So much, then, by way of proof that the method of establishing laws in science is exactly the same as that pursued in common life. Let us now turn to another matter (though really it is but another phase of the same question), and that is, the method by which, from the relations of certain phenomena, we prove that some stand in the position of causes towards the others.
I want to put the case clearly before you, and I will therefore show you what I mean by another familiar example. I will suppose that one of you, on coming down in the morning to the parlour of your house, finds that a tea-pot and some spoons which had been left in the room on the previous evening are gone,--the window is open, and you observe the mark of a dirty hand on the window-frame, and perhaps, in addition to that, you notice the impress of a hob-nailed shoe on the gravel outside. All these phenomena have struck your attention instantly, and before two seconds have passed you say, "Oh, somebody has broken open the window, entered the room, and run off with the spoons and the tea-pot!" That speech is out of your mouth in a moment. And you will probably add, "I know there has; I am quite sure of it!" You mean to say exactly what you know; but in reality you are giving expression to what is, in all essential particulars, an hypothesis. You do not know it at all; it is nothing but an hypothesis rapidly framed in your own mind. And it is an hypothesis founded on a long train of inductions and deductions.
What are those inductions and deductions, and how have you got at this hypothesis? You have observed, in the first place, that the window is open; but by a train of reasoning involving many inductions and deductions, you have probably arrived long before at the general law--and a very good one it is--that windows do not open of themselves; and you therefore conclude that something has opened the window. A second general law that you have arrived at in the same way is, that tea-pots and spoons do not go out of a window spontaneously, and you are satisfied that, as they are not now where you left them, they have been removed. In the third place, you look at the marks on the window-sill, and the shoe-marks outside, and you say that in all previous experience the former kind of mark has never been produced by anything else but the hand of a human being; and the same experience shows that no other animal but man at present wears shoes with hob-nails in them such as would produce the marks in the gravel. I do not know, even if we could discover any of those "missing links" that are talked about, that they would help us to any other conclusion! At any rate the law which states our present experience is strong enough for my present purpose. You next reach the conclusion, that as these kinds of marks have not been left by any other animals than men, or are liable to be formed in any other way than by a man's hand and shoe, the marks in question have been formed by a man in that way. You have, further, a general law, founded on observation and experience, and that, too, is, I am sorry to say, a very universal and unimpeachable one,--that some men are thieves; and you assume at once from all these premisses--and that is what constitutes your hypothesis--that the man who made the marks outside and on the window-sill, opened the window, got into the room, and stole your tea-pot and spoons. You have now arrived at a vera causa;--you have assumed a cause which, it is plain, is competent to produce all the phenomena you have observed. You can explain all these phenomena only by the hypothesis of a thief. But that is a hypothetical conclusion, of the justice of which you have no absolute proof at all; it is only rendered highly probable by a series of inductive and deductive reasonings.
I suppose your first action, assuming that you are a man of ordinary common sense, and that you have established this hypothesis to your own satisfaction, will very likely be to go off for the police, and set them on the track of the burglar, with the view to the recovery of your property. But just as you are starting with this object, some person comes in, and on learning what you are about, says, "My good friend, you are going on a great deal too fast. How do you know that the man who really made the marks took the spoons? It might have been a monkey that took them, and the man may have merely looked in afterwards." You would probably reply, "Well, that is all very well, but you see it is contrary to all experience of the way tea-pots and spoons are abstracted; so that, at any rate, your hypothesis is less probable than mine." While you are talking the thing over in this way, another friend arrives, one of that good kind of people that I was talking of a little while ago. And he might say, "Oh, my dear sir, you are certainly going on a great deal too fast. You are most presumptuous. You admit that all these occurrences took place when you were fast asleep, at a time when you could not possibly have known anything about what was taking place. How do you know that the laws of Nature are not suspended during the night? It may be that there has been some kind of supernatural interference in this case." In point of fact, he declares that your hypothesis is one of which you cannot at all demonstrate the truth, and that you are by no means sure that the laws of Nature are the same when you are asleep as when you are awake.
