VII

THE PERPETUATION OF LIVING BEINGS, HEREDITARY TRANSMISSION AND VARIATION.

The inquiry which we undertook, at our last meeting, into the state of our knowledge of the causes of the phenomena of organic nature,—of the past and of the present,—resolved itself into two subsidiary inquiries: the first was, whether we know anything, either historically or experimentally, of the mode of origin of living beings; the second subsidiary inquiry was, whether, granting the origin, we know anything about the perpetuation and modifications of the forms of organic beings. The reply which I had to give to the first question was altogether negative, and the chief result of my last lecture was, that, neither historically nor experimentally, do we at present know anything whatsoever about the origin of living forms. We saw that, historically, we are not likely to know anything about it, although we may perhaps learn something experimentally; but that at present we are an enormous distance from the goal I indicated.

I now, then, take up the next question, What do we know of the reproduction, the perpetuation, and the modifications of the forms of living beings, supposing that we have put the question as to their origination on one side, and have assumed that at present the causes of their origination are beyond us, and that we know nothing about them? Upon this question the state of our knowledge is extremely different; it is exceedingly large: and, if not complete, our experience is certainly most extensive. It would be impossible to lay it all before you, and the most I can do, or need do to-night, is to take up the principal points and put them before you with such prominence as may subserve the purposes of our present argument.

The method of the perpetuation of organic beings is of two kinds,—the asexual and the sexual. In the first the perpetuation takes place from and by a particular act of an individual organism, which sometimes may not be classed as belonging to any sex at all. In the second case, it is in consequence of the mutual action and inter-action of certain portions of the organisms of usually two distinct individuals—the male and the female. The cases of asexual perpetuation are by no means so common as the cases of sexual perpetuation; and they are by no means so common in the animal as in the vegetable world. You are all probably familiar with the fact, as a matter of experience, that you can propagate plants by means of what are called “cuttings”; for example, that by taking a cutting from a geranium plant, and rearing it properly, by supplying it with light and warmth and nourishment from the earth, it grows up and takes the form of its parent, having all the properties and peculiarities of the original plant.

Sometimes this process, which the gardener performs artificially, takes place naturally; that is to say, a little bulb, or portion of the plant, detaches itself, drops off, and becomes capable of growing as a separate thing. That is the case with many bulbous plants, which throw off in this way secondary bulbs, which are lodged in the ground and become developed into plants. This is an asexual process, and from it results the repetition or reproduction of the form of the original being from which the bulb proceeds.

Among animals the same thing takes place. Among the lower forms of animal life, the infusorial animalculæ we have already spoken of throw off certain portions, or break themselves up in various directions, sometimes transversely or sometimes longitudinally; or they may give off buds, which detach themselves and develop into their proper forms. There is the common fresh-water Polype, for instance, which multiplies itself in this way. Just in the same way as the gardener is able to multiply and reproduce the peculiarities and characters of particular plants by means of cuttings, so can the physiological experimentalist,—as was shown by the Abbé Trembley many years ago,—so can he do the same thing with many of the lower forms of animal life. M. de Trembley showed that you could take a polype and cut it into two, or four, or many pieces, mutilating it in all directions, and the pieces would still grow up and reproduce completely the original form of the animal. These are all cases of asexual multiplication, and there are other instances, and still more extraordinary ones, in which this process takes place naturally, in a more hidden, a more recondite kind of way. You are all of you familiar with that little green insect, the Aphis or blight, as it is called. These little animals, during a very considerable part of their existence, multiply themselves by means of a kind of internal budding, the buds being developed into essentially asexual animals, which are neither male nor female; they become converted into young Aphides, which repeat the process, and their offspring after them, and so on again; you may go on for nine or ten, or even twenty or more successions; and there is no very good reason to say how soon it might terminate, or how long it might not go on if the proper conditions of warmth and nourishment were kept up.

Sexual reproduction is quite a distinct matter. Here, in all these cases, what is required is the detachment of two portions of the parental organisms, which portions we know as the egg or the spermatozoon. In plants it is the ovule and the pollen-grain, as in the flowering plants, or the ovule and the antherozooid, as in the flowerless. Among all forms of animal life, the spermatozoa proceed from the male sex, and the egg is the product of the female. Now, what is remarkable about this mode of reproduction is this, that the egg by itself, or the spermatozoa by themselves, are unable to assume the parental form; but if they be brought into contact with one another, the effect of the mixture of organic substances proceeding from two sources appears to confer an altogether new vigour to the mixed product. This process is brought about, as we all know, by the sexual intercourse of the two sexes, and is called the act of impregnation. The result of this act on the part of the male and female is, that the formation of a new being is set up in the ovule or egg; this ovule or egg soon begins to be divided and subdivided, and to be fashioned into various complex organisms, and eventually to develop into the form of one of its parents, as I explained in the first lecture. These are the processes by which the perpetuation of organic beings is secured. Why there should be the two modes—why this reinvigoration should be required on the part of the female element we do not know; but it is most assuredly the fact, and it is presumable, that, however long the process of asexual multiplication could be continued,—I say there is good reason to believe that it would come to an end if a new commencement were not obtained by a conjunction of the two sexual elements.

That character which is common to these two distinct processes is this, that, whether we consider the reproduction, or perpetuation, or modification of organic beings as they take place asexually, or as they may take place sexually,—in either case, I say, the offspring has a constant tendency to assume, speaking generally, the character of the parent. As I said just now, if you take a slip of a plant, and tend it with care, it will eventually grow up and develop into a plant like that from which it had sprung; and this tendency is so strong that, as gardeners know, this mode of multiplying by means of cuttings is the only secure mode of propagating very many varieties of plants; the peculiarity of the primitive stock seems to be better preserved if you propagate it by means of a slip than if you resort to the sexual mode.

