secret of thy knowledge, as thou didst before the creation of the firmament, the division of light from darkness; let the children of thy spirit, placed in their firmament, make their light shine upon the earth, mark the division of night and day, and announce the revolution of the times; for the old order

is passed, and the new arises; the night is spent, the day is come forth; and thou shalt crown the year with thy blessing, when thou shalt send forth laborers into thy harvest sown by other hands than theirs; when thou shalt send forth new laborers to new seedtimes, whereof the harvest shall be not yet."

THE PHYSICAL BASIS OF LIFE

THOMAS HENRY HUXLEY

[From an address delivered in Edinburgh in 1868; and published in "Lay Sermons," 1870.]

In order to make the title of this discourse generally intelligible, I have translated the term "Protoplasm," which is the scientific name of the substance of which I am about to speak, by the words "the physical basis of life." I suppose that, to many, the idea that there is such a thing as a physical basis, or matter, of life may be novelso widely spread is the conception of life as a something which works through matter, but is independent of it; and even those who are aware that matter and life are inseparably connected, may not be prepared for the conclusion plainly suggested by the phrase, "the physical basis or matter of life," that there is some one kind of matter which is common to all living beings, and that their endless diversities are bound together by a physical, as well as an ideal, unity. In fact, when first apprehended, such a doctrine as this appears almost shocking to common

sense.

What, truly, can seem to be more obviously different from one another, in faculty, in form, and in substance, than the various kinds of living beings? What community of faculty can there be between the brightlycolored lichen, which so nearly resembles a mere mineral incrustation of the bare rock on which it grows, and the painter, to whom it is instinct with beauty, or the botanist, whom it feeds with knowledge?

Again, think of the microscopic fungusa mere infinitesimal ovoid particle, which finds space and duration enough to multiply into countless millions in the body of a liv

ing fly; and then of the wealth of foliage, the luxuriance of flower and fruit, which lies between this bald sketch of a plant and the giant pine of California, towering to the dimensions of a cathedral spire, or the Indian fig, which covers acres with its profound shadow, and endures while nations and empires come and go around its vast circumference. Or, turning to the other half of the world of life, picture to yourselves the great Finner whale, hugest of beasts that live, or have lived, disporting his eighty or ninety feet of bone, muscle, and blubber, with easy roll, among waves in which the stoutest ship that ever left dockyard would flounder hopelessly; and contrast him with the invisible animalcules-mere gelatinous specks, multitudes of which could, in fact, dance upon the point of a needle with the same ease as the angels of the Schoolmen could, in imagination. With these images before your minds, you may well ask, what community of form, or structure, is there between the animalcule and the whale; or between the fungus and the fig-tree? And, à fortiori, between all four?

Finally, if we regard substance, or material composition, what hidden bond can connect the flower which a girl wears in her hair and the blood which courses through her youthful veins; or, what is there in common between the dense and resisting mass of the oak, or the strong fabric of the tortoise, and those broad disks of glassy jelly which may be seen pulsating through the waters of a calm sea, but which drain away to mere films in the hand which raises them out of their element?

Such objections as these must, I think, arise in the mind of every one who ponders, for the first time, upon the conception of a single physical basis of life underlying all the diversities of vital existence; but I pro

pose to demonstrate to you that, notwithstanding these apparent difficulties, a threefold unity-namely, a unity of power or faculty, a unity of form, and a unity of substantial composition-does pervade the whole living world.

No very abstruse argumentation is needed, in the first place, to prove that the powers, or faculties, of all kinds of living matter, diverse as they may be in degree, are substantially similar in kind.

Goethe has condensed a survey of all powers of mankind into the well-known epigram:

"Warum treibt sich das Volk so und schreit? Es will sich ernähren, Kinder zeugen, und die nähren so gut es vermag.

