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by Thomas H. Huxley
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The authors whom I quote say that they "cannot express" the excessive minuteness of the granules in question, and they estimate their diameter at less than 1/200000 of an inch. Under the highest powers of the microscope, at present applicable, such specks are hardly discernible. Nevertheless, particles of this size are massive when compared to physical molecules; whence there is no reason to doubt that each, small as it is, may have a molecular structure sufficiently complex to give rise to the phenomena of life. And, as a matter of fact, by patient watching of the place at which these infinitesimal living particles were discharged, our observers assured themselves of their growth and development into new monads. In about four hours from their being set free, they had attained a sixth of the length of the parent, with the characteristic cilia, though at first they were quite motionless; and, in four hours more, they had attained the dimensions and exhibited all the activity of the adult. These inconceivably minute particles are therefore the germs of the Heteromita; and from the dimensions of these germs it is easily shown that the body formed by conjugation may, at a low estimate, have given exit to thirty thousand of them; a result of a matrimonial process whereby the contracting parties, without a metaphor, "become one flesh," enough to make a Malthusian despair of the future of the Universe.

I am not aware that the investigators from whom I have borrowed this history have endeavoured to ascertain whether their monads take solid nutriment or not; so that though they help us very much to fill up the blanks in the history of my Heteromita, their observations throw no light on the problem we are trying to solve—Is it an animal or is it a plant?

Undoubtedly it is possible to bring forward very strong arguments in favour of regarding Heteromita as a plant.

For example, there is a Fungus, an obscure and almost microscopic mould, termed Peronospora infestans. Like many other Fungi, the Peronosporoe are parasitic upon other plants; and this particular Peronospora happens to have attained much notoriety and political importance, in a way not without a parallel in the career of notorious politicians, namely, by reason of the frightful mischief it has done to mankind. For it is this Fungus which is the cause of the potato disease; and, therefore, Peronospora infestans (doubtless of exclusively Saxon origin, though not accurately known to be so) brought about the Irish famine. The plants afflicted with the malady are found to be infested by a mould, consisting of fine tubular filaments, termed hyphoe, which burrow through the substance of the potato plant, and appropriate to themselves the substance of their host; while, at the same time, directly or indirectly, they set up chemical changes by which even its woody framework becomes blackened, sodden, and withered.

In structure, however, the Peronospora is as much a mould as the common Penicillium; and just as the Penicillium multiplies by the breaking up of its hyphoe into separate rounded bodies, the spores; so, in the Peronospora, certain of the hyphoe grow out into the air through the interstices of the superficial cells of the potato plant, and develop spores. Each of these hyphoe usually gives off several branches. The ends of the branches dilate and become closed sacs, which eventually drop off as spores. The spores falling on some part of the same potato plant, or carried by the wind to another, may at once germinate, throwing out tubular prolongations which become hyphoe, and burrow into the substance of the plant attacked. But, more commonly, the contents of the spore divide into six or eight separate portions. The coat of the spore gives way, and each portion then emerges as an independent organism, which has the shape of a bean, rather narrower at one end than the other, convex on one side, and depressed or concave on the opposite. From the depression, two long and delicate cilia proceed, one shorter than the other, and directed forwards. Close to the origin of these cilia, in the substance of the body, is a regularly pulsating, contractile vacuole. The shorter cilium vibrates actively, and effects the locomotion of the organism, while the other trails behind; the whole body rolling on its axis with its pointed end forwards.

The eminent botanist, De Bary, who was not thinking of our problem, tells us, in describing the movements of these "Zoospores," that, as they swim about, "Foreign bodies are carefully avoided, and the whole movement has a deceptive likeness to the voluntary changes of place which are observed in microscopic animals."

After swarming about in this way in the moisture on the surface of a leaf or stem (which, film though it may be, is an ocean to such a fish) for half an hour, more or less, the movement of the zoospore becomes slower, and is limited to a slow turning upon its axis, without change of place. It then becomes quite quiet, the cilia disappear, it assumes a spherical form, and surrounds itself with a distinct, though delicate, membranous coat. A protuberance then grows out from one side of the sphere, and rapidly increasing in length, assumes the character of a hypha. The latter penetrates into the substance of the potato plant, either by entering a stomate, or by boring through the wall of an epidermic cell, and ramifies, as a mycelium, in the substance of the plant, destroying the tissues with which it comes in contact. As these processes of multiplication take place very rapidly, millions of spores are soon set free from a single infested plant; and, from their minuteness, they are readily transported by the gentlest breeze. Since, again, the zoospores set free from each spore, in virtue of their powers of locomotion, swiftly disperse themselves over the surface, it is no wonder that the infection, once started, soon spreads from field to field, and extends its ravages over a whole country.

However, it does not enter into my present plan to treat of the potato disease, instructively as its history bears upon that of other epidemics; and I have selected the case of the Peroganspora simply because it affords an example of an organism, which, in one stage of its existence, is truly a "Monad," indistinguishable by any important character from our Heteromita, and extraordinarily like it in some respects. And yet this "Monad" can be traced, step by step, through the series of metamorphoses which I have described, until it assumes the features of an organism, which is as much a plant as is an oak or an elm.

Moreover, it would be possible to pursue the analogy farther. Under certain circumstances, a process of conjugation takes place in the Peronospora. Two separate portions of its protoplasm become fused together, surround themselves with a thick coat and give rise to a sort of vegetable egg called an oospore. After a period of rest, the contents of the oospore break up into a number of zoospores like those already described, each of which, after a period of activity, germinates in the ordinary way. This process obviously corresponds with the conjugation and subsequent setting free of germs in the Heteromita.

But it may be said that the Peronospora is, after all, a questionable sort of plant; that it seems to be wanting in the manufacturing power, selected as the main distinctive character of vegetable life; or, at any rate, that there is no proof that it does not get its protein matter ready made from the potato plant.

Let us, therefore, take a case which is not open to these objections.

There are some small plants known to botanists as members of the genus Colcochaete, which, without being truly parasitic, grow upon certain water-weeds, as lichens grow upon trees. The little plant has the form of an elegant green star, the branching arms of which are divided into cells. Its greenness is due to its chlorophyll, and it undoubtedly has the manufacturing power in full degree, decomposing carbonic acid and setting oxygen free, under the influence of sunlight. But the protoplasmic contents of some of the cells of which the plant is made up occasionally divide, by a method similar to that which effects the division of the contents of the Peronospora spore; and the severed portions are then set free as active monad-like zoospores. Each is oval and is provided at one extremity with two long active cilia. Propelled by these, it swims about for a longer or shorter time, but at length comes to a state of rest and gradually grows into a Coleochaete. Moreover, as in the Peronospora, conjugation may take place and result in an oospore; the contents of which divide and are set free as monadiform germs.

If the whole history of the zoospores of Peronospora and of Coleochaete were unknown, they would undoubtedly be classed among "Monads" with the same right as Heteromita; why then may not Heteromita be a plant, even though the cycle of forms through which it passes shows no terms quite so complex as those which occur in Peronospora and Coleochaete? And, in fact, there are some green organisms, in every respect characteristically plants, such as Chlamydomonas, and the common Volvox, or so-called "Globe animalcule," which run through a cycle of forms of just the same simple character as those of Heteromita.

The name of Chlamydomonas is applied to certain microscopic green bodies, each of which consists of a protoplasmic central substance invested by a structureless sac. The latter contains cellulose, as in ordinary plants; and the chlorophyll which gives the green colour enables the Chlamydomonas to decompose carbonic acid and fix carbon as they do. Two long cilia protrude through the cell-wall, and effect the rapid locomotion of this "monad," which, in all respects except its mobility, is characteristically a plant. Under ordinary circumstances, the Chlamydomonas multiplies by simple fission, each splitting into two or into four parts, which separate and become independent organisms. Sometimes, however, the Chlamydomonas divides into eight parts, each of which is provided with four instead of two cilia. These "zoospores" conjugate in pairs, and give rise to quiescent bodies, which multiply by division, find eventually pass into the active state.

Thus, so far as outward form and the general character of the cycle of modifications, through which the organism passes in the course of its life, are concerned, the resemblance between Chlamydomonas and Heteromita is of the closest description. And on the face of the matter there is no ground for refusing to admit that Heteromita may be related to Chlamydomonas, as the colourless fungus is to the green alga. Volvox may be compared to a hollow sphere, the wall of which is made up of coherent Chlamydomonads; and which progresses with a rotating motion effected by the paddling of the multitudinous pairs of cilia which project from its surface. Each Volvox-monad, moreover, possesses a red pigment spot, like the simplest form of eye known among animals. The methods of fissive multiplication and of conjugation observed in the monads of this locomotive globe are essentially similar to those observed in Chlamydomonas; and, though a hard battle has been fought over it, Volvox is now finally surrendered to the Botanists.

Thus there is really no reason why Heteromita may not be a plant; and this conclusion would be very satisfactory, if it were not equally easy to show that there is really no reason why it should not be an animal. For there are numerous organisms presenting the closest resemblance to Heteromita, and, like it, grouped under the general name of "Monads," which, nevertheless, can be observed to take in solid nutriment, and which, therefore, have a virtual, if not an actual, mouth and digestive cavity, and thus come under Cuvier's definition of an animal. Numerous forms of such animals have been described by Ehrenberg, Dujardin, H. James Clark, and other writers on the Infusoria. Indeed, in another infusion of hay in which my Heteromita lens occurred, there were innumerable such infusorial animalcules belonging to the well-known species Colpoda cucullus.[6]

[Footnote 6: Excellently described by Stein, almost all of whose statements I have verified.]