Again, in experiments upon the lower animals, such as the polype, to which I have referred, it is most extraordinary that, although cut up into various pieces, each particular piece will grow up into the form of the primitive stock; the head, if separated, will reproduce the body and the tail; and if you cut off the tail, you will find that that will reproduce the body and all the rest of the members, without in any way deviating from the plan of the organism from which these portions have been detached. And so far does this go, that some experimentalists have carefully examined the lower orders of animals,—among them the Abbé Spallanzani, who made a number of experiments upon snails and salamanders,—and have found that they might mutilate them to an incredible extent; that you might cut off the jaw or the greater part of the head, or the leg or the tail, and repeat the experiment several times, perhaps, cutting off the same member again and again; and yet each of those types would be reproduced according to the primitive type: nature making no mistake, never putting on a fresh kind of leg, or head, or tail, but always tending to repeat and to return to the primitive type.

It is the same in sexual reproduction: it is a matter of perfectly common experience, that the tendency on the part of the offspring always is, speaking broadly, to reproduce the form of the parents. The proverb has it that the thistle does not bring forth grapes; so, among ourselves, there is always a likeness, more or less marked and distinct, between children and their parents. That is a matter of familiar and ordinary observation. We notice the same thing occurring in the cases of the domestic animals—dogs, for instance, and their offspring. In all these cases of propagation and perpetuation, there seems to be a tendency in the offspring to take the characters of the parental organisms. To that tendency a special name is given—and as I may very often use it, I will write it up here on this blackboard that you may remember it—it is called Atavism; it expresses this tendency to revert to the ancestral type, and comes from the Latin word atavus, ancestor.

Well, this Atavismwhich I shall speak of, is, as I said before, one of the most marked and striking tendencies of organic beings; but, side by side with this hereditary tendency there is an equally distinct and remarkable tendency to variation. The tendency to reproduce the original stock has, as it were, its limits, and side by side with it there is a tendency to vary in certain directions, as if there were two opposing powers working upon the organic being, one tending to take it in a straight line, and the other tending to make it diverge from that straight line, first to one side and then to the other.

So that you see these two tendencies need not precisely contradict one another, as the ultimate result may not always be very remote from what would have been the case if the line had been quite straight.

This tendency to variation is less marked in that mode of propagation which takes place asexually; it is in that mode that the minor characters of animal and vegetable structures are most completely preserved. Still, it will happen sometimes, that the gardener, when he has planted a cutting of some favourite plant, will find, contrary to his expectation, that the slip grows up a little different from the primitive stock—that it produces flowers of a different colour or make, or some deviation in one way or another. This is what is called the “sporting” of plants.

In animals the phenomena of asexual propagation are so obscure, that at present we cannot be said to know much about them; but if we turn to that mode of perpetuation which results from the sexual process, then we find variation a perfectly constant occurrence, to a certain extent; and, indeed, I think that a certain amount of variation from the primitive stock is the necessary result of the method of sexual propagation itself; for, inasmuch as the thing propagated proceeds from two organisms of different sexes and different makes and temperaments, and as the offspring is to be either of one sex or the other, it is quite clear that it cannot be an exact diagonal of the two, or it would be of no sex at all; it cannot be an exact intermediate form between that of each of its parents—it must deviate to one side or the other. You do not find that the male follows the precise type of the male parent, nor does the female always inherit the precise characteristics of the mother,—there is always a proportion of the female character in the male offspring, and of the male character in the female offspring. That must be quite plain to all of you who have looked at all attentively on your own children or those of your neighbours; you will have noticed how very often it may happen that the son shall exhibit the maternal type of character, or the daughter possess the characteristics of the father’s family. There are all sorts of intermixtures and intermediate conditions between the two, where complexion, or beauty, or fifty other different peculiarities belonging to either side of the house, are reproduced in other members of the same family. Indeed, it is sometimes to be remarked in this kind of variation, that the variety belongs, strictly speaking, to neither of the immediate parents; you will see a child in a family who is not like either its father or its mother; but some old person who knew its grandfather or grandmother, or, it may be, an uncle, or, perhaps, even a more distant relative, will see a great similarity between the child and one of these. In this way it constantly happens that the characteristic of some previous member of the family comes out and is reproduced and recognized in the most unexpected manner.

But apart from that matter of general experience, there are some cases which put that curious mixture in a very clear light. You are aware that the offspring of the Ass and the Horse, or rather of the he-Ass and the Mare, is what is called a Mule; and, on the other hand, the offspring of the Stallion and the she-Ass is what is called a Hinny. It is a very rare thing in this country to see a Hinny. I never saw one myself; but they have been very carefully studied. Now, the curious thing is this, that although you have the same elements in the experiment in each case, the offspring is entirely different in character, according as the male influence comes from the Ass or the Horse. Where the Ass is the male, as in the case of the Mule, you find that the head is like that of the Ass, that the ears are long, the tail is tufted at the end, the feet are small, and the voice is an unmistakable bray; these are all points of similarity to the Ass; but, on the other hand, the barrel of the body and the cut of the neck are much more like those of the Mare. Then, if you look at the Hinny,—the result of the union of the Stallion and the she-Ass, then you find it is the Horse that has the predominance; that the head is more like that of the Horse, the ears are shorter, the legs coarser, and the type is altogether altered; while the voice, instead of being a bray, is the ordinary neigh of the Horse. Here, you see, is a most curious thing: you take exactly the same elements, Ass and Horse, but you combine the sexes in a different manner, and the result is modified accordingly. You have in this case, however, a result which is not general and universal—there is usually an important preponderance, but not always on the same side.

Here, then, is one intelligible, and, perhaps, necessary cause of variation: the fact, that there are two sexes sharing in the production of the offspring, and that the share taken by each is different and variable, not only for each combination, but also for different members of the same family.

