, or in repeating anything by rote, he is generally forced to go back to recover the habitual train of thought: so P. Huber found it was with a caterpillar, which makes a very complicated hammock; for if he took a caterpillar which had completed its hammock up to, say, the sixth stage of construction, and put it into a hammock completed up only to the third stage, the caterpillar simply re-performed the fourth, fifth, and sixth stages of construction. If, however, a caterpillar were taken out of a hammock made up, for instance, to the third stage, and were put into one finished up to the sixth stage, so that much of its work was already done for it, far from feeling the benefit of this, it was much embarrassed, and, in order to complete its hammock, seemed forced to start from the third stage, where it had left off, and thus tried to complete the already finished work.

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假定任何习惯性的活动被遗传——可以指出，有时确有这种情形发生——那么原为习性和原为本能之间，就变得密切相似，难分难解。如果莫扎特不是在三岁时经过极少的练习就能弹奏钢琴，而是全然没有练习就能弹奏一曲，那么可以说他的弹奏确实是出于本能的了。但是假定大多数本能是由一个世代中的习性得来的，然后遗传给后继世代，则是大错特错。能够清楚表明，我们所熟知的最奇异的本能，如蜜蜂和许多蚁的本能，不可能是由习性得来的。

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If we suppose any habitual action to become inherited—and I think it can be shown that this does sometimes happen—then the resemblance between what originally was a habit and an instinct becomes so close as not to be distinguished. If Mozart, instead of playing the pianoforte at three years old with wonderfully little practice, had played a tune with no practice at all, he might truly be said to have done so instinctively. But it would be the most serious error to suppose that the greater number of instincts have been acquired by habit in one generation, and then transmitted by inheritance to succeeding generations. It can be clearly shown that the most wonderful instincts with which we are acquainted, namely, those of the hive-bee and of many ants, could not possibly have been thus acquired.

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人们普遍承认，本能对于各个物种在现今生活条件之下的利益，有如肉体构造一样重要。在多变的生活条件下，本能的微小变异有利于物种，至少是可能的；如果能够指出，本能确曾发生过些许变异，我看自然选择就不难把本能的变异保存下来并继续累积到任何有利的程度。我相信，一切最复杂的和奇异的本能就是这样起源的。使用或习性引起肉体构造的变异，并使之增强，而不使用使之缩小或消失，我并不怀疑本能也是这样的。但我认为，习性的效果，同所谓本能偶发变异的自然选择效果相比是次要的。也就是，产生身体构造的微小偏差有一些未知原因，也产生变异，叫本能偶发变异。

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It will be universally admitted that instincts are as important as corporeal structure for the welfare of each species, under its present conditions of life. Under changed conditions of life, it is at least possible that slight modifications of instinct might be profitable to a species; and if it can be shown that instincts do vary ever so little, then I can see no difficulty in natural selection preserving and continually accumulating variations of instinct to any extent that may be profitable. It is thus, as I believe, that all the most complex and wonderful instincts have originated. As modifications of corporeal structure arise from, and are increased by, use or habit, and are diminished or lost by disuse, so I do not doubt it has been with instincts. But I believe that the effects of habit are of quite subordinate importance to the effects of the natural selection of what may be called accidental variations of instincts;—that is of variations produced by the same unknown causes which produce slight deviations of bodily structure.

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除非经过有益的变异积少成多，缓慢而逐渐的积累，否则复杂的本能不可能通过自然选择而产生。因此，像身体构造的情形一样，我们在自然界中所寻求的不应是获得每一复杂本能的实际过渡诸级，这些只有在各物种的直系祖先里才能找到，但应当从旁系世系里去寻求这些分级的蛛丝马迹，至少能够指出某种分级是可能的；而这一点肯定能够办到。考虑到除了欧洲北美洲以外，动物本能还极少被观察过，并且灭绝物种的本能更是一无所知，我感到惊异的是，最复杂本能所赖以完成的分级能够广泛被发现。“自然界里没有飞跃”的准则适用于本能的力度不亚于身体器官。同一物种在生命的不同时期或一年中的不同季节，或处于不同的环境条件下等等而具有不同的本能，这就往往会促进本能的变化；在这种情况下，自然选择会把这种或那种本能保存下来。可以证明，同一物种中本能的多样性在自然界中也是存在的。

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No complex instinct can possibly be produced through natural selection, except by the slow and gradual accumulation of numerous, slight, yet profitable, variations. Hence, as in the case of corporeal structures, we ought to find in nature, not the actual transitional gradations by which each complex instinct has been acquired—for these could be found only in the lineal ancestors of each species—but we ought to find in the collateral lines of descent some evidence of such gradations; or we ought at least to be able to show that gradations of some kind are possible; and this we certainly can do. I have been surprised to find, making allowance for the instincts of animals having been but little observed except in Europe and North America, and for no instinct being known amongst extinct species, how very generally gradations, leading to the most complex instincts, can be discovered. The canon of “Natura non facit saltum” applies with almost equal force to instincts as to bodily organs. Changes of instinct may sometimes be facilitated by the same species having different instincts at different periods of life, or at different seasons of the year, or when placed under different circumstances, etc.; in which case either one or the other instinct might be preserved by natural selection. And such instances of diversity of instinct in the same species can be shown to occur in nature.

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还有，像身体构造那样，各物种的本能都是为了自己的利益，据我们所能判断的，它从来没有为了其他物种的独享利益而产生过，这和我的理论也是符合的。我知道一个极有力的事例，一种动物的活动从表面看来完全是为了别种动物的利益，这就是蚜虫自愿把甜的分泌物供给蚂蚁：这样做是出于自愿，可由下列事实来说明。我把羊蹄酸模（dock-plant）上的所有蚂蚁从十来只蚜虫堆里搬走，数小时内不让回来。过了这段时间，我确实觉得蚜虫要进行分泌了。我用放大镜观察了一些时候，没有一个分泌的，于是，我尽力模仿蚂蚁用触角的样子，用一根毛轻轻地触动抚摩，但还没有一只蚜虫分泌。随后我让一只蚂蚁去接近它们，从它热切跑动的样子看来，它好像立刻觉得发现了丰盛的食物；于是用触角去拨蚜虫的腹部，先是这一只，然后那一只；蚜虫一接触到触角，即刻举起腹部，分泌出一滴澄清的甜液，蚂蚁慌忙吞食了。甚至十分幼小的蚜虫也有这样的动作，可见这种活动是本能，而不是经验所致。但是，排泄物极黏，被取去也许对于蚜虫是便利的，所以分泌本能大概不是专为蚂蚁的利益。虽然我不相信世界上任何动物会为了其他物种的独享利益而从事活动，然而各物种却试图利用其他物种的本能，正像利用其他物种的虚弱身体构造一样。这样，某些本能就不能看作是绝对完善的；但是详细讨论这一点以及其他类似之点，并非必不可少，所以这里就不赘述了。

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Again as in the case of corporeal structure, and conformably with my theory, the instinct of each species is good for itself, but has never, as far as we can judge, been produced for the exclusive good of others. One of the strongest instances of an animal apparently performing an action for the sole good of another, with which I am acquainted, is that of aphides voluntarily yielding their sweet excretion to ants: that they do so voluntarily, the following facts show. I removed all the ants from a group of about a dozen aphides on a dock-plant, and prevented their attendance during several hours. After this interval, I felt sure that the aphides would want to excrete. I watched them for some time through a lens, but not one excreted; I then tickled and stroked them with a hair in the same manner, as well as I could, as the ants do with their antennae; but not one excreted. Afterwards I allowed an ant to visit them, and it immediately seemed, by its eager way of running about, to be well aware what a rich flock it had discovered; it then began to play with its antennae on the abdomen first of one aphis and then of another; and each aphis, as soon as it felt the antennae, immediately lifted up its abdomen and excreted a limpid drop of sweet juice, which was eagerly devoured by the ant. Even the quite young aphides behaved in this manner, showing that the action was instinctive, and not the result of experience. But as the excretion is extremely viscid, it is probably a convenience to the aphides to have it removed; and therefore probably the aphides do not instinctively excrete for the sole good of the ants. Although I do not believe that any animal in the world performs an action for the exclusive good of another of a distinct species, yet each species tries to take advantage of the instincts of others, as each takes advantage of the weaker bodily structure of others. So again, in some few cases, certain instincts cannot be considered as absolutely perfect; but as details on this and other such points are not indispensable, they may be here passed over.

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本能在自然状态下有某种程度的变异以及这些变异的遗传，既然是自然选择的作用所不可少的，那就应该尽量举出许多事例来，但是篇幅限制，无法这样做。我只能断言，本能确实是变异的——例如迁徙的本能，不但在范围和方向上能变异，而且也会完全消失。鸟巢也是如此，变异部分发生于对选定的位置以及居住地性质和气温的依赖度，但常常由于全然未知的原因而发生变异。奥杜邦曾举出几个显著的例子，说明美国北方和南方同一物种的鸟巢有所不同。对于敌害的恐惧必然是一种本能品质，从未离巢的雏鸟身上可以看到，但这种恐惧可由经验或因看见其他动物对于同一敌害的恐惧而强化。对于人类的恐惧，如我在他处指出的，栖息在荒岛上的各种动物是慢慢获得的。甚至英国也可以看到这样的事例，即一切大型鸟比小型鸟更怕人，因为更多地遭受过人的迫害。英国的大型鸟更怕人，可以稳妥地归于这个原因；在无人岛，大型鸟并不比小型鸟更怕人；喜鹊（magpie）在英国很警惕，但在挪威却很驯顺，埃及的羽冠乌鸦（hooded crow）也是不怕人的。

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As some degree of variation in instincts under a state of nature, and the inheritance of such variations, are indispensable for the action of natural selection, as many instances as possible ought to have been here given; but want of space prevents me. I can only assert, that instincts certainly do vary—for instance, the migratory instinct, both in extent and direction, and in its total loss. So it is with the nests of birds, which vary partly in dependence on the situations chosen, and on the nature and temperature of the country inhabited, but often from causes wholly unknown to us: Audubon has given several remarkable cases of differences in nests of the same species in the northern and southern United States. Fear of any particular enemy is certainly an instinctive quality, as may be seen in nestling birds, though it is strengthened by experience, and by the sight of fear of the same enemy in other animals. But fear of man is slowly acquired, as I have elsewhere shown, by various animals inhabiting desert islands; and we may see an instance of this, even in England, in the greater wildness of all our large birds than of our small birds; for the large birds have been most persecuted by man. We may safely attribute the greater wildness of our large birds to this cause; for in uninhabited islands large birds are not more fearful than small; and the magpie, so wary in England, is tame in Norway, as is the hooded crow in Egypt.

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有许多事实可以证明，自然状态下产生的同类动物的脾气极多样化。还有若干个案可以举出，表明某些物种偶发的奇特习性若对这个物种有利，就会通过自然选择产生新的本能。但我十分清楚，泛泛而谈，没有详细的事实，在读者的心目中只会当耳旁风。我只好重复保证，不说没有可靠证据的话。

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That the general disposition of individuals of the same species, born in a state of nature, is extremely diversified, can be shown by a multitude of facts. Several cases also, could be given, of occasional and strange habits in certain species, which might, if advantageous to the species, give rise, through natural selection, to quite new instincts. But I am well aware that these general statements, without facts given in detail, can produce but a feeble effect on the reader’s mind. I can only repeat my assurance, that I do not speak without good evidence.

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简略考察一下家养的若干例子，则自然状态下本能遗传变异的可能性将加强，甚至出现大的可能性。由此可见习性和所谓偶发变异的选择，在改变家养动物精神品质上分别发生的作用。有许多奇异而真实的例子可以说明，与某种心境或某一时期有关的各种不同脾气嗜好以及怪癖都是遗传的。但是让我们看看众所熟知的几种狗的例子。毫无疑问，幼小的指示犬第一次带出去，有时就能够指示猎物的所在，甚至能够援助别的狗（我曾亲见这动人的情形）；寻回犬（retriever）确实在某种程度上可以把寻回的特性遗传下去；牧羊犬并不跑在羊群之内，而有在羊群周围环跑的倾向。幼小动物不依靠经验而有这些活动，同时各个体又差不多以同一方式进行，并且都欣然且不知道目的地去进行——幼小的指示犬并不知道指示方向是在帮助主人，有如白蝴蝶并不知道为什么要在圆白菜叶子上产卵一样——我无法看出这些活动在本质上与真正的本能有什么区别。如果看见一种狼，在幼小而且没有受过任何训练时，一旦嗅出猎物，先站着一动不动，随后又用特别的步法慢慢爬过去；又看见另一种狼环绕鹿群追逐，却不直冲，以便把鹿赶到远的地点去，必然要把这活动叫作本能。所谓家养下的本能，的确远不及自然的本能那么固定；但是其所蒙受的选择作用也极不严格，而且是在较不固定的生活条件下，在无比短暂的时间内传递下来的。

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The possibility, or even probability, of inherited variations of instinct in a state of nature will be strengthened by briefly considering a few cases under domestication. We shall thus also be enabled to see the respective parts which habit and the selection of so-called accidental variations have played in modifying the mental qualities of our domestic animals. A number of curious and authentic instances could be given of the inheritance of all shades of disposition and tastes, and likewise of the oddest tricks, associated with certain frames of mind or periods of time. But let us look to the familiar case of the several breeds of dogs: it cannot be doubted that young pointers (I have myself seen a striking instance) will sometimes point and even back other dogs the very first time that they are taken out; retrieving is certainly in some degree inherited by retrievers; and a tendency to run round, instead of at, a flock of sheep, by shepherd-dogs. I cannot see that these actions, performed without experience by the young, and in nearly the same manner by each individual, performed with eager delight by each breed, and without the end being known,—for the young pointer can no more know that he points to aid his master, than the white butterfly knows why she lays her eggs on the leaf of the cabbage,—I cannot see that these actions differ essentially from true instincts. If we were to see one kind of wolf, when young and without any training, as soon as it scented its prey, stand motionless like a statue, and then slowly crawl forward with a peculiar gait; and another kind of wolf rushing round, instead of at, a herd of deer, and driving them to a distant point, we should assuredly call these actions instinctive. Domestic instincts, as they may be called, are certainly far less fixed or invariable than natural instincts; but they have been acted on by far less rigorous selection, and have been transmitted for an incomparably shorter period, under less fixed conditions of life.