Full-sized specimens of this animalcule attain a length of between 1/300 or 1/400 of an inch, so that it may have ten times the length and a thousand times the mass of a Heteromita. In shape, it is not altogether unlike Heteromita. The small end, however, is not produced into one long cilium, but the general surface of the body is covered with small actively vibrating ciliary organs, which are only longest at the small end. At the point which answers to that from which the two cilia arise in Heteromita, there is a conical depression, the mouth; and, in young specimens, a tapering filament, which reminds one of the posterior cilium of Heteromita, projects from this region.

The body consists of a soft granular protoplasmic substance, the middle of which is occupied by a large oval mass called the "nucleus"; while, at its hinder end, is a "contractile vacuole," conspicuous by its regular rhythmic appearances and disappearances. Obviously, although the Colpoda is not a monad, it differs from one only in subordinate details. Moreover, under certain conditions, it becomes quiescent, incloses itself in a delicate case or cyst, and then divides into two, four, or more portions, which are eventually set free and swim about as active Colpodoe.

But this creature is an unmistakable animal, and full-sized Colpodoe may be fed as easily as one feeds chickens. It is only needful to diffuse very finely ground carmine through the water in which they live, and, in a very short time, the bodies of the Colpodoe are stuffed with the deeply-coloured granules of the pigment.

And if this were not sufficient evidence of the animality of Colpoda, there comes the fact that it is even more similar to another well-known animalcule, Paramoecium, than it is to a monad. But Paramoecium is so huge a creature compared with those hitherto discussed—it reaches 1/120 of an inch or more in length—that there is no difficulty in making out its organisation in detail; and in proving that it is not only an animal, but that it is an animal which possesses a somewhat complicated organisation. For example, the surface layer of its body is different in structure from the deeper parts. There are two contractile vacuoles, from each of which radiates a system of vessel-like canals; and not only is there a conical depression continuous with a tube, which serve as mouth and gullet, but the food ingested takes a definite course, and refuse is rejected from a definite region. Nothing is easier than to feed these animals, and to watch the particles of indigo or carmine accumulate at the lower end of the gullet. From this they gradually project, surrounded by a ball of water, which at length passes with a jerk, oddly simulating a gulp, into the pulpy central substance of the body, there to circulate up one side and down the other, until its contents are digested and assimilated. Nevertheless, this complex animal multiplies by division, as the monad does, and, like the monad, undergoes conjugation. It stands in the same relation to Heteromita on the animal side, as Coleochaete does on the plant side. Start from either, and such an insensible series of gradations leads to the monad that it is impossible to say at any stage of the progress where the line between the animal and the plant must be drawn.

There is reason to think that certain organisms which pass through a monad stage of existence, such as the Myxomycetes, are, at one time of their lives, dependent upon external sources for their protein matter, or are animals; and, at another period, manufacture it, or are plants. And seeing that the whole progress of modern investigation is in favour of the doctrine of continuity, it is a fair and probable speculation—though only a speculation—that, as there are some plants which can manufacture protein out of such apparently intractable mineral matters as carbonic acid, water, nitrate of ammonia, metallic and earthy salts; while others need to be supplied with their carbon and nitrogen in the somewhat less raw form of tartrate of ammonia and allied compounds; so there may be yet others, as is possibly the case with the true parasitic plants, which can only manage to put together materials still better prepared—still more nearly approximated to protein—until we arrive at such organisms as the Psorospermioe and the Panhistophyton, which are as much animal as vegetable in structure, but are animal in their dependence on other organisms for their food.

The singular circumstance observed by Meyer, that the Torula of yeast, though an indubitable plant, still flourishes most vigorously when supplied with the complex nitrogenous substance, pepsin; the probability that the Peronospora is nourished directly by the protoplasm of the potato-plant; and the wonderful facts which have recently been brought to light respecting insectivorous plants, all favour this view; and tend to the conclusion that the difference between animal and plant is one of degree rather than of kind, and that the problem whether, in a given case, an organism is an animal or a plant, may be essentially insoluble.



VII

A LOBSTER; OR, THE STUDY OF ZOOLOGY

[1861]

Natural history is the name familiarly applied to the study of the properties of such natural bodies as minerals, plants, and animals; the sciences which embody the knowledge man has acquired upon these subjects are commonly termed Natural Sciences, in contradistinction to other so- called "physical" sciences; and those who devote themselves especially to the pursuit of such sciences have been and are commonly termed "Naturalists."

Linnaeus was a naturalist in this wide sense, and his "Systema Naturae" was a work upon natural history, in the broadest acceptation of the term; in it, that great methodising spirit embodied all that was known in his time of the distinctive characters of minerals, animals, and plants. But the enormous stimulus which Linnaeus gave to the investigation of nature soon rendered it impossible that any one man should write another "Systema Naturae," and extremely difficult for any one to become even a naturalist such as Linnaeus was.

Great as have been the advances made by all the three branches of science, of old included under the title of natural history, there can be no doubt that zoology and botany have grown in an enormously greater ratio than mineralogy; and hence, as I suppose, the name of "natural history" has gradually become more and more definitely attached to these prominent divisions of the subject, and by "naturalist" people have meant more and more distinctly to imply a student of the structure and function of living beings.

However this may be, it is certain that the advance of knowledge has gradually widened the distance between mineralogy and its old associates, while it has drawn zoology and botany closer together; so that of late years it has been found convenient (and indeed necessary) to associate the sciences which deal with vitality and all its phenomena under the common head of "biology"; and the biologists have come to repudiate any blood-relationship with their foster-brothers, the mineralogists.

Certain broad laws have a general application throughout both the animal and the vegetable worlds, but the ground common to these kingdoms of nature is not of very wide extent, and the multiplicity of details is so great, that the student of living beings finds himself obliged to devote his attention exclusively either to the one or the other. If he elects to study plants, under any aspect, we know at once what to call him. He is a botanist, and his science is botany. But if the investigation of animal life be his choice, the name generally applied to him will vary according to the kind of animals he studies, or the particular phenomena of animal life to which he confines his attention. If the study of man is his object, he is called an anatomist, or a physiologist, or an ethnologist; but if he dissects animals, or examines into the mode in which their functions are performed, he is a comparative anatomist or comparative physiologist. If he turns his attention to fossil animals, he is a palaeontologist. If his mind is more particularly directed to the specific description, discrimination, classification, and distribution of animals, he is termed a zoologist.

For the purpose of the present discourse, however, I shall recognise none of these titles save the last, which I shall employ as the equivalent of botanist, and I shall use the term zoology is denoting the whole doctrine of animal life, in contradistinction to botany, which signifies the whole doctrine of vegetable life.

Employed in this sense, zoology, like botany, is divisible into three great but subordinate sciences, morphology, physiology, and distribution, each of which may, to a very great extent, be studied independently of the other.

Zoological morphology is the doctrine of animal form or structure. Anatomy is one of its branches; development is another; while classification is the expression of the relations which different animals bear to one another, in respect of their anatomy and their development.

Zoological distribution is the study of animals in relation to the terrestrial conditions which obtain now, or have obtained at any previous epoch of the earth's history.

Zoological physiology, lastly, is the doctrine of the functions or actions of animals. It regards animal bodies as machines impelled by certain forces, and performing an amount of work which can be expressed in terms of the ordinary forces of nature. The final object of physiology is to deduce the facts of morphology, on the one hand, and those of distribution on the other, from the laws of the molecular forces of matter.

Such is the scope of zoology. But if I were to content myself with the enunciation of these dry definitions, I should ill exemplify that method of teaching this branch of physical science, which it is my chief business to-night to recommend. Let us turn away then from abstract definitions. Let us take some concrete living thing, some animal, the commoner the better, and let us see how the application of common sense and common logic to the obvious facts it presents, inevitably leads us into all these branches of zoological science.

I have before me a lobster. When I examine it, what appears to be the most striking character it presents? Why, I observe that this part which we call the tail of the lobster, is made up of six distinct hard rings and a seventh terminal piece. If I separate one of the middle rings, say the third, I find it carries upon its under surface a pair of limbs or appendages, each of which consists of a stalk and two terminal pieces. So that I can represent a transverse section of the ring and its appendages upon the diagram board in this way.

If I now take the fourth ring, I find it has the same structure, and so have the fifth and the second; so that, in each of these divisions of the tail, I find parts which correspond with one another, a ring and two appendages; and in each appendage a stalk and two end pieces. These corresponding parts are called, in the technical language of anatomy, "homologous parts." The ring of the third division is the "homologue" of the ring of the fifth, the appendage of the former is the homologue of the appendage of the latter. And, as each division exhibits corresponding parts in corresponding places, we say that all the divisions are constructed upon the same plan. But now let us consider the sixth division. It is similar to, and yet different from, the others. The ring is essentially the same as in the other divisions; but the appendages look at first as if they were very different; and yet when we regard them closely, what do we find? A stalk and two terminal divisions, exactly as in the others, but the stalk is very short and very thick, the terminal divisions are very broad and flat, and one of them is divided into two pieces.

I may say, therefore, that the sixth segment is like the others in plan, but that it is modified in its details.

The first segment is like the others, so far as its ring is concerned, and though its appendages differ from any of those yet examined in the simplicity of their structure, parts corresponding with the stem and one of the divisions of the appendages of the other segments can be readily discerned in them.