Secondly, there is a variation, to a certain extent,—though in all probability the influence of this cause has been very much exaggerated—but there is no doubt that variation is produced, to a certain extent, by what are commonly known as external conditions,—such as temperature, food, warmth, and moisture. In the long run, every variation depends, in some sense, upon external conditions, seeing that everything has a cause of its own. I use the term “external conditions” now in the sense in which it is ordinarily employed: certain it is, that external conditions have a definite effect. You may take a plant which has single flowers, and by dealing with the soil, and nourishment, and so on, you may by-and-by convert single flowers into double flowers, and make thorns shoot out into branches. You may thicken or make various modifications in the shape of the fruit. In animals, too, you may produce analogous changes in this way, as in the case of that deep bronze colour which persons rarely lose after having passed any length of time in tropical countries. You may also alter the development of the muscles very much, by dint of training; all the world knows that exercise has a great effect in this way; we always expect to find the arm of a blacksmith hard and wiry, and possessing a large development of the brachial muscles. No doubt, training, which is one of the forms of external conditions, converts what are originally only instructions, teachings, into habits, or, in other words, into organizations, to a great extent; but this second cause of variation cannot be considered to be by any means a large one. The third cause that I have to mention, however, is a very extensive one. It is one that, for want of a better name, has been called “spontaneous variation”; which means that when we do not know anything about the cause of phenomena, we call it spontaneous. In the orderly chain of causes and effects in this world, there are very few things of which it can be said with truth that they are spontaneous. Certainly not in these physical matters,—in these there is nothing of the kind,—everything depends on previous conditions. But when we cannot trace the cause of phenomena, we call them spontaneous.

Of these variations, multitudinous as they are, but little is known with perfect accuracy, I will mention to you some two or three cases, because they are very remarkable in themselves, and also because I shall want to use them afterwards. Réaumur, a famous French naturalist, a great many years ago, in an essay which he wrote upon the art of hatching chickens,—which was indeed a very curious essay,—had occasion to speak of variations and monstrosities. One very remarkable case had come under his notice of a variation in the form of a human member, in the person of a Maltese, of the name of Gratio Kelleia, who was born with six fingers upon each hand, and the like number of toes to each of his feet. That was a case of spontaneous variation. Nobody knows why he was born with that number of fingers and toes, and as we don’t know, we call it a case of “spontaneous” variation. There is another remarkable case also. I select these, because they happen to have been observed and noted very carefully at the time. It frequently happens that a variation occurs, but the persons who notice it do not take any care in noting down the particulars, until at length, when inquiries come to be made, the exact circumstances are forgotten; and hence, multitudinous as may be such “spontaneous” variations, it is exceedingly difficult to get at the origin of them.

The second case is one of which you may find the whole details in the “Philosophical Transactions” for the year 1813, in a paper communicated by Colonel Humphreys to the President of the Royal Society,—“On a new Variety in the Breed of Sheep,” giving an account of a very remarkable breed of sheep, which at one time was well known in the northern states of America, and which went by the name of the Ancon or the Otter breed of sheep. In the year 1791, there was a farmer of the name of Seth Wright in Massachusetts, who had a flock of sheep, consisting of a ram and, I think, of some twelve or thirteen ewes. Of this flock of ewes, one at the breeding-time bore a lamb which was very singularly formed; it had a very long body, very short legs, and those legs were bowed! I will tell you by-and-by how this singular variation in the breed of sheep came to be noted, and to have the prominence that it now has. For the present, I mention only these two cases; but the extent of variation in the breed of animals is perfectly obvious to any one who has studied natural history with ordinary attention, or to any person who compares animals with others of the same kind. It is strictly true that there are never any two specimens which are exactly alike; however similar, they will always differ in some certain particular.

Now let us go back to Atavism,—to the hereditary tendency I spoke of. What will come of a variation when you breed from it, when Atavism comes, if I may say so, to intersect variation? The two cases of which I have mentioned the history, give a most excellent illustration of what occurs. Gratio Kelleia, the Maltese, married when he was twenty-two years of age, and, as I suppose there were no six-fingered ladies in Malta, he married an ordinary five-fingered person. The result of that marriage was four children; the first, who was christened Salvator, had six fingers and six toes, like his father; the second was George, who had five fingers and toes, but one of them was deformed, showing a tendency to variation; the third was Andrè; he had five fingers and five toes, quite perfect; the fourth was a girl, Marie; she had five fingers and five toes, but her thumbs were deformed, showing a tendency toward the sixth.

These children grew up, and when they came to adult years, they all married, and of course it happened that they all married five-fingered and five-toed persons. Now let us see what were the results. Salvator had four children; they were two boys, a girl, and another boy: the first two boys and the girl were six-fingered and six-toed like their grandfather; the fourth boy had only five fingers and five toes. George had only four children: there were two girls with six fingers and six toes; there was one girl with six fingers and five toes on the right side, and five fingers and five toes on the left side, so that she was half and half. The last, a boy, had five fingers and five toes. The third, Andrè, you will recollect, was perfectly well-formed, and he had many children whose hands and feet were all regularly developed. Marie, the last, who, of course, married a man who had only five fingers, had four children: the first, a boy, was born with six toes, but the other three were normal.

Now observe what very extraordinary phenomena are presented here. You have an accidental variation arising from what you may call a monstrosity; you have that monstrosity tendency or variation diluted in the first instance by an admixture with a female of normal construction, and you would naturally expect that, in the results of such an union, the monstrosity, if repeated, would be in equal proportion with the normal type; that is to say, that the children would be half and half, some taking the peculiarity of the father, and the others being of the purely normal type of the mother; but you see we have a great preponderance of the abnormal type. Well, this comes to be mixed once more with the pure, the normal type, and the abnormal is again produced in large proportion, notwithstanding the second dilution. Now what would have happened if these abnormal types had intermarried with each other; that is to say, suppose the two boys of Salvator had taken it into their heads to marry their first cousins, the two first girls of George, their uncle? You will remember that these are all of the abnormal type of their grandfather. The result would probably have been, that their offspring would have been in every case a further development of that abnormal type. You see it is only in the fourth, in the person of Marie, that the tendency, when it appears but slightly in the second generation, is washed out in the third, while the progeny of Andrè, who escaped in the first instance, escape altogether.

We have in this case a good example of nature’s tendency to the perpetuation of a variation. Here it is certainly a variation which carried with it no use or benefit; and yet you see the tendency to perpetuation may be so strong, that, notwithstanding a great admixture of pure blood, the variety continues itself up to the third generation, which is largely marked with it. In this case, as I have said, there was no means of the second generation intermarrying with any but five-fingered persons, and the question naturally suggests itself, What would have been the result of such marriage? Réaumur narrates this case only as far as the third generation. Certainly it would have been an exceedingly curious thing if we could have traced this matter any further; had the cousins intermarried, a six-fingered variety of the human race might have been set up.