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不同品种的狗进行杂交时，即能很好地看出这家养下的本能、习性以及脾气的遗传性是何等强烈，并且混合得多么奇妙。例如众所周知，长驱跑狗与斗牛狗杂交，可影响勇敢性和顽强性至许多世代；牧羊犬与长驱跑狗杂交，则使全族都得到捕捉野兔的倾向。这家养下的本能，如用上述杂交方法来试验，是与自然的本能相类似的；自然的本能也按照同样的方式奇异地混合在一起，而且长期表现出其祖代任何一方的本能的痕迹。例如，勒鲁瓦（Le Roy）描述过一条狗，曾祖父是狼；它只有一点表示了野生祖先的痕迹，即呼唤它时，不是直线走向主人。

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How strongly these domestic instincts, habits, and dispositions are inherited, and how curiously they become mingled, is well shown when different breeds of dogs are crossed. Thus it is known that a cross with a bull-dog has affected for many generations the courage and obstinacy of greyhounds; and a cross with a greyhound has given to a whole family of shepherd-dogs a tendency to hunt hares. These domestic instincts, when thus tested by crossing, resemble natural instincts, which in a like manner become curiously blended together, and for a long period exhibit traces of the instincts of either parent: for example, Le Roy describes a dog, whose great-grandfather was a wolf, and this dog showed a trace of its wild parentage only in one way, by not coming in a straight line to his master when called.

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家养下的本能有时被说成为完全由长期的强迫养成的习性所遗传下来的动作，我看这是不正确的。从未有人会想象去教或者曾经教过翻飞鸽去翻飞——据我所见到的，幼鸽从不曾见过鸽的翻飞，却会翻飞。我们相信，曾经有过一只鸽子表现了这种奇怪习性的微小倾向，并且在连续的世代中，经过对最好的个体的长期选择，才造成像今日那样的翻飞鸽；据布伦特（Brent）先生说，格拉斯哥附近的家养翻飞鸽，一飞到十八英寸高就要翻筋斗。假如未曾有过一只狗自然具有指示方向的倾向，是否会有人想到训练狗去指示方向是存疑的；人们知道这种倾向有时会出现，我就看见过一次，见于纯种的里。指示方向的最初倾向一旦出现，此后在每一世代中有计划选择和强迫训练的遗传效果，将会很快大功告成；而且无意识选择至今仍在继续进行，每一个人虽然本意不在改进品种，但总试图获得最善于指示方向和狩猎的狗。另一方面，在某些情形下，仅仅习性一项已经足够了；没有动物比小野兔更难以驯服的了，也几乎没有动物比小家兔更驯顺的了。但我很难设想家兔仅仅为了驯服性才被选择下来；所以从极野的到极驯服的性质的遗传变化，必须全部归因于习性和长久持续的严格圈养。

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Domestic instincts are sometimes spoken of as actions which have become inherited solely from long-continued and compulsory habit, but this, I think, is not true. No one would ever have thought of teaching, or probably could have taught, the tumbler-pigeon to tumble,—an action which, as I have witnessed, is performed by young birds, that have never seen a pigeon tumble. We may believe that some one pigeon showed a slight tendency to this strange habit, and that the long-continued selection of the best individuals in successive generations made tumblers what they now are; and near Glasgow there are house-tumblers, as I hear from Mr. Brent, which cannot fly eighteen inches high without going head over heels. It may be doubted whether any one would have thought of training a dog to point, had not some one dog naturally shown a tendency in this line; and this is known occasionally to happen, as I once saw in a pure terrier. When the first tendency was once displayed, methodical selection and the inherited effects of compulsory training in each successive generation would soon complete the work; and unconscious selection is still at work, as each man tries to procure, without intending to improve the breed, dogs which will stand and hunt best. On the other hand, habit alone in some cases has sufficed; no animal is more difficult to tame than the young of the wild rabbit; scarcely any animal is tamer than the young of the tame rabbit; but I do not suppose that domestic rabbits have ever been selected for tameness; and I presume that we must attribute the whole of the inherited change from extreme wildness to extreme tameness, simply to habit and long-continued close confinement.

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自然的本能在家养状况下可以消失：最显著的例子见于少孵蛋或不喜孵蛋的鸡品种。仅仅由于司空见惯，我们才看不出家养动物的心理曾经有过多么普遍的变化。对于人类的亲爱已经成了狗的本能，这是毋庸置疑的。一切狼、狐、胡狼（jackal）以及猫属的物种即使驯养后，也极渴望攻击鸡、羊和猪；火地和澳洲这些地方的未开化人不养狗，因为曾发现这种倾向在家里养的小狗身上是不能矫正的。另一方面，已经文明化了的狗，甚至在十分幼小的时候，也很少必要去教其不要攻击鸡、羊和猪！无疑会偶尔攻击一下子，于是就要遭一顿打；如果还不能矫正，就会被弄死；那个习性，通过某种程度的选择，也许协同地靠遗传使家狗文明化了。另一方面，小鸡完全出于习性，已经消失了原本无疑惧怕猫狗的本能；而小雉鸡尽管是由母鸡抚养的，却是明显具有这种本能的。倒不是小鸡失去了一切惧怕，而只是失去了对于猫狗的惧怕，因为，母鸡发出报告危险的叫声，小鸡便从母鸡的翼下跑开（小火鸡尤其如此），躲到四周的草丛里或林子里去了。这显然是本能的动作，便于母鸟飞走，就如我们在野生的陆栖鸟类里所看到的那样。但是小鸡还保留着这种本能，在家养状况下已经没有用处，因为母鸡由于不使用的缘故，已经几乎失掉了飞翔能力。

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Natural instincts are lost under domestication: a remarkable instance of this is seen in those breeds of fowls which very rarely or never become “broody,” that is, never wish to sit on their eggs. Familiarity alone prevents our seeing how universally and largely the minds of our domestic animals have been modified by domestication. It is scarcely possible to doubt that the love of man has become instinctive in the dog. All wolves, foxes, jackals, and species of the cat genus, when kept tame, are most eager to attack poultry, sheep, and pigs; and this tendency has been found incurable in dogs which have been brought home as puppies from countries, such as Tierra del Fuego and Australia, where the savages do not keep these domestic animals. How rarely, on the other hand, do our civilised dogs, even when quite young, require to be taught not to attack poultry, sheep, and pigs! No doubt they occasionally do make an attack, and are then beaten; and if not cured, they are destroyed; so that habit, with some degree of selection, has probably concurred in civilising by inheritance our dogs. On the other hand, young chickens have lost, wholly by habit, that fear of the dog and cat which no doubt was originally instinctive in them, in the same way as it is so plainly instinctive in young pheasants, though reared under a hen. It is not that chickens have lost all fear, but fear only of dogs and cats, for if the hen gives the danger-chuckle, they will run (more especially young turkeys) from under her, and conceal themselves in the surrounding grass or thickets; and this is evidently done for the instinctive purpose of allowing, as we see in wild ground-birds, their mother to fly away. But this instinct retained by our chickens has become useless under domestication, for the mother-hen has almost lost by disuse the power of flight.

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因此，可以断定，动物在家养下可以获得新的本能；而失去自然的本能，这一部分是由于习性，一部分是由于人类在连续世代中选择和累积了特殊的精神习性和精神活动，而它们的最初发生，我们无知地看作是出于偶然的原因。在某些情形下，只是强制的习性一项，已足以产生这种遗传的心理变化；在另外一些情形下，强制的习性就不能发生作用，一切都是有计划选择和无意识选择的结果。但是在大多数情形下，习性和选择大概是双管齐下的。

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Hence, we may conclude, that domestic instincts have been acquired and natural instincts have been lost partly by habit, and partly by man selecting and accumulating during successive generations, peculiar mental habits and actions, which at first appeared from what we must in our ignorance call an accident. In some cases compulsory habit alone has sufficed to produce such inherited mental changes; in other cases compulsory habit has done nothing, and all has been the result of selection, pursued both methodically and unconsciously; but in most cases, probably, habit and selection have acted together.

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我们只要考察少数事例，大概就能很好地理解本能在自然状态下如何由于选择作用而改变的。我只选择三个例子，其余的可能要以后著书讨论了——即，杜鹃在别种鸟巢里下蛋的本能，某些蚂蚁蓄奴的本能，以及蜜蜂造蜂房的本能。学者们已经把后两种本能，一般地而且恰当地列为一切已知本能中最奇异的了。

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We shall, perhaps, best understand how instincts in a state of nature have become modified by selection, by considering a few cases. I will select only three, out of the several which I shall have to discuss in my future work,— namely, the instinct which leads the cuckoo to lay her eggs in other birds’ nests; the slave-making instinct of certain ants; and the comb-making power of the hive-bee: these two latter instincts have generally, and most justly, been ranked by naturalists as the most wonderful of all known instincts.

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现在已经有共识，杜鹃的这种本能的比较直接的决定性原因，是其并不每日下蛋，而是隔二差三下蛋一次；所以，如果自己筑巢，自己孵蛋，则最先下的蛋便须搁置一些时间后才能得到孵抱，同一个巢里就会有不同龄期的蛋和小鸟了。这样，下蛋和孵蛋的过程就会漫长而不方便，特别是雌鸟在很早就要迁徙，而最初孵化的小鸟势必要由雄鸟单独哺养。但是美洲杜鹃就处于这样的困境；她自己筑巢，而且要在同一时期内产蛋和照顾相继孵化的幼鸟。有人说美洲杜鹃有时也在别种鸟巢里下蛋，但我从权威的布留尔（Brewer）博士那里听到，这不对。不过，我可以举出各种鸟类偶尔在别种鸟巢里下蛋的若干事例。现在假定欧洲杜鹃的古代祖先也有美洲杜鹃的习性，但偶尔也在别种鸟巢里下蛋。如果这种偶尔的习性有利于老鸟，如果小鸟由于利用了其他物种的误养本能，比起母鸟哺养更为强壮——母鸟必须同时照顾不同龄期的蛋和小鸟，不免受到拖累——那么老鸟或寄养的小鸟都会受益。以此类推，我可以相信，这样哺养起来的小鸟大概就会遗传母鸟那种偶然的奇特习性，倾向于把蛋下在别种的鸟巢里，这样就能够成功哺养幼鸟。我相信杜鹃的奇异本能会由这种连续过程而产生出来。补充一句，格雷博士等人说，欧洲杜鹃并未完全失去母爱和对后代的关怀。

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It is now commonly admitted that the more immediate and final cause of the cuckoo’s instinct is, that she lays her eggs, not daily, but at intervals of two or three days; so that, if she were to make her own nest and sit on her own eggs, those first laid would have to be left for some time unincubated, or there would be eggs and young birds of different ages in the same nest. If this were the case, the process of laying and hatching might be inconveniently long, more especially as she has to migrate at a very early period; and the first hatched young would probably have to be fed by the male alone. But the American cuckoo is in this predicament; for she makes her own nest and has eggs and young successively hatched, all at the same time. It has been asserted that the American cuckoo occasionally lays her eggs in other birds’ nests; but I hear on the high authority of Dr. Brewer, that this is a mistake. Nevertheless, I could give several instances of various birds which have been known occasionally to lay their eggs in other birds’ nests. Now let us suppose that the ancient progenitor of our European cuckoo had the habits of the American cuckoo; but that occasionally she laid an egg in another bird’s nest. If the old bird profited by this occasional habit, or if the young were made more vigorous by advantage having been taken of the mistaken maternal instinct of another bird, than by their own mother’s care, encumbered as she can hardly fail to be by having eggs and young of different ages at the same time; then the old birds or the fostered young would gain an advantage. And analogy would lead me to believe, that the young thus reared would be apt to follow by inheritance the occasional and aberrant habit of their mother, and in their turn would be apt to lay their eggs in other birds’ nests, and thus be successful in rearing their young. By a continued process of this nature, I believe that the strange instinct of our cuckoo could be, and has been, generated. I may add that, according to Dr. Gray and to some other observers, the European cuckoo has not utterly lost all maternal love and care for her own offspring.

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鸟类偶尔会把蛋下在同种别种的鸟巢里这种习性，在鸡科里并非不普通，并且可以解释近缘鸵鸟群的奇特本能的来源。至少是美国种的个案，若干母鸵鸟共同先在一个巢里，然后在另一个巢里下一些蛋，由雄鸟去孵。这种本能或可以解释为雌鸟下蛋很多，但如杜鹃一样，隔二差三才下一次。然而美洲鸵鸟的这种本能，还没有达到完善；因为有多得出奇的蛋都散落在地上，我在一天的游猎中，就拾得了不下二十个丢弃的蛋。

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The occasional habit of birds laying their eggs in other birds’ nests, either of the same or of a distinct species, is not very uncommon with the Gallinaceae; and this perhaps explains the origin of a singular instinct in the allied group of ostriches. For several hen ostriches, at least in the case of the American species, unite and lay first a few eggs in one nest and then in another; and these are hatched by the males. This instinct may probably be accounted for by the fact of the hens laying a large number of eggs; but, as in the case of the cuckoo, at intervals of two or three days. This instinct, however, of the American ostrich has not as yet been perfected; for a surprising number of eggs lie strewed over the plains, so that in one day’s hunting I picked up no less than twenty lost and wasted eggs.

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许多蜂是寄生的，总是把卵产在别种的蜂巢里，这个个案比杜鹃更令人瞩目；这种蜂随着寄生习性，不但改变了本能，而且改变了构造；它们不具有采集花粉的器具，如果要为幼蜂贮藏食料，这是必不可少的。泥蜂科（Sphecidae；形似胡蜂）的某些物种同样也是寄生的；法布尔最近提出充分的理由认为：一种小唇沙蜂（Tachytes nigra）虽然通常都自己造巢，而且为幼虫储蓄麻痹了的食物，但发现别种泥蜂所造储蓄有食物的巢，便会加以利用，而变成临时的寄生者。这种情形和杜鹃的假设情形是一样的，我觉得如果一种临时的习性对于物种有利，同时被害的蜂类不会因巢和储蓄的食物被无情夺取而遭到灭绝，自然选择就不难把它永久化。

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Many bees are parasitic, and always lay their eggs in the nests of bees of other kinds. This case is more remarkable than that of the cuckoo; for these bees have not only their instincts but their structure modified in accordance with their parasitic habits; for they do not possess the pollen-collecting apparatus which would be necessary if they had to store food for their own young. Some species, likewise, of Sphegidae (wasp-like insects) are parasitic on other species; and M. Fabre has lately shown good reason for believing that although the Tachytes nigra generally makes its own burrow and stores it with paralysed prey for its own larvae to feed on, yet that when this insect finds a burrow already made and stored by another sphex, it takes advantage of the prize, and becomes for the occasion parasitic. In this case, as with the supposed case of the cuckoo, I can see no difficulty in natural selection making an occasional habit permanent, if of advantage to the species, and if the insect whose nest and stored food are thus feloniously appropriated, be not thus exterminated.