Thus it appears that the lobster's tail is composed of a series of segments which are fundamentally similar, though each presents peculiar modifications of the plan common to all. But when I turn to the forepart of the body I see, at first, nothing but a great shield-like shell, called technically the "carapace," ending in front in a sharp spine, on either side of which are the curious compound eyes, set upon the ends of stout movable stalks. Behind these, on the under side of the body, are two pairs of long feelers, or antennae, followed by six pairs of jaws folded against one another over the mouth, and five pairs of legs, the foremost of these being the great pinchers, or claws, of the lobster.

It looks, at first, a little hopeless to attempt to find in this complex mass a series of rings, each with its pair of appendages, such as I have shown you in the abdomen, and yet it is not difficult to demonstrate their existence. Strip off the legs, and you will find that each pair is attached to a very definite segment of the under wall of the body; but these segments, instead of being the lower parts of free rings, as in the tail, are such parts of rings which are all solidly united and bound together; and the like is true of the jaws, the feelers, and the eye- stalks, every pair of which is borne upon its own special segment. Thus the conclusion is gradually forced upon us, that the body of the lobster is composed of as many rings as there are pairs of appendages, namely, twenty in all, but that the six hindmost rings remain free and movable, while the fourteen front rings become firmly soldered together, their backs forming one continuous shield—the carapace.

Unity of plan, diversity in execution, is the lesson taught by the study of the rings of the body, and the same instruction is given still more emphatically by the appendages. If I examine the outermost jaw I find it consists of three distinct portions, an inner, a middle, and an outer, mounted upon a common stem; and if I compare this jaw with the legs behind it, or the jaws in front of it, I find it quite easy to see, that, in the legs, it is the part of the appendage which corresponds with the inner division, which becomes modified into what we know familiarly as the "leg," while the middle division disappears, and the outer division is hidden under the carapace. Nor is it more difficult to discern that, in the appendages of the tail, the middle division appears again and the outer vanishes; while, on the other hand, in the foremost jaw, the so- called mandible, the inner division only is left; and, in the same way, the parts of the feelers and of the eye-stalks can be identified with those of the legs and jaws.

But whither does all this tend? To the very remarkable conclusion that a unity of plan, of the same kind as that discoverable in the tail or abdomen of the lobster, pervades the whole organisation of its skeleton, so that I can return to the diagram representing any one of the rings of the tail, which I drew upon the board, and by adding a third division to each appendage, I can use it as a sort of scheme or plan of any ring of the body. I can give names to all the parts of that figure, and then if I take any segment of the body of the lobster, I can point out to you exactly, what modification the general plan has undergone in that particular segment; what part has remained movable, and what has become fixed to another; what has been excessively developed and metamorphosed and what has been suppressed.

But I imagine I hear the question, How is all this to be tested? No doubt it is a pretty and ingenious way of looking at the structure of any animal; but is it anything more? Does Nature acknowledge, in any deeper way, this unity of plan we seem to trace?

The objection suggested by these questions is a very valid and important one, and morphology was in an unsound state so long as it rested upon the mere perception of the analogies which obtain between fully formed parts. The unchecked ingenuity of speculative anatomists proved itself fully competent to spin any number of contradictory hypotheses out of the same facts, and endless morphological dreams threatened to supplant scientific theory.

Happily, however, there is a criterion of morphological truth, and a sure test of all homologies. Our lobster has not always been what we see it; it was once an egg, a semifluid mass of yolk, not so big as a pin's head, contained in a transparent membrane, and exhibiting not the least trace of any one of those organs, the multiplicity and complexity of which, in the adult, are so surprising. After a time, a delicate patch of cellular membrane appeared upon one face of this yolk, and that patch was the foundation of the whole creature, the clay out of which it would be moulded. Gradually investing the yolk, it became subdivided by transverse constrictions into segments, the forerunners of the rings of the body. Upon the ventral surface of each of the rings thus sketched out, a pair of bud-like prominences made their appearance—the rudiments of the appendages of the ring. At first, all the appendages were alike, but, as they grew, most of them became distinguished into a stem and two terminal divisions, to which, in the middle part of the body, was added a third outer division; and it was only at a later period, that by the modification, or absorption, of certain of these primitive constituents, the limbs acquired their perfect form.

Thus the study of development proves that the doctrine of unity of plan is not merely a fancy, that it is not merely one way of looking at the matter, but that it is the expression of deep-seated natural facts. The legs and jaws of the lobster may not merely be regarded as modifications of a common type,—in fact and in nature they are so,—the leg and the jaw of the young animal being, at first, indistinguishable.

These are wonderful truths, the more so because the zoologist finds them to be of universal application. The investigation of a polype, of a snail, of a fish, of a horse, or of a man, would have led us, though by a less easy path, perhaps, to exactly the same point. Unity of plan everywhere lies hidden under the mask of diversity of structure—the complex is everywhere evolved out of the simple. Every animal has at first the form of an egg, and every animal and every organic part, in reaching its adult state, passes through conditions common to other animals and other adult parts; and this leads me to another point. I have hitherto spoken as if the lobster were alone in the world, but, as I need hardly remind you, there are myriads of other animal organisms. Of these, some, such as men, horses, birds, fishes, snails, slugs, oysters, corals, and sponges, are not in the least like the lobster. But other animals, though they may differ a good deal from the lobster, are yet either very like it, or are like something that is like it. The cray fish, the rock lobster, and the prawn, and the shrimp, for example, however different, are yet so like lobsters, that a child would group them as of the lobster kind, in contradistinction to snails and slugs; and these last again would form a kind by themselves, in contradistinction to cows, horses, and sheep, the cattle kind.

But this spontaneous grouping into "kinds" is the first essay of the human mind at classification, or the calling by a common name of those things that are alike, and the arranging them in such a manner as best to suggest the sum of their likenesses and unlikenesses to other things.

Those kinds which include no other subdivisions than the sexes, or various breeds, are called, in technical language, species. The English lobster is a species, our cray fish is another, our prawn is another. In other countries, however, there are lobsters, cray fish, and prawns, very like ours, and yet presenting sufficient differences to deserve distinction. Naturalists, therefore, express this resemblance and this diversity by grouping them as distinct species of the same "genus." But the lobster and the cray fish, though belonging to distinct genera, have many features in common, and hence are grouped together in an assemblage which is called a family. More distant resemblances connect the lobster with the prawn and the crab, which are expressed by putting all these into the same order. Again, more remote, but still very definite, resemblances unite the lobster with the woodlouse, the king crab, the water flea, and the barnacle, and separate them from all other animals; whence they collectively constitute the larger group, or class, Crustacea. But the Crustacea exhibit many peculiar features in common with insects, spiders, and centipedes, so that these are grouped into the still larger assemblage or "province" Articulata; and, finally, the relations which these have to worms and other lower animals, are expressed by combining the whole vast aggregate into the sub-kingdom of Annulosa.

If I had worked my way from a sponge instead of a lobster, I should have found it associated, by like ties, with a great number of other animals into the sub-kingdom Protozoa; if I had selected a fresh-water polype or a coral, the members of what naturalists term the sub-kingdom Coelenterata, would have grouped themselves around my type; had a snail been chosen, the inhabitants of all univalve and bivalve, land and water, shells, the lamp shells, the squids, and the sea-mat would have gradually linked themselves on to it as members of the same sub-kingdom of Mollusca; and finally, starting from man, I should have been compelled to admit first, the ape, the rat, the horse, the dog, into the same class; and then the bird, the crocodile, the turtle, the frog, and the fish, into the same sub-kingdom of Vertebrata.

And if I had followed out all these various lines of classification fully, I should discover in the end that there was no animal, either recent or fossil, which did not at once fall into one or other of these sub-kingdoms. In other words, every animal is organised upon one or other of the five, or more, plans, the existence of which renders our classification possible. And so definitely and precisely marked is the structure of each animal, that, in the present state of our knowledge, there is not the least evidence to prove that a form, in the slightest degree transitional between any of the two groups Vertebrata, Annulosa, Mollusca, and Coelenterata, either exists, or has existed, during that period of the earth's history which is recorded by the geologist.[1] Nevertheless, you must not for a moment suppose, because no such transitional forms are known, that the members of the sub-kingdoms are disconnected from, or independent of, one another. On the contrary, in their earliest condition they are all similar, and the primordial germs of a man, a dog, a bird, a fish, a beetle, a snail, and a polype are, in no essential structural respects, distinguishable.

[Footnote 1: The different grouping necessitated by later knowledge does not affect the principle of the argument.—1894.]

In this broad sense, it may with truth be said, that all living animals, and all those dead faunae which geology reveals, are bound together by an all-pervading unity of organisation, of the same character, though not equal in degree, to that which enables us to discern one and the same plan amidst the twenty different segments of a lobster's body. Truly it has been said, that to a clear eye the smallest fact is a window through which the Infinite may be seen.

Turning from these purely morphological considerations, let us now examine into the manner in which the attentive study of the lobster impels us into other lines of research.

Lobsters are found in all the European seas; but on the opposite shores of the Atlantic and in the seas of the southern hemisphere they do not exist. They are, however, represented in these regions by very closely allied, but distinct forms—the Homarus Americanus and the Homarus Capensis: so that we may say that the European has one species of Homuarus; the American, another; the African, another; and thus the remarkable facts of geographical distribution begin to dawn upon us.