To show you that this supposition is by no means an unreasonable one, let me now point out what took place in the case of Seth Wright’s sheep, where it happened to be a matter of moment to him to obtain a breed or raise a flock of sheep like that accidental variety that I have described—and I will tell you why. In that part of Massachusetts where Seth Wright was living, the fields were separated by fences, and the sheep, which were very active and robust, would roam abroad, and without much difficulty jump over these fences into other people’s farms. As a matter of course, this exuberant activity on the part of the sheep constantly gave rise to all sorts of quarrels, bickerings, and contentions among the farmers of the neighbourhood; so it occurred to Seth Wright, who was, like his successors, more or less ’cute, that if he could get a stock of sheep like those with the bandy legs, they would not be able to jump over the fences so readily; and he acted upon that idea. He killed his old ram, and as soon as the young one arrived at maturity, he bred altogether from it. The result was even more striking than in the human experiment which I mentioned just now. Colonel Humphreys testifies that it always happened that the offspring were either pure Ancons or pure ordinary sheep; that in no case was there any mixing of the Ancons with the others. In consequence of this, in the course of a very few years, the farmer was able to get a very considerable flock of this variety, and a large number of them were spread throughout Massachusetts. Most unfortunately, however—I suppose it was because they were so common—nobody took enough notice of them to preserve their skeletons; and although Colonel Humphreys states that he sent a skeleton to the President of the Royal Society at the same time that he forwarded his paper, I am afraid that the variety has entirely disappeared; for a short time after these sheep had become prevalent in that district, the Merino sheep were introduced; and as their wool was much more valuable, and as they were a quiet race of sheep, and showed no tendency to trespass or jump over fences, the Otter breed of sheep, the wool of which was inferior to that of the Merino, was gradually allowed to die out.

You see that these facts illustrate perfectly well what may be done if you take care to breed from stocks that are similar to each other. After having got a variation, if, by crossing a variation with the original stock, you multiply that variation, and then take care to keep that variation distinct from the original stock, and make them breed together,—then you may almost certainly produce a race whose tendency to continue the variation is exceedingly strong.

This is what is called “selection”; and it is by exactly the same process as that by which Seth Wright bred his Ancon sheep, that our breeds of cattle, dogs, and fowls, are obtained. There are some possibilities of exception, but still, speaking broadly, I may say that this is the way in which all our varied races of domestic animals have arisen; and you must understand that it is not one peculiarity or one characteristic alone in which animals may vary. There is not a single peculiarity or characteristic of any kind, bodily or mental, in which offspring may not vary to a certain extent from the parent and other animals.

Among ourselves this is well known. The simplest physical peculiarity is mostly reproduced. I know a case of a woman who has the lobe of one of her ears a little flattened. An ordinary observer might scarcely notice it, and yet every one of her children has an approximation to the same peculiarity to some extent. If you look at the other extreme, too, the gravest diseases, such as gout, scrofula, and consumption, may be handed down with just the same certainty and persistence as we noticed in the perpetuation of the bandy legs of the Ancon sheep.

However, these facts are best illustrated in animals, and the extent of the variation, as is well known, is very remarkable in dogs. For example, there are some dogs very much smaller than others; indeed, the variation is so enormous that probably the smallest dog would be about the size of the head of the largest; there are very great variations in the structural forms not only of the skeleton but also in the shape of the skull, and in the proportions of the face and the disposition of the teeth.

The Pointer, the Retriever, Bulldog, and the Terrier, differ very greatly, and yet there is every reason to believe that every one of these races has arisen from the same source,—that all the most important races have arisen by this selective breeding from accidental variation.

A still more striking case of what may be done by selective breeding, and it is a better case, because there is no chance of that partial infusion of error to which I alluded, has been studied very carefully by Mr. Darwin,—the case of the domestic pigeons. I dare say there may be some among you who may be pigeon fanciers, and I wish you to understand that in approaching the subject, I would speak with all humility and hesitation, as I regret to say that I am not a pigeon fancier. I know it is a great art and mystery, and a thing upon which a man must not speak lightly; but I shall endeavour, as far as my understanding goes, to give you a summary of the published and unpublished information which I have gained from Mr. Darwin.

Among the enormous variety,—I believe there are somewhere about a hundred and fifty kinds of pigeons,—there are four kinds which may be selected as representing the extremest divergences of one kind from another. Their names are the Carrier, the Pouter, the Fantail, and the Tumbler. In these large diagrams that I have here they are each represented in their relative sizes to each other. This first one is the Carrier; you will notice this large excrescence on its beak; it has a comparatively small head; there is a bare space round the eyes; it has a long neck, a very long beak, very strong legs, large feet, long wings, and so on. The second one is the Pouter, a very large bird, with very long legs and beak. It is called the Pouter because it is in the habit of causing its gullet to swell up by inflating it with air. I should tell you that all pigeons have a tendency to do this at times, but in the Pouter it is carried to an enormous extent. The birds appear to be quite proud of their power of swelling and puffing themselves out in this way; and I think it is about as droll a sight as you can well see to look at a cage full of these pigeons puffing and blowing themselves out in this ridiculous manner.

This diagram is a representation of the third kind I mentioned—the Fantail. It is, you see, a small bird, with exceedingly small legs and a very small beak. It is most curiously distinguished by the size and extent of its tail, which, instead of containing twelve feathers, may have many more,—say thirty, or even more—I believe there are some with as many as forty-two. This bird has a curious habit of spreading out the feathers of its tail in such a way that they reach forward, and touch its head; and if this can be accomplished, I believe it is looked upon as a point of great beauty.

