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雙語《物種起源》 第七章 本能

所屬教程:譯林版·物種起源

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2022年06月28日

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CHAPTER VII INSTINCT

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

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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:—

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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?

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.

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.

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.

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.

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.

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.

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.

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.

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.

第七章 本能

本能與習(xí)性可比,但起源不同——本能的分級——蚜蟲和蟻——本能是變異的——家養(yǎng)的本能,其起源——杜鵑、鴕鳥以及寄生蜂的自然本能——蓄奴蟻——蜜蜂,營造蜂房的本能——自然選擇學(xué)說應(yīng)用于本能的難點(diǎn)——中性或不育的昆蟲——提要

本能問題原本可以納入前面的章節(jié)。但我想,單獨(dú)討論比較方便,尤其是蜜蜂筑巢的本能是如此奇妙,在許多讀者看來大概是一個足以推翻我的全部學(xué)說的難點(diǎn)。我先要聲明一點(diǎn),就是我不準(zhǔn)備討論智力的起源,就如我未曾討論生命本身的起源一樣。我們所要討論的,只是同綱動物中本能的多樣性,以及其他精神品質(zhì)的多樣性。

我并不想給本能下任何定義。容易闡明,這一術(shù)語通常包含著若干不同的精神活動;但是,說本能促使杜鵑遷徙并把蛋下在別種鳥巢里,每一個人都知道這是什么意義。我們自己需要經(jīng)驗(yàn)才能完成的一種活動,而被一種沒有經(jīng)驗(yàn)的動物,特別是被幼小動物所完成,并且許多個體并不知道為了什么目的卻按照同一方式去完成時,一般就稱為本能。但是我能闡明,這些性狀沒有一個是普遍的。如于貝爾(Pierre Huber)所說的,甚至在自然系統(tǒng)中低級的動物里,小量的判斷或理性也常發(fā)生作用。

弗·居維葉(Frederick Cuvier)等老一輩玄學(xué)者曾把本能與習(xí)性加以比較。我想,這一比較對于完成本能活動時的心理狀態(tài),提供了精確的概念,但不一定涉及它的起源。許多習(xí)慣性活動是在無意識下進(jìn)行的,甚至不少直接與我們有意識的意志相反!然而意志和理性可以改變它們。習(xí)性容易與其他習(xí)性,與一定的時期,與身體的狀態(tài)相聯(lián)系。習(xí)性一經(jīng)獲得,常常終生保持不變。還可以指出本能和習(xí)性之間的其他若干類似點(diǎn)。有如反復(fù)歌唱一首名曲,在本能里也是一種活動有節(jié)奏地隨著另一活動;如果一個人歌唱時,或在反復(fù)背誦東西時被打斷了,一般地他就被迫走回頭路,恢復(fù)已經(jīng)成為習(xí)慣的思路。于貝爾發(fā)現(xiàn)能制造很復(fù)雜繭床的毛毛蟲(caterpillar)就是如此;如果在完成構(gòu)造第六階段時把它抓出,放在只完成構(gòu)造第三階段的繭床里,這個毛毛蟲僅重筑第四、第五、第六個階段的構(gòu)造。然而,如果把完成構(gòu)造第三階段的毛毛蟲,放在已完成構(gòu)造第六階段的繭床里,那么工作已大都完成了,可是它并沒有從中感到受益,反而不知所措,并且為了完成繭床,它似乎不得不從構(gòu)造第三階段開始,它是在這里放下的,就這樣試圖去完成已經(jīng)完成了的工作。

假定任何習(xí)慣性的活動被遺傳——可以指出,有時確有這種情形發(fā)生——那么原為習(xí)性和原為本能之間,就變得密切相似,難分難解。如果莫扎特不是在三歲時經(jīng)過極少的練習(xí)就能彈奏鋼琴,而是全然沒有練習(xí)就能彈奏一曲,那么可以說他的彈奏確實(shí)是出于本能的了。但是假定大多數(shù)本能是由一個世代中的習(xí)性得來的,然后遺傳給后繼世代,則是大錯特錯。能夠清楚表明,我們所熟知的最奇異的本能,如蜜蜂和許多蟻的本能,不可能是由習(xí)性得來的。

人們普遍承認(rèn),本能對于各個物種在現(xiàn)今生活條件之下的利益,有如肉體構(gòu)造一樣重要。在多變的生活條件下,本能的微小變異有利于物種,至少是可能的;如果能夠指出,本能確曾發(fā)生過些許變異,我看自然選擇就不難把本能的變異保存下來并繼續(xù)累積到任何有利的程度。我相信,一切最復(fù)雜的和奇異的本能就是這樣起源的。使用或習(xí)性引起肉體構(gòu)造的變異,并使之增強(qiáng),而不使用使之縮小或消失,我并不懷疑本能也是這樣的。但我認(rèn)為,習(xí)性的效果,同所謂本能偶發(fā)變異的自然選擇效果相比是次要的。也就是,產(chǎn)生身體構(gòu)造的微小偏差有一些未知原因,也產(chǎn)生變異,叫本能偶發(fā)變異。

除非經(jīng)過有益的變異積少成多,緩慢而逐漸的積累,否則復(fù)雜的本能不可能通過自然選擇而產(chǎn)生。因此,像身體構(gòu)造的情形一樣,我們在自然界中所尋求的不應(yīng)是獲得每一復(fù)雜本能的實(shí)際過渡諸級,這些只有在各物種的直系祖先里才能找到,但應(yīng)當(dāng)從旁系世系里去尋求這些分級的蛛絲馬跡,至少能夠指出某種分級是可能的;而這一點(diǎn)肯定能夠辦到。考慮到除了歐洲北美洲以外,動物本能還極少被觀察過,并且滅絕物種的本能更是一無所知,我感到驚異的是,最復(fù)雜本能所賴以完成的分級能夠廣泛被發(fā)現(xiàn)?!白匀唤缋餂]有飛躍”的準(zhǔn)則適用于本能的力度不亞于身體器官。同一物種在生命的不同時期或一年中的不同季節(jié),或處于不同的環(huán)境條件下等等而具有不同的本能,這就往往會促進(jìn)本能的變化;在這種情況下,自然選擇會把這種或那種本能保存下來。可以證明,同一物種中本能的多樣性在自然界中也是存在的。

還有,像身體構(gòu)造那樣,各物種的本能都是為了自己的利益,據(jù)我們所能判斷的,它從來沒有為了其他物種的獨(dú)享利益而產(chǎn)生過,這和我的理論也是符合的。我知道一個極有力的事例,一種動物的活動從表面看來完全是為了別種動物的利益,這就是蚜蟲自愿把甜的分泌物供給螞蟻:這樣做是出于自愿,可由下列事實(shí)來說明。我把羊蹄酸模(dock-plant)上的所有螞蟻從十來只蚜蟲堆里搬走,數(shù)小時內(nèi)不讓回來。過了這段時間,我確實(shí)覺得蚜蟲要進(jìn)行分泌了。我用放大鏡觀察了一些時候,沒有一個分泌的,于是,我盡力模仿螞蟻用觸角的樣子,用一根毛輕輕地觸動撫摩,但還沒有一只蚜蟲分泌。隨后我讓一只螞蟻去接近它們,從它熱切跑動的樣子看來,它好像立刻覺得發(fā)現(xiàn)了豐盛的食物;于是用觸角去撥蚜蟲的腹部,先是這一只,然后那一只;蚜蟲一接觸到觸角,即刻舉起腹部,分泌出一滴澄清的甜液,螞蟻慌忙吞食了。甚至十分幼小的蚜蟲也有這樣的動作,可見這種活動是本能,而不是經(jīng)驗(yàn)所致。但是,排泄物極黏,被取去也許對于蚜蟲是便利的,所以分泌本能大概不是專為螞蟻的利益。雖然我不相信世界上任何動物會為了其他物種的獨(dú)享利益而從事活動,然而各物種卻試圖利用其他物種的本能,正像利用其他物種的虛弱身體構(gòu)造一樣。這樣,某些本能就不能看作是絕對完善的;但是詳細(xì)討論這一點(diǎn)以及其他類似之點(diǎn),并非必不可少,所以這里就不贅述了。