Again, if we examine the contents of the earth's crust, we shall find in the latter of those deposits, which have served as the great burying grounds of past ages, numberless lobster-like animals, but none so similar to our living lobster as to make zoologists sure that they belonged even to the same genus. If we go still further back in time, we discover, in the oldest rocks of all, the remains of animals, constructed on the same general plan as the lobster, and belonging to the same great group of Crustacea; but for the most part totally different from the lobster, and indeed from any other living form of crustacean; and thus we gain a notion of that successive change of the animal population of the globe, in past ages, which is the most striking fact revealed by geology.

Consider, now, where our inquiries have led us. We studied our type morphologically, when we determined its anatomy and its development, and when comparing it, in these respects, with other animals, we made out its place in a system of classification. If we were to examine every animal in a similar manner, we should establish a complete body of zoological morphology.

Again, we investigated the distribution of our type in space and in time, and, if the like had been done with every animal, the sciences of geographical and geological distribution would have attained their limit.

But you will observe one remarkable circumstance, that, up to this point, the question of the life of these organisms has not come under consideration. Morphology and distribution might be studied almost as well, if animals and plants were a peculiar kind of crystals, and possessed none of those functions which distinguish living beings so remarkably. But the facts of morphology and distribution have to be accounted for, and the science, the aim of which it is to account for them, is Physiology.

Let us return to our lobster once more. If we watched the creature in its native element, we should see it climbing actively the submerged rocks, among which it delights to live, by means of its strong legs; or swimming by powerful strokes of its great tail, the appendages of the sixth joint of which are spread out into a broad fan-like Propeller: seize it, and it will show you that its great claws are no mean weapons of offence; suspend a piece of carrion among its haunts, and it will greedily devour it, tearing and crushing the flesh by means of its multitudinous jaws.

Suppose that we had known nothing of the lobster but as an inert mass, an organic crystal, if I may use the phrase, and that we could suddenly see it exerting all these powers, what wonderful new ideas and new questions would arise in our minds! The great new question would be, "How does all this take place?" the chief new idea would be, the idea of adaptation to purpose,—the notion, that the constituents of animal bodies are not mere unconnected parts, but organs working together to an end. Let us consider the tail of the lobster again from this point of view. Morphology has taught us that it is a series of segments composed of homologous parts, which undergo various modifications—beneath and through which a common plan of formation is discernible. But if I look at the same part physiologically, I see that it is a most beautifully constructed organ of locomotion, by means of which the animal can swiftly propel itself either backwards or forwards.

But how is this remarkable propulsive machine made to perform its functions? If I were suddenly to kill one of these animals and to take out all the soft parts, I should find the shell to be perfectly inert, to have no more power of moving itself than is possessed by the machinery of a mill when disconnected from its steam-engine or water-wheel. But if I were to open it, and take out the viscera only, leaving the white flesh, I should perceive that the lobster could bend and extend its tail as well as before. If I were to cut off the tail, I should cease to find any spontaneous motion in it; but on pinching any portion of the flesh, I should observe that it underwent a very curious change—each fibre becoming shorter and thicker. By this act of contraction, as it is termed, the parts to which the ends of the fibre are attached are, of course, approximated; and according to the relations of their points of attachment to the centres of motions of the different rings, the bending or the extension of the tail results. Close observation of the newly- opened lobster would soon show that all its movements are due to the same cause—the shortening and thickening of these fleshy fibres, which are technically called muscles.

Here, then, is a capital fact. The movements of the lobster are due to muscular contractility. But why does a muscle contract at one time and not at another? Why does one whole group of muscles contract when the lobster wishes to extend his tail, and another group when he desires to bend it? What is it originates, directs, and controls the motive power?

Experiment, the great instrument for the ascertainment of truth in physical science, answers this question for us. In the head of the lobster there lies a small mass of that peculiar tissue which is known as nervous substance. Cords of similar matter connect his brain of the lobster, directly or indirectly, with the muscles. Now, if these communicating cords are cut, the brain remaining entire, the power of exerting what we call voluntary motion in the parts below the section is destroyed; and, on the other hand, if, the cords remaining entire, the brain mass be destroyed, the same voluntary mobility is equally lost. Whence the inevitable conclusion is, that the power of originating these motions resides in the brain and is propagated along the nervous cords.

In the higher animals the phenomena which attend this transmission have been investigated, and the exertion of the peculiar energy which resides in the nerves has been found to be accompanied by a disturbance of the electrical state of their molecules.

If we could exactly estimate the signification of this disturbance; if we could obtain the value of a given exertion of nerve force by determining the quantity of electricity, or of heat, of which it is the equivalent; if we could ascertain upon what arrangement, or other condition of the molecules of matter, the manifestation of the nervous and muscular energies depends (and doubtless science will some day or other ascertain these points), physiologists would have attained their ultimate goal in this direction; they would have determined the relation of the motive force of animals to the other forms of force found in nature; and if the same process had been successfully performed for all the operations which are carried on in, and by, the animal frame, physiology would be perfect, and the facts of morphology and distribution would be deducible from the laws which physiologists had established, combined with those determining the condition of the surrounding universe.

There is not a fragment of the organism of this humble animal whose study would not lead us into regions of thought as large as those which I have briefly opened up to you; but what I have been saying, I trust, has not only enabled you to form a conception of the scope and purport of zoology, but has given you an imperfect example of the manner in which, in my opinion, that science, or indeed any physical science, may be best taught. The great matter is, to make teaching real and practical, by fixing the attention of the student on particular facts; but at the same time it should be rendered broad and comprehensive, by constant reference to the generalisations of which all particular facts are illustrations. The lobster has served as a type of the whole animal kingdom, and its anatomy and physiology have illustrated for us some of the greatest truths of biology. The student who has once seen for himself the facts which I have described, has had their relations explained to him, and has clearly comprehended them, has, so far, a knowledge of zoology, which is real and genuine, however limited it may be, and which is worth more than all the mere reading knowledge of the science he could ever acquire. His zoological information is, so far, knowledge and not mere hearsay.

And if it were nay business to fit you for the certificate in zoological science granted by this department, I should pursue a course precisely similar in principle to that which I have taken to-night. I should select a fresh-water sponge, a fresh-water polype or a Cyanoea, a fresh-water mussel, a lobster, a fowl, as types of the five primary divisions of the animal kingdom. I should explain their structure very fully, and show how each illustrated the great principles of zoology. Having gone very carefully and fully over this ground, I should feel that you had a safe foundation, and I should then take you in the same way, but less minutely, over similarly selected illustrative types of the classes; and then I should direct your attention to the special forms enumerated under the head of types, in this syllabus, and to the other facts there mentioned.

That would, speaking generally, be my plan. But I have undertaken to explain to you the best mode of acquiring and communicating a knowledge of zoology, and you may therefore fairly ask me for a more detailed and precise account of the manner in which I should propose to furnish you with the information I refer to.

My own impression is, that the best model for all kinds of training in physical science is that afforded by the method of teaching anatomy, in use in the medical schools. This method consists of three elements— lectures, demonstrations, and examinations.

The object of lectures is, in the first place, to awaken the attention and excite the enthusiasm of the student; and this, I am sure, may be effected to a far greater extent by the oral discourse and by the personal influence of a respected teacher than in any other way. Secondly, lectures have the double use of guiding the student to the salient points of a subject, and at the same time forcing him to attend to the whole of it, and not merely to that part which takes his fancy. And lastly, lectures afford the student the opportunity of seeking explanations of those difficulties which will, and indeed ought to, arise in the course of his studies.

What books shall I read? is a question constantly put by the student to the teacher. My reply usually is, "None: write your notes out carefully and fully; strive to understand them thoroughly; come to me for the explanation of anything you cannot understand; and I would rather you did not distract your mind by reading." A properly composed course of lectures ought to contain fully as much matter as a student can assimilate in the time occupied by its delivery; and the teacher should always recollect that his business is to feed, and not to cram the intellect. Indeed, I believe that a student who gains from a course of lectures the simple habit of concentrating his attention upon a definitely limited series of facts, until they are thoroughly mastered, has made a step of immeasurable importance.

But, however good lectures may be, and however extensive the course of reading by which they are followed up, they are but accessories to the great instrument of scientific teaching—demonstration. If I insist unweariedly, nay fanatically, upon the importance of physical science as an educational agent, it is because the study of any branch of science, if properly conducted, appears to me to fill up a void left by all other means of education. I have the greatest respect and love for literature; nothing would grieve me more than to see literary training other than a very prominent branch of education: indeed, I wish that real literary discipline were far more attended to than it is; but I cannot shut my eyes to the fact, that there is a vast difference between men who have had a purely literary, and those who have had a sound scientific, training.

Seeking for the cause of this difference, I imagine I can find it in the fact that, in the world of letters, learning and knowledge are one, and books are the source of both; whereas in science, as in life, learning and knowledge are distinct, and the study of things, and not of books, is the source of the latter.

All that literature has to bestow may be obtained by reading and by practical exercise in writing and in speaking; but I do not exaggerate when I say, that none of the best gifts of science are to be won by these means. On the contrary, the great benefit which a scientific education bestows, whether is training or as knowledge, is dependent upon the extent to which the mind of the student is brought into immediate contact with facts—upon the degree to which he learns the habit of appealing directly to Nature, and of acquiring through his senses concrete images of those properties of things, which are, and always will be, but approximatively expressed in human language. Our way of looking at Nature, and of speaking about her, varies from year to year; but a fact once seen, a relation of cause and effect, once demonstratively apprehended, are possessions which neither change nor pass away, but, on the contrary, form fixed centres, about which other truths aggregate by natural affinity.