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蓄奴的本能。——这种奇妙的本能，是由于贝尔最初在红褐蚁（Formica［Polyerges］rufescens）身上发现的，他是一位比他著名的父亲更为优秀的观察者。这种蚂蚁绝对依赖奴隶而生活；没有奴隶的帮助，这个物种一年之内一定灭绝。雄蚁和能育的雌蚁不从事任何工作，工蚁即不育的雌蚁虽然在捕捉奴隶上极为卖力勇敢，但不做其他任何工作。它们不能营造自己的巢，也不能哺喂自己的幼虫。在老巢已不适用，势必迁徙的时候，是由奴蚁来决定迁徙的事情，并且实际上把主人们衔在颚间搬走。主人们十分的不中用，当于贝尔捉了三十个关起来而没有一个奴蚁时，虽然那里放着它们最喜爱的丰富食物，而且有自己的幼虫和蛹刺激它们工作，它们还是无所事事；它们甚至不会自己吃东西，许多蚂蚁就此饿毙。于贝尔随后放进一个奴蚁——黑蚁（F. fusca），她即刻开始工作，喂哺和拯救那些生存者；并且营造了几间虫房，来照料幼虫，一切都整得井井有条。有什么比这十分肯定的事实更为奇异的呢？如果我们不知道任何其他蓄奴的蚁类，大概就无法想象如此奇异的本能是怎样完善的。

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Slave-making instinct.—This remarkable instinct was first discovered in the Formica (Polyerges) rufescens by Pierre Huber, a better observer even than his celebrated father. This ant is absolutely dependent on its slaves; without their aid, the species would certainly become extinct in a single year. The males and fertile females do no work. The workers or sterile females, though most energetic and courageous in capturing slaves, do no other work. They are incapable of making their own nests, or of feeding their own larvae. When the old nest is found inconvenient, and they have to migrate, it is the slaves which determine the migration, and actually carry their masters in their jaws. So utterly helpless are the masters, that when Huber shut up thirty of them without a slave, but with plenty of the food which they like best, and with their larvae and pupae to stimulate them to work, they did nothing; they could not even feed themselves, and many perished of hunger. Huber then introduced a single slave (F. fusca), and she instantly set to work, fed and saved the survivors; made some cells and tended the larvae, and put all to rights. What can be more extraordinary than these well-ascertained facts? If we had not known of any other slave-making ant, it would have been hopeless to have speculated how so wonderful an instinct could have been perfected.

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血蚁（Formica sanguinea）同样蓄奴，也是于贝尔最初发现的。这个物种见于英格兰南部，大英博物馆史密斯（F. Smith）先生研究过它的习性。关于这个问题以及其他问题，我深深感激他的帮助。虽然我充分相信于贝尔和史密斯先生的叙述，但仍然以怀疑的心情来处理这个问题，任何人对于蓄奴的这种异常丑恶本能的存在有所怀疑，大概都得谅解。因此，我愿意稍微详细地谈谈我的观察。我曾掘开十四个血蚁巢，都发现了若干奴蚁。奴种的雄蚁和能育的雌蚁，只见于它们自己固有的群中，在血蚁巢中从未看见过。黑色奴蚁，不及红色主人的一半大，外貌上对比强烈。蚁巢被微微扰动时，奴蚁偶尔跑出外边来，像主人一样十分激动，并且保家卫国；当蚁巢被扰动得很厉害，幼虫和蛹暴露出来的时候，奴蚁和主人一齐奋发地把它们运送到安全的地方。因此，奴蚁显然是熟门熟路。在连续三个年头的六月和七月里，我在萨里郡和萨塞克斯郡，曾对几个蚁巢观察了几个小时，从来没有看到一个奴蚁自蚁巢里走出走进。在这些月份里，奴蚁的数目很少，因此我想数目多的时候，行动大概就不同了；但史密斯先生告诉我说，五月、六月以及八月间，在萨里和汉普郡，他在各种不同的时间内注意观察了蚁巢，虽然八月份奴蚁的数目很多，但也不曾看到它们走出走进蚁巢。因此，他认为它们是严格的家奴。而主人却不然，经常看到它们不断地搬运着造巢材料和各种食物。然而今年七月里，我遇见一个奴蚁特别多的蚁群，观察到有少数奴蚁和主人混在一起离巢，沿着同一条路向着约二十五码远的一株高大欧洲赤松前进，它们一齐爬到树上去，大概是为了找寻蚜虫或胭脂虫的。于贝尔有过许多观察的机会，他说，瑞士的奴蚁在造蚁巢的时候常常和主人一起干，而在早晚间则单独看管门户。于贝尔还明确地说，奴蚁的主要职务是搜寻蚜虫。两个国家里的主奴两蚁的普通习性如此不同，大概仅仅由于在瑞士被捕捉的奴蚁数目比英格兰多。

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Formica sanguinea was likewise first discovered by P. Huber to be a slave-making ant. This species is found in the southern parts of England, and its habits have been attended to by Mr. F. Smith, of the British Museum, to whom I am much indebted for information on this and other subjects. Although fully trusting to the statements of Huber and Mr. Smith, I tried to approach the subject in a sceptical frame of mind, as any one may well be excused for doubting the truth of so extraordinary and odious an instinct as that of making slaves. Hence I will give the observations which I have myself made, in some little detail. I opened fourteen nests of F. sanguinea, and found a few slaves in all. Males and fertile females of the slave-species are found only in their own proper communities, and have never been observed in the nests of F. sanguinea. The slaves are black and not above half the size of their red masters, so that the contrast in their appearance is very great. When the nest is slightly disturbed, the slaves occasionally come out, and like their masters are much agitated and defend the nest: when the nest is much disturbed and the larvae and pupae are exposed, the slaves work energetically with their masters in carrying them away to a place of safety. Hence, it is clear, that the slaves feel quite at home. During the months of June and July, on three successive years, I have watched for many hours several nests in Surrey and Sussex, and never saw a slave either leave or enter a nest. As, during these months, the slaves are very few in number, I thought that they might behave differently when more numerous; but Mr. Smith informs me that he has watched the nests at various hours during May, June and August, both in Surrey and Hampshire, and has never seen the slaves, though present in large numbers in August, either leave or enter the nest. Hence he considers them as strictly household slaves. The masters, on the other hand, may be constantly seen bringing in materials for the nest, and food of all kinds. During the present year, however, in the month of July, I came across a community with an unusually large stock of slaves, and I observed a few slaves mingled with their masters leaving the nest, and marching along the same road to a tall Scotch-fir-tree, twenty-five yards distant, which they ascended together, probably in search of aphides or cocci. According to Huber, who had ample opportunities for observation, in Switzerland the slaves habitually work with their masters in making the nest, and they alone open and close the doors in the morning and evening; and, as Huber expressly states, their principal office is to search for aphides. This difference in the usual habits of the masters and slaves in the two countries, probably depends merely on the slaves being captured in greater numbers in Switzerland than in England.

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有一天，我有幸看到了血蚁搬巢，于贝尔描述过，主人们谨慎地把奴蚁带在颚间，这真是极有趣的奇观。另一天，大约有二十只蓄奴蚁在同一地点猎取东西，而显然不是找寻食物，这引起了我的注意。它们走近一种奴蚁——独立的黑蚁群，并且遭到猛烈的反击。有时候三个奴蚁揪住蓄奴血蚁的腿不放，蓄奴蚁残忍地弄死了小抵抗者，并且把尸体拖到二十九码远的巢中去当食物，但得不到一个蛹来培养为奴。于是我从另一个巢里掘出一小团黑蚁的蛹，放在邻近战场的一处空地上。于是这班暴君热切地把它们捉住并且拖走，大概以为毕竟是在最后的战役中获胜了。

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One day I fortunately chanced to witness a migration from one nest to another, and it was a most interesting spectacle to behold the masters carefully carrying, as Huber has described, their slaves in their jaws. Another day my attention was struck by about a score of the slave-makers haunting the same spot, and evidently not in search of food; they approached and were vigorously repulsed by an independent community of the slave species (F. fusca); sometimes as many as three of these ants clinging to the legs of the slave-making F. sanguinea. The latter ruthlessly killed their small opponents, and carried their dead bodies as food to their nest, twenty-nine yards distant; but they were prevented from getting any pupae to rear as slaves. I then dug up a small parcel of the pupae of F. fusca from another nest, and put them down on a bare spot near the place of combat; they were eagerly seized, and carried off by the tyrants, who perhaps fancied that, after all, they had been victorious in their late combat.

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同时，我在同一场所放下另一物种——黄蚁（F. flava）的一小团蛹，其上还有几只小黄蚁攀附在蚁巢破片上。如史密斯先生所描述的，这个物种有时会沦落为奴，但很少见。这种蚁虽然这么小，但极勇敢，我看到过它们凶猛地攻击别种蚁。有一个事例，我惊奇地看见蓄奴血蚁巢下有一块石头，底下是一个独立的黄蚁群；我偶然地扰动这两个巢，小蚂蚁就以惊人的勇气去攻击它们的大邻居。当时我渴望确定血蚁能否辨别常捉做奴隶的黑蚁的蛹与很少捉拿的小型而凶猛的黄蚁的蛹，明显地它们确能立刻辨别；因为我们看见遇到黑蚁的蛹时，它们即刻热切地去捉，而遇到黄蚁的蛹，甚至遇到其巢的泥土时，便惊慌失措，赶紧跑开；但是，大约经过一刻钟，当这种小黄蚁都爬走之后，它们才鼓起勇气，把蛹搬走。

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At the same time I laid on the same place a small parcel of the pupae of another species, F. flava, with a few of these little yellow ants still clinging to the fragments of the nest. This species is sometimes, though rarely, made into slaves, as has been described by Mr. Smith. Although so small a species, it is very courageous, and I have seen it ferociously attack other ants. In one instance I found to my surprise an independent community of F. flava under a stone beneath a nest of the slave-making F. sanguinea; and when I had accidentally disturbed both nests, the little ants attacked their big neighbours with surprising courage. Now I was curious to ascertain whether F. sanguinea could distinguish the pupae of F. fusca, which they habitually make into slaves, from those of the little and furious F. flava, which they rarely capture, and it was evident that they did at once distinguish them: for we have seen that they eagerly and instantly seized the pupae of F. fusca, whereas they were much terrified when they came across the pupae, or even the earth from the nest of F. flava, and quickly ran away; but in about a quarter of an hour, shortly after all the little yellow ants had crawled away, they took heart and carried off the pupae.

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一天傍晚，我看见另一群血蚁，发现若干这种蚁拖着黑蚁的尸体（可以看出不是迁徙）和无数的蛹回巢。我跟着背着战利品鱼贯而行的蚁追踪前去，大约有四十码之远，到了一处密集的石南科灌木丛（heath），我看到最后一个血蚁出现，拖着一个蛹，但无法在密丛中找到被蹂躏的蚁巢。然而那巢一定就在附近，因为有两三只黑蚁极度张皇地冲出来，有一只嘴里还衔着自己的蛹一动不动地停留在石南的小枝顶上，在破碎的家上方。

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One evening I visited another community of F. sanguinea, and found a number of these ants entering their nest, carrying the dead bodies of F. fusca (showing that it was not a migration) and numerous pupae. I traced the returning file burthened with booty, for about forty yards, to a very thick clump of heath, whence I saw the last individual of F. sanguinea emerge, carrying a pupa; but I was not able to find the desolated nest in the thick heath. The nest, however, must have been close at hand, for two or three individuals of F. fusca were rushing about in the greatest agitation, and one was perched motionless with its own pupa in its mouth on the top of a spray of heath over its ravaged home.

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这些都是关于蓄奴的奇异本能的事实，无须我来证实。让我们看一看，血蚁的本能习性和欧洲大陆上的红褐蚁的鲜明对照。后一种不会筑巢，不会决定自己的迁徙，不会为自己和幼蚁采集食物，甚至不会自己吃东西：完全依赖无数的奴蚁。血蚁则不然，拥有很少的奴蚁，初夏时奴蚁是极少的。主人决定在何时何地营造新蚁巢，并且迁徙的时候，还衔着奴蚁走。瑞士和英格兰的奴蚁似乎都专门照顾幼蚁，主人单独做捕捉奴蚁的远征。瑞士的奴蚁和主人一齐工作，搬运材料回去造巢；主奴共同地，但主要是奴蚁在照顾它们的蚜虫，并进行所谓的挤乳；这样，主奴都为本群采集食物。在英格兰，通常是主人单独出去搜寻筑巢材料，为它们自己、奴蚁和幼蚁搜寻食物。所以，我国奴蚁为主人所服的劳役，比在瑞士少得多。

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Such are the facts, though they did not need confirmation by me, in regard to the wonderful instinct of making slaves. Let it be observed what a contrast the instinctive habits of F. sanguinea present with those of the F. rufescens. The latter does not build its own nest, does not determine its own migrations, does not collect food for itself or its young, and cannot even feed itself: it is absolutely dependent on its numerous slaves. Formica sanguinea, on the other hand, possesses much fewer slaves, and in the early part of the summer extremely few. The masters determine when and where a new nest shall be formed, and when they migrate, the masters carry the slaves. Both in Switzerland and England the slaves seem to have the exclusive care of the larvae, and the masters alone go on slave-making expeditions. In Switzerland the slaves and masters work together, making and bringing materials for the nest: both, but chiefly the slaves, tend, and milk as it may be called, their aphides; and thus both collect food for the community. In England the masters alone usually leave the nest to collect building materials and food for themselves, their slaves and larvae. So that the masters in this country receive much less service from their slaves than they do in Switzerland.

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血蚁的本能靠什么步骤发生，我不愿妄加臆测。但是，据我所看到的，不蓄奴的蚁如果有其他物种的蛹散落在蚁巢近旁，也要把这些蛹拖进去，所以这些本来贮作食物的蛹可能发育起来；这样无意识地被养育起来的外来蚁将会追随自己的固有本能，做它们所能做的工作。如果它们的存在证明对于捕获它们的物种有用——如果捕捉工蚁比自己生育工蚁对于这个物种更有利——那么，本是采集蚁蛹供食用的这种习性，大概会因自然选择而加强，并且变为永久的，以达到非常不同的蓄奴目的。本能一旦获得，即使它的应用范围远不及英国的血蚁（如我们所看到的，这种蚁在依赖奴蚁的帮助上比瑞士的同一物种为少），我看自然选择也不难增强和改变这种本能——始终假定每一个变异对于物种都有用处——直到形成一种像红褐蚁那样卑鄙地依靠奴隶来生活的蚁类。

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By what steps the instinct of F. sanguinea originated I will not pretend to conjecture. But as ants, which are not slave-makers, will, as I have seen, carry off pupae of other species, if scattered near their nests, it is possible that pupae originally stored as food might become developed; and the ants thus unintentionally reared would then follow their proper instincts, and do what work they could. If their presence proved useful to the species which had seized them—if it were more advantageous to this species to capture workers than to procreate them—the habit of collecting pupae originally for food might by natural selection be strengthened and rendered permanent for the very different purpose of raising slaves. When the instinct was once acquired, if carried out to a much less extent even than in our British F. sanguinea, which, as we have seen, is less aided by its slaves than the same species in Switzerland, I can see no difficulty in natural selection increasing and modifying the instinct—always supposing each modification to be of use to the species—until an ant was formed as abjectly dependent on its slaves as is the Formica rufescens.