本能在自然狀態(tài)下有某種程度的變異以及這些變異的遺傳,既然是自然選擇的作用所不可少的,那就應(yīng)該盡量舉出許多事例來,但是篇幅限制,無法這樣做。我只能斷言,本能確實(shí)是變異的——例如遷徙的本能,不但在范圍和方向上能變異,而且也會完全消失。鳥巢也是如此,變異部分發(fā)生于對選定的位置以及居住地性質(zhì)和氣溫的依賴度,但常常由于全然未知的原因而發(fā)生變異。奧杜邦曾舉出幾個顯著的例子,說明美國北方和南方同一物種的鳥巢有所不同。對于敵害的恐懼必然是一種本能品質(zhì),從未離巢的雛鳥身上可以看到,但這種恐懼可由經(jīng)驗(yàn)或因看見其他動物對于同一敵害的恐懼而強(qiáng)化。對于人類的恐懼,如我在他處指出的,棲息在荒島上的各種動物是慢慢獲得的。甚至英國也可以看到這樣的事例,即一切大型鳥比小型鳥更怕人,因?yàn)楦嗟卦馐苓^人的迫害。英國的大型鳥更怕人,可以穩(wěn)妥地歸于這個原因;在無人島,大型鳥并不比小型鳥更怕人;喜鵲(magpie)在英國很警惕,但在挪威卻很馴順,埃及的羽冠烏鴉(hooded crow)也是不怕人的。

有許多事實(shí)可以證明,自然狀態(tài)下產(chǎn)生的同類動物的脾氣極多樣化。還有若干個案可以舉出,表明某些物種偶發(fā)的奇特習(xí)性若對這個物種有利,就會通過自然選擇產(chǎn)生新的本能。但我十分清楚,泛泛而談,沒有詳細(xì)的事實(shí),在讀者的心目中只會當(dāng)耳旁風(fēng)。我只好重復(fù)保證,不說沒有可靠證據(jù)的話。

簡略考察一下家養(yǎng)的若干例子,則自然狀態(tài)下本能遺傳變異的可能性將加強(qiáng),甚至出現(xiàn)大的可能性。由此可見習(xí)性和所謂偶發(fā)變異的選擇,在改變家養(yǎng)動物精神品質(zhì)上分別發(fā)生的作用。有許多奇異而真實(shí)的例子可以說明,與某種心境或某一時期有關(guān)的各種不同脾氣嗜好以及怪癖都是遺傳的。但是讓我們看看眾所熟知的幾種狗的例子。毫無疑問,幼小的指示犬第一次帶出去,有時就能夠指示獵物的所在,甚至能夠援助別的狗(我曾親見這動人的情形);尋回犬(retriever)確實(shí)在某種程度上可以把尋回的特性遺傳下去;牧羊犬并不跑在羊群之內(nèi),而有在羊群周圍環(huán)跑的傾向。幼小動物不依靠經(jīng)驗(yàn)而有這些活動,同時各個體又差不多以同一方式進(jìn)行,并且都欣然且不知道目的地去進(jìn)行——幼小的指示犬并不知道指示方向是在幫助主人,有如白蝴蝶并不知道為什么要在圓白菜葉子上產(chǎn)卵一樣——我無法看出這些活動在本質(zhì)上與真正的本能有什么區(qū)別。如果看見一種狼,在幼小而且沒有受過任何訓(xùn)練時,一旦嗅出獵物,先站著一動不動,隨后又用特別的步法慢慢爬過去;又看見另一種狼環(huán)繞鹿群追逐,卻不直沖,以便把鹿趕到遠(yuǎn)的地點(diǎn)去,必然要把這活動叫作本能。所謂家養(yǎng)下的本能,的確遠(yuǎn)不及自然的本能那么固定;但是其所蒙受的選擇作用也極不嚴(yán)格,而且是在較不固定的生活條件下,在無比短暫的時間內(nèi)傳遞下來的。

不同品種的狗進(jìn)行雜交時,即能很好地看出這家養(yǎng)下的本能、習(xí)性以及脾氣的遺傳性是何等強(qiáng)烈,并且混合得多么奇妙。例如眾所周知,長驅(qū)跑狗與斗牛狗雜交,可影響勇敢性和頑強(qiáng)性至許多世代;牧羊犬與長驅(qū)跑狗雜交,則使全族都得到捕捉野兔的傾向。這家養(yǎng)下的本能,如用上述雜交方法來試驗(yàn),是與自然的本能相類似的;自然的本能也按照同樣的方式奇異地混合在一起,而且長期表現(xiàn)出其祖代任何一方的本能的痕跡。例如,勒魯瓦(Le Roy)描述過一條狗,曾祖父是狼;它只有一點(diǎn)表示了野生祖先的痕跡,即呼喚它時,不是直線走向主人。

家養(yǎng)下的本能有時被說成為完全由長期的強(qiáng)迫養(yǎng)成的習(xí)性所遺傳下來的動作,我看這是不正確的。從未有人會想象去教或者曾經(jīng)教過翻飛鴿去翻飛——據(jù)我所見到的,幼鴿從不曾見過鴿的翻飛,卻會翻飛。我們相信,曾經(jīng)有過一只鴿子表現(xiàn)了這種奇怪習(xí)性的微小傾向,并且在連續(xù)的世代中,經(jīng)過對最好的個體的長期選擇,才造成像今日那樣的翻飛鴿;據(jù)布倫特(Brent)先生說,格拉斯哥附近的家養(yǎng)翻飛鴿,一飛到十八英寸高就要翻筋斗。假如未曾有過一只狗自然具有指示方向的傾向,是否會有人想到訓(xùn)練狗去指示方向是存疑的;人們知道這種傾向有時會出現(xiàn),我就看見過一次,見于純種的里。指示方向的最初傾向一旦出現(xiàn),此后在每一世代中有計劃選擇和強(qiáng)迫訓(xùn)練的遺傳效果,將會很快大功告成;而且無意識選擇至今仍在繼續(xù)進(jìn)行,每一個人雖然本意不在改進(jìn)品種,但總試圖獲得最善于指示方向和狩獵的狗。另一方面,在某些情形下,僅僅習(xí)性一項(xiàng)已經(jīng)足夠了;沒有動物比小野兔更難以馴服的了,也幾乎沒有動物比小家兔更馴順的了。但我很難設(shè)想家兔僅僅為了馴服性才被選擇下來;所以從極野的到極馴服的性質(zhì)的遺傳變化,必須全部歸因于習(xí)性和長久持續(xù)的嚴(yán)格圈養(yǎng)。

自然的本能在家養(yǎng)狀況下可以消失:最顯著的例子見于少孵蛋或不喜孵蛋的雞品種。僅僅由于司空見慣,我們才看不出家養(yǎng)動物的心理曾經(jīng)有過多么普遍的變化。對于人類的親愛已經(jīng)成了狗的本能,這是毋庸置疑的。一切狼、狐、胡狼(jackal)以及貓屬的物種即使馴養(yǎng)后,也極渴望攻擊雞、羊和豬;火地和澳洲這些地方的未開化人不養(yǎng)狗,因?yàn)樵l(fā)現(xiàn)這種傾向在家里養(yǎng)的小狗身上是不能矯正的。另一方面,已經(jīng)文明化了的狗,甚至在十分幼小的時候,也很少必要去教其不要攻擊雞、羊和豬!無疑會偶爾攻擊一下子,于是就要遭一頓打;如果還不能矯正,就會被弄死;那個習(xí)性,通過某種程度的選擇,也許協(xié)同地靠遺傳使家狗文明化了。另一方面,小雞完全出于習(xí)性,已經(jīng)消失了原本無疑懼怕貓狗的本能;而小雉雞盡管是由母雞撫養(yǎng)的,卻是明顯具有這種本能的。倒不是小雞失去了一切懼怕,而只是失去了對于貓狗的懼怕,因?yàn)椋鸽u發(fā)出報告危險的叫聲,小雞便從母雞的翼下跑開(小火雞尤其如此),躲到四周的草叢里或林子里去了。這顯然是本能的動作,便于母鳥飛走,就如我們在野生的陸棲鳥類里所看到的那樣。但是小雞還保留著這種本能,在家養(yǎng)狀況下已經(jīng)沒有用處,因?yàn)槟鸽u由于不使用的緣故,已經(jīng)幾乎失掉了飛翔能力。