Therefore, the great business of the scientific teacher is, to imprint the fundamental, irrefragable facts of his science, not only by words upon the mind, but by sensible impressions upon the eye, and ear, and touch of the student, in so complete a manner, that every term used, or law enunciated, should afterwards call up vivid images of the particular structural, or other, facts which furnished the demonstration of the law, or the illustration of the term.

Now this important operation can only be achieved by constant demonstration, which may take place to a certain imperfect extent during a lecture, but which ought also to be carried on independently, and which should be addressed to each individual student, the teacher endeavouring, not so much to show a thing to the learner, as to make him see it for himself.

I am well aware that there are great practical difficulties in the way of effectual zoological demonstrations. The dissection of animals is not altogether pleasant, and requires much time; nor is it easy to secure an adequate supply of the needful specimens. The botanist has here a great advantage; his specimens are easily obtained, are clean and wholesome, and can be dissected in a private house as well as anywhere else; and hence, I believe, the fact, that botany is so much more readily and better taught than its sister science. But, be it difficult or be it easy, if zoological science is to be properly studied, demonstration, and, consequently, dissection, must be had. Without it, no man can have a really sound knowledge of animal organisation.

A good deal may be done, however, without actual dissection on the student's part, by demonstration upon specimens and preparations; and in all probability it would not be very difficult, were the demand sufficient, to organise collections of such objects, sufficient for all the purposes of elementary teaching, at a comparatively cheap rate. Even without these, much might be effected, if the zoological collections, which are open to the public, were arranged according to what has been termed the "typical principle"; that is to say, if the specimens exposed to public view were so selected that the public could learn something from them, instead of being, as at present, merely confused by their multiplicity. For example, the grand ornithological gallery at the British Museum contains between two and three thousand species of birds, and sometimes five or six specimens of a species. They are very pretty to look at, and some of the cases are, indeed, splendid; but I will undertake to say, that no man but a professed ornithologist has ever gathered much information from the collection. Certainly, no one of the tens of thousands of the general public who have walked through that gallery ever knew more about the essential peculiarities of birds when he left the gallery than when he entered it. But if, somewhere in that vast hall, there were a few preparations, exemplifying the leading structural peculiarities and the mode of development of a common fowl; if the types of the genera, the leading modifications in the skeleton, in the plumage at various ages, in the mode of nidification, and the like, among birds, were displayed; and if the other specimens were put away in a place where the men of science, to whom they are alone useful, could have free access to them, I can conceive that this collection might become a great instrument of scientific education.

The last implement of the teacher to which I have adverted is examination—a means of education now so thoroughly understood that I need hardly enlarge upon it. I hold that both written and oral examinations are indispensable, and, by requiring the description of specimens, they may be made to supplement demonstration.

Such is the fullest reply the time at my disposal will allow me to give to the question—how may a knowledge of zoology be best acquired and communicated?

But there is a previous question which may be moved, and which, in fact, I know many are inclined to move. It is the question, why should teachers be encouraged to acquire a knowledge of this, or any other branch of physical science? What is the use, it is said, of attempting to make physical science a branch of primary education? Is it not probable that teachers, in pursuing such studies, will be led astray from the acquirement of more important but less attractive knowledge? And, even if they can learn something of science without prejudice to their usefulness, what is the good of their attempting to instil that knowledge into boys whose real business is the acquisition of reading, writing, and arithmetic?

These questions are, and will be, very commonly asked, for they arise from that profound ignorance of the value and true position of physical science, which infests the minds of the most highly educated and intelligent classes of the community. But if I did not feel well assured that they are capable of being easily and satisfactorily answered; that they have been answered over and over again; and that the time will come when men of liberal education will blush to raise such questions—I should be ashamed of my position here to-night. Without doubt, it is your great and very important function to carry out elementary education; without question, anything that should interfere with the faithful fulfilment of that duty on your part would be a great evil; and if I thought that your acquirement of the elements of physical science, and your communication of those elements to your pupils, involved any sort of interference with your proper duties, I should be the first person to protest against your being encouraged to do anything of the kind.

But is it true that the acquisition of such a knowledge of science as is proposed, and the communication of that knowledge, are calculated to weaken your usefulness? Or may I not rather ask, is it possible for you to discharge your functions properly without these aids?

What is the purpose of primary intellectual education? I apprehend that its first object is to train the young in the use of those tools wherewith men extract knowledge from the ever-shifting succession of phenomena which pass before their eyes; and that its second object is to inform them of the fundamental laws which have been found by experience to govern the course of things, so that they may not be turned out into the world naked, defenceless, and a prey to the events they might control.

A boy is taught to read his own and other languages, in order that he may have access to infinitely wider stores of knowledge than could ever be opened to him by oral intercourse with his fellow men; he learns to write, that his means of communication with the rest of mankind may be indefinitely enlarged, and that he may record and store up the knowledge he acquires. He is taught elementary mathematics, that he may understand all those relations of number and form, upon which the transactions of men, associated in complicated societies, are built, and that he may have some practice in deductive reasoning.

All these operations of reading, writing, and ciphering, are intellectual tools, whose use should, before all things, be learned, and learned thoroughly; so that the youth may be enabled to make his life that which it ought to be, a continual progress in learning and in wisdom.

But, in addition, primary education endeavours to fit a boy out with a certain equipment of positive knowledge. He is taught the great laws of morality; the religion of his sect; so much history and geography as will tell him where the great countries of the world are, what they are, and how they have become what they are.

Without doubt all these are most fitting and excellent things to teach a boy; I should be very sorry to omit any of them from any scheme of primary intellectual education. The system is excellent, so far as it goes.

But if I regard it closely, a curious reflection arises. I suppose that, fifteen hundred years ago, the child of any well-to-do Roman citizen was taught just these same things; reading and writing in his own, and, perhaps, the Greek tongue; the elements of mathematics; and the religion, morality, history, and geography current in his time. Furthermore, I do not think I err in affirming, that, if such a Christian Roman boy, who had finished his education, could be transplanted into one of our public schools, and pass through its course of instruction, he would not meet with a single unfamiliar line of thought; amidst all the new facts he would have to learn, not one would suggest a different mode of regarding the universe from that current in his own time.

And yet surely there is some great difference between the civilisation of the fourth century and that of the nineteenth, and still more between the intellectual habits and tone of thought of that day and this?

And what has made this difference? I answer fearlessly—The prodigious development of physical science within the last two centuries.

Modern civilisation rests upon physical science; take away her gifts to our own country, and our position among the leading nations of the world is gone to-morrow; for it is physical science only that makes intelligence and moral energy stronger than brute force.

The whole of modern thought is steeped in science; it has made its way into the works of our best poets, and even the mere man of letters, who affects to ignore and despise science, is unconsciously impregnated with her spirit, and indebted for his best products to her methods. I believe that the greatest intellectual revolution mankind has yet seen is now slowly taking place by her agency. She is teaching the world that the ultimate court of appeal is observation and experiment, and not authority; she is teaching it to estimate the value of evidence; she is creating a firm and living faith in the existence of immutable moral and physical laws, perfect obedience to which is the highest possible aim of an intelligent being.

But of all this your old stereotyped system of education takes no note. Physical science, its methods, its problems, and its difficulties, will meet the poorest boy at every turn, and yet we educate him in such a manner that he shall enter the world as ignorant of the existence of the methods and facts of science as the day he was born. The modern world is full of artillery; and we turn out our children to do battle in it, equipped with the shield and sword of an ancient gladiator.

Posterity will cry shame on us if we do not remedy this deplorable state of things. Nay, if we live twenty years longer, our own consciences will cry shame on us.

It is my firm conviction that the only way to remedy it is to make the elements of physical science an integral part of primary education. I have endeavoured to show you how that may be done for that branch of science which it is my business to pursue; and I can but add, that I should look upon the day when every schoolmaster throughout this land was a centre of genuine, however rudimentary, scientific knowledge, as an epoch in the history of the country.

But let me entreat you to remember my last words. Addressing myself to you, as teachers, I would say, mere book learning in physical science is a sham and a delusion—what you teach, unless you wish to be impostors, that you must first know; and real knowledge in science means personal acquaintance with the facts, be they few or many.[2]

[Footnote 2: It has been suggested to me that these words may be taken to imply a discouragement on my part of any sort of scientific instruction which does not give an acquaintance with the facts at first hand. But this is not my meaning. The ideal of scientific teaching is, no doubt, a system by which the scholar sees every fact for himself, and the teacher supplies only the explanations. Circumstances, however, do not often allow of the attainment of that ideal, and we must put up with the next best system—one in which the scholar takes a good deal on trust from a teacher, who, knowing the facts by his own knowledge, can describe them with so much vividness as to enable his audience to form competent ideas concerning them. The system which I repudiate is that which allows teachers who have not come into direct contact with the leading facts of a science to pass their second-hand information on. The scientific virus, like vaccine lymph, if passed through too long a succession of organisms, will lose all its effect in protecting the young against the intellectual epidemics to which they are exposed.

[The remarks on p. 222 applied to the Natural History Collection of the British Museum in 1861. The visitor to the Natural History Museum in 1894 need go no further than the Great Hall to see the realisation of my hopes by the present Director.]]