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蜜蜂营造蜂房的本能。——这个问题不拟详加讨论，而只是把我所得到的结论的纲要说一说。凡是考察过蜂巢的精巧构造的人，看到如此美妙地适应它的目的而不热烈赞赏，必定是愚钝不堪。听到数学家说蜜蜂已实际解决了深奥的问题，把蜂房造成适当的形状，来容纳最大可能容量的蜜，而在建造中则用最小限度的贵重蜡质。有人说，一个熟练的工人，用合适的工具和度量衡，也很难造出正形的蜡质蜂房来，但是一群蜜蜂却能在黑暗的蜂箱内把它造成。随便你说这是什么本能都可以，乍一看似乎是不可思议的，如何能造出所有必要的角和面，甚至如何能觉察出做工正确。但是这难点并不像最初看来那样大；我想可以证明，这一切美妙的工作都是来自几种简单的本能。

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Cell-making instinct of the hive-Bee.—I will not here enter on minute details on this subject, but will merely give an outline of the conclusions at which I have arrived. He must be a dull man who can examine the exquisite structure of a comb, so beautifully adapted to its end, without enthusiastic admiration. We hear from mathematicians that bees have practically solved a recondite problem, and have made their cells of the proper shape to hold the greatest possible amount of honey, with the least possible consumption of precious wax in their construction. It has been remarked that a skilful workman, with fitting tools and measures, would find it very difficult to make cells of wax of the true form, though this is perfectly effected by a crowd of bees working in a dark hive. Grant whatever instincts you please, and it seems at first quite inconceivable how they can make all the necessary angles and planes, or even perceive when they are correctly made. But the difficulty is not nearly so great as it at first appears: all this beautiful work can be shown, I think, to follow from a few very simple instincts.

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我研究这个问题，是受沃特豪斯先生的引导。他阐明，蜂房的形状和邻接蜂房的存在有密切关系；下述观点大概只能看作是他的理论的修正。让我们看看伟大的分级原理，看看自然是否向我们揭示了其工作方法。这个简短系列的一端有大黄蜂，用旧茧来贮蜜，有时候在茧壳上添加蜡质短管，而且同样也会做出分隔的、很不规则的圆形蜡质蜂房。这系列的另一端则有蜜蜂的蜂房，排列为双层：每一个蜂房，众所周知，都是六面柱体，六边的底边倾斜地联合成三个菱形所组成的倒角锥体。菱形都有一定的角度，并且在蜂巢的一面，一个蜂房的角锥形底部的三条边，正好构成了反面的三个连接蜂房的底部。这一系列里，处于极完美的蜜蜂蜂房和简单的大黄蜂蜂房之间的，还有墨西哥蜂（Melipona domestica）的蜂房，于贝尔曾经仔细描述过和绘制过。墨西哥蜂的身体构造介于蜜蜂和大黄蜂之间，但与后者关系比较接近；能营造差不多规则的蜡质蜂巢，圆柱形蜂房，在里面孵化幼蜂，此外还有一些用作贮蜜的大型蜡质蜂房。这些大型的蜂房接近球状，大小差不多相等，并且聚集成不规则的一堆。这里要注意的要点是，蜂房总是营造得彼此很靠近，如果完全成为球状时，蜡壁势必就要交切或串通；但是从来不会如此，因为这种蜂会在有交切倾向的球状蜂房之间把蜡壁造成平面的。因此，每个蜂房都是由外方的球状部分和两三个或更多平面构成的，这要看这个蜂房与两三个或更多的蜂房相连接来决定。一个蜂房连接三个蜂房时，由于球形是差不多大小的，这种情形常常而且必然发生，所以三个平面连合成为一个角锥体；据于贝尔说，这种角锥体明显与蜜蜂蜂房的三边角锥形底部十分相像。这里和蜜蜂蜂房一样，任何蜂房的三个平面必然成为所连接的三个蜂房的构成部分。墨西哥蜂用这种营造方法，显然可以节省蜡；因为连接蜂房之间的平面壁并不是双层的，其厚薄和外面的球状部分相同，然而每一个平面壁却构成了两个蜂房的共同部分。

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I was led to investigate this subject by Mr. Waterhouse, who has shown that the form of the cell stands in close relation to the presence of adjoining cells; and the following view may, perhaps, be considered only as a modification of his theory. Let us look to the great principle of gradation, and see whether Nature does not reveal to us her method of work. At one end of a short series we have humble-bees, which use their old cocoons to hold honey, sometimes adding to them short tubes of wax, and likewise making separate and very irregular rounded cells of wax. At the other end of the series we have the cells of the hive-bee, placed in a double layer: each cell, as is well known, is an hexagonal prism, with the basal edges of its six sides bevelled so as to join on to a pyramid, formed of three rhombs. These rhombs have certain angles, and the three which form the pyramidal base of a single cell on one side of the comb, enter into the composition of the bases of three adjoining cells on the opposite side. In the series between the extreme perfection of the cells of the hive-bee and the simplicity of those of the humble-bee, we have the cells of the Mexican Melipona domestica, carefully described and figured by Pierre Huber. The Melipona itself is intermediate in structure between the hive and humble bee, but more nearly related to the latter: it forms a nearly regular waxen comb of cylindrical cells, in which the young are hatched, and, in addition, some large cells of wax for holding honey. These latter cells are nearly spherical and of nearly equal sizes, and are aggregated into an irregular mass. But the important point to notice, is that these cells are always made at that degree of nearness to each other, that they would have intersected or broken into each other, if the spheres had been completed; but this is never permitted, the bees building perfectly flat walls of wax between the spheres which thus tend to intersect. Hence each cell consists of an outer spherical portion and of two, three, or more perfectly flat surfaces, according as the cell adjoins two, three or more other cells. When one cell comes into contact with three other cells, which, from the spheres being nearly of the same size, is very frequently and necessarily the case, the three flat surfaces are united into a pyramid; and this pyramid, as Huber has remarked, is manifestly a gross imitation of the three-sided pyramidal basis of the cell of the hive-bee. As in the cells of the hive-bee, so here, the three plane surfaces in any one cell necessarily enter into the construction of three adjoining cells. It is obvious that the Melipona saves wax by this manner of building; for the flat walls between the adjoining cells are not double, but are of the same thickness as the outer spherical portions, and yet each flat portion forms a part of two cells.

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考虑这个个案时，我觉得如果墨西哥蜂在一定的彼此距离间营造球状蜂房，并且造成一样大小，同时对称排列成双层，那么这构造就会像蜜蜂巢一样完美了。所以我写信给剑桥的米勒（Miller）教授，根据他的复信，我写出了以下的叙述，这位几何学家惠读了，并且告诉我说，这是完全正确的：

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Reflecting on this case, it occurred to me that if the Melipona had made its spheres at some given distance from each other, and had made them of equal sizes and had arranged them symmetrically in a double layer, the resulting structure would probably have been as perfect as the comb of the hive-bee. Accordingly I wrote to Professor Miller, of Cambridge, and this geometer has kindly read over the following statement, drawn up from his information, and tells me that it is strictly correct:—

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设若干同等大小的球，球心在两个平行层上；每一个球的球心与同层中围绕它的六个球的球心相距等于或稍微小于半径×，即半径×1.41421；并且与别一平行层中连接的球的球心相距也如上；于是，如果把这双层每两个球的交接面都画出来，就会形成一个双层六面柱体，其互相衔接的面都是由三个菱形所组成的角锥形底部联结而成的；这个角锥形与六面柱体的边所成的角，与经过精密测量的蜜蜂蜂房的角完全相等。

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If a number of equal spheres be described with their centres placed in two parallel layers; with the centre of each sphere at the distance of radius , or radius×1.41421 (or at some lesser distance), from the centres of the six surrounding spheres in the same layer; and at the same distance from the centres of the adjoining spheres in the other and parallel layer; then, if planes of intersection between the several spheres in both layers be formed, there will result a double layer of hexagonal prisms united together by pyramidal bases formed of three rhombs; and the rhombs and the sides of the hexagonal prisms will have every angle identically the same with the best measurements which have been made of the cells of the hive-bee.

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因此可以稳妥地断定，如果能把墨西哥蜂的并不很奇异的已有本能稍微改变一下，便能造出像蜜蜂那样巧夺天工的蜂房。我们必须假定，墨西哥蜂有能力来营造真正球状的和大小相等的蜂房；鉴于已经能够在一定程度上做到这点，鉴于还有许多昆虫也能够在树木上造成多么完美的圆柱形孔穴，分明是依据一个固定的点旋转而成的，这就没有什么值得奇怪的了。必须假定，墨西哥蜂能把蜂房排列在水平层上，而其圆柱形蜂房就是这样排列的。必须进一步假定，当几只工蜂工友分别营造球状蜂房时，能好歹正确判断彼此应当距离多远，而这是最困难的事；不过，已经能判断距离了，所以总是能使球状蜂房有某种程度的交切；然后把交切点用完全的平面连接起来。必须再进一步假定，六面柱体由同层连接球体的交接面形成之后，可以任意延长六面柱体的长度，使之符合仓储蜂蜜的要求，而这一点不难；就像粗鲁的大黄蜂给旧茧的圆孔增加蜡质圆管一样的。本来并不奇异的本能——不比指导鸟类造巢的本能更奇异，经过这样的变异之后，我相信蜜蜂通过自然选择就获得了难以模仿的营造能力。

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Hence we may safely conclude that if we could slightly modify the instincts already possessed by the Melipona, and in themselves not very wonderful, this bee would make a structure as wonderfully perfect as that of the hive-bee. We must suppose the Melipona to make her cells truly spherical, and of equal sizes; and this would not be very surprising, seeing that she already does so to a certain extent, and seeing what perfectly cylindrical burrows in wood many insects can make, apparently by turning round on a fixed point. We must suppose the Melipona to arrange her cells in level layers, as she already does her cylindrical cells; and we must further suppose, and this is the greatest difficulty, that she can somehow judge accurately at what distance to stand from her fellow-labourers when several are making their spheres; but she is already so far enabled to judge of distance, that she always describes her spheres so as to intersect largely; and then she unites the points of intersection by perfectly flat surfaces. We have further to suppose, but this is no difficulty, that after hexagonal prisms have been formed by the intersection of adjoining spheres in the same layer, she can prolong the hexagon to any length requisite to hold the stock of honey; in the same way as the rude humble-bee adds cylinders of wax to the circular mouths of her old cocoons. By such modifications of instincts in themselves not very wonderful,—hardly more wonderful than those which guide a bird to make its nest,—I believe that the hive-bee has acquired, through natural selection, her inimitable architectural powers.

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这种理论可用试验来证明。照特盖特迈耶（Tegetmeier）先生的榜样，我把两个蜂巢分开，中间放一块长而厚的方形蜡版：蜜蜂随即开始在蜡版上凿掘圆形的小凹穴；向深处凿掘这些小穴时，逐渐使它们拓宽，变成约莫蜂房直径的浅盆形，看起来恰似真正球状或者球的一部分。下面的情形是极有趣的：凡是几只蜂彼此靠近开始凿掘盆形凹穴时，相互之间的距离恰使盆形凹穴得到上述宽度（大约相当于一个普通蜂房的宽度），并且在深度上达到这些盆形凹穴所构成的球体直径的六分之一，这时盆形凹穴的边便交切，或彼此串通。一遇到这种情形，即停止往深处凿掘，并且开始在盆边之间的交切处造起平面的蜡壁，所以，每一个六面柱体并不是像普通蜂房那样，建筑在三边角锥体的直边上面，而是建造在一个平滑盆形的扇形边上面的。

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But this theory can be tested by experiment. Following the example of Mr. Tegetmeier, I separated two combs, and put between them a long, thick, square strip of wax: the bees instantly began to excavate minute circular pits in it; and as they deepened these little pits, they made them wider and wider until they were converted into shallow basins, appearing to the eye perfectly true or parts of a sphere, and of about the diameter of a cell. It was most interesting to me to observe that wherever several bees had begun to excavate these basins near together, they had begun their work at such a distance from each other, that by the time the basins had acquired the above stated width (i.e. about the width of an ordinary cell), and were in depth about one sixth of the diameter of the sphere of which they formed a part, the rims of the basins intersected or broke into each other. As soon as this occurred, the bees ceased to excavate, and began to build up flat walls of wax on the lines of intersection between the basins, so that each hexagonal prism was built upon the festooned edge of a smooth basin, instead of on the straight edges of a three-sided pyramid as in the case of ordinary cells.

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然后我把一块薄而狭的涂有朱红色、其边如刃的蜡片放进蜂箱里去，以代替以前所用的方形厚蜡版。于是蜜蜂即刻一如既往地在蜡片的两面开始凿掘一些彼此接近的盆形小穴。但蜡片太薄，如果盆形小穴的底掘得像上述试验一样深，两面便要彼此串通了。然而蜜蜂并不会让这种情形发生，及时停止了开掘；于是那些盆形小穴，掘得深一点时，便出现了平的底，这等由剩下未被咬去的一小薄片朱红色蜡所形成的平底，根据目测，正好位于蜡片正反面的盆形小穴之间的想象上的交切面处。部分地方只咬去一点点，其他地方则是在对面的盆形小穴之间留下大片菱形板，不是自然状态的东西，所以不能精巧地完成工作。蜂在朱红色蜡片的两面，浑圆地咬去蜡质，并使盆形加深，其工作速度想必是差不多的，这是为了能够成功地在交切面处停止工作，而在盆形小穴之间留下平面。

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I then put into the hive, instead of a thick, square piece of wax, a thin and narrow, knife-edged ridge, coloured with vermilion. The bees instantly began on both sides to excavate little basins near to each other, in the same way as before; but the ridge of wax was so thin, that the bottoms of the basins, if they had been excavated to the same depth as in the former experiment, would have broken into each other from the opposite sides. The bees, however, did not suffer this to happen, and they stopped their excavations in due time; so that the basins, as soon as they had been a little deepened, came to have flat bottoms; and these flat bottoms, formed by thin little plates of the vermilion wax having been left ungnawed, were situated, as far as the eye could judge, exactly along the planes of imaginary intersection between the basins on the opposite sides of the ridge of wax. In parts, only little bits, in other parts, large portions of a rhombic plate had been left between the opposed basins, but the work, from the unnatural state of things, had not been neatly performed. The bees must have worked at very nearly the same rate on the opposite sides of the ridge of vermilion wax, as they circularly gnawed away and deepened the basins on both sides, in order to have succeeded in thus leaving flat plates between the basins, by stopping work along the intermediate planes or planes of intersection.