因此,可以斷定,動物在家養(yǎng)下可以獲得新的本能;而失去自然的本能,這一部分是由于習(xí)性,一部分是由于人類在連續(xù)世代中選擇和累積了特殊的精神習(xí)性和精神活動,而它們的最初發(fā)生,我們無知地看作是出于偶然的原因。在某些情形下,只是強(qiáng)制的習(xí)性一項(xiàng),已足以產(chǎn)生這種遺傳的心理變化;在另外一些情形下,強(qiáng)制的習(xí)性就不能發(fā)生作用,一切都是有計劃選擇和無意識選擇的結(jié)果。但是在大多數(shù)情形下,習(xí)性和選擇大概是雙管齊下的。

我們只要考察少數(shù)事例,大概就能很好地理解本能在自然狀態(tài)下如何由于選擇作用而改變的。我只選擇三個例子,其余的可能要以后著書討論了——即,杜鵑在別種鳥巢里下蛋的本能,某些螞蟻蓄奴的本能,以及蜜蜂造蜂房的本能。學(xué)者們已經(jīng)把后兩種本能,一般地而且恰當(dāng)?shù)亓袨橐磺幸阎灸苤凶钇娈惖牧恕?/p>

現(xiàn)在已經(jīng)有共識,杜鵑的這種本能的比較直接的決定性原因,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,則最先下的蛋便須擱置一些時間后才能得到孵抱,同一個巢里就會有不同齡期的蛋和小鳥了。這樣,下蛋和孵蛋的過程就會漫長而不方便,特別是雌鳥在很早就要遷徙,而最初孵化的小鳥勢必要由雄鳥單獨(dú)哺養(yǎng)。但是美洲杜鵑就處于這樣的困境;她自己筑巢,而且要在同一時期內(nèi)產(chǎn)蛋和照顧相繼孵化的幼鳥。有人說美洲杜鵑有時也在別種鳥巢里下蛋,但我從權(quán)威的布留爾(Brewer)博士那里聽到,這不對。不過,我可以舉出各種鳥類偶爾在別種鳥巢里下蛋的若干事例?,F(xiàn)在假定歐洲杜鵑的古代祖先也有美洲杜鵑的習(xí)性,但偶爾也在別種鳥巢里下蛋。如果這種偶爾的習(xí)性有利于老鳥,如果小鳥由于利用了其他物種的誤養(yǎng)本能,比起母鳥哺養(yǎng)更為強(qiáng)壯——母鳥必須同時照顧不同齡期的蛋和小鳥,不免受到拖累——那么老鳥或寄養(yǎng)的小鳥都會受益。以此類推,我可以相信,這樣哺養(yǎng)起來的小鳥大概就會遺傳母鳥那種偶然的奇特習(xí)性,傾向于把蛋下在別種的鳥巢里,這樣就能夠成功哺養(yǎng)幼鳥。我相信杜鵑的奇異本能會由這種連續(xù)過程而產(chǎn)生出來。補(bǔ)充一句,格雷博士等人說,歐洲杜鵑并未完全失去母愛和對后代的關(guān)懷。

鳥類偶爾會把蛋下在同種別種的鳥巢里這種習(xí)性,在雞科里并非不普通,并且可以解釋近緣鴕鳥群的奇特本能的來源。至少是美國種的個案,若干母鴕鳥共同先在一個巢里,然后在另一個巢里下一些蛋,由雄鳥去孵。這種本能或可以解釋為雌鳥下蛋很多,但如杜鵑一樣,隔二差三才下一次。然而美洲鴕鳥的這種本能,還沒有達(dá)到完善;因?yàn)橛卸嗟贸銎娴牡岸忌⒙湓诘厣?,我在一天的游獵中,就拾得了不下二十個丟棄的蛋。

許多蜂是寄生的,總是把卵產(chǎn)在別種的蜂巢里,這個個案比杜鵑更令人矚目;這種蜂隨著寄生習(xí)性,不但改變了本能,而且改變了構(gòu)造;它們不具有采集花粉的器具,如果要為幼蜂貯藏食料,這是必不可少的。泥蜂科(Sphecidae;形似胡蜂)的某些物種同樣也是寄生的;法布爾最近提出充分的理由認(rèn)為:一種小唇沙蜂(Tachytes nigra)雖然通常都自己造巢,而且為幼蟲儲蓄麻痹了的食物,但發(fā)現(xiàn)別種泥蜂所造儲蓄有食物的巢,便會加以利用,而變成臨時的寄生者。這種情形和杜鵑的假設(shè)情形是一樣的,我覺得如果一種臨時的習(xí)性對于物種有利,同時被害的蜂類不會因巢和儲蓄的食物被無情奪取而遭到滅絕,自然選擇就不難把它永久化。

蓄奴的本能?!@種奇妙的本能,是由于貝爾最初在紅褐蟻(Formica[Polyerges]rufescens)身上發(fā)現(xiàn)的,他是一位比他著名的父親更為優(yōu)秀的觀察者。這種螞蟻絕對依賴奴隸而生活;沒有奴隸的幫助,這個物種一年之內(nèi)一定滅絕。雄蟻和能育的雌蟻不從事任何工作,工蟻即不育的雌蟻雖然在捕捉奴隸上極為賣力勇敢,但不做其他任何工作。它們不能營造自己的巢,也不能哺喂自己的幼蟲。在老巢已不適用,勢必遷徙的時候,是由奴蟻來決定遷徙的事情,并且實(shí)際上把主人們銜在顎間搬走。主人們十分的不中用,當(dāng)于貝爾捉了三十個關(guān)起來而沒有一個奴蟻時,雖然那里放著它們最喜愛的豐富食物,而且有自己的幼蟲和蛹刺激它們工作,它們還是無所事事;它們甚至不會自己吃東西,許多螞蟻就此餓斃。于貝爾隨后放進(jìn)一個奴蟻——黑蟻(F. fusca),她即刻開始工作,喂哺和拯救那些生存者;并且營造了幾間蟲房,來照料幼蟲,一切都整得井井有條。有什么比這十分肯定的事實(shí)更為奇異的呢?如果我們不知道任何其他蓄奴的蟻類,大概就無法想象如此奇異的本能是怎樣完善的。

血蟻(Formica sanguinea)同樣蓄奴,也是于貝爾最初發(fā)現(xiàn)的。這個物種見于英格蘭南部,大英博物館史密斯(F. Smith)先生研究過它的習(xí)性。關(guān)于這個問題以及其他問題,我深深感激他的幫助。雖然我充分相信于貝爾和史密斯先生的敘述,但仍然以懷疑的心情來處理這個問題,任何人對于蓄奴的這種異常丑惡本能的存在有所懷疑,大概都得諒解。因此,我愿意稍微詳細(xì)地談?wù)勎业挠^察。我曾掘開十四個血蟻巢,都發(fā)現(xiàn)了若干奴蟻。奴種的雄蟻和能育的雌蟻,只見于它們自己固有的群中,在血蟻巢中從未看見過。黑色奴蟻,不及紅色主人的一半大,外貌上對比強(qiáng)烈。蟻巢被微微擾動時,奴蟻偶爾跑出外邊來,像主人一樣十分激動,并且保家衛(wèi)國;當(dāng)蟻巢被擾動得很厲害,幼蟲和蛹暴露出來的時候,奴蟻和主人一齊奮發(fā)地把它們運(yùn)送到安全的地方。因此,奴蟻顯然是熟門熟路。在連續(xù)三個年頭的六月和七月里,我在薩里郡和薩塞克斯郡,曾對幾個蟻巢觀察了幾個小時,從來沒有看到一個奴蟻?zhàn)韵伋怖镒叱鲎哌M(jìn)。在這些月份里,奴蟻的數(shù)目很少,因此我想數(shù)目多的時候,行動大概就不同了;但史密斯先生告訴我說,五月、六月以及八月間,在薩里和漢普郡,他在各種不同的時間內(nèi)注意觀察了蟻巢,雖然八月份奴蟻的數(shù)目很多,但也不曾看到它們走出走進(jìn)蟻巢。因此,他認(rèn)為它們是嚴(yán)格的家奴。而主人卻不然,經(jīng)??吹剿鼈儾粩嗟匕徇\(yùn)著造巢材料和各種食物。然而今年七月里,我遇見一個奴蟻特別多的蟻群,觀察到有少數(shù)奴蟻和主人混在一起離巢,沿著同一條路向著約二十五碼遠(yuǎn)的一株高大歐洲赤松前進(jìn),它們一齊爬到樹上去,大概是為了找尋蚜蟲或胭脂蟲的。于貝爾有過許多觀察的機(jī)會,他說,瑞士的奴蟻在造蟻巢的時候常常和主人一起干,而在早晚間則單獨(dú)看管門戶。于貝爾還明確地說,奴蟻的主要職務(wù)是搜尋蚜蟲。兩個國家里的主奴兩蟻的普通習(xí)性如此不同,大概僅僅由于在瑞士被捕捉的奴蟻數(shù)目比英格蘭多。