Weiter bringt es kein Mensch, stell' er sich wie er auch will." 1

In physiological language this means, that all the multifarious and complicated activities of man are comprehensible under three categories. Either they are immediately directed towards the maintenance and development of the body, or they effect transitory changes in the relative positions of parts of the body, or they tend towards the continuance of the species. Even those manifestations of intellect, of feeling, and of will, which we rightly name the higher faculties, are not excluded from this classification, inasmuch as to every one but the subject of them, they are known only as transitory changes in the relative positions of parts of the body. Speech, gesture, and every other form of human action are, in the long run, resolvable into muscular contraction, and muscular contraction is but a transitory change in the relative positions of the parts of a muscle. But the scheme which is large enough to embrace the activities of the highest form of life, covers all those of the lower creatures. The lowest plant, or animalcule, feeds, grows, and reproduces its kind. In addition, all animals manifest those transitory changes of form which we class under irritability and contractility; and it is more than probable that when the vegetable world is thoroughly explored, we shall find all plants in possession of the same powers, at

1 Why does the populace rush so and make clamor? It wishes to eat, bring forth children, and feed these as well as it may. No man can do better, strive how he will.

one time or other of their existence. I am not now alluding to such phenomena, at once rare and conspicuous, as those exhibited by the leaflets of the sensitive plants, or the stamens of the barberry, but to much more widely spread, and at the same time, more subtle and hidden, manifestations of vegetable contractility. You are doubtless aware that the common nettle owes its stinging property to the innumerable stiff and needle-like, though exquisitely delicate, hairs which cover its surface. Each stingingneedle tapers from a broad base to a slender summit, which, though rounded at the end, is of such microscopic fineness that it readily penetrates, and breaks off in, the skin. The whole hair consists of a very delicate outer case of wood, closely applied to the inner surface of which is a layer of semifluid matter, full of innumerable granules of extreme minuteness. This semi-fluid lining is protoplasm, which thus constitutes a kind of bag, full of a limpid liquid, and roughly corresponding in form with the interior of the hair which it fills. When viewed with a sufficiently high magnifying power, the protoplasmic layer of the nettle hair is seen to be in a condition of unceasing activity. Local contractions of the whole thickness of its substance pass slowly and gradually from point to point, and give rise to the appearance of progressive waves, just as the bending of successive stalks of corn by a breeze produces the apparent billows of a corn-field.

But, in addition to these movements, and independently of them, the granules are driven, in relatively rapid streams, through channels in the protoplasm which seem to have a considerable amount of persistence. Most commonly, the currents in adjacent parts of the protoplasm take similar directions; and, thus, there is a general stream up one side of the hair and down the other. But this does not prevent the existence of partial currents which take different routes; and sometimes trains of granules may be seen coursing swiftly in opposite directions within a twenty-thousandth of an inch of one another; while, occasionally, opposite streams come into direct collision, and, after a longer or shorter struggle, one predominates. The cause of these currents seems to lie in contractions of the protoplasm which bounds the channels in which they flow, but which are so minute that the best microscopes show only their effects, and not themselves.

The spectacle afforded by the wonderful energies prisoned within the compass of the microscopic hair of a plant, which we commonly regard as a merely passive organism, is not easily forgotten by one who has watched its display, continued hour after hour, without pause or sign of weakening. The possible complexity of many other organic forms, seemingly as simple as the protoplasm of the nettle, dawns upon one; and the comparison of such a protoplasm to a body with an internal circulation, which has been put forward by an eminent physiologist, loses much of its startling character. Currents similar to those of the hairs of the nettle have been observed in a great multitude of very different plants, and weighty authorities have suggested that they probably occur, in more or less perfection, in all young vegetable cells. If such be the case, the wonderful noonday silence of a tropical forest is, after all, due only to the dullness of our hearing; and could our ears catch the murmur of these tiny Maelstroms, as they whirl in the innumerable myriads of living cells which constitute each tree, we should be stunned, as with the roar of a great city.

Among the lower plants, it is the rule rather than the exception, that contractility should be still more openly manifested at some periods of their existence. The protoplasm of Alga and Fungi becomes, under many circumstances, partially, or completely, freed from its woody case, and exhibits movements of its whole mass, or is propelled by the contractility of one, or more, hair-like prolongations of its body, which are called vibratile cilia. And, so far as the conditions of the manifestation of the phenomena of contractility have yet been studied, they are the same for the plant as for the animal. Heat and electric shocks influence both, and in the same way, though it may be in different degrees. It is by no means my intention to suggest that there is no difference in faculty between the lowest plant and the highest, or between plants and animals. But the difference between the powers of the lowest plant, or animal, and those of the highest, is one of degree, not of kind, and depends, as MilneEdwards long ago so well pointed out, upon the extent to which the principle of the division of labor is carried out in the living economy. In the lowest organism all parts are competent to perform all functions, and one and the same portion of protoplasm

may successfully take on the function of feeding, moving, or reproducing apparatus. In the highest, on the contrary, a great number of parts combine to perform each function, each part doing its allotted share of the work with great accuracy and efficiency, but being useless for any other purpose.