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鉴于薄蜡片十分柔软，我想，蜂在蜡片两面工作时，不难觉察到什么时候咬到适当的薄度，于是停止工作。在普通的蜂巢里，我认为蜂在两面的工作速度，并不永远能实现完全相等；我注意过一个刚开始营造的蜂房底部上半完工的菱形板，其一面稍为凹进，我想象这是这面掘得太快的缘故，另一面则凸出，因为这面工作得慢一些。在一个著名事例里，我把这蜂巢放回蜂箱里去，让蜂继续工作一个短时间，然后再检查蜂房，发现菱形板已经完工，并且已经完全平了：蜡片是极薄的，所以绝对不可能是从凸的一面把蜡咬去，做成上述的样子；我猜测这种情形大概是站在反面的蜂，把可塑而温暖的蜡恰到好处地推压弯曲到中间板处（我试验过，很容易做），这样就找平了。

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Considering how flexible thin wax is, I do not see that there is any difficulty in the bees, whilst at work on the two sides of a strip of wax, perceiving when they have gnawed the wax away to the proper thinness, and then stopping their work. In ordinary combs it has appeared to me that the bees do not always succeed in working at exactly the same rate from the opposite sides; for I have noticed half-completed rhombs at the base of a just-commenced cell, which were slightly concave on one side, where I suppose that the bees had excavated too quickly, and convex on the opposed side, where the bees had worked less quickly. In one well-marked instance, I put the comb back into the hive, and allowed the bees to go on working for a short time, and again examined the cell, and I found that the rhombic plate had been completed, and had become perfectly flat: it was absolutely impossible, from the extreme thinness of the little rhombic plate, that they could have effected this by gnawing away the convex side; and I suspect that the bees in such cases stand in the opposed cells and push and bend the ductile and warm wax (which as I have tried is easily done) into its proper intermediate plane, and thus flatten it.

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从朱红蜡片的试验可以看出，若要建造一堵蜡质的薄壁，蜂便彼此站在一定距离，以同等的速度凿掘下去，并且努力做成同等大小的球状空室，但永远不会让空室彼此串通，这样就可造适当形状的蜂房。检查一下正在建造的蜂巢边缘，一眼就可看出首先在蜂巢的周围造一堵粗糙的围墙缘边，然后从两面对咬，加深每一个蜂房时，总是绕圈工作。并不在同一时间内营造任一蜂房三边角锥形的整个底部，而是看情况先搞定位于正在建造的极端边缘的一两块菱形板；并且在没有营造六面壁之前，绝不完成菱形板上部的边。这些叙述有些和大名鼎鼎的老于贝尔所说的有所不同，但我相信是正确的；如果有篇幅，我将阐明这符合我的理论。

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From the experiment of the ridge of vermilion wax, we can clearly see that if the bees were to build for themselves a thin wall of wax, they could make their cells of the proper shape, by standing at the proper distance from each other, by excavating at the same rate, and by endeavouring to make equal spherical hollows, but never allowing the spheres to break into each other. Now bees, as may be clearly seen by examining the edge of a growing comb, do make a rough, circumferential wall or rim all round the comb; and they gnaw into this from the opposite sides, always working circularly as they deepen each cell. They do not make the whole three-sided pyramidal base of any one cell at the same time, but only the one rhombic plate which stands on the extreme growing margin, or the two plates, as the case may be; and they never complete the upper edges of the rhombic plates, until the hexagonal walls are commenced. Some of these statements differ from those made by the justly celebrated elder Huber, but I am convinced of their accuracy; and if I had space, I could show that they are conformable with my theory.

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于贝尔说，最初的第一个蜂房是从侧面相平行的蜡质小壁凿掘造出来的，就我所看到的，这一叙述并不严格正确。最初着手的经常是一个小蜡兜，但这里我不拟详论。我们知道，在蜂房的构造里，凿掘起着何等重要的作用；但如果设想蜂不能在适当的位置——即沿着两个连接的球形体之间的交切面——营造粗糙的蜡壁，就是极大的错误。我有几件标本明显指出是能够这样做的。甚至在环绕着建造中的蜂巢周围的粗糙边缘即蜡壁上，有时候也可观察到弯曲的情形，所在的位置相当于未来蜂房的菱形底面。但在一切场合中，粗糙的蜡壁是靠大口咬掉两面的蜡而完成的。蜂的这种营造方法是奇妙的；总是把最初的粗糙墙壁，造得比最后要留下的蜂房的极薄的壁厚十倍乃至二十倍。要理解它们的工作方法，可以假定泥水匠首先用水泥堆起一堵宽阔基墙，然后在近地面处的两侧把水泥同等地削去，直到中间部分形成一堵光滑而很薄的墙壁；泥水匠总是把削去的水泥堆在墙壁的顶上，还要加入新水泥。于是，薄壁就这样不断地垒上去，但上面总是有一个厚大的顶盖。一切蜂房，无论刚开始营造还是已经完成的，上面都有这样一个坚固的蜡盖，因此，蜂能够聚集在蜂巢上爬来爬去，而不会把薄六面壁损坏。壁的厚度只有约四百分之一英寸，菱形底片大约比其厚一倍。用上述这样特别的营造方法，可以极端地省蜡，同时还能不断地使蜂巢加固。

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Huber’s statement that the very first cell is excavated out of a little parallel-sided wall of wax, is not, as far as I have seen, strictly correct; the first commencement having always been a little hood of wax; but I will not here enter on these details. We see how important a part excavation plays in the construction of the cells; but it would be a great error to suppose that the bees cannot build up a rough wall of wax in the proper position—that is, along the plane of intersection between two adjoining spheres. I have several specimens showing clearly that they can do this. Even in the rude circumferential rim or wall of wax round a growing comb, flexures may sometimes be observed, corresponding in position to the planes of the rhombic basal plates of future cells. But the rough wall of wax has in every case to be finished off, by being largely gnawed away on both sides. The manner in which the bees build is curious; they always make the first rough wall from ten to twenty times thicker than the excessively thin finished wall of the cell, which will ultimately be left. We shall understand how they work, by supposing masons first to pile up a broad ridge of cement, and then to begin cutting it away equally on both sides near the ground, till a smooth, very thin wall is left in the middle; the masons always piling up the cut-away cement, and adding fresh cement, on the summit of the ridge. We shall thus have a thin wall steadily growing upward; but always crowned by a gigantic coping. From all the cells, both those just commenced and those completed, being thus crowned by a strong coping of wax, the bees can cluster and crawl over the comb without injuring the delicate hexagonal walls, which are only about one four-hundredth of an inch in thickness; the plates of the pyramidal basis being about twice as thick. By this singular manner of building, strength is continually given to the comb, with the utmost ultimate economy of wax.

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大批蜜蜂聚集一起工作，乍看这对于理解蜂房的营造方式会增加困难；一只蜂在一个蜂房工作一个短时间后，便到另一个蜂房，所以，如于贝尔所说的，甚至第一个蜂房开始营造时就有二十只蜂在工作。我用实践的方法阐明了这一事实：用朱红色的极薄熔蜡涂在一个蜂房六面壁的边上，或者涂在一个营造着的蜂巢围墙的极端边缘上，结果必定看到蜂把这颜色极细腻地分布开去——细腻得就像画家布色——有颜色的蜡从涂抹的地方一星一星地取去，放到周围蜂房扩大着的边缘上去。营造的工作在多蜂之间似乎有某种平衡分配，彼此本能地站在同一相对距离上，都试图凿掘相等的球形，然后，建造起或者说留下不咬球形之间的交切面。说起来实在是奇异，有时会遇到困难，例如两个蜂巢成角度相遇时，往往把已成的蜂房彻底拆掉，用不同的方法重造，而有时候再现拆去的形状。

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It seems at first to add to the difficulty of understanding how the cells are made, that a multitude of bees all work together; one bee after working a short time at one cell going to another, so that, as Huber has stated, a score of individuals work even at the commencement of the first cell. I was able practically to show this fact, by covering the edges of the hexagonal walls of a single cell, or the extreme margin of the circumferential rim of a growing comb, with an extremely thin layer of melted vermilion wax; and I invariably found that the colour was most delicately diffused by the bees—as delicately as a painter could have done with his brush—by atoms of the coloured wax having been taken from the spot on which it had been placed, and worked into the growing edges of the cells all round. The work of construction seems to be a sort of balance struck between many bees, all instinctively standing at the same relative distance from each other, all trying to sweep equal spheres, and then building up, or leaving ungnawed, the planes of intersection between these spheres. It was really curious to note in cases of difficulty, as when two pieces of comb met at an angle, how often the bees would entirely pull down and rebuild in different ways the same cell, sometimes recurring to a shape which they had at first rejected.

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蜂遇到可以各就各位进行工作的一处地方，例如，一块木片上，木片恰好处于向下建造的一个蜂巢的中部之下，那么这蜂巢势必就要营造在木片的一面——在这种情况下，蜂便会筑起新的六面体一堵墙的基础，突出于已经完成的蜂房之外，位置严格规定。只要蜂能彼此站在适当的距离并且与最后完成的蜂房墙壁保持适当的距离，掘造了想象的球形体，就足以在两个邻接的球形体之间造起中间蜡壁来；但据我所看到的，非到那蜂房和邻接的几个蜂房大都造成之后，从不咬去和修光蜂房的角。蜂在一定环境条件下，能在两个刚开始营造的蜂房中间把一堵粗糙的壁建立在适当位置上，这种能力是重要的；因为这与一项事实有关，最初看来它似乎可以推翻上述理论；这事实就是，黄蜂最外边缘上的一些蜂房也常常是严格的六边形；但这里没有篇幅讨论这一问题。我并不觉得单独一个昆虫（例如黄蜂蜂后）营造六边形的蜂房会有什么大困难，只要能在同时开始的两三个巢房的内侧和外侧交互地工作，始终与刚开工的蜂房部件保持适当的距离，掘造球形或圆筒形，并且建造起中间的平壁。甚至可以想象，一个昆虫固定于一点开始构筑蜂房，然后移动出去，先到一点，然后到另外五个点，到中心点的相对距离和点之间的距离恰到好处，可以打造诸交切面，构筑孤立的六边形。但我不知道这种个案有没有观察到过，而且构筑单个六边形也没有什么好处，因为这就比构筑圆柱体需要更多的建筑材料。

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When bees have a place on which they can stand in their proper positions for working,—for instance, on a slip of wood, placed directly under the middle of a comb growing downwards so that the comb has to be built over one face of the slip—in this case the bees can lay the foundations of one wall of a new hexagon, in its strictly proper place, projecting beyond the other completed cells. It suffices that the bees should be enabled to stand at their proper relative distances from each other and from the walls of the last completed cells, and then, by striking imaginary spheres, they can build up a wall intermediate between two adjoining spheres; but, as far as I have seen, they never gnaw away and finish off the angles of a cell till a large part both of that cell and of the adjoining cells has been built. This capacity in bees of laying down under certain circumstances a rough wall in its proper place between two just-commenced cells, is important, as it bears on a fact, which seems at first quite subversive of the foregoing theory; namely, that the cells on the extreme margin of wasp-combs are sometimes strictly hexagonal; but I have not space here to enter on this subject. Nor does there seem to me any great difficulty in a single insect (as in the case of a queen-wasp) making hexagonal cells, if she work alternately on the inside and outside of two or three cells commenced at the same time, always standing at the proper relative distance from the parts of the cells just begun, sweeping spheres or cylinders, and building up intermediate planes. It is even conceivable that an insect might, by fixing on a point at which to commence a cell, and then moving outside, first to one point, and then to five other points, at the proper relative distances from the central point and from each other, strike the planes of intersection, and so make an isolated hexagon: but I am not aware that any such case has been observed; nor would any good be derived from a single hexagon being built, as in its construction more materials would be required than for a cylinder.