有一天,我有幸看到了血蟻搬巢,于貝爾描述過,主人們謹(jǐn)慎地把奴蟻帶在顎間,這真是極有趣的奇觀。另一天,大約有二十只蓄奴蟻在同一地點(diǎn)獵取東西,而顯然不是找尋食物,這引起了我的注意。它們走近一種奴蟻——獨(dú)立的黑蟻群,并且遭到猛烈的反擊。有時候三個奴蟻揪住蓄奴血蟻的腿不放,蓄奴蟻殘忍地弄死了小抵抗者,并且把尸體拖到二十九碼遠(yuǎn)的巢中去當(dāng)食物,但得不到一個蛹來培養(yǎng)為奴。于是我從另一個巢里掘出一小團(tuán)黑蟻的蛹,放在鄰近戰(zhàn)場的一處空地上。于是這班暴君熱切地把它們捉住并且拖走,大概以為畢竟是在最后的戰(zhàn)役中獲勝了。

同時,我在同一場所放下另一物種——黃蟻(F. flava)的一小團(tuán)蛹,其上還有幾只小黃蟻攀附在蟻巢破片上。如史密斯先生所描述的,這個物種有時會淪落為奴,但很少見。這種蟻雖然這么小,但極勇敢,我看到過它們兇猛地攻擊別種蟻。有一個事例,我驚奇地看見蓄奴血蟻巢下有一塊石頭,底下是一個獨(dú)立的黃蟻群;我偶然地擾動這兩個巢,小螞蟻就以驚人的勇氣去攻擊它們的大鄰居。當(dāng)時我渴望確定血蟻能否辨別常捉做奴隸的黑蟻的蛹與很少捉拿的小型而兇猛的黃蟻的蛹,明顯地它們確能立刻辨別;因?yàn)槲覀兛匆娪龅胶谙伒挠紩r,它們即刻熱切地去捉,而遇到黃蟻的蛹,甚至遇到其巢的泥土?xí)r,便驚慌失措,趕緊跑開;但是,大約經(jīng)過一刻鐘,當(dāng)這種小黃蟻都爬走之后,它們才鼓起勇氣,把蛹搬走。

一天傍晚,我看見另一群血蟻,發(fā)現(xiàn)若干這種蟻拖著黑蟻的尸體(可以看出不是遷徙)和無數(shù)的蛹回巢。我跟著背著戰(zhàn)利品魚貫而行的蟻?zhàn)粉櫱叭?,大約有四十碼之遠(yuǎn),到了一處密集的石南科灌木叢(heath),我看到最后一個血蟻出現(xiàn),拖著一個蛹,但無法在密叢中找到被蹂躪的蟻巢。然而那巢一定就在附近,因?yàn)橛袃扇缓谙仒O度張皇地沖出來,有一只嘴里還銜著自己的蛹一動不動地停留在石南的小枝頂上,在破碎的家上方。

這些都是關(guān)于蓄奴的奇異本能的事實(shí),無須我來證實(shí)。讓我們看一看,血蟻的本能習(xí)性和歐洲大陸上的紅褐蟻的鮮明對照。后一種不會筑巢,不會決定自己的遷徙,不會為自己和幼蟻采集食物,甚至不會自己吃東西:完全依賴無數(shù)的奴蟻。血蟻則不然,擁有很少的奴蟻,初夏時奴蟻是極少的。主人決定在何時何地營造新蟻巢,并且遷徙的時候,還銜著奴蟻?zhàn)?。瑞士和英格蘭的奴蟻似乎都專門照顧幼蟻,主人單獨(dú)做捕捉奴蟻的遠(yuǎn)征。瑞士的奴蟻和主人一齊工作,搬運(yùn)材料回去造巢;主奴共同地,但主要是奴蟻在照顧它們的蚜蟲,并進(jìn)行所謂的擠乳;這樣,主奴都為本群采集食物。在英格蘭,通常是主人單獨(dú)出去搜尋筑巢材料,為它們自己、奴蟻和幼蟻搜尋食物。所以,我國奴蟻為主人所服的勞役,比在瑞士少得多。

血蟻的本能靠什么步驟發(fā)生,我不愿妄加臆測。但是,據(jù)我所看到的,不蓄奴的蟻如果有其他物種的蛹散落在蟻巢近旁,也要把這些蛹拖進(jìn)去,所以這些本來貯作食物的蛹可能發(fā)育起來;這樣無意識地被養(yǎng)育起來的外來蟻將會追隨自己的固有本能,做它們所能做的工作。如果它們的存在證明對于捕獲它們的物種有用——如果捕捉工蟻比自己生育工蟻對于這個物種更有利——那么,本是采集蟻蛹供食用的這種習(xí)性,大概會因自然選擇而加強(qiáng),并且變?yōu)橛谰玫模赃_(dá)到非常不同的蓄奴目的。本能一旦獲得,即使它的應(yīng)用范圍遠(yuǎn)不及英國的血蟻(如我們所看到的,這種蟻在依賴奴蟻的幫助上比瑞士的同一物種為少),我看自然選擇也不難增強(qiáng)和改變這種本能——始終假定每一個變異對于物種都有用處——直到形成一種像紅褐蟻那樣卑鄙地依靠奴隸來生活的蟻類。

蜜蜂營造蜂房的本能。——這個問題不擬詳加討論,而只是把我所得到的結(jié)論的綱要說一說。凡是考察過蜂巢的精巧構(gòu)造的人,看到如此美妙地適應(yīng)它的目的而不熱烈贊賞,必定是愚鈍不堪。聽到數(shù)學(xué)家說蜜蜂已實(shí)際解決了深奧的問題,把蜂房造成適當(dāng)?shù)男螤睿瑏砣菁{最大可能容量的蜜,而在建造中則用最小限度的貴重蠟質(zhì)。有人說,一個熟練的工人,用合適的工具和度量衡,也很難造出正形的蠟質(zhì)蜂房來,但是一群蜜蜂卻能在黑暗的蜂箱內(nèi)把它造成。隨便你說這是什么本能都可以,乍一看似乎是不可思議的,如何能造出所有必要的角和面,甚至如何能覺察出做工正確。但是這難點(diǎn)并不像最初看來那樣大;我想可以證明,這一切美妙的工作都是來自幾種簡單的本能。