Well, now, you cannot at the moment answer that kind of reasoning. You feel that your worthy friend has you somewhat at a disadvantage. You will feel perfectly convinced in your own mind, however, that you are quite right, and you say to him, "My good friend, I can only be guided by the natural probabilities of the case, and if you will be kind enough to stand aside and permit me to pass, I will go and fetch the police." Well, we will suppose that your journey is successful, and that by good luck you meet with a policeman; that eventually the burglar is found with your property on his person, and the marks correspond to his hand and to his boots. Probably any jury would consider those facts a very good experimental verification of your hypothesis, touching the cause of the abnormal phenomena observed in your parlour, and would act accordingly.
Now, in this suppositious case, I have taken phenomena of a very common kind, in order that you might see what are the different steps in an ordinary process of reasoning, if you will only take the trouble to analyse it carefully. All the operations I have described, you will see, are involved in the mind of any man of sense in leading him to a conclusion as to the course he should take in order to make good a robbery and punish the offender. I say that you are led, in that case, to your conclusion by exactly the same train of reasoning as that which a man of science pursues when he is endeavouring to discover the origin and laws of the most occult phenomena. The process is, and always must be, the same; and precisely the same mode of reasoning was employed by Newton and Laplace in their endeavours to discover and define the causes of the movements of the heavenly bodies, as you, with your own common sense, would employ to detect a burglar. The only difference is, that the nature of the inquiry being more abstruse, every step has to be most carefully watched, so that there may not be a single crack or flaw in your hypothesis. A flaw or crack in many of the hypotheses of daily life may be of little or no moment as affecting the general correctness of the conclusions at which we may arrive; but, in a scientific inquiry, a fallacy, great or small, is always of importance, and is sure to be in the long run constantly productive of mischievous, if not fatal results.
Do not allow yourselves to be misled by the common notion that an hypothesis is untrustworthy simply because it is an hypothesis. It is often urged, in respect to some scientific conclusion, that, after all, it is only an hypothesis. But what more have we to guide us in nine-tenths of the most important affairs of daily life than hypotheses, and often very ill-based ones? So that in science, where the evidence of an hypothesis is subjected to the most rigid examination, we may rightly pursue the same course. You may have hypotheses and hypotheses. A man may say, if he likes, that the moon is made of green cheese: that is an hypothesis. But another man, who has devoted a great deal of time and attention to the subject, and availed himself of the most powerful telescopes and the results of the observations of others, declares that in his opinion it is probably composed of materials very similar to those of which our own earth is made up: and that is also only an hypothesis. But I need not tell you that there is an enormous difference in the value of the two hypotheses. That one which is based on sound scientific knowledge is sure to have a corresponding value; and that which is a mere hasty random guess is likely to have but little value. Every great step in our progress in discovering causes has been made in exactly the same way as that which I have detailed to you. A person observing the occurrence of certain facts and phenomena asks, naturally enough, what process, what kind of operation known to occur in Nature applied to the particular case, will unravel and explain the mystery? Hence you have the scientific hypothesis; and its value will be proportionate to the care and completeness with which its basis had been tested and verified. It is in these matters as in the commonest affairs of practical life: the guess of the fool will be folly, while the guess of the wise man will contain wisdom. In all cases, you see that the value of the result depends on the patience and faithfulness with which the investigator applies to his hypothesis every possible kind of verification.
I dare say I may have to return to this point by and by; but having dealt thus far with our logical methods, I must now turn to something which, perhaps, you may consider more interesting, or, at any rate, more tangible. But in reality there are but few things that can be more important for you to understand than the mental processes and the means by which we obtain scientific conclusions and theories. [*] Having granted that the inquiry is a proper one, and having determined on the nature of the methods we are to pursue and which only can lead to success, I must now turn to the consideration of our knowledge of the nature of the processes which have resulted in the present condition of organic nature.
- Those who wish to study fully the doctrines of which I have endeavoured to give some rough-and-ready illustrations, must read Mr. John Stuart Mill's System of Logic.
Here, let me say at once, lest some of you misunderstand me, that I have extremely little to report. The question of how the present condition of organic nature came about, resolves itself into two questions. The first is: How has organic or living matter commenced its existence? And the second is: How has it been perpetuated? On the second question I shall have more to say hereafter. But on the first one, what I now have to say will be for the most part of a negative character.