On the other hand, notwithstanding all the fundamental resemblances which exist between the powers of the protoplasm in plants and in animals, they present a striking difference (to which I shall advert more at length presently), in the fact that plants can manufacture fresh protoplasm out of mineral compounds, whereas animals are obliged to procure it ready made, and hence, in the long run, depend upon plants. Upon what condition this difference in the powers of the two great divisions of the world of life depends, nothing is at present known.

With such qualifications as arise out of the last-mentioned fact, it may be truly said that the acts of all living things are fundamentally one. Is any such unity predicable of their forms? Let us seek in easily verified facts for a reply to this question. If a drop of blood be drawn by pricking one's finger, and viewed with proper precautions, and under a sufficiently high microscopic power, there will be seen, among the innumerable multitude of little, circular, discoidal bodies, or corpuscles, which float in it and give it its color, a comparatively small number of colorless corpuscles, of somewhat larger size and very irregular shape. If the drop of blood be kept at the temperature of the body, these colorless corpuscles will be seen to exhibit a marvellous activity, changing their forms with great rapidity, drawing in and thrusting out prolongations of their substance, and creeping about as if they were independent organisms.

The substance which is thus active is a mass of protoplasm, and its activity differs in detail, rather than in principle, from that of the protoplasm of the nettle. Under sundry circumstances the corpuscle dies and becomes distended into a round mass, in the midst of which is seen a smaller spherical body, which existed, but was more or less hidden, in the living corpuscle, and is called its nucleus. Corpuscles of essentially similar structure are to be found in the skin, in the lining of the mouth, and scattered through the whole framework of

the body. Nay, more: in the earliest condition of the human organism, in that state in which it has but just become distinguishable from the egg in which it arises, it is nothing but an aggregation of such corpuscles, and every organ of the body was, once, no more than such an aggregation.

Thus a nucleated mass of protoplasm turns out to be what may be termed the structural unit of the human body. As a matter of fact, the body, in its earliest state, is a mere multiple of such units; and in its perfect condition, it is a multiple of such units, variously modified.

But does the formula which expresses the essential structural character of the highest animal cover all the rest, as the statement of its powers and faculties covered that of all others? Very nearly. Beast and fowl, reptile and fish, mollusk, worm, and polyp, are all composed of structural units of the same character, namely, masses of protoplasm with a nucleus. There are sundry very low animals, each of which, structurally, is a mere colorless blood-corpuscle, leading an independent life.

But, at the very bottom of the animal scale, even this simplicity becomes simplified, and all the phenomena of life are manifested by a particle of protoplasm without a nucleus. Nor are such organisms insignificant by reason of their want of complexity. It is a fair question whether the protoplasm of those simplest forms of life, which people an immense extent of the bottom of the sea, would not outweigh that of all the higher living beings which inhabit the land put together. And in ancient times, no less than at the present day, such living beings as these have been the greatest of rock builders.

What has been said of the animal world is no less true of plants. Imbedded in the protoplasm at the broad, or attached, end of the nettle hair, there lies a spheroidal nucleus. Careful examination further proves that the whole substance of the nettle is made up of a repetition of such masses of nucleated protoplasm, each contained in a wooden case, which is modified in form, sometimes into a woody fiber, sometimes into a duct or spiral vessel, sometimes into a pollen grain, or an ovule. Traced back to its earliest state, the nettle arises as the man does, in a particle of nucleated protoplasm. And in the lowest plants, as in the lowest animals, a single mass of such protoplasm may constitute the whole plant,

or the protoplasm may exist without a nucleus.

Under these circumstances it may well be asked, how is one mass of non-nucleated protoplasm to be distinguished from another? why call one "plant" and the other "animal"?