But here is the last great variety,—the Tumbler; and of that great variety, one of the principal kinds, and one most prized, is the specimen represented here—the short-faced Tumbler. Its beak, you see, is reduced to a mere nothing. Just compare the beak of this one and that of the first one, the Carrier—I believe the orthodox comparison of the head and beak of a thoroughly well-bred Tumbler is to stick an oat into a cherry, and that will give you the proper relative proportions of the beak and head. The feet and legs are exceedingly small, and the bird appears to be quite a dwarf when placed side by side with this great Carrier.

These are differences enough in regard to their external appearance; but these differences are by no means the whole or even the most important of the differences which obtain between these birds. There is hardly a single point of their structure which has not become more or less altered; and to give you an idea of how extensive these alterations are, I have here some very good skeletons, for which I am indebted to my friend Mr. Tegetmeier, a great authority in these matters; by means of which, if you examine them by-and-by, you will be able to see the enormous difference in their bony structures.

I had the privilege, some time ago, of access to some important MSS. of Mr. Darwin, who, I may tell you, has taken very great pains and spent much valuable time and attention on the investigation of these variations, and getting together all the facts that bear upon them. I obtained from these MSS. the following summary of the differences between the domestic breeds of pigeons; that is to say, a notification of the various points in which their organization differs. In the first place, the back of the skull may differ a good deal, and the development of the bones of the face may vary a great deal; the back varies a good deal; the shape of the lower jaw varies; the tongue varies very greatly, not only in correlation to the length and size of the beak, but it seems also to have a kind of independent variation of its own. Then the amount of naked skin round the eyes, and at the base of the beak, may vary enormously; so may the length of the eyelids, the shape of the nostrils, and the length of the neck. I have already noticed the habit of blowing out the gullet, so remarkable in the Pouter, and comparatively so in the others. There are great differences, too, in the size of the female and the male, the shape of the body, the number and width of the processes of the ribs, the development of the ribs, and the size, shape, and development of the breastbone. We may notice, too,—and I mention the fact because it has been disputed by what is assumed to be high authority,—the variation in the number of the sacral vertebræ. The number of these varies from eleven to fourteen, and that without any diminution in the number of the vertebræ of the back or of the tail. Then the number and position of the tail-feathers may vary enormously, and so may the number of the primary and secondary feathers of the wings. Again, the length of the feet and of the beak,—although they have no relation to each other, yet appear to go together,—that is, you have a long beak wherever you have long feet. There are differences also in the periods of the acquirement of the perfect plumage,—the size and shape of the eggs,—the nature of flight, and the powers of flight,—so-called “homing” birds having enormous flying powers;[52] while, on the other hand, the little Tumbler is so called because of its extraordinary faculty of turning head over heels in the air, instead of pursuing a distinct course. And, lastly, the dispositions and voices of the birds may vary. Thus the case of the pigeons shows you that there is hardly a single particular,—whether of instinct, or habit, or bony structure, or of plumage,—of either the internal economy or the external shape, in which some variation or change may not take place, which, by selective breeding, may become perpetuated, and form the foundation of, and give rise to, a new race.

If you carry in your mind’s eye these four varieties of pigeons, you will bear with you as good a notion as you can have, perhaps, of the enormous extent to which a deviation from a primitive type may be carried by means of this process of selective breeding.

VIII

THE CONDITIONS OF EXISTENCE AS AFFECTING THE PERPETUATION OF LIVING BEINGS.

In the last Lecture I endeavoured to prove to you that, while, as a general rule, organic beings tend to reproduce their kind, there is in them, also, a constantly recurring tendency to vary—to vary to a greater or to a less extent. Such a variety, I pointed out to you, might arise from causes which we do not understand; we therefore called it spontaneous; and it might come into existence as a definite and marked thing, without any gradations between itself and the form which preceded it. I further pointed out, that such a variety having once arisen, might be perpetuated to some extent, and indeed to a very marked extent, without any direct interference, or without any exercise of that process which we called selection. And then I stated further, that by such selection, when exercised artificially—if you took care to breed only from those forms which presented the same peculiarities of any variety which had arisen in this manner—the variation might be perpetuated, as far as we can see, indefinitely.

The next question, and it is an important one for us, is this: Is there any limit to the amount of variation from the primitive stock which can be produced by this process of selective breeding? In considering this question, it will be useful to class the characteristics, in respect of which organic beings vary, under two heads: we may consider structural characteristics, and we may consider physiological characteristics.

In the first place, as regards structural characteristics, I endeavoured to show you, by the skeletons which I had upon the table, and by reference to a great many well-ascertained facts, that the different breeds of Pigeons, the Carriers, Pouters, and Tumblers, might vary in any of their internal and important structural characters to a very great degree; not only might there be changes in the proportions of the skull, and the characters of the feet and beaks, and so on; but that there might be an absolute difference in the number of the vertebræ of the back, as in the sacral vertebræ of the Pouter; and so great is the extent of the variation in these and similar characters that I pointed out to you, by reference to the skeletons and the diagrams, that these extreme varieties may absolutely differ more from one another in their structural characters than do what naturalists call distinct Species of pigeons; that is to say, that they differ so much in structure that there is a greater difference between the Pouter and the Tumbler than there is between such wild and distinct forms as the Rock Pigeon or the Ring Pigeon, or the Ring Pigeon and the Stock Dove; and indeed the differences are of greater value than this, for the structural differences between these domesticated pigeons are such as would be admitted by a naturalist, supposing he knew nothing at all about their origin, to entitle them to constitute even distinct genera.

As I have used this term Species, and shall probably use it a good deal, I had better perhaps devote a word or two to explaining what I mean by it.

Animals and plants are divided into groups, which become gradually smaller, beginning with a Kingdom, which is divided into Sub-Kingdoms; then come the smaller divisions called Provinces; and so on from a Province to a Class, from a Class to an Order, from Orders to Families, and from these to Genera, until we come at length to the smallest groups of animals which can be defined one from the other by constant characters, which are not sexual; and these are what naturalists call Species in practice, whatever they may do in theory.