If you consider what kind of evidence we can have upon this matter, it will resolve itself into two kinds. We may have historical evidence and we may have experimental evidence. It is, for example, conceivable, that inasmuch as the hardened mud which forms a considerable portion of the thickness of the earth's crust contains faithful records of the past forms of life, and inasmuch as these differ more and more as we go further down,--it is possible and conceivable that we might come to some particular bed or stratum which should contain the remains of those creatures with which organic life began upon the earth. And if we did so, and if such forms of organic life were preservable, we should have what I would call historical evidence of the mode in which organic life began upon this planet. Many persons will tell you, and indeed you will find it stated in many works on geology, that this has been done, and that we really possess such a record; there are some who imagine that the earliest forms of life of which we have as yet discovered any record, are in truth the forms in which animal life began upon the globe. The grounds on which they base that supposition are these:--That if you go through the enormous thickness of the earth's crust and get down to the older rocks, the higher vertebrate animals--the quadrupeds, birds, and fishes--cease to be found; beneath them you find only the invertebrate animals; and in the deepest and lowest rocks those remains become scantier and scantier, not in any very gradual progression, however, until, at length, in what are supposed to be the oldest rocks, the animal remains which are found are almost always confined to four forms--Oldhamia, whose precise nature is not known, whether plant or animal; Lingula, a kind of mollusc; Trilobites, a crustacean animal, having the same essential plan of construction, though differing in many details from a lobster or crab; and Hymenocaris, which is also a crustacean. So that you have all the Fauna reduced, at this period, to four forms: one a kind of animal or plant that we know nothing about, and three undoubted animals--two crustaceans and one mollusc.
I think, considering the organisation of these mollusca and crustacea, and looking at their very complex nature, that it does indeed require a very strong imagination to conceive that these were the first created of all living things. And you must take into consideration the fact that we have not the slightest proof that these which we call the oldest beds are really so: I repeat, we have not the slightest proof of it. When you find in some places that in an enormous thickness of rocks there are but very scanty traces of life, or absolutely none at all; and that in other parts of the world rocks of the very same formation are crowded with the records of living forms, I think it is impossible to place any reliance on the supposition, or to feel one's self justified in supposing that these are the forms in which life first commenced. I have not time here to enter upon the technical grounds upon which I am led to this conclusion,--that could hardly be done properly in half a dozen lectures on that part alone:--I must content myself with saying that I do not at all believe that these are the oldest forms of life.
I turn to the experimental side to see what evidence we have there. To enable us to say that we know anything about the experimental origination of organisation and life, the investigator ought to be able to take inorganic matters, such as carbonic acid, ammonia, water, and salines, in any sort of inorganic combination, and be able to build them up into protein matter, and then that protein matter ought to begin to live in an organic form. That, nobody has done as yet, and I suspect it will be a long while before anybody does do it. But the thing is by no means so impossible as it looks; for the researches of modern chemistry have shown us--I won't say the road towards it, but, if I may so say, they have shown the finger-post pointing to the road that may lead to it.
It is not many years ago--and you must recollect that Organic Chemistry is a young science, not above a couple of generations old, you must not expect too much of it,--it is not many years ago since it was said to be perfectly impossible to fabricate any organic compound; that is to say, any non-mineral compound which is to be found in an organised being. It remained so for a very long period; but it is now a considerable number of years since a distinguished foreign chemist contrived to fabricate urea, a substance of a very complex character, which forms one of the waste products of animal structures. And of late years a number of other compounds, such as butyric acid, and others, have been added to the list. I need not tell you that chemistry is an enormous distance from the goal I indicate; all I wish to point out to you is, that it is by no means safe to say that that goal may not be reached one day. It may be that it is impossible for us to produce the conditions requisite to the origination of life; but we must speak modestly about the matter, and recollect that Science has put her foot upon the bottom round of the ladder. Truly he would be a bold man who would venture to predict where she will be fifty years hence.
