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 vertebrae of the back, as in the sacral vertebrae 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.3

3 I lay stress here on the ‘practical’ signification of “Species.” Whether a physiological test between species exist or not, it is hardly ever applicable by the practical naturalist.

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 habit 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 of 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 degrees — s............