VIII

BIOGENESIS AND ABIOGENESIS

(THE PRESIDENTIAL ADDRESS TO THE BRITISH ASSOCIATION FOR THE ADVANCEMENT OF SCIENCE FOR 1870)

It has long been the custom for the newly installed President of the British Association for the Advancement of Science to take advantage of the elevation of the position in which the suffrages of his colleagues had, for the time, placed him, and, casting his eyes around the horizon of the scientific world, to report to them what could be seen from his watch-tower; in what directions the multitudinous divisions of the noble army of the improvers of natural knowledge were marching; what important strongholds of the great enemy of us all, ignorance, had been recently captured; and, also, with due impartiality, to mark where the advanced posts of science had been driven in, or a long-continued siege had made no progress.

I propose to endeavour to follow this ancient precedent, in a manner suited to the limitations of my knowledge and of my capacity. I shall not presume to attempt a panoramic survey of the world of science, nor even to give a sketch of what is doing in the one great province of biology, with some portions of which my ordinary occupations render me familiar. But I shall endeavour to put before you the history of the rise and progress of a single biological doctrine; and I shall try to give some notion of the fruits, both intellectual and practical, which we owe, directly or indirectly, to the working out, by seven generations of patient and laborious investigators, of the thought which arose, more than two centuries ago, in the mind of a sagacious and observant Italian naturalist.

It is a matter of everyday experience that it is difficult to prevent many articles of food from becoming covered with mould; that fruit, sound enough to all appearance, often contains grubs at the core; that meat, left to itself in the air, is apt to putrefy and swarm with maggots. Even ordinary water, if allowed to stand in an open vessel, sooner or later becomes turbid and full of living matter.

The philosophers of antiquity, interrogated as to the cause of these phenomena, were provided with a ready and a plausible answer. It did not enter their minds even to doubt that these low forms of life were generated in the matters in which they made their appearance. Lucretius, who had drunk deeper of the scientific spirit than any poet of ancient or modern times except Goethe, intends to speak as a philosopher, rather than as a poet, when he writes that "with good reason the earth has gotten the name of mother, since all things are produced out of the earth. And many living creatures, even now, spring out of the earth, taking form by the rains and the heat of the sun."[1] The axiom of ancient science, "that the corruption of one thing is the birth of another," had its popular embodiment in the notion that a seed dies before the young plant springs from it; a belief so widespread and so fixed, that Saint Paul appeals to it in one of the most splendid outbursts of his fervid eloquence:—

"Thou fool, that which thou sowest is not quickened, except it die."[2]

[Footnote 1: It is thus that Mr. Munro renders

"Linquitur, ut merito maternum nomen adepta Terra sit, e terra quoniam sunt cuncta creata. Multaque nunc etiam exsistant animalia terris Imbribus et calido solis concreta vapore."

De Rerum Natura, lib. v. 793-796.

But would not the meaning of the last line be better rendered "Developed in rain-water and in the warm vapours raised by the sun"?]

[Footnote 2: 1 Corinthians xv. 36.]

The proposition that life may, and does, proceed from that which has no life, then, was held alike by the philosophers, the poets, and the people, of the most enlightened nations, eighteen hundred years ago; and it remained the accepted doctrine of learned and unlearned Europe, through the Middle Ages, down even to the seventeenth century.

It is commonly counted among the many merits of our great countryman, Harvey, that he was the first to declare the opposition of fact to venerable authority in this, as in other matters; but I can discover no justification for this widespread notion. After careful search through the "Exercitationes de Generatione," the most that appears clear to me is, that Harvey believed all animals and plants to spring from what he terms a "primordium vegetale," a phrase which may nowadays be rendered "a vegetative germ"; and this, he says, is "oviforme," or "egg-like"; not, he is careful to add, that it necessarily has the shape of an egg, but because it has the constitution and nature of one. That this "primordium oviforme" must needs, in all cases, proceed from a living parent is nowhere expressly maintained by Harvey, though such an opinion may be thought to be implied in one or two passages; while, on the other hand, he does, more than once, use language which is consistent only with a full belief in spontaneous or equivocal generation.[3] In fact, the main concern of Harvey's wonderful little treatise is not with generation, in the physiological sense, at all, but with development; and his great object is the establishment of the doctrine of epigenesis.

[Footnote 3: See the following passage in Exercitatio I.:—"Item sponte nascentia dicuntur; non quod ex putredine oriunda sint, sed quod casu, naturae sponte, et aequivoca (ut aiunt) generatione, a parentibus sui dissimilibus proveniant." Again, in De Uteri Membranis:—"In cunctorum viventium generatione (sicut diximus) hoc solenne est, ut ortum ducunt a primordio aliquo, quod tum materiam tum elficiendi potestatem in se habet: sitque, adeo id, ex quo et a quo quicquid nascitur, ortum suum ducat. Tale primordium in animalibus (sive ab aliis generantibus proveniant, sive sponte, aut ex putredine nascentur) est humor in tunica, aliquaaut putami ne conclusus." Compare also what Redi has to say respecting Harvey's opinions, Esperienze, p. 11.]

The first distinct enunciation of the hypothesis that all living matter has sprung from pre-existing living matter, came from a contemporary, though a junior, of Harvey, a native of that country, fertile in men great in all departments of human activity, which was to intellectual Europe, in the sixteenth and seventeenth centuries, what Germany is in the nineteenth. It was in Italy, and from Italian teachers, that Harvey received the most important part of his scientific education. And it was a student trained in the same schools, Francesco Redi—a man of the widest knowledge and most versatile abilities, distinguished alike as scholar, poet, physician, and naturalist—who, just two hundred and two years ago, published his "Esperienze intorno alla Generazione degl' Insetti," and gave to the world the idea, the growth of which it is my purpose to trace. Redi's book went through five editions in twenty years; and the extreme simplicity of his experiments, and the clearness of his arguments, gained for his views, and for their consequences, almost universal acceptance.

Redi did not trouble himself much with speculative considerations, but attacked particular cases of what was supposed to be "spontaneous generation" experimentally. Here are dead animals, or pieces of meat, says he; I expose them to the air in hot weather, and in a few days they swarm with maggots. You tell me that these are generated in the dead flesh; but if I put similar bodies, while quite fresh, into a jar, and tie some fine gauze over the top of the jar, not a maggot makes its appearance, while the dead substances, nevertheless, putrefy just in the same way as before. It is obvious, therefore, that the maggots are not generated by the corruption of the meat; and that the cause of their formation must be a something which is kept away by gauze. But gauze will not keep away aeriform bodies, or fluids. This something must, therefore, exist in the form of solid particles too big to get through the gauze. Nor is one long left in doubt what these solid particles are; for the blowflies, attracted by the odour of the meat, swarm round the vessel, and, urged by a powerful but in this case misleading instinct, lay eggs out of which maggots are immediately hatched, upon the gauze. The conclusion, therefore, is unavoidable; the maggots are not generated by the meat, but the eggs which give rise to them are brought through the air by the flies.

These experiments seem almost childishly simple, and one wonders how it was that no one ever thought of them before. Simple as they are, however, they are worthy of the most careful study, for every piece of experimental work since done, in regard to this subject, has been shaped upon the model furnished by the Italian philosopher. As the results of his experiments were the same, however varied the nature of the materials he used, it is not wonderful that there arose in Redi's mind a presumption, that, in all such cases of the seeming production of life from dead matter, the real explanation was the introduction of living germs from without into that dead matter.[4] And thus the hypothesis that living matter always arises by the agency of pre-existing living matter, took definite shape; and had, henceforward, a right to be considered and a claim to be refuted, in each particular case, before the production of living matter in any other way could be admitted by careful reasoners. It will be necessary for me to refer to this hypothesis so frequently, that, to save circumlocution, I shall call it the hypothesis of Biogenesis; and I shall term the contrary doctrine—that living matter may be produced by not living matter—the hypothesis of Abiogenesis.

[Footnote 4: "Pure contentandomi sempre in questa ed in ciascuna altro cosa, da ciascuno piu savio, la dove io difettuosamente parlassi, esser corretto; non tacero, che per molte osservazioni molti volti da me fatte, mi sento inclinato a credere che la terra, da quelle prime piante, e da quei primi animali in poi, che ella nei primi giorni del mondo produsse per comandemento del sovrano ed omnipotente Fattore, non abbia mai piu prodotto da se medesima ne erba ne albero, ne animale alcuno perfetto o imperfetto che ei se fosse; e che tutto quello, che ne' tempi trapassati e nato e che ora nascere in lei, o da lei veggiamo, venga tutto dalla semenza reale e vera delle piante, e degli animali stessi, i quali col mezzo del proprio seme la loro spezie conservano. E se bene tutto giorno scorghiamo da' cadaveri degli animali, e da tutte quante le maniere dell' erbe, e de' fiori, e dei frutti imputriditi, e corrotti nascere vermi infiniti—

'Nonne vides quaecunque mora, fluidoque calore Corpora tabescunt in parva animalia verti'—

Io mi sento, dico, inclinato, a credere che tutti quei vermi si generino dal seme paterno; e che le carni, e l' erbe, e l' altre cose tutte putrefatte, o putrefattibili non facciano altra parte, ne abbiano altro ufizio nella generazione degl' insetti, se non d'apprestare un luogo o un nido proporzionato, in cui dagli animali nel tempo della figliatura sieno portati, e partoriti i vermi, o l' uova o l' altre semenze dei vermi, i quali tosto che nati sono, trovano in esso nido un sufficiente alimento abilissimo per nutricarsi: e se in quello non son portate dalle madri queste suddette semenze, niente mai, e replicatamente niente, vi s' ingegneri e nasca."—REDI, Esperienze, pp. 14-16.]