There is another inquiry which bears indirectly upon this question, and upon which I must say a few words. You are all of you aware of the phenomena of what is called spontaneous generation. Our forefathers, down to the seventeenth century, or thereabouts, all imagined, in perfectly good faith, that certain vegetable and animal forms gave birth, in the process of their decomposition, to insect life. Thus, if you put a piece of meat in the sun, and allowed it to putrefy, they conceived that the grubs which soon began to appear were the result of the action of a power of spontaneous generation which the meat contained. And they could give you receipts for making various animal and vegetable preparations which would produce particular kinds of animals. A very distinguished Italian naturalist, named Redi, took up the question, at a time when everybody believed in it; among others our own great Harvey, the discoverer of the circulation of the blood. You will constantly find his name quoted, however, as an opponent of the doctrine of spontaneous generation; but the fact is, and you will see it if you will take the trouble to look into his works, Harvey believed it as profoundly as any man of his time; but he happened to enunciate a very curious proposition--that every living thing came from an egg; he did not mean to use the word in the sense in which we now employ it, he only meant to say that every living thing originated in a little rounded particle of organised substance; and it is from this circumstance, probably, that the notion of Harvey having opposed the doctrine originated. Then came Redi, and he proceeded to upset the doctrine in a very simple manner. He merely covered the piece of meat with some very fine gauze, and then he exposed it to the same conditions. The result of this was that no grubs or insects were produced; he proved that the grubs originated from the insects who came and deposited their eggs in the meat, and that they were hatched by the heat of the sun. By this kind of inquiry he thoroughly upset the doctrine of spontaneous generation, for his time at least.
Then came the discovery and application of the microscope to scientific inquiries, which showed to naturalists that besides the organisms which they already knew as living beings and plants, there were an immense number of minute things which could be obtained apparently almost at will from decaying vegetable and animal forms. Thus, if you took some ordinary black pepper or some hay, and steeped it in water, you would find in the course of a few days that the water had become impregnated with an immense number of animalcules swimming about in all directions. From facts of this kind naturalists were led to revive the theory of spontaneous generation. They were headed here by an English naturalist,--Needham,--and afterwards in France by the learned Buffon. They said that these things were absolutely begotten in the water of the decaying substances out of which the infusion was made. It did not matter whether you took animal or vegetable matter, you had only to steep it in water and expose it, and you would soon have plenty of animalcules. They made an hypothesis about this which was a very fair one. They said, this matter of the animal world, or of the higher plants, appears to be dead, but in reality it has a sort of dim life about it, which, if it is placed under fair conditions, will cause it to break up into the forms of these little animalcules, and they will go through their lives in the same way as the animal or plant of which they once formed a part.
The question now became very hotly debated. Spallanzani, an Italian naturalist, took up opposite views to those of Needham and Buffon, and by means of certain experiments he showed that it was quite possible to stop the process by boiling the water, and closing the vessel in which it was contained. "Oh!" said his opponents; "but what do you know you may be doing when you heat the air over the water in this way? You may be destroying some property of the air requisite for the spontaneous generation of the animalcules."
However, Spallanzani's views were supposed to be upon the right side, and those of the others fell into discredit; although the fact was that Spallanzani had not made good his views. Well, then, the subject continued to be revived from time to time, and experiments were made by several persons; but these experiments were not altogether satisfactory. It was found that if you put an infusion in which animalcules would appear if it were exposed to the air into a vessel and boiled it, and then sealed up the mouth of the vessel, so that no air, save such as had been heated to 212°, could reach its contents, that then no animalcules would be found; but if you took the same vessel and exposed the infusion to the air, then you would get animalcules. Furthermore, it was found that if you connected the mouth of the vessel with a red-hot tube in such a way that the air would have to pass through the tube before reaching the infusion, that then you would get no animalcules. Yet another thing was noticed: if you took two flasks containing the same kind of infusion, and left one entirely exposed to the air, and in the mouth of the other placed a ball of cotton wool, so that the air would have to filter itself through it before reaching the infusion, that then, although you might have plenty of animalcules in the first flask, you would certainly obtain none from the second.