我研究這個問題,是受沃特豪斯先生的引導(dǎo)。他闡明,蜂房的形狀和鄰接蜂房的存在有密切關(guān)系;下述觀點(diǎn)大概只能看作是他的理論的修正。讓我們看看偉大的分級原理,看看自然是否向我們揭示了其工作方法。這個簡短系列的一端有大黃蜂,用舊繭來貯蜜,有時候在繭殼上添加蠟質(zhì)短管,而且同樣也會做出分隔的、很不規(guī)則的圓形蠟質(zhì)蜂房。這系列的另一端則有蜜蜂的蜂房,排列為雙層:每一個蜂房,眾所周知,都是六面柱體,六邊的底邊傾斜地聯(lián)合成三個菱形所組成的倒角錐體。菱形都有一定的角度,并且在蜂巢的一面,一個蜂房的角錐形底部的三條邊,正好構(gòu)成了反面的三個連接蜂房的底部。這一系列里,處于極完美的蜜蜂蜂房和簡單的大黃蜂蜂房之間的,還有墨西哥蜂(Melipona domestica)的蜂房,于貝爾曾經(jīng)仔細(xì)描述過和繪制過。墨西哥蜂的身體構(gòu)造介于蜜蜂和大黃蜂之間,但與后者關(guān)系比較接近;能營造差不多規(guī)則的蠟質(zhì)蜂巢,圓柱形蜂房,在里面孵化幼蜂,此外還有一些用作貯蜜的大型蠟質(zhì)蜂房。這些大型的蜂房接近球狀,大小差不多相等,并且聚集成不規(guī)則的一堆。這里要注意的要點(diǎn)是,蜂房總是營造得彼此很靠近,如果完全成為球狀時,蠟壁勢必就要交切或串通;但是從來不會如此,因?yàn)檫@種蜂會在有交切傾向的球狀蜂房之間把蠟壁造成平面的。因此,每個蜂房都是由外方的球狀部分和兩三個或更多平面構(gòu)成的,這要看這個蜂房與兩三個或更多的蜂房相連接來決定。一個蜂房連接三個蜂房時,由于球形是差不多大小的,這種情形常常而且必然發(fā)生,所以三個平面連合成為一個角錐體;據(jù)于貝爾說,這種角錐體明顯與蜜蜂蜂房的三邊角錐形底部十分相像。這里和蜜蜂蜂房一樣,任何蜂房的三個平面必然成為所連接的三個蜂房的構(gòu)成部分。墨西哥蜂用這種營造方法,顯然可以節(jié)省蠟;因?yàn)檫B接蜂房之間的平面壁并不是雙層的,其厚薄和外面的球狀部分相同,然而每一個平面壁卻構(gòu)成了兩個蜂房的共同部分。

考慮這個個案時,我覺得如果墨西哥蜂在一定的彼此距離間營造球狀蜂房,并且造成一樣大小,同時對稱排列成雙層,那么這構(gòu)造就會像蜜蜂巢一樣完美了。所以我寫信給劍橋的米勒(Miller)教授,根據(jù)他的復(fù)信,我寫出了以下的敘述,這位幾何學(xué)家惠讀了,并且告訴我說,這是完全正確的:

設(shè)若干同等大小的球,球心在兩個平行層上;每一個球的球心與同層中圍繞它的六個球的球心相距等于或稍微小于半徑×,即半徑×1.41421;并且與別一平行層中連接的球的球心相距也如上;于是,如果把這雙層每兩個球的交接面都畫出來,就會形成一個雙層六面柱體,其互相銜接的面都是由三個菱形所組成的角錐形底部聯(lián)結(jié)而成的;這個角錐形與六面柱體的邊所成的角,與經(jīng)過精密測量的蜜蜂蜂房的角完全相等。

因此可以穩(wěn)妥地斷定,如果能把墨西哥蜂的并不很奇異的已有本能稍微改變一下,便能造出像蜜蜂那樣巧奪天工的蜂房。我們必須假定,墨西哥蜂有能力來營造真正球狀的和大小相等的蜂房;鑒于已經(jīng)能夠在一定程度上做到這點(diǎn),鑒于還有許多昆蟲也能夠在樹木上造成多么完美的圓柱形孔穴,分明是依據(jù)一個固定的點(diǎn)旋轉(zhuǎn)而成的,這就沒有什么值得奇怪的了。必須假定,墨西哥蜂能把蜂房排列在水平層上,而其圓柱形蜂房就是這樣排列的。必須進(jìn)一步假定,當(dāng)幾只工蜂工友分別營造球狀蜂房時,能好歹正確判斷彼此應(yīng)當(dāng)距離多遠(yuǎn),而這是最困難的事;不過,已經(jīng)能判斷距離了,所以總是能使球狀蜂房有某種程度的交切;然后把交切點(diǎn)用完全的平面連接起來。必須再進(jìn)一步假定,六面柱體由同層連接球體的交接面形成之后,可以任意延長六面柱體的長度,使之符合倉儲蜂蜜的要求,而這一點(diǎn)不難;就像粗魯?shù)拇簏S蜂給舊繭的圓孔增加蠟質(zhì)圓管一樣的。本來并不奇異的本能——不比指導(dǎo)鳥類造巢的本能更奇異,經(jīng)過這樣的變異之后,我相信蜜蜂通過自然選擇就獲得了難以模仿的營造能力。

這種理論可用試驗(yàn)來證明。照特蓋特邁耶(Tegetmeier)先生的榜樣,我把兩個蜂巢分開,中間放一塊長而厚的方形蠟版:蜜蜂隨即開始在蠟版上鑿掘圓形的小凹穴;向深處鑿掘這些小穴時,逐漸使它們拓寬,變成約莫蜂房直徑的淺盆形,看起來恰似真正球狀或者球的一部分。下面的情形是極有趣的:凡是幾只蜂彼此靠近開始鑿掘盆形凹穴時,相互之間的距離恰使盆形凹穴得到上述寬度(大約相當(dāng)于一個普通蜂房的寬度),并且在深度上達(dá)到這些盆形凹穴所構(gòu)成的球體直徑的六分之一,這時盆形凹穴的邊便交切,或彼此串通。一遇到這種情形,即停止往深處鑿掘,并且開始在盆邊之間的交切處造起平面的蠟壁,所以,每一個六面柱體并不是像普通蜂房那樣,建筑在三邊角錐體的直邊上面,而是建造在一個平滑盆形的扇形邊上面的。

然后我把一塊薄而狹的涂有朱紅色、其邊如刃的蠟片放進(jìn)蜂箱里去,以代替以前所用的方形厚蠟版。于是蜜蜂即刻一如既往地在蠟片的兩面開始鑿掘一些彼此接近的盆形小穴。但蠟片太薄,如果盆形小穴的底掘得像上述試驗(yàn)一樣深,兩面便要彼此串通了。然而蜜蜂并不會讓這種情形發(fā)生,及時停止了開掘;于是那些盆形小穴,掘得深一點(diǎn)時,便出現(xiàn)了平的底,這等由剩下未被咬去的一小薄片朱紅色蠟所形成的平底,根據(jù)目測,正好位于蠟片正反面的盆形小穴之間的想象上的交切面處。部分地方只咬去一點(diǎn)點(diǎn),其他地方則是在對面的盆形小穴之間留下大片菱形板,不是自然狀態(tài)的東西,所以不能精巧地完成工作。蜂在朱紅色蠟片的兩面,渾圓地咬去蠟質(zhì),并使盆形加深,其工作速度想必是差不多的,這是為了能夠成功地在交切面處停止工作,而在盆形小穴之間留下平面。

鑒于薄蠟片十分柔軟,我想,蜂在蠟片兩面工作時,不難覺察到什么時候咬到適當(dāng)?shù)谋《?,于是停止工作。在普通的蜂巢里,我認(rèn)為蜂在兩面的工作速度,并不永遠(yuǎn)能實(shí)現(xiàn)完全相等;我注意過一個剛開始營造的蜂房底部上半完工的菱形板,其一面稍為凹進(jìn),我想象這是這面掘得太快的緣故,另一面則凸出,因?yàn)檫@面工作得慢一些。在一個著名事例里,我把這蜂巢放回蜂箱里去,讓蜂繼續(xù)工作一個短時間,然后再檢查蜂房,發(fā)現(xiàn)菱形板已經(jīng)完工,并且已經(jīng)完全平了:蠟片是極薄的,所以絕對不可能是從凸的一面把蠟咬去,做成上述的樣子;我猜測這種情形大概是站在反面的蜂,把可塑而溫暖的蠟恰到好處地推壓彎曲到中間板處(我試驗(yàn)過,很容易做),這樣就找平了。