If in a state of nature you find any two groups of living beings, which are separated one from the other by some constantly-recurring characteristic, I don’t care how slight and trivial, so long as it is defined and constant, and does not depend on sexual peculiarities, then all naturalists agree in calling them two species; that is what is meant by the use of the word species—that is to say, it is, for the practical naturalist, a mere question of structural differences.[53]

We have seen now—to repeat this point once more, and it is very essential that we should rightly understand it—we have seen that breeds, known to have been derived from a common stock by selection, may be as different in their structure from the original stock as species may be distinct from each other.

But is the like true of the physiological characteristics of animals? Do the physiological differences of varieties amount in degree to those observed between forms which naturalists call distinct species? This is a most important point for us to consider.

As regards the great majority of physiological characteristics, there is no doubt that they are capable of being developed, increased, and modified by selection.

There is no doubt that breeds may be made as different as species in many physiological characters. I have already pointed out to you very briefly the different habits of the breeds of Pigeons, all of which depend upon their physiological peculiarities,—as the peculiar habit of tumbling, in the Tumbler,—the peculiarities of flight, in the “homing” birds,—the strange habit of spreading out the tail, and walking in a peculiar fashion, in the Fantail,—and, lastly, the habit of blowing out the gullet, so characteristic of the Pouter. These are all due to physiological modifications, and in all these respects these birds differ as much from each other as any two ordinary species do.

So with Dogs in their habits and instincts. It is a physiological peculiarity which leads the Greyhound to chase its prey by sight,—that enables the Beagle to track it by the scent,—that impels the Terrier to its rat-hunting propensity,—and that leads the Retriever to its habits of retrieving. These habits and instincts are all the results of physiological differences and peculiarities, which have been developed from a common stock, at least there is every reason to believe so. But it is a most singular circumstance, that while you may run through almost the whole series of physiological processes, without finding a check to your argument, you come at last to a point where you do find a check, and that is in the reproductive processes. For there is a most singular circumstance in respect to natural species—at least about some of them—and it would be sufficient for the purposes of this argument, if it were true of only one of them, but there is, in fact, a great number of such cases—and that is, that similar as they may appear to be to mere races or breeds, they present a marked peculiarity in the reproductive process. If you breed from the male and female of the same race, you of course have offspring of the like kind, and if you make the offspring breed together, you obtain the same result, and if you breed from these again, you will still have the same kind of offspring; there is no check. But if you take members of two distinct species, however similar they may be to each other, and make them breed together, you will find a check, with some modifications and exceptions, however, which I shall speak of presently. If you cross two such species with each other, then,—although you may get offspring in the case of the first cross, yet, if you attempt to breed from the products of that crossing, which are what are called Hybrids—that is, if you couple a male and a female hybrid—then the result is that in ninety-nine cases out of a hundred you will get no offspring at all: there will be no result whatsoever.

The reason of this is quite obvious in some cases; the male hybrids, although possessing all the external appearances and characteristics of perfect animals, are physiologically imperfect and deficient in the structural parts of the reproductive elements necessary to generation. It is said to be invariably the case with the male mule, the cross between the Ass and the Mare; and hence it is, that, although crossing the Horse with the Ass is easy enough, and is constantly done, as far as I am aware, if you take two mules, a male and a female, and endeavour to breed from them, you get no offspring whatever; no generation will take place. This is what is called the sterility of the hybrids between two distinct species.

You see that this is a very extraordinary circumstance; one does not see why it should be. The common teleological explanation is, that it is to prevent the impurity of the blood resulting from the crossing of one species with another, but you see it does not in reality do anything of the kind. There is nothing in this fact that hybrids cannot breed with each other, to establish such a theory; there is nothing to prevent the Horse breeding with the Ass, or the Ass with the Horse. So that this explanation breaks down, as a great many explanations of this kind do, that are only founded on mere assumptions.

Thus you see that there is a great difference between “mongrels,” which are crosses between distinct races, and “hybrids,” which are crosses between distinct species. The mongrels are, so far as we know, fertile with one another. But between species, in many cases, you cannot succeed in obtaining even the first cross: at any rate it is quite certain that the hybrids are often absolutely infertile one with another.

Here is a feature, then, great or small as it may be, which distinguishes natural species of animals. Can we find any approximation to this in the different races known to be produced by selective breeding from a common stock? Up to the present time the answer to that question is absolutely a negative one. As far as we know at present, there is nothing approximating to this check. In crossing the breeds between the Fantail and the Pouter, the Carrier and the Tumbler, or any other variety or race you may name—so far as we know at present—there is no difficulty in breeding together the mongrels. Take the Carrier and the Fantail, for instance, and let them represent the Horse and the Ass in the case of distinct species; then you have, as the result of their breeding, the Carrier-Fantail mongrel,—we will say the male and female mongrel,—and, as far as we know, these two when crossed would not be less fertile than the original cross, or than Carrier with Carrier. Here, you see, is a physiological contrast between the races produced by selective modification and natural species. I shall inquire into the value of this fact, and of some modifying circumstances by and by; for the present I merely put it broadly before you.

But while considering this question of the limitations of species, a word must be said about what is called Recurrence—the tendency of races which have been developed by selective breeding from varieties to return to their primitive type. This is supposed by many to put an absolute limit to the extent of selective and all other variations. People say, “It is all very well to talk about producing these different races, but you know very well that if you turned all these birds wild, these Pouters, and Carriers, and so on, they would all return to their primitive stock.” This is very commonly assumed to be a fact, and it is an argument that is commonly brought forward as conclusive; but if you will take the trouble to inquire into it rather closely, I think you will find that it is not worth very much. The first question of course is, Do they thus return to the primitive stock? And commonly as the thing is assumed and accepted, it is extremely difficult to get anything like good evidence of it. It is constantly said, for example, that if domesticated Horses are turned wild, as they have been in some parts of Asia Minor and South America, that they return at once to the primitive stock from which they were bred. But the first answer that you make to this assumption is, to ask who knows what the primitive stock was; and the second answer is, that in that case the wild Horses of Asia Minor ought to be exactly like the wild Horses of South America. If they are both like the same thing, they ought manifestly to be like each other! The best authorities, however, tell you that it is quite different. The wild Horse of Asia is said to be of a dun colour, with a largish head, and a great many other peculiarities; while the best authorities on the wild Horses of South America tell you that there is no similarity between their wild Horses and those of Asia Minor; the cut of their heads is very different, and they are commonly chestnut or bay-coloured. It is quite clear, therefore, that as by these facts there ought to have been two primitive stocks, they go for nothing in support of the assumption that races recur to one primitive stock, and so far as this evidence is concerned, it falls to the ground.