In the seventeenth century, as I have said, the latter was the dominant view, sanctioned alike by antiquity and by authority; and it is interesting to observe that Redi did not escape the customary tax upon a discoverer of having to defend himself against the charge of impugning the authority of the Scriptures;[5] for his adversaries declared that the generation of bees from the carcase of a dead lion is affirmed, in the Book of Judges, to have been the origin of the famous riddle with which Samson perplexed the Philistines:—

Out of the eater came forth meat, And out of the strong came forth sweetness.

[Footnote 5: "Molti, e molti altri ancora vi potrei annoverare, se non fossi chiamato a rispondere alle rampogne di alcuni, che bruscamente mi rammentano cio, che si legge nel capitolo quattordicesimo del sacrosanto Libro de' giudici ... "—REDI, loc. cit. p. 45.]

Against all odds, however, Redi, strong with the strength of demonstrable fact, did splendid battle for Biogenesis; but it is remarkable that he held the doctrine in a sense which, if he lead lived in these times, would have infallibly caused him to be classed among the defenders of "spontaneous generation." "Omne vivum ex vivo," "no life without antecedent life," aphoristically sums up Redi's doctrine; but he went no further. It is most remarkable evidence of the philosophic caution and impartiality of his mind, that although he had speculatively anticipated the manner in which grubs really are deposited in fruits and in the galls of plants, he deliberately admits that the evidence is insufficient to bear him out; and he therefore prefers the supposition that they are generated by a modification of the living substance of the plants themselves. Indeed, he regards these vegetable growths as organs, by means of which the plant gives rise to an animal, and looks upon this production of specific animals as the final cause of the galls and of, at any rate, some fruits. And he proposes to explain the occurrence of parasites within the animal body in the same way.[6]

[Footnote 6: The passage (Esperienze, p. 129) is worth quoting in full:—

"Se dovessi palesarvi il mio sentimento crederei che i frutti, i legumi, gli alberi e le foglie, in due maniere inverminassero. Una, perche venendo i bachi per di fuora, e cercando l' alimento, col rodere ci aprono la strada, ed arrivano alla piu interna midolla de' frutti e de' legni. L'altra maniera si e, che io per me stimerei, che non fosse gran fatto disdicevole il credere, che quell' anima o quella virtu, la quale genera i fiori ed i frutti nelle piante viventi, sia quella stessa che generi ancora i bachi di esse piante. E chi sa, forse, che molti frutti degli alberi non sieno prodotti, non per un fine primario e principale, ma bensi per un uffizio secondario e servile, destinato alla generazione di que' vermi, servendo a loro in vece di matrice, in cui dimorino un prefisso e determinato tempo; il quale arrivato escan fuora a godere il sole.

"Io m' immagino, che questo mio pensiero non vi parra totalmento un paradosso; mentro farete riflessione a quelle tanto sorte di galle, di gallozzole, di coccole, di ricci, di calici, di cornetti ed i lappole, che son produtte dalle quercel, dalle farnie, da' cerri, da' sugheri, da' leeci e da altri simili alberi de ghianda; imperciocche in quello gallozzole, e particolarmente nelle piu grosse, che si chiamano coronati, ne' ricci capelluti, che ciuffoli da' nostri contadini son detti; nei ricci legnosi del cerro, ne' ricci stellati della quercia, nelle galluzze della foglia del leccio si vede evidentissimamente, che la prima e principale intenzione della natura e formare dentro di quelle un animale volante; vedendosi nel centro della gallozzola un uovo, che col crescere e col maturarsi di essa gallozzola va crescendo e maturando anch' egli, e cresce altresi a suo tempo quel verme, che nell' uovo si racchiude; il qual verme, quando la gallozzola e finita di maturare e che e venuto il termine destinato al suo nascimento, diventa, di verme che era, una mosca.... Io vi confesso ingenuamente, che prima d'aver fatte queste mie esperienze intorno alla generazione degl' insetti mi dava a credere, o per dir meglio sospettava, che forse la gallozzola nascesse, perche arrivando la mosca nel tempo della primavera, e facendo una piccolissima fessura ne' rami piu teneri della quercia, in quella fessura nascondesse uno de suoi semi, il quale fosse cagione che sbocciasse fuora la gallozzola; e che mai non si vedessero galle o gallozzole o ricci o cornetti o calici o coccole, se non in que' rami, ne' quali le mosche avessero depositate le loro semenze; e mi dava ad intendere, che le gallozzole fossero una malattia cagionata nelle querce dalle punture delle mosche, in quella giusa stessa che dalle punture d'altri animaletti simiglievoli veggiamo crescere de' tumori ne' corpi degli animali."]

It is of great importance to apprehend Redi's position rightly; for the lines of thought he laid down for us are those upon which naturalists have been working ever since. Clearly, he held Biogenesis as against Abiogenesis; and I shall immediately proceed, in the first place, to inquire how far subsequent investigation has borne him out in so doing.

But Redi also thought that there were two modes of Biogenesis. By the one method, which is that of common and ordinary occurrence, the living parent gives rise to offspring which passes through the same cycle of changes as itself—like gives rise to like; and this has been termed Homogenesis. By the other mode, the living parent was supposed to give rise to offspring which passed through a totally different series of states from those exhibited by the parent, and did not return into the cycle of the parent; this is what ought to be called Heterogenesis, the offspring being altogether, and permanently, unlike the parent. The term Heterogenesis, however, has unfortunately been used in a different sense, and M. Milne-Edwards has therefore substituted for it Xenogenesis, which means the generation of something foreign. After discussing Redi's hypothesis of universal Biogenesis, then, I shall go on to ask how far the growth of science justifies his other hypothesis of Xenogenesis.

The progress of the hypothesis of Biogenesis was triumphant and unchecked for nearly a century. The application of the microscope to anatomy in the hands of Grew, Leeuwenhoek, Swammerdam, Lyonnet, Vallisnieri, Reaurnur, and other illustrious investigators of nature of that day, displayed such a complexity of organisation in the lowest and minutest forms, and everywhere revealed such a prodigality of provision for their multiplication by germs of one sort or another, that the hypothesis of Abiogenesis began to appear not only untrue, but absurd; and, in the middle of the eighteenth century, when Needham and Buffon took up the question, it was almost universally discredited.[7]

[Footnote 7: Needham, writing in 1750, says:—

"Les naturalistes modernes s'accordent unaninement a etablir, comme une verite certaine, que toute plante vient do sa semence specifique, tout animal d'un oeuf ou de quelque chose d'analogue preexistant dans la plante, ou dans l'animal de meme espece qui l'a produit."—Nouvelles Observations, p. 169.

"Les naturalistes out generalemente cru que les animaux microscopiques etaient engendres par des oeufs transportes dans l'air, ou deposes dans des eaux dormantes par des insectes volans."—Ibid. p. 176.]

But the skill of the microscope makers of the eighteenth century soon reached its limit. A microscope magnifying 400 diameters was a chef d'oeuvre of the opticians of that day; and, at the same time, by no means trustworthy. But a magnifying power of 400 diameters, even when definition reaches the exquisite perfection of our modern achromatic lenses, hardly suffices for the mere discernment of the smallest forms of life. A speck, only 1/25th of an inch in diameter, has, at ten inches from the eye, the same apparent size as an object 1/10000th of an inch in diameter, when magnified 400 times; but forms of living matter abound, the diameter of which is not more than 1/40000th of an inch. A filtered infusion of hay, allowed to stand for two days, will swarm with living things among which, any which reaches the diameter of a human red blood- corpuscle, or about 1/3200th of an inch, is a giant. It is only by bearing these facts in mind, that we can deal fairly with the remarkable statements and speculations put forward by Buffon and Needham in the middle of the eighteenth century.

When a portion of any animal or vegetable body is infused in water, it gradually softens and disintegrates; and, as it does so, the water is found to swarm with minute active creatures, the so-called Infusorial Animalcules, none of which can be seen, except by the aid of the microscope; while a large proportion belong to the category of smallest things of which I have spoken, and which must have looked like mere dots and lines under the ordinary microscopes of the eighteenth century.

Led by various theoretical considerations which I cannot now discuss, but which looked promising enough in the lights of their time, Buffon and Needham doubted the applicability of Redi's hypothesis to the infusorial animalcules, and Needham very properly endeavoured to put the question to an experimental test. He said to himself, If these infusorial animalcules come from germs, their germs must exist either in the substance infused, or in the water with which the infusion is made, or in the superjacent air. Now the vitality of all germs is destroyed by heat. Therefore, if I boil the infusion, cork it up carefully, cementing the cork over with mastic, and then heat the whole vessel by heaping hot ashes over it, I must needs kill whatever germs are present. Consequently, if Redi's hypothesis hold good, when the infusion is taken away and allowed to cool, no animalcules ought to be developed in it; whereas, if the animalcules are not dependent on pre-existing germs, but are generated from the infused substance, they ought, by and by, to make their appearance. Needham found that, under the circumstances in which he made his experiments, animalcules always did arise in the infusions, when a sufficient time had elapsed to allow for their development.