從朱紅蠟片的試驗(yàn)可以看出,若要建造一堵蠟質(zhì)的薄壁,蜂便彼此站在一定距離,以同等的速度鑿掘下去,并且努力做成同等大小的球狀空室,但永遠(yuǎn)不會讓空室彼此串通,這樣就可造適當(dāng)形狀的蜂房。檢查一下正在建造的蜂巢邊緣,一眼就可看出首先在蜂巢的周圍造一堵粗糙的圍墻緣邊,然后從兩面對咬,加深每一個蜂房時,總是繞圈工作。并不在同一時間內(nèi)營造任一蜂房三邊角錐形的整個底部,而是看情況先搞定位于正在建造的極端邊緣的一兩塊菱形板;并且在沒有營造六面壁之前,絕不完成菱形板上部的邊。這些敘述有些和大名鼎鼎的老于貝爾所說的有所不同,但我相信是正確的;如果有篇幅,我將闡明這符合我的理論。

于貝爾說,最初的第一個蜂房是從側(cè)面相平行的蠟質(zhì)小壁鑿掘造出來的,就我所看到的,這一敘述并不嚴(yán)格正確。最初著手的經(jīng)常是一個小蠟兜,但這里我不擬詳論。我們知道,在蜂房的構(gòu)造里,鑿掘起著何等重要的作用;但如果設(shè)想蜂不能在適當(dāng)?shù)奈恢谩囱刂鴥蓚€連接的球形體之間的交切面——營造粗糙的蠟壁,就是極大的錯誤。我有幾件標(biāo)本明顯指出是能夠這樣做的。甚至在環(huán)繞著建造中的蜂巢周圍的粗糙邊緣即蠟壁上,有時候也可觀察到彎曲的情形,所在的位置相當(dāng)于未來蜂房的菱形底面。但在一切場合中,粗糙的蠟壁是靠大口咬掉兩面的蠟而完成的。蜂的這種營造方法是奇妙的;總是把最初的粗糙墻壁,造得比最后要留下的蜂房的極薄的壁厚十倍乃至二十倍。要理解它們的工作方法,可以假定泥水匠首先用水泥堆起一堵寬闊基墻,然后在近地面處的兩側(cè)把水泥同等地削去,直到中間部分形成一堵光滑而很薄的墻壁;泥水匠總是把削去的水泥堆在墻壁的頂上,還要加入新水泥。于是,薄壁就這樣不斷地壘上去,但上面總是有一個厚大的頂蓋。一切蜂房,無論剛開始營造還是已經(jīng)完成的,上面都有這樣一個堅(jiān)固的蠟蓋,因此,蜂能夠聚集在蜂巢上爬來爬去,而不會把薄六面壁損壞。壁的厚度只有約四百分之一英寸,菱形底片大約比其厚一倍。用上述這樣特別的營造方法,可以極端地省蠟,同時還能不斷地使蜂巢加固。

大批蜜蜂聚集一起工作,乍看這對于理解蜂房的營造方式會增加困難;一只蜂在一個蜂房工作一個短時間后,便到另一個蜂房,所以,如于貝爾所說的,甚至第一個蜂房開始營造時就有二十只蜂在工作。我用實(shí)踐的方法闡明了這一事實(shí):用朱紅色的極薄熔蠟涂在一個蜂房六面壁的邊上,或者涂在一個營造著的蜂巢圍墻的極端邊緣上,結(jié)果必定看到蜂把這顏色極細(xì)膩地分布開去——細(xì)膩得就像畫家布色——有顏色的蠟從涂抹的地方一星一星地取去,放到周圍蜂房擴(kuò)大著的邊緣上去。營造的工作在多蜂之間似乎有某種平衡分配,彼此本能地站在同一相對距離上,都試圖鑿掘相等的球形,然后,建造起或者說留下不咬球形之間的交切面。說起來實(shí)在是奇異,有時會遇到困難,例如兩個蜂巢成角度相遇時,往往把已成的蜂房徹底拆掉,用不同的方法重造,而有時候再現(xiàn)拆去的形狀。

蜂遇到可以各就各位進(jìn)行工作的一處地方,例如,一塊木片上,木片恰好處于向下建造的一個蜂巢的中部之下,那么這蜂巢勢必就要營造在木片的一面——在這種情況下,蜂便會筑起新的六面體一堵墻的基礎(chǔ),突出于已經(jīng)完成的蜂房之外,位置嚴(yán)格規(guī)定。只要蜂能彼此站在適當(dāng)?shù)木嚯x并且與最后完成的蜂房墻壁保持適當(dāng)?shù)木嚯x,掘造了想象的球形體,就足以在兩個鄰接的球形體之間造起中間蠟壁來;但據(jù)我所看到的,非到那蜂房和鄰接的幾個蜂房大都造成之后,從不咬去和修光蜂房的角。蜂在一定環(huán)境條件下,能在兩個剛開始營造的蜂房中間把一堵粗糙的壁建立在適當(dāng)位置上,這種能力是重要的;因?yàn)檫@與一項(xiàng)事實(shí)有關(guān),最初看來它似乎可以推翻上述理論;這事實(shí)就是,黃蜂最外邊緣上的一些蜂房也常常是嚴(yán)格的六邊形;但這里沒有篇幅討論這一問題。我并不覺得單獨(dú)一個昆蟲(例如黃蜂蜂后)營造六邊形的蜂房會有什么大困難,只要能在同時開始的兩三個巢房的內(nèi)側(cè)和外側(cè)交互地工作,始終與剛開工的蜂房部件保持適當(dāng)?shù)木嚯x,掘造球形或圓筒形,并且建造起中間的平壁。甚至可以想象,一個昆蟲固定于一點(diǎn)開始構(gòu)筑蜂房,然后移動出去,先到一點(diǎn),然后到另外五個點(diǎn),到中心點(diǎn)的相對距離和點(diǎn)之間的距離恰到好處,可以打造諸交切面,構(gòu)筑孤立的六邊形。但我不知道這種個案有沒有觀察到過,而且構(gòu)筑單個六邊形也沒有什么好處,因?yàn)檫@就比構(gòu)筑圓柱體需要更多的建筑材料。

自然選擇僅僅靠構(gòu)造或本能的微小變異的積累才發(fā)揮作用,而各個變異都對個體在其生活條件下是有利的。所以可以合理地發(fā)問:變異了的建筑本能所經(jīng)歷的漫長而級進(jìn)的連續(xù)階段,都趨向現(xiàn)今那樣完善的建筑規(guī)劃,對于蜜蜂祖先,曾起過怎樣有利的作用呢?我想,解答這個問題并不困難:我們知道,蜂為了采足花蜜,常常受到很大壓力。特蓋特邁耶先生告訴我說,實(shí)驗(yàn)已經(jīng)證明,蜜蜂分泌一磅蠟須消耗十二到十五磅干糖;所以一個蜂箱里的蜜蜂為了分泌營造蜂巢所必需的蠟,必須采集并消耗大量的液狀花蜜。還有,許多蜂在分泌的過程中,勢必有許多天不能工作。大量蜂蜜的貯藏,對于維持大群蜂的冬季生活是必不可缺少的;并且我們知道,蜂群的安全主要決定于大量的蜂得以維持。因此,大大節(jié)省蜂蜜,從而省蠟,必定是任何蜂族成功的重要因素。當(dāng)然,其成功還可能決定于寄生物等敵害的數(shù)量,決定于截然不同的原因,所以根本不取決于蜜蜂所能采集的蜜量。但是,讓我們假定采集蜜量的能力決定了任何一處地方大黃蜂的數(shù)量,這倒是常常發(fā)生;讓我們進(jìn)一步假定,那蜂群度過了冬季,結(jié)果就需要貯藏蜂蜜:在這種情形下,如果其本能有微小的變異,使得蠟房造得靠近些,略略彼此相切,無疑會有利于這批土蜂;一堵公共的壁即使僅連接兩個蜂房,也會節(jié)省少許蠟。因此,如果蜂房造得日益整齊,相互日益靠近,并且像墨西哥蜂的蜂房那樣聚集在一起,就會不斷地日益有利于這種大黃蜂;因?yàn)檫@樣各蜂房的大部分界壁將會用作鄰接蜂房的界壁,就可以大大省蠟。還有,由于同樣的原因,如果墨西哥蜂能把蜂房造得比現(xiàn)在接近些,并且在各方面都更規(guī)則些,這于己有利;因?yàn)椋缥覀兯吹降?,蜂房的球形面會完全消失,代以平面;而墨西哥蜂所造的蜂巢就會達(dá)到蜜蜂巢那樣完善的地步。在建造上超越這種完善的階段,自然選擇便不能再起作用;因?yàn)閾?jù)我們所知,蜜蜂巢在節(jié)蠟方面絕對是完美的。