Suppose for a moment that it were so, and that domesticated races, when turned wild, did return to some common condition, I cannot see that this would prove much more than that similar conditions are likely to produce similar results; and that when you take back domesticated animals into what we call natural conditions, you do exactly the same thing as if you carefully undid all the work you had gone through, for the purpose of bringing the animal from its wild to its domesticated state. I do not see anything very wonderful in the fact, if it took all that trouble to get it from a wild state, that it should go back into its original state as soon as you removed the conditions which produced the variation to the domesticated form. There is an important fact, however, forcibly brought forward by Mr. Darwin, which has been noticed in connection with the breeding of domesticated pigeons; and it is, that however different these breeds of pigeons may be from each other, and we have already noticed the great differences in these breeds, that if, among any of those variations, you chance to have a blue pigeon turn up, it will be sure to have the black bars across the wings, which are characteristic of the original wild stock, the Rock Pigeon.

Now, this is certainly a very remarkable circumstance; but I do not see myself how it tells very strongly either one way or the other. I think, in fact, that this argument in favour of recurrence to the primitive type might prove a great deal too much for those who so constantly bring it forward. For example, Mr. Darwin has very forcibly urged, that nothing is commoner than if you examine a dun horse—and I had an opportunity of verifying this illustration lately, while in the islands of the West Highlands, where there are a great many dun horses—to find that horse exhibit a long black stripe down his back, very often stripes on his shoulder, and very often stripes on his legs. I, myself, saw a pony of this description a short time ago, in a baker’s cart, near Rothesay, in Bute: it had the long stripe down the back, and stripes on the shoulders and legs, just like those of the Ass, the Quagga, and the Zebra. Now, if we interpret the theory of recurrence as applied to this case, might it not be said that here was a case of a variation exhibiting the characters and conditions of an animal occupying something like an intermediate position between the Horse, the Ass, the Quagga, and the Zebra, and from which these had been developed? In the same way with regard even to Man. Every anatomist will tell you that there is nothing commoner, in dissecting the human body, than to meet with what are called muscular variations—that is, if you dissect two bodies very carefully, you will probably find that the modes of attachment and insertion of the muscles are not exactly the same in both, there being great peculiarities in the mode in which the muscles are arranged; and it is very singular, that in some dissections of the human body you will come upon arrangements of the muscles very similar indeed to the same parts in the Apes. Is the conclusion in that case to be, that this is like the black bars in the case of the Pigeon, and that it indicates a recurrence to the primitive type from which the animals have been probably developed? Truly, I think that the opponents of modification and variation had better leave the argument of recurrence alone, or it may prove altogether too strong for them.

To sum up,—the evidence as far as we have gone is against the argument as to any limit to divergences, so far as structure is concerned; and in favour of a physiological limitation. By selective breeding we can produce structural divergences as great as those of species, but we cannot produce equal physiological divergences. For the present I leave the question there.

Now, the next problem that lies before us—and it is an extremely important one—is this: Does this selective breeding occur in nature? Because, if there is no proof of it, all that I have been telling you goes for nothing in accounting for the origin of species. Are natural causes competent to play the part of selection in perpetuating varieties? Here we labour under very great difficulties. In the last lecture I had occasion to point out to you the extreme difficulty of obtaining evidence even of the first origin of those varieties which we know to have occurred in domesticated animals. I told you, that almost always the origin of these varieties is overlooked, so that I could only produce two or three cases, as that of Gratio Kelleia and of the Ancon sheep. People forget, or do not take notice of them until they come to have a prominence; and if that is true of artificial cases, under our own eyes, and in animals in our own care, how much more difficult it must be to have at first hand good evidence of the origin of varieties in nature! Indeed, I do not know that it is possible by direct evidence to prove the origin of a variety in nature, or to prove selective breeding; but I will tell you what we can prove—and this comes to the same thing—that varieties exist in nature within the limits of species, and, what is more, that when a variety has come into existence in nature, there are natural causes and conditions, which are amply competent to play the part of a selective breeder; and although that is not quite the evidence that one would like to have—though it is not direct testimony—yet it is exceeding good and exceedingly powerful evidence in its way.

As to the first point, of varieties existing among natural species, I might appeal to the universal experience of every naturalist, and of any person who has ever turned any attention at all to the characteristics of plants and animals in a state of nature; but I may as well take a few definite cases, and I will begin with Man himself.

I am one of those who believe that, at present, there is no evidence whatever for saying, that mankind sprang originally from any more than a single pair; I must say, that I cannot see any good ground whatever, or even any tenable sort of evidence, for believing that there is more than one species of Man. Nevertheless, as you know, just as there are numbers of varieties in animals, so there are remarkable varieties of men. I speak not merely of those broad and distinct variations which you see at a glance. Everybody, of course, knows the difference between a Negro and a white man, and can tell a Chinaman from an Englishman. They each have peculiar characteristics of colour and physiognomy; but you must recollect that the characters of these races go very far deeper—they extend to the bony structure, and to the characters of that most important of all organs to us—the brain; so that, among men belonging to different races, or even within the same race, one man shall have a brain a third, or half, or even seventy per cent bigger than another; and if you take the whole range of human brains, you will find a variation in some cases of a hundred per cent. Apart from these variations in the size of the brain, the characters of the skull vary. Thus if I draw the figures of a Mongul and of a Negro head on the blackboard, in the case of the last the breadth would be about seven-tenths, and in the other it would be nine-tenths of the total length. So that you see there is abundant evidence of variation among men in their natural condition. And if you turn to other animals there is just the same thing. The fox, for example, which has a very large geographical distribution all over Europe, and parts of Asia, and on the American Continent, varies greatly. There are mostly large foxes in the North, and smaller ones in the South. In Germany alone, the foresters reckon some eight different sorts.