These experiments, you see, all tended towards one conclusion--that the infusoria were developed from little minute spores or eggs which were constantly floating in the atmosphere, and which lose their power of germination if subjected to heat. But one observer now made another experiment, which seemed to go entirely the other way, and puzzled him altogether. He took some of this boiled infusion that I have been speaking of, and by the use of a mercurial bath--a kind of trough used in laboratories--he deftly inverted a vessel containing the infusion into the mercury, so that the latter reached a little beyond the level of the mouth of the inverted vessel. You see that he thus had a quantity of the infusion shut off from any possible communication with the outer air by being inverted upon a bed of mercury.
He then prepared some pure oxygen and nitrogen gases, and passed them by means of a tube going from the outside of the vessel, up through the mercury into the infusion; so that he thus had it exposed to a perfectly pure atmosphere of the same constituents as the external air. Of course, he expected he would get no infusorial animalcules at all in that infusion; but, to his great dismay and discomfiture, he found he almost always did get them.
Furthermore, it has been found that experiments made in the manner described above answer well with most infusions; but that if you fill the vessel with boiled milk, and then stop the neck with cotton-wool, you will have infusoria. So that yousee there were two experiments that brought you to one kind of conclusion, and three to another; which was a most unsatisfactory state of things to arrive at in a scientific inquiry.
Some few years after this, the question began to be very hotly discussed in France. There was M. Pouchet, a professor at Rouen, a very learned man, but certainly not a very rigid experimentalist. He published a number of experiments of his own, some of which were very ingenious, to show that if you went to work in a proper way, there was a truth in the doctrine of spontaneous generation. Well, it was one of the most fortunate things in the world that M. Pouchet took up this question, because it induced a distinguished French chemist, M. Pasteur, to take up the question on the other side; and he has certainly worked it out in the most perfect manner. I am glad to say, too, that he has published his researches in time to enable me to give you an account of them. He verified all the experiments which I have just mentioned to you--and then finding those extraordinary anomalies, as in the case of the mercury bath and the milk, he set himself to work to discover their nature. In the case of milk he found it to be a question of temperature. Milk in a fresh state is slightly alkaline; and it is a very curious circumstance, but this very slight degree of alkalinity seems to have the effect of preserving the organisms which fall into it from the air from being destroyed at a temperature of 212°, which is the boiling point. But if you raise the temperature 10° when you boil it, the milk behaves like everything else; and if the air with which it comes in contact, after being boiled at this temperature, is passed through a red-hot tube, you will not get a trace of organisms.
He then turned his attention to the mercury bath, and found on examination that the surface of the mercury was almost always covered with a very fine dust. He found that even the mercury itself was positively full of organic matters; that from being constantly exposed to the air, it had collected an immense number of these infusorial organisms from the air. Well, under these circumstances he felt that the case was quite clear, and that the mercury was not what it had appeared to M. Schwann to be,--a bar to the admission of these organisms; but that, in reality, it acted as a reservoir from which the infusion was immediately supplied with the large quantity that had so puzzled him.
But not content with explaining the experiments of others, M. Pasteur went to work to satisfy himself completely. He said to himself: "If my view is right, and if, in point of fact, all these appearances of spontaneous generation are altogether due to the falling of minute germs suspended in the atmosphere,--why, I ought not only to be able to show the germs, but I ought to be able to catch and sow them, and produce the resulting organisms." He, accordingly, constructed a very ingenious apparatus to enable him to accomplish the trapping of the "germ dust" in the air. He fixed in the window of his room a glass tube, in the centre of which he had placed a ball of gun-cotton, which, as you all know, is ordinary cotton-wool, which, from having been steeped in strong acid, is converted into a substance of great explosive power. It is also soluble in alcohol and ether. One end of the glass tube was, of course, open to the external air; and at the other end of it he placed an aspirator, a contrivance for causing a current of the external air to pass through the tube. He kept this apparatus going for four-and-twenty hours, and then removed the dusted gun-cotton, and dissolved it in alcohol and ether. He then allowed this to stand for a few hours, and the result was, that a very fine dust was gradually deposited at the bottom of it. That dust, on being transferred to the stage of a microscope, was found to contain an enormous number of starch grains. You know that the materials of our food and the greater portion of plants are composed of starch, and we are constantly making use of it in a variety of ways, so that there is always a quantity of it suspended in the air. It is these starch grains which form many of those bright specks that we see dancing in a ray of light sometimes. But besides these, M. Pasteur found also an immense number of other organic substances such as spores of fungi, which had been floating about in the air and had got caged in this way.