因此,我認(rèn)為,一切既知本能中最奇異的本能——蜜蜂的本能,可以根據(jù)自然選擇利用了簡單本能之無數(shù)的、連續(xù)發(fā)生的微小變異來解釋;自然選擇曾經(jīng)緩慢、逐步完善地使得蜂在雙層上掘造彼此保持一定距離的、同等大小的球形體,并且沿著交切面筑起和鑿掘蠟壁。當(dāng)然,蜂不會知道自己在彼此保持一定距離掘造球形體,正如它們不會知道六面柱體與底部的菱形板是什么角度。自然選擇過程的動力在于節(jié)蠟;各蜂群在蠟的分泌上消耗最少的蜜,得到了最大的成功,并且把新獲得的節(jié)約本能遺傳給了新蜂群,以便在生存斗爭中獲得成功的最大機(jī)會。

無疑還可用許多極難解釋的本能來反對自然選擇學(xué)說——例如有些本能,我們不知道是怎樣起源的;有些本能,我們不知道有中間級進(jìn)存在;有些本能看上去很不重要,自然選擇不大會發(fā)生作用;有些本能在自然系統(tǒng)相距甚遠(yuǎn)的動物里竟幾乎相同,所以不能用共同祖先的遺傳來說明其相似性,結(jié)果只好相信這些本能是通過自然選擇而獨(dú)立獲得的。我不預(yù)備在這里討論這些個例子,而僅僅討論一個特別的難點(diǎn),起初我認(rèn)為這個難點(diǎn)是難以克服的,并且實(shí)際上對于我的整個理論是致命的。我所指的就是昆蟲社會里的中性即不育的雌蟲;這些中性蟲在本能和構(gòu)造上常與雄蟲以及能育的雌蟲有很大的差異,可是由于不育,卻不能繁殖同類。

這個問題很值得詳細(xì)討論,但這里只舉一個個案,即不育的工蟻。工蟻怎么會變?yōu)椴挥膫€體是個難點(diǎn),但不比構(gòu)造上任何顯著變異更難于解釋;可以證明,自然狀態(tài)下某些昆蟲以及別種節(jié)足動物偶爾也會變?yōu)椴挥?;如果這種昆蟲是社會性的,而且每年生下若干能工作但不能生殖的個體對于群體有利的話,那我認(rèn)為不難理解這是由于自然選擇的作用。但必須跳過這種初步的難點(diǎn)不談。最大的難點(diǎn)在于工蟻與雄蟻和能育的雌蟻在構(gòu)造上有巨大的差異,如工蟻具有不同形狀的胸部,缺翅膀,有時沒有眼睛,并且具有不同的本能。單以本能而論,蜜蜂可以極好地證明工蜂與完全的雌蜂之間有驚人的差異。如果工蟻或別種中性蟲原是正常的動物,那我就會毫不遲疑地假定,一切性狀都是通過自然選擇慢慢獲得的;這就是說,由于生下的個體構(gòu)造上都具有微小的有利變異,又都遺傳給了后代;而且后代又發(fā)生變異,又被選擇,如此等等,不一而足。但是工蟻和雙親之間的差異很大,又是絕對不育的,所以絕不可能把歷代獲得的構(gòu)造上或本能上的變異遺傳給后代。于是可以設(shè)問:這怎么能符合自然選擇的學(xué)說呢?

首先,請記住,家養(yǎng)生物和自然狀態(tài)下的生物里,構(gòu)造的各種各樣差異是與一定年齡或性別相關(guān)的,這方面有無數(shù)的案例。差異不但與性別相關(guān),而且與生殖系統(tǒng)活躍的那一短暫時期相關(guān),例如,許多鳥類的求婚羽,雄三文魚鉤曲的顎,都是這種情形。公牛經(jīng)人工去勢后,不同品種的角甚至相關(guān)地表現(xiàn)了微小的差異;某些品種的去勢公牛,與同品種的公牝雙方比較,犄角比其他品種更長。因此,我認(rèn)為任何性狀變得與昆蟲社會里某些成員的不育狀態(tài)相關(guān),并不存在多大難點(diǎn);難點(diǎn)在于理解這種構(gòu)造上的相關(guān)變異如何因自然選擇作用而慢慢累積起來。

這個難點(diǎn)表面上看來是難以克服的,可是只要記住選擇作用可以應(yīng)用于個體也可以應(yīng)用于全族,而且可以由此如愿以償,那么難點(diǎn)便會縮小,或者如我所相信的,便會消除。比如,一棵味道好的蔬菜煮熟吃了,該個體就消滅了。可是園藝家播下同種蔬菜的種子,信心十足地期望收獲差不多的變種。養(yǎng)牛者喜歡肉和脂肪交織成大理石紋的樣子,牲口已經(jīng)屠殺了,但是養(yǎng)牛者有信心繼續(xù)找到同樣的牛。我對于選擇的力量也是信心十足,并不懷疑總是產(chǎn)生異常長角的去勢公牛的品種,可以慢慢培養(yǎng),只要仔細(xì)觀察什么樣的公牛和牝牛個體交配才能產(chǎn)生最長角的去勢公牛;雖然沒有一只去勢的牛曾經(jīng)繁殖過同類。我想社會性的昆蟲也是如此:與同群某些成員的不育狀態(tài)相關(guān)的構(gòu)造、本能上的輕微變異,對于群體有利,結(jié)果能育的雄體和雌體得到了繁生,并把這種產(chǎn)生具有同樣變異的不育成員的傾向,傳遞給了能育的后代。我認(rèn)為,這一過程重復(fù)過了許多次,直到同一物種的能育雌體和不育雌體之間產(chǎn)生了巨大的差異量,就像我們在許多種社會性昆蟲里所見到的那樣。

但我們還沒有談及登峰造極的難點(diǎn):有幾種蟻的中性蟲不但與能育的雌蟲和雄蟲有所差異,而且彼此之間也有差異,有時差異甚至到了讓人幾乎難以置信的程度,并且因此被分成兩個級(castes),甚至三個級。還有,這些級一般并不彼此逐漸重疊,而是區(qū)別得十分清楚,有如同屬的兩個物種,同科的兩個屬。例如,埃西頓(Eciton)行軍蟻的中性工蟻和兵蟻具有大相徑庭的顎和本能:隱角蟻(Cryptocerus)只有一個級的工蟻,頭上生有一種奇異的盾,用途不清楚;墨西哥的蜜蟻(Myrmecocystus)有一個級的工蟻從不離巢,由另一個級的工蟻喂食,腹部發(fā)育得很大,能分泌出一種蜜汁,以代替蚜蟲所排泄的東西;蚜蟲或者可以被稱為蟻乳牛,歐洲的蟻常把它們守衛(wèi)圈禁起來。