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自然选择仅仅靠构造或本能的微小变异的积累才发挥作用，而各个变异都对个体在其生活条件下是有利的。所以可以合理地发问：变异了的建筑本能所经历的漫长而级进的连续阶段，都趋向现今那样完善的建筑规划，对于蜜蜂祖先，曾起过怎样有利的作用呢？我想，解答这个问题并不困难：我们知道，蜂为了采足花蜜，常常受到很大压力。特盖特迈耶先生告诉我说，实验已经证明，蜜蜂分泌一磅蜡须消耗十二到十五磅干糖；所以一个蜂箱里的蜜蜂为了分泌营造蜂巢所必需的蜡，必须采集并消耗大量的液状花蜜。还有，许多蜂在分泌的过程中，势必有许多天不能工作。大量蜂蜜的贮藏，对于维持大群蜂的冬季生活是必不可缺少的；并且我们知道，蜂群的安全主要决定于大量的蜂得以维持。因此，大大节省蜂蜜，从而省蜡，必定是任何蜂族成功的重要因素。当然，其成功还可能决定于寄生物等敌害的数量，决定于截然不同的原因，所以根本不取决于蜜蜂所能采集的蜜量。但是，让我们假定采集蜜量的能力决定了任何一处地方大黄蜂的数量，这倒是常常发生；让我们进一步假定，那蜂群度过了冬季，结果就需要贮藏蜂蜜：在这种情形下，如果其本能有微小的变异，使得蜡房造得靠近些，略略彼此相切，无疑会有利于这批土蜂；一堵公共的壁即使仅连接两个蜂房，也会节省少许蜡。因此，如果蜂房造得日益整齐，相互日益靠近，并且像墨西哥蜂的蜂房那样聚集在一起，就会不断地日益有利于这种大黄蜂；因为这样各蜂房的大部分界壁将会用作邻接蜂房的界壁，就可以大大省蜡。还有，由于同样的原因，如果墨西哥蜂能把蜂房造得比现在接近些，并且在各方面都更规则些，这于己有利；因为，如我们所看到的，蜂房的球形面会完全消失，代以平面；而墨西哥蜂所造的蜂巢就会达到蜜蜂巢那样完善的地步。在建造上超越这种完善的阶段，自然选择便不能再起作用；因为据我们所知，蜜蜂巢在节蜡方面绝对是完美的。

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As natural selection acts only by the accumulation of slight modifications of structure or instinct, each profitable to the individual under its conditions of life, it may reasonably be asked, how a long and graduated succession of modified architectural instincts, all tending towards the present perfect plan of construction, could have profited the progenitors of the hive-bee? I think the answer is not difficult: it is known that bees are often hard pressed to get sufficient nectar; and I am informed by Mr. Tegetmeier that it has been experimentally found that no less than from twelve to fifteen pounds of dry sugar are consumed by a hive of bees for the secretion of each pound of wax; so that a prodigious quantity of fluid nectar must be collected and consumed by the bees in a hive for the secretion of the wax necessary for the construction of their combs. Moreover, many bees have to remain idle for many days during the process of secretion. A large store of honey is indispensable to support a large stock of bees during the winter; and the security of the hive is known mainly to depend on a large number of bees being supported. Hence the saving of wax by largely saving honey must be a most important element of success in any family of bees. Of course the success of any species of bee may be dependent on the number of its parasites or other enemies, or on quite distinct causes, and so be altogether independent of the quantity of honey which the bees could collect. But let us suppose that this latter circumstance determined, as it probably often does determine, the numbers of a humble-bee which could exist in a country; and let us further suppose that the community lived throughout the winter, and consequently required a store of honey: there can in this case be no doubt that it would be an advantage to our humble-bee, if a slight modification of her instinct led her to make her waxen cells near together, so as to intersect a little; for a wall in common even to two adjoining cells, would save some little wax. Hence it would continually be more and more advantageous to our humble-bee, if she were to make her cells more and more regular, nearer together, and aggregated into a mass, like the cells of the Melipona; for in this case a large part of the bounding surface of each cell would serve to bound other cells, and much wax would be saved. Again, from the same cause, it would be advantageous to the Melipona, if she were to make her cells closer together, and more regular in every way than at present; for then, as we have seen, the spherical surfaces would wholly disappear, and would all be replaced by plane surfaces; and the Melipona would make a comb as perfect as that of the hive-bee. Beyond this stage of perfection in architecture, natural selection could not lead; for the comb of the hive-bee, as far as we can see, is absolutely perfect in economising wax.

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因此，我认为，一切既知本能中最奇异的本能——蜜蜂的本能，可以根据自然选择利用了简单本能之无数的、连续发生的微小变异来解释；自然选择曾经缓慢、逐步完善地使得蜂在双层上掘造彼此保持一定距离的、同等大小的球形体，并且沿着交切面筑起和凿掘蜡壁。当然，蜂不会知道自己在彼此保持一定距离掘造球形体，正如它们不会知道六面柱体与底部的菱形板是什么角度。自然选择过程的动力在于节蜡；各蜂群在蜡的分泌上消耗最少的蜜，得到了最大的成功，并且把新获得的节约本能遗传给了新蜂群，以便在生存斗争中获得成功的最大机会。

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Thus, as I believe, the most wonderful of all known instincts, that of the hive-bee, can be explained by natural selection having taken advantage of numerous, successive, slight modifications of simpler instincts; natural selection having by slow degrees, more and more perfectly, led the bees to sweep equal spheres at a given distance from each other in a double layer, and to build up and excavate the wax along the planes of intersection. The bees, of course, no more knowing that they swept their spheres at one particular distance from each other, than they know what are the several angles of the hexagonal prisms and of the basal rhombic plates. The motive power of the process of natural selection having been economy of wax; that individual swarm which wasted least honey in the secretion of wax, having succeeded best, and having transmitted by inheritance its newly acquired economical instinct to new swarms, which in their turn will have had the best chance of succeeding in the struggle for existence.

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无疑还可用许多极难解释的本能来反对自然选择学说——例如有些本能，我们不知道是怎样起源的；有些本能，我们不知道有中间级进存在；有些本能看上去很不重要，自然选择不大会发生作用；有些本能在自然系统相距甚远的动物里竟几乎相同，所以不能用共同祖先的遗传来说明其相似性，结果只好相信这些本能是通过自然选择而独立获得的。我不预备在这里讨论这些个例子，而仅仅讨论一个特别的难点，起初我认为这个难点是难以克服的，并且实际上对于我的整个理论是致命的。我所指的就是昆虫社会里的中性即不育的雌虫；这些中性虫在本能和构造上常与雄虫以及能育的雌虫有很大的差异，可是由于不育，却不能繁殖同类。

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No doubt many instincts of very difficult explanation could be opposed to the theory of natural selection,—cases, in which we cannot see how an instinct could possibly have originated; cases, in which no intermediate gradations are known to exist; cases of instinct of apparently such trifling importance, that they could hardly have been acted on by natural selection; cases of instincts almost identically the same in animals so remote in the scale of nature, that we cannot account for their similarity by inheritance from a common parent, and must therefore believe that they have been acquired by independent acts of natural selection. I will not here enter on these several cases, but will confine myself to one special difficulty, which at first appeared to me insuperable, and actually fatal to my whole theory. I allude to the neuters or sterile females in insect-communities: for these neuters often differ widely in instinct and in structure from both the males and fertile females, and yet, from being sterile, they cannot propagate their kind.

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这个问题很值得详细讨论，但这里只举一个个案，即不育的工蚁。工蚁怎么会变为不育的个体是个难点，但不比构造上任何显著变异更难于解释；可以证明，自然状态下某些昆虫以及别种节足动物偶尔也会变为不育；如果这种昆虫是社会性的，而且每年生下若干能工作但不能生殖的个体对于群体有利的话，那我认为不难理解这是由于自然选择的作用。但必须跳过这种初步的难点不谈。最大的难点在于工蚁与雄蚁和能育的雌蚁在构造上有巨大的差异，如工蚁具有不同形状的胸部，缺翅膀，有时没有眼睛，并且具有不同的本能。单以本能而论，蜜蜂可以极好地证明工蜂与完全的雌蜂之间有惊人的差异。如果工蚁或别种中性虫原是正常的动物，那我就会毫不迟疑地假定，一切性状都是通过自然选择慢慢获得的；这就是说，由于生下的个体构造上都具有微小的有利变异，又都遗传给了后代；而且后代又发生变异，又被选择，如此等等，不一而足。但是工蚁和双亲之间的差异很大，又是绝对不育的，所以绝不可能把历代获得的构造上或本能上的变异遗传给后代。于是可以设问：这怎么能符合自然选择的学说呢？

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The subject well deserves to be discussed at great length, but I will here take only a single case, that of working or sterile ants. How the workers have been rendered sterile is a difficulty; but not much greater than that of any other striking modification of structure; for it can be shown that some insects and other articulate animals in a state of nature occasionally become sterile; and if such insects had been social, and it had been profitable to the community that a number should have been annually born capable of work, but incapable of procreation, I can see no very great difficulty in this being effected by natural selection. But I must pass over this preliminary difficulty. The great difficulty lies in the working ants differing widely from both the males and the fertile females in structure, as in the shape of the thorax and in being destitute of wings and sometimes of eyes, and in instinct. As far as instinct alone is concerned, the prodigious difference in this respect between the workers and the perfect females, would have been far better exemplified by the hive-bee. If a working ant or other neuter insect had been an animal in the ordinary state, I should have unhesitatingly assumed that all its characters had been slowly acquired through natural selection; namely, by an individual having been born with some slight profitable modification of structure, this being inherited by its offspring, which again varied and were again selected, and so onwards. But with the working ant we have an insect differing greatly from its parents, yet absolutely sterile; so that it could never have transmitted successively acquired modifications of structure or instinct to its progeny. It may well be asked how is it possible to reconcile this case with the theory of natural selection?

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首先，请记住，家养生物和自然状态下的生物里，构造的各种各样差异是与一定年龄或性别相关的，这方面有无数的案例。差异不但与性别相关，而且与生殖系统活跃的那一短暂时期相关，例如，许多鸟类的求婚羽，雄三文鱼钩曲的颚，都是这种情形。公牛经人工去势后，不同品种的角甚至相关地表现了微小的差异；某些品种的去势公牛，与同品种的公牝双方比较，犄角比其他品种更长。因此，我认为任何性状变得与昆虫社会里某些成员的不育状态相关，并不存在多大难点；难点在于理解这种构造上的相关变异如何因自然选择作用而慢慢累积起来。

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First, let it be remembered that we have innumerable instances, both in our domestic productions and in those in a state of nature, of all sorts of differences of structure which have become correlated to certain ages, and to either sex. We have differences correlated not only to one sex, but to that short period alone when the reproductive system is active, as in the nuptial plumage of many birds, and in the hooked jaws of the male salmon. We have even slight differences in the horns of different breeds of cattle in relation to an artificially imperfect state of the male sex; for oxen of certain breeds have longer horns than in other breeds, in comparison with the horns of the bulls or cows of these same breeds. Hence I can see no real difficulty in any character having become correlated with the sterile condition of certain members of insect-communities: the difficulty lies in understanding how such correlated modifications of structure could have been slowly accumulated by natural selection.

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这个难点表面上看来是难以克服的，可是只要记住选择作用可以应用于个体也可以应用于全族，而且可以由此如愿以偿，那么难点便会缩小，或者如我所相信的，便会消除。比如，一棵味道好的蔬菜煮熟吃了，该个体就消灭了。可是园艺家播下同种蔬菜的种子，信心十足地期望收获差不多的变种。养牛者喜欢肉和脂肪交织成大理石纹的样子，牲口已经屠杀了，但是养牛者有信心继续找到同样的牛。我对于选择的力量也是信心十足，并不怀疑总是产生异常长角的去势公牛的品种，可以慢慢培养，只要仔细观察什么样的公牛和牝牛个体交配才能产生最长角的去势公牛；虽然没有一只去势的牛曾经繁殖过同类。我想社会性的昆虫也是如此：与同群某些成员的不育状态相关的构造、本能上的轻微变异，对于群体有利，结果能育的雄体和雌体得到了繁生，并把这种产生具有同样变异的不育成员的倾向，传递给了能育的后代。我认为，这一过程重复过了许多次，直到同一物种的能育雌体和不育雌体之间产生了巨大的差异量，就像我们在许多种社会性昆虫里所见到的那样。

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This difficulty, though appearing insuperable, is lessened, or, as I believe, disappears, when it is remembered that selection may be applied to the family, as well as to the individual, and may thus gain the desired end. Thus, a well-flavoured vegetable is cooked, and the individual is destroyed; but the horticulturist sows seeds of the same stock, and confidently expects to get nearly the same variety; breeders of cattle wish the flesh and fat to be well marbled together; the animal has been slaughtered, but the breeder goes with confidence to the same family. I have such faith in the powers of selection, that I do not doubt that a breed of cattle, always yielding oxen with extraordinarily long horns, could be slowly formed by carefully watching which individual bulls and cows, when matched, produced oxen with the longest horns; and yet no one ox could ever have propagated its kind. Thus I believe it has been with social insects: a slight modification of structure, or instinct, correlated with the sterile condition of certain members of the community, has been advantageous to the community: consequently the fertile males and females of the same community flourished, and transmitted to their fertile offspring a tendency to produce sterile members having the same modification. And I believe that this process has been repeated, until that prodigious amount of difference between the fertile and sterile females of the same species has been produced, which we see in many social insects.

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但我们还没有谈及登峰造极的难点：有几种蚁的中性虫不但与能育的雌虫和雄虫有所差异，而且彼此之间也有差异，有时差异甚至到了让人几乎难以置信的程度，并且因此被分成两个级（castes），甚至三个级。还有，这些级一般并不彼此逐渐重叠，而是区别得十分清楚，有如同属的两个物种，同科的两个属。例如，埃西顿（Eciton）行军蚁的中性工蚁和兵蚁具有大相径庭的颚和本能：隐角蚁（Cryptocerus）只有一个级的工蚁，头上生有一种奇异的盾，用途不清楚；墨西哥的蜜蚁（Myrmecocystus）有一个级的工蚁从不离巢，由另一个级的工蚁喂食，腹部发育得很大，能分泌出一种蜜汁，以代替蚜虫所排泄的东西；蚜虫或者可以被称为蚁乳牛，欧洲的蚁常把它们守卫圈禁起来。

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But we have not as yet touched on the climax of the difficulty; namely, the fact that the neuters of several ants differ, not only from the fertile females and males, but from each other, sometimes to an almost incredible degree, and are thus divided into two or even three castes. The castes, moreover, do not generally graduate into each other, but are perfectly well defined; being as distinct from each other, as are any two species of the same genus, or rather as any two genera of the same family. Thus in Eciton, there are working and soldier neuters, with jaws and instincts extraordinarily different: in Cryptocerus, the workers of one caste alone carry a wonderful sort of shield on their heads, the use of which is quite unknown: in the Mexican Myrmecocystus, the workers of one caste never leave the nest; they are fed by the workers of another caste, and they have an enormously developed abdomen which secretes a sort of honey, supplying the place of that excreted by the aphides, or the domestic cattle as they may be called, which our European ants guard or imprison.