Of the tiger, no one supposes that there is more than one species; they extend from the hottest parts of Bengal, into the dry, cold, bitter steppes of Siberia, into a latitude of 50°,—so that they may even prey upon the reindeer. These tigers have exceedingly different characteristics, but still they all keep their general features, so that there is no doubt as to their being tigers. The Siberian tiger has a thick fur, a small mane, and a longitudinal stripe down the back, while the tigers of Java and Sumatra differ in many important respects from the tigers of Northern Asia. So lions vary; so birds vary; and so, if you go further back and lower down in creation, you find that fishes vary. In different streams, in the same country even, you will find the trout to be quite different to each other and easily recognizable by those who fish in the particular streams. There is the same differences in leeches; leech collectors can easily point out to you the differences and the peculiarities which you yourself would probably pass by; so with fresh-water mussels; so, in fact, with every animal you can mention.

In plants there is the same kind of variation. Take such a case even as the common bramble. The botanists are all at war about it; some of them wanting to make out that there are many species of it, and others maintaining that they are but many varieties of one species; and they cannot settle to this day which is a species and which is a variety!

So that there can be no doubt whatsoever that any plant and any animal may vary in nature; that varieties may arise in the way I have described,—as spontaneous varieties,—and that those varieties may be perpetuated in the same way that I have shown you spontaneous varieties are perpetuated; I say, therefore, that there can be no doubt as to the origin and perpetuation of varieties in nature.

But the question now is:—Does selection take place in nature? is there anything like the operation of man in exercising selective breeding, taking place in nature? You will observe that, at present, I say nothing about species; I wish to confine myself to the consideration of the production of those natural races which everybody admits to exist. The question is, whether in nature there are causes competent to produce races, just in the same way as man is able to produce, by selection, such races of animals as we have already noticed.

When a variety has arisen, the Conditions of Existence are such as to exercise an influence which is exactly comparable to that of artificial selection. By Conditions of Existence I mean two things,—there are conditions which are furnished by the physical, the inorganic world, and there are conditions of existence which are furnished by the organic world. There is, in the first place, Climate; under that head I include only temperature and the varied amount of moisture of particular places. In the next place there is what is technically called Station, which means—given the climate, the particular kind of place in which an animal or a plant lives or grows; for example, the station of a fish is in the water, of a fresh-water fish in fresh water; the station of a marine fish is in the sea, and a marine animal may have a station higher or deeper. So again with land animals: the differences in their stations are those of different soils and neighbourhoods; some being best adapted to a calcareous, and others to an arenaceous soil. The third condition of existence is Food, by which I mean food in the broadest sense, the supply of the materials necessary to the existence of an organic being; in the case of a plant the inorganic matters, such as carbonic acid, water, ammonia, and the earthy salts or salines; in the case of the animal the inorganic and organic matters, which we have seen they require; then these are all, at least the two first, what we may call the inorganic or physical conditions of existence. Food takes a mid-place, and then come the organic conditions; by which I mean the conditions which depend upon the state of the rest of the organic creation, upon the number and kind of living beings, with which an animal is surrounded. You may class these under two heads: there are organic beings, which operate as opponents, and there are organic beings which operate as helpers to any given organic creature. The opponents may be of two kinds: there are the indirect opponents, which are what we may call rivals; and there are the direct opponents, those which strive to destroy the creature; and these we call enemies. By rivals I mean, of course, in the case of plants, those which require for their support the same kind of soil and station, and, among animals, those which require the same kind of station, or food, or climate; those are the indirect opponents; the direct opponents are, of course, those which prey upon an animal or vegetable. The helpers may also be regarded as direct and indirect: in the case of a carnivorous animal, for example, a particular herbaceous plant may in multiplying be an indirect helper, by enabling the herbivora on which the carnivore preys to get more food, and thus to nourish the carnivore more abundantly; the direct helper may be best illustrated by reference to some parasitic creature, such as the tape-worm. The tape-worm exists in the human intestines, so that the fewer there are of men the fewer there will be of tape-worms, other things being alike. It is a humiliating reflection, perhaps, that we may be classed as direct helpers to the tape-worm, but the fact is so: we can all see that if there were no men there would be no tape-worms.

It is extremely difficult to estimate, in a proper way, the importance and the working of the Conditions of Existence. I do not think there were any of us who had the remotest notion of properly estimating them until the publication of Mr. Darwin’s work, which has placed them before us with remarkable clearness; and I must endeavour, as far as I can in my own fashion, to give you some notion of how they work. We shall find it easiest to take a simple case, and one as free as possible from every kind of complication.

I will suppose, therefore, that all the habitable part of this globe—the dry land, amounting to about 51,000,000 square miles,—I will suppose that the whole of that dry land has the same climate, and that it is composed of the same kind of rock or soil, so that there will be the same station everywhere; we thus get rid of the peculiar influence of different climates and stations. I will then imagine that there shall be but one organic being in the world, and that shall be a plant. In this we start fair. Its food is to be carbonic acid, water and ammonia, and the saline matters in the soil, which are, by the supposition, everywhere alike. We take one single plant, with no opponents, no helpers, and no rivals; it is to be a “fair field, and no favour.” Now, I will ask you to imagine further that it shall be a plant which shall produce every year fifty seeds, which is a very moderate number for a plant to produce; and that, by the action of the winds and currents, these seeds shall be equally and gradually distributed over the whole surface of the land. I want you now to trace out what will occur, and you will observe that I am not talking fallaciously any more than a mathematician does when he expounds his problem. If you show that the conditions of your problem are such as may actually occur in nature and do not transgress any of the known laws of nature in working out your proposition, then you are as safe in the conclusion you arrive at as is the mathematician in arriving at the solution of his problem. In science, the only way of getting rid of the complications with which a subject of this kind is environed, is to work in this deductive method. What will be the result, then? I will suppose that every plant requires one square foot of ground to live upon; and the result will be that, in the course of nine years, the plant will have occupied every single available spot in the whole globe! I have chalked upon the blackboard the figures by which I arrive at the result:—