He went farther, and said to himself, "If these really are the things that give rise to the appearance of spontaneous generation, I ought to be able to take a ball of this dusted gun-cotton and put it into one of my vessels, containing that boiled infusion which has been kept away from the air, and in which no infusoria are at present developed, and then, if I am right, the introduction of this gun-cotton will give rise to organisms."
Accordingly, he took one of these vessels of infusion, which had been kept eighteen months, without the least appearance of life in it, and by a most ingenious contrivance, he managed to break it open and introduce such a ball of gun-cotton, without allowing the infusion or the cotton ball to come into contact with any air but that which had been subjected to a red heat, and in twenty-four hours he had the satisfaction of finding all the indications of what had been hitherto called spontaneous generation. He had succeeded in catching the germs and developing organisms in the way ho had anticipated.
It now struck him that the truth of his conclusions might be demonstrated without all the apparatus he had employed. To do this, he took some decaying animal or vegetable substance, such as urine, which is an extremely decomposable substance, or the juice of yeast, or perhaps some other artificial preparation, and filled a vessel having a long tubular neck with it. He then boiled the liquid and bent that long neck into an S shape or zig-zag, leaving it open at the end. The infusion then gave no trace of any appearance of spontaneous generation, however long it might be left, as all the germs in the air were deposited in the beginning of the bent neck. He then cut the tube close to the vessel, and allowed the ordinary air to have free and direct access; and the result of that was the appearance of organisms in it, as soon as the infusion had been allowed to stand long enough to allow of the growth of those it received from the air, which was about forty-eight hours. The result of M. Pasteur's experiments proved, therefore, in the most conclusive manner, that all the appearances of spontaneous generation arose from nothing more than the deposition of the germs of organisms which were constantly floating in the air.
To this conclusion, however, the objection was made, that if that were the cause, then the air would contain such an enormous number of these germs, that it would be a continual fog. But M. Pasteur replied that they are not there in anything like the number we might suppose, and that an exaggerated view has been held on that subject; he showed that the chances of animal or vegetable life appearing in infusions, depend entirely on the conditions under which they are exposed. If they are exposed to the ordinary atmosphere around us, why, of course, you may have organisms appearing early. But, on the other hand, if they are exposed to air at a great height, or in some very quiet cellar, you will often not find a single trace of life.
So that M. Pasteur arrived at last at the clear and definite result, that all these appearances are like the case of the worms in the piece of meat, which was refuted by Redi, simply germs carried by the air and deposited in the liquids in which they afterwards appear. For my own part, I conceive that, with the particulars of M. Pasteur's experiments before us, we cannot fail to arrive at his conclusions; and that the doctrine of spontaneous generation has received a final coup de grâce.
You, of course, understand that all this in no way interferes with the possibility of the fabrication of organic matters by the direct method to which I have referred, remote as that possibility may be.
IV. 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 non-sexual 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 interaction of certain portions of the organisms of usually two distinct individuals,--the male and the female. The cases of non-sexual 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 a non-sexual 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 non-sexual 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 non-sexual 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 organs, 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 re-invigoration 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 non-sexual 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 non-sexually, 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 black-board 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 Atavism which 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 non-sexually; 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 non-sexual 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 recognised 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 organisations, 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 Humphrey 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 giving rise to what you may call a monstrosity; you have that monstrosity 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 earned 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 trespasser 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 organisation 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 vertebrae. The number of these varies from eleven to fourteen, and that without any diminution in the number of the vertebrae 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; [*] 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 direct 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.
- The "Carrier", I learn from Mr. Tegetmeier, does not carry; a high-bred bird of this breed being but a poor flier. The birds which fly long distances, and come home--"homing" birds-and are consequently used as carriers, are not "carriers" in the fancy sense.
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.