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如果不承认这种奇异而十分确实的事实可以瞬间颠覆我的理论，人们必然会想，我对自然选择的原理过于信心饱满了。如果中性虫只有一个级，我相信它与能育的雄虫和雌虫之间的差异是通过自然选择得到的，在这种比较简单的情形里，根据普通变异类推，我们可以断言，各种连续的、微小的、有利的变异，最初并非发生于同一窝中的所有中性虫，而只发生于少数的中性虫；经过长期持续选择能够产生极多的具有有利变异的中性虫的能育亲种，一切中性虫最终就都会具有所需的性状。按照这种观点，我们应该在同一巢中偶尔发现那些表现有构造分级的同种中性虫；实际我们是发现了，鉴于欧洲以外的中性昆虫很少仔细检查过，甚至可以说并不稀罕。史密斯先生曾阐明，有几种英国蚁的中性虫彼此在大小方面，有时在颜色方面，表现了惊人的差异；并且在两极端的类型之间，有时可由同巢中的一些个体连接起来：笔者就比较过这种完美的级进情形。有时可以看到，大形或者小形的工蚁数目最多；或者大形和小形两种都多，而中间形的数目却稀少。黄蚁有大工蚁和小工蚁，中间形的工蚁有一些；如史密斯先生所观察的，在这个物种里，大工蚁有单眼（ocelli），虽小但能够清楚辨认；而小工蚁的单眼则是残迹。仔细解剖了几只工蚁标本之后，我能确定小工蚁的眼睛根本不发育，远非单单用其小比例所能解释；并且我充分相信，虽然我不敢很肯定地断言，中间形工蚁的单眼正好处在中间状态。所以，一个巢内有两群不育工蚁，不但在大小上，并且在视觉器官上，都表现了差异，然而有少数中间状态的成员连接起来。我再补充几句题外的话，如果小工蚁对于蚁群最有利，产生越来越多小工蚁的雄蚁和雌蚁不断被选择，最后所有的工蚁都具有那种形态了。于是就形成了一个蚁种，其中性虫差不多就像褐蚁属（Myrmica）工蚁那样。褐蚁属的工蚁甚至连残迹的单眼都没有，尽管这个属的雄蚁和雌蚁都生有很发达的单眼。

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It will indeed be thought that I have an overweening confidence in the principle of natural selection, when I do not admit that such wonderful and well-established facts at once annihilate my theory. In the simpler case of neuter insects all of one caste or of the same kind, which have been rendered by natural selection, as I believe to be quite possible, different from the fertile males and females,—in this case, we may safely conclude from the analogy of ordinary variations, that each successive, slight, profitable modification did not probably at first appear in all the individual neuters in the same nest, but in a few alone; and that by the long-continued selection of the fertile parents which produced most neuters with the profitable modification, all the neuters ultimately came to have the desired character. On this view we ought occasionally to find neuter-insects of the same species, in the same nest, presenting gradations of structure; and this we do find, even often, considering how few neuter-insects out of Europe have been carefully examined. Mr. F. Smith has shown how surprisingly the neuters of several British ants differ from each other in size and sometimes in colour; and that the extreme forms can sometimes be perfectly linked together by individuals taken out of the same nest: I have myself compared perfect gradations of this kind. It often happens that the larger or the smaller sized workers are the most numerous; or that both large and small are numerous, with those of an intermediate size scanty in numbers. Formica flava has larger and smaller workers, with some of intermediate size; and, in this species, as Mr. F. Smith has observed, the larger workers have simple eyes (ocelli), which though small can be plainly distinguished, whereas the smaller workers have their ocelli rudimentary. Having carefully dissected several specimens of these workers, I can affirm that the eyes are far more rudimentary in the smaller workers than can be accounted for merely by their proportionally lesser size; and I fully believe, though I dare not assert so positively, that the workers of intermediate size have their ocelli in an exactly intermediate condition. So that we here have two bodies of sterile workers in the same nest, differing not only in size, but in their organs of vision, yet connected by some few members in an intermediate condition. I may digress by adding, that if the smaller workers had been the most useful to the community, and those males and females had been continually selected, which produced more and more of the smaller workers, until all the workers had come to be in this condition; we should then have had a species of ant with neuters very nearly in the same condition with those of Myrmica. For the workers of Myrmica have not even rudiments of ocelli, though the male and female ants of this genus have well- developed ocelli.

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我再举一例：同一物种的不同级的中性虫之间，我满有信心地期望可以找到重要构造的中间分级，欣然利用史密斯先生所提供的取自西非驱逐蚁（Anomma）同巢中的许多标本。我不举实际的测量数字，只做一个严格精确的说明，读者大概就能最好地了解这工蚁之间的差异量；差异就好比看到一群建筑工人，其中有许多是五英尺四英寸高，还有许多是十六英尺高；但我们必须再假定那大个儿工人的头比小个儿工人不止大三倍，却要大四倍，而颚则要大近五倍。再者，大小不同的工蚁的颚不仅在形状上大有差异，而且牙齿的形状和数目也相差悬殊。但重要的事实却是，虽然工蚁可以依大小分为不同的等级，却不知不觉地彼此级进重叠，其构造大不相同的颚也是这样。关于后面一点我有把握，卢伯克先生曾用描图器把我所解剖的几种大小不同的工蚁的颚逐一作图。

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I may give one other case: so confidently did I expect to find gradations in important points of structure between the different castes of neuters in the same species, that I gladly availed myself of Mr. F. Smith’s offer of numerous specimens from the same nest of the driver ant (Anomma) of West Africa. The reader will perhaps best appreciate the amount of difference in these workers, by my giving not the actual measurements, but a strictly accurate illustration: the difference was the same as if we were to see a set of workmen building a house of whom many were five feet four inches high, and many sixteen feet high; but we must suppose that the larger workmen had heads four instead of three times as big as those of the smaller men, and jaws nearly five times as big. The jaws, moreover, of the working ants of the several sizes differed wonderfully in shape, and in the form and number of the teeth. But the important fact for us is, that though the workers can be grouped into castes of different sizes, yet they graduate insensibly into each other, as does the widely-different structure of their jaws. I speak confidently on this latter point, as Mr. Lubbock made drawings for me with the camera lucida of the jaws which I had dissected from the workers of the several sizes.

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根据摆在我面前的这些事实，我相信自然选择由于作用于能育的蚁双亲，便可以形成一个物种，专门产生体形大而具有某一形状的颚的中性虫，或者专门产生体形小而构造大不相同的颚的中性虫；最后，这是登峰造极的难点，一群工蚁具有一种大小和构造，另一群工蚁具有不同大小和构造，同时存在——最先形成的是一个级进的系列，就像驱逐蚁的情形那样，然后，由于自然选择支持生育它们的双亲，就产生了越来越多最有利于蚁群的两极端类型，最后具有中间构造的个体不再产生。

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With these facts before me, I believe that natural selection, by acting on the fertile parents, could form a species which should regularly produce neuters, either all of large size with one form of jaw, or all of small size with jaws having a widely different structure; or lastly, and this is our climax of difficulty, one set of workers of one size and structure, and simultaneously another set of workers of a different size and structure;—a graduated series having been first formed, as in the case of the driver ant, and then the extreme forms, from being the most useful to the community, having been produced in greater and greater numbers through the natural selection of the parents which generated them; until none with an intermediate structure were produced.

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我认为，奇异事实就是这样发生的，同一巢里生存的、区别分明的不育工蚁两级，不但彼此之间大不相同，并且和双亲之间也大不相同。我们可以看出工蚁的生成对于蚁社会有多大的用处，与社会分工对于文明人的用处同理。由于蚁是用遗传的本能和遗传的工具或武器来工作的，而不用学得的知识和制造的器具，所以完美分工只能通过工蚁不育来实施。如果它们能育，就会杂交，其本能和构造就会混杂。我认为，大自然通过自然选择在蚁群中实施了这一令人惊叹的分工。但是必须坦白承认，我虽然完全相信自然选择，若不是有这等中性虫个案让我心悦诚服，决不会料到这一原理是如此高度有效。所以，为了阐明自然选择的力量，并且因为这是我的理论所遭到的特别严重的难点，我对于这个个案做了稍多的但挂一漏万的讨论。而且这个个案也很有趣，证明了动物同植物一样，把无数的微小的必须称为偶发的构造变异，只要是稍微有利的就累积下来，没有锻炼或习性参加作用，任何量的变异都能实现。蚁群的不育成员的锻炼，或者习性，或者意愿，丝毫也不可能影响专事遗留后代的能育成员的构造或者本能。我觉得奇怪的是，至今没有人提出用这种中性虫的演示个案去反对众所熟知的拉马克（Lamarck）的学说。

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Thus, as I believe, the wonderful fact of two distinctly defined castes of sterile workers existing in the same nest, both widely different from each other and from their parents, has originated. We can see how useful their production may have been to a social community of insects, on the same principle that the division of labour is useful to civilised man. As ants work by inherited instincts and by inherited tools or weapons, and not by acquired knowledge and manufactured instruments, a perfect division of labour could be effected with them only by the workers being sterile; for had they been fertile, they would have intercrossed, and their instincts and structure would have become blended. And nature has, as I believe, effected this admirable division of labour in the communities of ants, by the means of natural selection. But I am bound to confess, that, with all my faith in this principle, I should never have anticipated that natural selection could have been efficient in so high a degree, had not the case of these neuter insects convinced me of the fact. I have, therefore, discussed this case, at some little but wholly insufficient length, in order to show the power of natural selection, and likewise because this is by far the most serious special difficulty, which my theory has encountered. The case, also, is very interesting, as it proves that with animals, as with plants, any amount of modification in structure can be effected by the accumulation of numerous, slight, and as we must call them accidental, variations, which are in any manner profitable, without exercise or habit having come into play. For no amount of exercise, or habit, or volition, in the utterly sterile members of a community could possibly have affected the structure or instincts of the fertile members, which alone leave descendants. I am surprised that no one has advanced this demonstrative case of neuter insects, against the well-known doctrine of Lamarck.

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提要。——本章勉力简要地指出了家养动物的精神品质要变异，且这种变异是遗传的。我又更简要地阐明本能在自然状态下也是轻微变异的。没有人会争辩本能对于各种动物有极端的重要性。所以我认为，在多变的生活条件下，自然选择不难把任何稍微有用的本能上的微小变异累积到任何程度。在许多情况下，习性或者用废大概也参加作用。我不敢说本章所举事实能在很大程度上巩固我的理论；但是根据我所能判断的，没有难解的个案可加以颠覆。另外，本能不总是绝对完善，而是易犯错误；没有一种本能可说是为了其他动物的独享利益而产生的，但各种动物都利用其他动物的本能；博物史上的格言“自然界里没有飞跃”，适用于身体构造也适用于本能，并且可用上述观点来清楚解释，别无他解——所有这些事实都确证了自然选择的学说。

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Summary.—I have endeavoured briefly in this chapter to show that the mental qualities of our domestic animals vary, and that the variations are inherited. Still more briefly I have attempted to show that instincts vary slightly in a state of nature. No one will dispute that instincts are of the highest importance to each animal. Therefore I can see no difficulty, under changing conditions of life, in natural selection accumulating slight modifications of instinct to any extent, in any useful direction. In some cases habit or use and disuse have probably come into play. I do not pretend that the facts given in this chapter strengthen in any great degree my theory; but none of the cases of difficulty, to the best of my judgment, annihilate it. On the other hand, the fact that instincts are not always absolutely perfect and are liable to mistakes;—that no instinct has been produced for the exclusive good of other animals, but that each animal takes advantage of the instincts of others;—that the canon in natural history, of “natura non facit saltum” is applicable to instincts as well as to corporeal structure, and is plainly explicable on the foregoing views, but is otherwise inexplicable,—all tend to corroborate the theory of natural selection.

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这个理论也得到其他几种关于本能的事实的加强；常见的个案如，密切近似的但不相同的物种，当天各一方并且生活在相当不同的生活条件下时，常常保持了几乎同样的本能。例如，根据遗传原理，我们能够理解，为什么南美鸫跟英国鸫的特别造巢方法一样，用泥来涂抹它们的巢；为什么北美洲的雄性鹪鹩（Troglodytes）像英国的雄性猫形鹪鹩（Kitty-wrens）那样地营造“雄鸟之巢”栖居——这种习性完全不像任何其他已知鸟类。最后，这可能是不合逻辑的演绎，但据我想象，这样说法最能令人满意，如小杜鹃把义兄弟逐出巢外，蓄奴蚁，姬蜂科（ichneumonidae）幼虫寄生在活的毛毛虫体内，不把这种本能看作是特别赋予或特别创造的，而看作是引导一切生物进化——即繁殖，变异，让最强者生存、最弱者死亡——的一般法则的小小结果。

51
-

This theory is, also, strengthened by some few other facts in regard to instincts; as by that common case of closely allied, but certainly distinct, species, when inhabiting distant parts of the world and living under considerably different conditions of life, yet often retaining nearly the same instincts. For instance, we can understand on the principle of inheritance, how it is that the thrush of South America lines its nest with mud, in the same peculiar manner as does our British thrush: how it is that the male wrens (Troglodytes) of North America, build “cock-nests,” to roost in, like the males of our distinct Kitty-wrens,—a habit wholly unlike that of any other known bird. Finally, it may not be a logical deduction, but to my imagination it is far more satisfactory to look at such instincts as the young cuckoo ejecting its foster-brothers,—ants making slaves,—the larvae of ichneumonidae feeding within the live bodies of caterpillars,—not as specially endowed or created instincts, but as small consequences of one general law, leading to the advancement of all organic beings, namely, multiply, vary, let the strongest live and the weakest die.

序号 英文/音标 中文解释 更多操作

Neuter

[’njuːtə(r)]

adj.中性的;不及物的;生殖器不完全的

overthrow

[’əʊvəθrəʊ]

v.推翻;打倒;颠覆;【体】高投

embraceable

[ɪm’breɪsəbl]

v.拥抱;包含;包围;接受;信奉

impel

[ɪm’pel]

v.驱使;激励

instinctive

[ɪn’stɪŋktɪv]

adj.本能的;天性的;直觉的

caterpillar

[’kætəpɪlə(r)]

n. 【定语】履带式...;