如果不承認(rèn)這種奇異而十分確實(shí)的事實(shí)可以瞬間顛覆我的理論,人們必然會想,我對自然選擇的原理過于信心飽滿了。如果中性蟲只有一個級,我相信它與能育的雄蟲和雌蟲之間的差異是通過自然選擇得到的,在這種比較簡單的情形里,根據(jù)普通變異類推,我們可以斷言,各種連續(xù)的、微小的、有利的變異,最初并非發(fā)生于同一窩中的所有中性蟲,而只發(fā)生于少數(shù)的中性蟲;經(jīng)過長期持續(xù)選擇能夠產(chǎn)生極多的具有有利變異的中性蟲的能育親種,一切中性蟲最終就都會具有所需的性狀。按照這種觀點(diǎn),我們應(yīng)該在同一巢中偶爾發(fā)現(xiàn)那些表現(xiàn)有構(gòu)造分級的同種中性蟲;實(shí)際我們是發(fā)現(xiàn)了,鑒于歐洲以外的中性昆蟲很少仔細(xì)檢查過,甚至可以說并不稀罕。史密斯先生曾闡明,有幾種英國蟻的中性蟲彼此在大小方面,有時在顏色方面,表現(xiàn)了驚人的差異;并且在兩極端的類型之間,有時可由同巢中的一些個體連接起來:筆者就比較過這種完美的級進(jìn)情形。有時可以看到,大形或者小形的工蟻數(shù)目最多;或者大形和小形兩種都多,而中間形的數(shù)目卻稀少。黃蟻有大工蟻和小工蟻,中間形的工蟻有一些;如史密斯先生所觀察的,在這個物種里,大工蟻有單眼(ocelli),雖小但能夠清楚辨認(rèn);而小工蟻的單眼則是殘跡。仔細(xì)解剖了幾只工蟻標(biāo)本之后,我能確定小工蟻的眼睛根本不發(fā)育,遠(yuǎn)非單單用其小比例所能解釋;并且我充分相信,雖然我不敢很肯定地斷言,中間形工蟻的單眼正好處在中間狀態(tài)。所以,一個巢內(nèi)有兩群不育工蟻,不但在大小上,并且在視覺器官上,都表現(xiàn)了差異,然而有少數(shù)中間狀態(tài)的成員連接起來。我再補(bǔ)充幾句題外的話,如果小工蟻對于蟻群最有利,產(chǎn)生越來越多小工蟻的雄蟻和雌蟻不斷被選擇,最后所有的工蟻都具有那種形態(tài)了。于是就形成了一個蟻種,其中性蟲差不多就像褐蟻屬(Myrmica)工蟻那樣。褐蟻屬的工蟻甚至連殘跡的單眼都沒有,盡管這個屬的雄蟻和雌蟻都生有很發(fā)達(dá)的單眼。

我再舉一例:同一物種的不同級的中性蟲之間,我滿有信心地期望可以找到重要構(gòu)造的中間分級,欣然利用史密斯先生所提供的取自西非驅(qū)逐蟻(Anomma)同巢中的許多標(biāo)本。我不舉實(shí)際的測量數(shù)字,只做一個嚴(yán)格精確的說明,讀者大概就能最好地了解這工蟻之間的差異量;差異就好比看到一群建筑工人,其中有許多是五英尺四英寸高,還有許多是十六英尺高;但我們必須再假定那大個兒工人的頭比小個兒工人不止大三倍,卻要大四倍,而顎則要大近五倍。再者,大小不同的工蟻的顎不僅在形狀上大有差異,而且牙齒的形狀和數(shù)目也相差懸殊。但重要的事實(shí)卻是,雖然工蟻可以依大小分為不同的等級,卻不知不覺地彼此級進(jìn)重疊,其構(gòu)造大不相同的顎也是這樣。關(guān)于后面一點(diǎn)我有把握,盧伯克先生曾用描圖器把我所解剖的幾種大小不同的工蟻的顎逐一作圖。

根據(jù)擺在我面前的這些事實(shí),我相信自然選擇由于作用于能育的蟻雙親,便可以形成一個物種,專門產(chǎn)生體形大而具有某一形狀的顎的中性蟲,或者專門產(chǎn)生體形小而構(gòu)造大不相同的顎的中性蟲;最后,這是登峰造極的難點(diǎn),一群工蟻具有一種大小和構(gòu)造,另一群工蟻具有不同大小和構(gòu)造,同時存在——最先形成的是一個級進(jìn)的系列,就像驅(qū)逐蟻的情形那樣,然后,由于自然選擇支持生育它們的雙親,就產(chǎn)生了越來越多最有利于蟻群的兩極端類型,最后具有中間構(gòu)造的個體不再產(chǎn)生。

我認(rèn)為,奇異事實(shí)就是這樣發(fā)生的,同一巢里生存的、區(qū)別分明的不育工蟻兩級,不但彼此之間大不相同,并且和雙親之間也大不相同。我們可以看出工蟻的生成對于蟻社會有多大的用處,與社會分工對于文明人的用處同理。由于蟻是用遺傳的本能和遺傳的工具或武器來工作的,而不用學(xué)得的知識和制造的器具,所以完美分工只能通過工蟻不育來實(shí)施。如果它們能育,就會雜交,其本能和構(gòu)造就會混雜。我認(rèn)為,大自然通過自然選擇在蟻群中實(shí)施了這一令人驚嘆的分工。但是必須坦白承認(rèn),我雖然完全相信自然選擇,若不是有這等中性蟲個案讓我心悅誠服,決不會料到這一原理是如此高度有效。所以,為了闡明自然選擇的力量,并且因?yàn)檫@是我的理論所遭到的特別嚴(yán)重的難點(diǎn),我對于這個個案做了稍多的但掛一漏萬的討論。而且這個個案也很有趣,證明了動物同植物一樣,把無數(shù)的微小的必須稱為偶發(fā)的構(gòu)造變異,只要是稍微有利的就累積下來,沒有鍛煉或習(xí)性參加作用,任何量的變異都能實(shí)現(xiàn)。蟻群的不育成員的鍛煉,或者習(xí)性,或者意愿,絲毫也不可能影響專事遺留后代的能育成員的構(gòu)造或者本能。我覺得奇怪的是,至今沒有人提出用這種中性蟲的演示個案去反對眾所熟知的拉馬克(Lamarck)的學(xué)說。

提要。——本章勉力簡要地指出了家養(yǎng)動物的精神品質(zhì)要變異,且這種變異是遺傳的。我又更簡要地闡明本能在自然狀態(tài)下也是輕微變異的。沒有人會爭辯本能對于各種動物有極端的重要性。所以我認(rèn)為,在多變的生活條件下,自然選擇不難把任何稍微有用的本能上的微小變異累積到任何程度。在許多情況下,習(xí)性或者用廢大概也參加作用。我不敢說本章所舉事實(shí)能在很大程度上鞏固我的理論;但是根據(jù)我所能判斷的,沒有難解的個案可加以顛覆。另外,本能不總是絕對完善,而是易犯錯誤;沒有一種本能可說是為了其他動物的獨(dú)享利益而產(chǎn)生的,但各種動物都利用其他動物的本能;博物史上的格言“自然界里沒有飛躍”,適用于身體構(gòu)造也適用于本能,并且可用上述觀點(diǎn)來清楚解釋,別無他解——所有這些事實(shí)都確證了自然選擇的學(xué)說。

這個理論也得到其他幾種關(guān)于本能的事實(shí)的加強(qiáng);常見的個案如,密切近似的但不相同的物種,當(dāng)天各一方并且生活在相當(dāng)不同的生活條件下時,常常保持了幾乎同樣的本能。例如,根據(jù)遺傳原理,我們能夠理解,為什么南美鶇跟英國鶇的特別造巢方法一樣,用泥來涂抹它們的巢;為什么北美洲的雄性鷦鷯(Troglodytes)像英國的雄性貓形鷦鷯(Kitty-wrens)那樣地營造“雄鳥之巢”棲居——這種習(xí)性完全不像任何其他已知鳥類。最后,這可能是不合邏輯的演繹,但據(jù)我想象,這樣說法最能令人滿意,如小杜鵑把義兄弟逐出巢外,蓄奴蟻,姬蜂科(ichneumonidae)幼蟲寄生在活的毛毛蟲體內(nèi),不把這種本能看作是特別賦予或特別創(chuàng)造的,而看作是引導(dǎo)一切生物進(jìn)化——即繁殖,變異,讓最強(qiáng)者生存、最弱者死亡——的一般法則的小小結(jié)果。

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