In much of his work Needham was associated with Buffon, and the results of their experiments fitted in admirably with the great French naturalist's hypothesis of "organic molecules," according to which, life is the indefeasible property of certain indestructible molecules of matter, which exist in all living things, and have inherent activities by which they are distinguished from not living matter. Each individual living organism is formed by their temporary combination. They stand to it in the relation of the particles of water to a cascade, or a whirlpool; or to a mould, into which the water is poured. The form of the organism is thus determined by the reaction between external conditions and the inherent activities of the organic molecules of which it is composed; and, as the stoppage of a whirlpool destroys nothing but a form, and leaves the molecules of the water, with all their inherent activities intact, so what we call the death and putrefaction of an animal, or of a plant, is merely the breaking up of the form, or manner of association, of its constituent organic molecules, which are then set free as infusorial animalcules.

It will be perceived that this doctrine is by no means identical with Abiogenesis, with which it is often confounded. On this hypothesis, a piece of beef, or a handful of hay, is dead only in a limited sense. The beef is dead ox, and the hay is dead grass; but the "organic molecules" of the beef or the hay are not dead, but are ready to manifest their vitality as soon as the bovine or herbaceous shrouds in which they are imprisoned are rent by the macerating action of water. The hypothesis therefore must be classified under Xenogenesis, rather than under Abiogenesis. Such as it was, I think it will appear, to those who will be just enough to remember that it was propounded before the birth of modern chemistry, and of the modern optical arts, to be a most ingenious and suggestive speculation.

But the great tragedy of Science—the slaying of a beautiful hypothesis by an ugly fact—which is so constantly being enacted under the eyes of philosophers, was played, almost immediately, for the benefit of Buffon and Needham.

Once more, an Italian, the Abbe Spallanzani, a worthy successor and representative of Redi in his acuteness, his ingenuity, and his learning, subjected the experiments and the conclusions of Needham to a searching criticism. It might be true that Needham's experiments yielded results such as he had described, but did they bear out his arguments? Was it not possible, in the first place, he had not completely excluded the air by his corks and mastic? And was it not possible, in the second place, that he had not sufficiently heated his infusions and the superjacent air? Spallanzani joined issue with the English naturalist on both these pleas, and he showed that if, in the first place, the glass vessels in which the infusions were contained were hermetically sealed by fusing their necks, and if, in the second place, they were exposed to the temperature of boiling water for three-quarters of an hour,[8] no animalcules ever made their appearance within them. It must be admitted that the experiments and arguments of Spallanzani furnish a complete and a crushing reply to those of Needham. But we all too often forget that it is one thing to refute a proposition, and another to prove the truth of a doctrine which, implicitly or explicitly, contradicts that proposition; and the advance of science soon showed that though Needham might be quite wrong, it did not follow that Spallanzani was quite right.

[Footnote 8: See Spallanzani, Opere, vi. pp. 42 and 51.]

Modern Chemistry, the birth of the latter half of the eighteenth century, grew apace, and soon found herself face to face with the great problems which biology had vainly tried to attack without her help. The discovery of oxygen led to the laying of the foundations of a scientific theory of respiration, and to an examination of the marvellous interactions of organic substances with oxygen. The presence of free oxygen appeared to be one of the conditions of the existence of life, and of those singular changes in organic matters which are known as fermentation and putrefaction. The question of the generation of the infusory animalcules thus passed into a new phase. For what might not have happened to the organic matter of the infusions, or to the oxygen of the air, in Spallanzani's experiments? What security was there that the development of life which ought to have taken place had not been checked or prevented by these changes?

The battle had to be fought again. It was needful to repeat the experiments under conditions which would make sure that neither the oxygen of the air, nor the composition of the organic matter, was altered in such a manner as to interfere with the existence of life.

Schulze and Schwann took up the question from this point of view in 1836 and 1837. The passage of air through red-hot glass tubes, or through strong sulphuric acid, does not alter the proportion of its oxygen, while it must needs arrest, or destroy, any organic matter which may be contained in the air. These experimenters, therefore, contrived arrangements by which the only air which should come into contact with a boiled infusion should be such as had either passed through red-hot tubes or through strong sulphuric acid. The result which they obtained was that an infusion so treated developed no living things, while, if the same infusion was afterwards exposed to the air, such things appeared rapidly and abundantly. The accuracy of these experiments has been alternately denied and affirmed. Supposing then, to be accepted, however, all that they really proved was that the treatment to which the air was subjected destroyed something that was essential to the development of life in the infusion. This "something" might be gaseous, fluid, or solid; that it consisted of germs remained only an hypothesis of greater or less probability.

Contemporaneously with these investigations a remarkable discovery was made by Cagniard de la Tour. He found that common yeast is composed of a vast accumulation of minute plants. The fermentation of must, or of wort, in the fabrication of wine and of beer, is always accompanied by the rapid growth and multiplication of these Toruloe. Thus, fermentation, in so far as it was accompanied by the development of microscopical organisms in enormous numbers, became assimilated to the decomposition of an infusion of ordinary animal or vegetable matter; and it was an obvious suggestion that the organisms were, in some way or other, the causes both of fermentation and of putrefaction. The chemists, with Berzelius and Liebig at their head, at first laughed this idea to scorn; but in 1843, a man then very young, who has since performed the unexampled feat of attaining to high eminence alike in Mathematics, Physics, and Physiology— I speak of the illustrious Helmholtz—reduced the matter to the test of experiment by a method alike elegant and conclusive. Helmholtz separated a putrefying or a fermenting liquid from one which was simply putrescible or fermentable by a membrane which allowed the fluids to pass through and become intermixed, but stopped the passage of solids. The result was, that while the putrescible or the fermentable liquids became impregnated with the results of the putrescence or fermentation which was going on on the other side of the membrane, they neither putrefied (in the ordinary way) nor fermented; nor were any of the organisms which abounded in the fermenting or putrefying liquid generated in them. Therefore the cause of the development of these organisms must lie in something which cannot pass through membranes; and as Helmholtz's investigations were long antecedent to Graham's researches upon colloids, his natural conclusion was that the agent thus intercepted must be a solid material. In point of fact, Helmholtz's experiments narrowed the issue to this: that which excites fermentation and putrefaction, and at the same time gives rise to living forms in a fermentable or putrescible fluid, is not a gas and is not a diffusible fluid; therefore it is either a colloid, or it is matter divided into very minute solid particles.

The researches of Schroeder and Dusch in 1854, and of Schroeder alone, in 1859, cleared up this point by experiments which are simply refinements upon those of Redi. A lump of cotton-wool is, physically speaking, a pile of many thicknesses of a very fine gauze, the fineness of the meshes of which depends upon the closeness of the compression of the wool. Now, Schroeder and Dusch found, that, in the case of all the putrefiable materials which they used (except milk and yolk of egg), an infusion boiled, and then allowed to come into contact with no air but such as had been filtered through cotton-wool, neither putrefied, nor fermented, nor developed living forms. It is hard to imagine what the fine sieve formed by the cotton-wool could have stopped except minute solid particles. Still the evidence was incomplete until it had been positively shown, first, that ordinary air does contain such particles; and, secondly, that filtration through cotton-wool arrests these particles and allows only physically pure air to pass. This demonstration has been furnished within the last year by the remarkable experiments of Professor Tyndall. It has been a common objection of Abiogenists that, if the doctrine of Biogeny is true, the air must be thick with germs; and they regard this as the height of absurdity. But nature occasionally is exceedingly unreasonable, and Professor Tyndall has proved that this particular absurdity may nevertheless be a reality. He has demonstrated that ordinary air is no better than a sort of stirabout of excessively minute solid particles; that these particles are almost wholly destructible by heat; and that they are strained off, and the air rendered optically pure, by its being passed through cotton-wool.

It remains yet in the order of logic, though not of history, to show that among these solid destructible particles, there really do exist germs capable of giving rise to the development of living forms in suitable menstrua. This piece of work was done by M. Pasteur in those beautiful researches which will ever render his name famous; and which, in spite of all attacks upon them, appear to me now, as they did seven years ago,[9] to be models of accurate experimentation and logical reasoning. He strained air through cotton-wool, and found, as Schroeder and Dusch had done, that it contained nothing competent to give rise to the development of life in fluids highly fitted for that purpose. But the important further links in the chain of evidence added by Pasteur are three. In the first place he subjected to microscopic examination the cotton-wool which had served as strainer, and found that sundry bodies clearly recognisable as germs, were among the solid particles strained off. Secondly, he proved that these germs were competent to give rise to living forms by simply sowing them in a solution fitted for their development. And, thirdly, he showed that the incapacity of air strained through cotton- wool to give rise to life, was not due to any occult change effected in the constituents of the air by the wool, by proving that the cotton-wool might be dispensed with altogether, and perfectly free access left between the exterior air and that in the experimental flask. If the neck of the flask is drawn out into a tube and bent downwards; and if, after the contained fluid has been carefully boiled, the tube is heated sufficiently to destroy any germs which may be present in the air which enters as the fluid cools, the apparatus may be left to itself for any time and no life will appear in the fluid. The reason is plain. Although there is free communication between the atmosphere laden with germs and the germless air in the flask, contact between the two takes place only in the tube; and as the germs cannot fall upwards, and there are no currents, they never reach the interior of the flask. But if the tube be broken short off where it proceeds from the flask, and free access be thus given to germs falling vertically out of the air, the fluid, which has remained clear and desert for months, becomes, in a few days, turbid and full of life.

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