Mozart

[’məʊtsɑːt]

n.莫扎特

inheritance

[ɪn’herɪtəns]

n.遗传;遗产;继承;继承物

acquaint

[ə’kweɪnt]

vt.使了解;使熟知;告知

modification

[ˌmɒdɪfɪ’keɪʃn]

n.修改;修正;修饰

continual

[kən’tɪnjuəl]

adj.不断的;频繁的;连续的

disuse

[dɪs’juːs]

n.废止

deviation

[ˌdiːvi’eɪʃn]

n.偏离;出轨;背弃

bodily

[’bɒdɪli]

adj.肉体的;身体的;具体的

gradual

[’ɡrædʒuəl]

adj.逐渐的;逐步的;平缓的

transitional

[træn’sɪʃənl]

adj.变化的;过渡期的;过渡性的

collateral

[kə’lætərəl]

adj.旁系的;并行的;附属的

descent

[dɪ’sent]

n.下降;下坡;家世;血统;侵袭;衰落;继承

allowance

[ə’laʊəns]

n.津贴

extinct

[ɪk’stɪŋkt]

adj.灭绝的;熄灭的;消亡的

voluntarily

[’vɒləntrəli]

adv.自动地;以自由意志

excretion

[ɪk’skriːʃn]

n.排泄;分泌;排泄物

felted

[’feltɪd]

v. 把 ... 制成毡(使 ... 粘结)

excrete

[ɪk’skriːt]

v.排泄;分泌

tickle

[’tɪkl]

vt.使发痒;逗乐;使快乐

antenna

[æn’tenə]

n.触角;天线

abdomen

[’æbdəmən]

n.腹部

liftable

[lɪftəbl]

a.1. 可以举起的

devour

[dɪ’vaʊə(r)]

v.吞食;毁灭;贪婪地阅读

migratory

[’maɪɡrətri]

adj.迁移的;流浪的

dependence

[dɪ’pendəns]

n.依赖;信赖;上瘾

inhabit

[ɪn’hæbɪt]

v.居住于;占据;栖息

wholly

[’həʊlli]

adv.完全地;全部地;一概

nestle

[’nesl]

v.依偎;(舒适地)安顿

persecute

[’pɜːsɪkjuːt]

vt.迫害;烦扰;为难

fearful

[’fɪəfl]

adj.担心的;可怕的;非常的

hood

[hʊd]

n.头巾;兜帽;遮罩;引擎盖

disposition

[ˌdɪspə’zɪʃn]

n.性情;倾向;安排;处置;控制;【计算机】 配置情况

diversify

[daɪ’vɜːsɪfaɪ]

vt.使多样化;使变化;使不同

advantageous

[ˌædvən’teɪdʒəs]

adj.有利的;有助的;有益的

feeble

[’fiːbl]

adj.虚弱的;无力的

assurance

[ə’ʃʊərəns]

n.保证，担保;确信;自信;保险

probability

[ˌprɒbə’bɪləti]

n.可能性

domestication

[dəˌmestɪ’keɪʃn]

n.驯服;教化

authentic

[ɔː’θentɪk]

adj.真正的;真实的;可信的;可靠的

pointer

[’pɔɪntə(r)]

n.指针;教鞭;(能指出猎物位置的)猎犬;暗示;主意

retrieve

[rɪ’triːv]

vt.恢复;挽回;取回

retriever

[rɪ’triːvə(r)]

n.能寻回猎物的犬;取回的人;复得者

prey

[preɪ]

n.牺牲者;被掠食者，猎物

motionless

[’məʊʃnləs]

adj.不动的;静止的

gait

[ɡeɪt]

n.步态;步法

herd

[hɜːd]

n. 【C】(牛、马、猪、象等)群;

invariable

[ɪn’veəriəbl]

adj.不变的;恒定的

mingle

[’mɪŋɡl]

vt.使混合;使结合

hare

[heə(r)]

n.野兔

great-grandfather

[’greɪt’grændˌfɑːðə]

n.曾祖父

compulsory

[kəm’pʌlsəri]

adj.强制的;势在必行的;义务的

tumble

[’tʌmbl]

v.暴跌;倒塌;跌倒;翻滚;弄乱;明白;突然发现

tumbler

[’tʌmblə(r)]

n.不倒翁;杂技演员;平底无脚酒杯;锁的制动栓

suffice

[sə’faɪs]

vi.足够;使满足

scarcely

[’skeəsli]

adv.几乎不;简直不;刚刚;决不

confinement

[kən’faɪnmənt]

n.拘禁;限制;分娩

fowl

[faʊl]

n.禽;禽肉

Familiarity

[fəˌmɪli’ærəti]

n.亲密;熟悉;精通;不拘礼节

jackal

[’dʒækl]

n.胡狼;豺狼;同谋;走狗

genus

[’dʒiːnəs]

n.【生物学】属;类;种;型

puppy

[’pʌpi]

n.小狗;自负的小伙子

savage

[’sævɪdʒ]

a. 野蛮的，未开化的;

concur

[kən’kɜː(r)]

v.意见相同;同时发生

pheasant

[’feznt]

n.雉;野鸡

thicket

[’θɪkɪt]

n.繁茂处;丛林;草丛

ignorance

[’ɪɡnərəns]

n.无知;愚昧

methodical

[mə’θɒdɪkl]

adj.有条不紊的;井然的

naturalist

[’nætʃrəlɪst]

n.自然主义者;博物学者

hatching

[’hætʃɪŋ]

n.阴影;影线

inconvenient

[ˌɪnkən’viːniənt]

adj.不方便的

hatch

[hætʃ]

n.孵化;舱口，舱口盖

predicament

[prɪ’dɪkəmənt]

n.困境

maternal

[mə’tɜːnl]

adj.母亲的;母系的;母方的;母性的

encumber

[ɪn’kʌmbə(r)]

v.妨害;阻害

utterly

[’ʌtəli:]

adv.完全;全然;绝对

offspring

[’ɒfsprɪŋ]

n.后代;子孙;产物

uncommon

[ʌn’kɒmən]

adj.不寻常的;不常见的;极度的

singular

[’sɪŋɡjələ(r)]

a. 非凡的；卓越的;

strew

[struː]

vt.散播;撒满;点缀

hunting

[’hʌntɪŋ]

n.打猎;搜寻【电子学】速度偏差.

pick

[pɪkt]

采摘，挑选；

accordance

[ə’kɔːdns]

n.一致;符合

larva

[’lɑːvə]

n.幼虫;幼体

felonious

[fə’ləʊniəs]

adj.重罪的

exterminate

[ɪk’stɜːmɪneɪt]

vt.消灭;扑灭;根绝

fertile

[’fɜːtaɪl]

adj.肥沃的;富饶的;能繁殖的;多产的;(创造力)丰富的

energetic

[ˌenə’dʒetɪk]

adj.精力旺盛的;有力的;能量的

incapable

[ɪn’keɪpəbl]

adj.无能力的;不胜任的

helpless

[’helpləs]

adj.无助的;无依靠的

pupa

[’pjuːpə]

n.蛹

perish

[’perɪʃ]

vi.毁灭;消失;腐烂

hopeless

[’həʊpləs]

adj.没有希望的;绝望的;无可救药的

agitate

[’ædʒɪteɪt]

v.煽动;搅动;焦虑

energetically

[ˌenə’dʒetɪkli]

adv.精力充沛地;积极地

marches

[mɑːtʃ]

1. n. (尤指英格兰与苏格兰或威尔士的)边界地区, 2.动词march的第三人称单数形式

ascend

[ə’send]

v.上升;攀登

coccus

[’kɒkəs]

n.小干果;球菌

ample

[’æmpl]

adj.丰富的;宽敞的;充足的

Switzerland

[’swɪtsələnd]

n.瑞士

habitual

[hə’bɪtʃuəl]

adj.惯常的;习惯的

expressly

[ɪk’spresli]

adv.清楚地;特意地;专门地

fortunate

[’fɔːtʃənət]

adj.幸运的;侥幸的;带来幸运的

spectacle

[’spektəkl]

n.景象;场面;奇观;壮观;公开展示;表相，假相

haunt

[hɔːnt]

vt. 常去，常到(某地);

repulse

[rɪ’pʌls]

n.击退;拒绝

ruthless

[’ruːθləs]

adj.残忍的;无情的

parcel

[’pɑːsl]

n.包裹;部分;一块(土地)

tyrant

[’taɪrənt]

n.暴君;专制的君主;专横的人

fancy

[’fænsi]

n. 【C】设想；幻想；空想;

ferocious

[fə’rəʊʃəs]

adj.残忍的;凶猛的;极度的

furious

[’fjʊəriəs]

adj.狂怒的;猛烈的

whereas

[ˌweər’æz]

conj.然而;鉴于

terrify

[’terɪfaɪ]

v.使害怕;使恐怖;威胁

desolate

[’desələt]

adj.荒凉的;荒芜的;孤单寂寞的

agitation

[ˌædʒɪ’teɪʃn]

n.激动;鼓动;搅动

perching

[’pɜːtʃɪŋ]

n.栖息;验布;铲软,

chiefly

[’tʃiːfli]

adv.主要地

unintentional

[ˌʌnɪn’tenʃənl]

adj.非故意的;无心的

abject

[’æbdʒekt]

adj.卑贱的;不幸的;可怜的

exquisite

[ɪk’skwɪzɪt]

adj.精挑细选的;精致的;细腻的;强烈的

enthusiastic

[ɪnˌθjuːzi’æstɪk]

adj.热心的;热情的;热烈的

mathematician

[ˌmæθəmə’tɪʃn]

n.数学家

skilful

[’skɪlfl]

adj.熟练的;巧妙的;技术好的.

adjoin

[ə’dʒɔɪn]

v.邻接;毗连

gradation

[ɡrə’deɪʃn]

n.渐变;阶段;等级;刻度;元音交替

irregular

[ɪ’reɡjələ(r)]

adj.不规则的;不整齐的;不合法的;不合规矩的

hexagonal

[heks’æɡənl]

adj.六角形的;六边的;底面为六角的

rhomb

[rɒm]

n.偏菱形(斜方形;斜方六面体)

pyramidal

[’pɪrəmɪdl]

adj.金字塔形的;角锥状的;锥体的

cylindrical

[sə’lɪndrɪkl]

adj.圆柱形的

spherical

[’sferɪkl]

adj.球的;球面的;球状的

aggregate

[’æɡrɪɡət]

adj.合计的;总的

intersect

[ˌɪntə’sekt]

v.贯穿;(和 ... )相交;交叉

manifest

[’mænɪfest]

vt.显示;证实;表露

imitation

[ˌɪmɪ’teɪʃn]

n.模仿;效法

thickness

[’θɪknəs]

n.厚度;浓度;层

Miller

[’mɪlə(r)]

n.磨坊主;碾磨工;【机】铣床;【昆】蛾

kindly

[’kaɪndli]

adj.和蔼的;温和的;爽快的

parallel

[’pærəlel]

adj.平行的

radius

[’reɪdiəs]

n.半径范围

lesser

[’lesə(r)]

adj.较少的;较小的;次要的

intersection

[ˌɪntə’sekʃn]

n.交集;十字路口;交叉点

burrow

[’bʌrəʊ]

n.(兔子等)所掘的地洞

prolong

[prə’lɒŋ]

vt.延长;拖延

hexagon

[’heksəɡən]

adj.六角形的

rude

[ruːd]

adj.粗鲁无礼的;原始的;未加工的;粗糙的;猛烈的

circular

[’sɜːkjələ(r)]

adj.圆形的;循环的

architectural

[ɑːkɪˈtekʧərəl]

【修】(体系)结构

deepen

[’diːpən]

vt. 使 ... 加深; 使 ... 强烈

basin

[’beɪsn]

n.脸盆;水池

diameter

[daɪ’æmɪtə(r)]

n.直径

width

[wɪdθ]

n.宽度;广度

festoon

[fe’stuːn]

v.以花彩装饰

excavate

[’ekskəveɪt]

v.挖掘;挖开;发掘(古物)

imaginary

[ɪ’mædʒɪnəri]

adj.想象的;虚构的

unnatural

[ʌn’nætʃrəl]

adj.不自然的;反常的;不近人情的

gnaw

[nɔː]

v.咬;折磨;侵蚀

convex

[’kɒn’veks]

adj.中凸的;凸圆的;凸面的

flatten

[’flætn]

vt.使变平;打倒

circumferential

[səˌkʌmfə’renʃəl]

adj.圆周的

commencement

[kə’mensmənt]

n.开始;毕业典礼

excavation

[ˌekskə’veɪʃn]

n.挖掘;发掘;挖掘的洞;发掘物

specimen

[’spesɪmən]

n.标本;样本

flexure

[’flekʃə]

n.屈曲;弯曲部分;褶缝

mason

[’meɪsn]

n.石匠;泥瓦匠

cement

[sɪ’ment]

n.水泥

grind

[ɡraɪnd]

v.磨;压迫;碾碎;磨得吱吱响;逐渐停顿

crown

[kraʊn]

n.王冠;王权;花冠

utmost

[’ʌtməʊst]

adj.极度的;最大限度的

invariably

[ɪn’veəriəbli]

adv.不变地;总是;一贯地

diffuse

[dɪ’fjuːz]

vt.扩散

atom

[’ætəm]

n.原子

recur

[rɪ’kɜː(r)]

vi.重现;再发生

alternate

[’ɔːltɜːnət]

v.交替;轮流

succession

[sək’seʃn]

n.连续;继承权;继位

progenitor

[prəʊ’dʒenɪtə(r)]

n.祖先;先驱

secretion

[sɪ’kriːʃn]

n.分泌;分泌物;藏匿

parasite

[’pærəsaɪt]

n.寄生虫;寄生生物;食客

swarm

[swɔːm]

n.群;蜂群

allude

[ə’luːd]

v.暗示;暗指

neuter

[’njuːtə(r)]

adj.中性的;不及物的;生殖器不完全的

exemplify

[ɪɡ’zemplɪfaɪ]

vt.例证;示范

correlate

[’kɒrəleɪt]

vt.使相互关联

reproductive

[ˌriːprə’dʌktɪv]

adj.复制的;生殖的

imperfect

[ɪm’pɜːfɪkt]

adj.不完美的;有缺点的;未完善的

breeder

[’briːdə(r)]

n.饲养动物的人;培育植物的人;【核】增殖(反应)堆

slaughter

[’slɔːtə(r)]

n.残杀;屠杀;大量杀戮

propagate

[’prɒpəɡeɪt]

v.繁殖;增殖;传播;传送

climax

[’klaɪmæks]

n.高潮;极点;层进法;【生】顶极群落;巅峰，最高峰

caste

[kɑːst]

n.印度的世袭阶级;(排他的)社会团体;种姓;等级制度

shield

[ʃiːld]

n.盾

secrete

[sɪ’kriːt]

vt.隐藏;隐匿

ocellus

[əʊ’seləs]

n.【昆】单眼;眼点

proportional

[prə’pɔːʃənl]

adj.成比例的;相称的

rudiment

[’ruːdɪmənt]

n. 【复】初步，基础，入门，基本原理;

avail

[ə’veɪl]

v.有利;有助于

Africa

[’æfrɪkə]

n.非洲

dissect

[dɪ’sekt]

v.解剖;切细;仔细研究;详细分析

simultaneous

[ˌsɪml’teɪniəs]

adj.同时发生的;同步的

labour

[ˈleɪbə]

n. 劳动;劳动力

manufacture

[ˌmænju’fæktʃə(r)]

v.(手工)制造

descendant

[dɪ’sendənt]

n.后裔;子孙;后代

demonstrative

[dɪ’mɒnstrətɪv]

adj.易流露感情的;论证的;【语法】指示的

liable

[’laɪəbl]

a. 有(法律)责任的；有义务的;

applicable

[ə’plɪkəbl]

adj.合适的

deduction

[dɪ’dʌkʃn]

n.扣除;扣除额;推论;推理;演绎

endow

[ɪn’daʊ]

vt.捐助;赋予;使具有某种品质

multiply

[’mʌltɪplaɪ]

vt.乘;增加

thrush

[θrʌʃ]

n.画眉;【动】鸫科;【医】鹅口疮

roost

[ruːst]

n.栖木;鸟窝;栖息处

cuckoo

[’kʊkuː]

n.杜鹃;杜鹃的叫声

eject

[i’dʒekt]

v.喷射;放逐;驱逐

larva

[’lɑːvə]

n.幼虫;幼体

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