The only reply is that, so far as form is concerned, plants and animals are not separable, and that, in many cases, it is a mere matter of convention whether we call a given organism an animal or a plant. There is a living body called Ethalium septicum, which appears upon decaying vegetable substances, and, in one of its forms, is common upon the surfaces of tan-pits. In this condition it is, to all intents and purposes, a fungus, and formerly was always regarded as such; but the remarkable investigations of De Bary have shown that, in another condition, the Ethalium is an actively locomotive creature, and takes in solid matters, upon which, apparently, it feeds, thus exhibiting the most characteristic feature of animality. Is this a plant; or is it an animal? Is it both; or is it neither? Some decide in favor of the last supposition, and establish an intermediate kingdom, a sort of biological No Man's Land for all these questionable forms. But, as it is admittedly impossible to draw any distinct boundary line between this no man's land and the vegetable world on the one hand, or the animal on the other, it appears to me that this proceeding merely doubles the difficulty which, before, was single.

Protoplasm, simple or nucleated, is the formal basis of all life. It is the clay of the potter: which, bake it and paint it as he will, remains clay, separated by artifice, and not by nature, from the commonest brick or sun-dried clod.

Thus it becomes clear that all living powers are cognate, and that all living forms are fundamentally of one character. The researches of the chemist have revealed a no less striking uniformity of material composition in living matter.

In perfect strictness, it is true that chemical investigation can tell us little or nothing, directly, of the composition of living matter, inasmuch as such matter must needs die in the act of analysis, and upon this very obvious ground, objections, which I confess seem to me to be somewhat frivolous, have been raised to the drawing of any conclusions whatever respecting the composition of actually living matter, from that of

the dead matter of life, which alone is accessible to us. But objectors of this class do not seem to reflect that it is also, in strictness, true that we know nothing about the composition of any body whatever, as it is. The statement that a crystal of calcspar consists of carbonate of lime, is quite true, if we only mean that, by appropriate processes, it may be resolved into carbonic acid and quicklime. If you pass the same carbonic acid over the very quicklime thus obtained, you will obtain carbonate of lime again; but it will not be calc-spar, nor anything like it. Can it, therefore, be said that chemical analysis teaches nothing about the chemical composition of calc-spar? Such a statement would be absurd; but it is hardly more so than the talk one occasionally hears about the uselessness of applying the results of chemical analysis to the living bodies which have yielded them.

To this

One fact, at any rate, is out of reach of such refinements, and this is, that all the forms of protoplasm which have yet been examined contain the four elements, carbon, hydrogen, oxygen, and nitrogen, in very complex union, and that they behave similarly towards several reagents. complex combination, the nature of which has never been determined with exactness, the name of Protein has been applied. And if we use this term with such caution as may properly arise out of our comparative ignorance of the things for which it stands, it may be truly said that all protoplasm is proteinaceous, or, as the white, or albumen, of an egg is one of the commonest examples of a nearly pure protein matter, we may say that all living matter is more or less albuminoid.

Perhaps it would not yet be safe to say that all forms of protoplasm are affected by the direct action of electric shocks; and yet the number of cases in which the contraction of protoplasm is shown to be affected by this agency increases every day.

Nor can it be affirmed with perfect confidence, that all forms of protoplasm are liable to undergo that peculiar coagulation at a temperature of 40°-50° Centigrade, which has been called "heat-stiffening," though Kühne's beautiful researches have proved this occurrence to take place in so many and such diverse living beings, that it is hardly rash to expect that the law holds good for all.

Enough has, perhaps, been said, to prove the existence of a general uniformity in the

character of the protoplasm, or physical basis, of life, in whatever group of living beings it may be studied. But it will be understood that this general uniformity by no means excludes any amount of special modifications of the fundamental substance. The mineral, carbonate of lime, assumes an immense diversity of characters, though no one doubts that, under all these Protean changes, is one and the same thing.

And now, what is the ultimate fate, and what the origin, of the matter of life?

Is it, as some of the older naturalists supposed, diffused throughout the universe in molecules, which are indestructible and unchangeable in themselves; but, in endless transmigration, unite in innumerable permutations, into the diversified forms of life we know? Or, is the matter of life composed of ordinary matter, differing from it only in the manner in which its atoms are aggregated? Is it built up of ordinary matter, and again resolved into ordinary matter when its work is done?

Modern science does not hesitate a moment between these alternatives. Physiology writes over the portals of life

"Debemur morti nos nostraque,"

991

with a profounder meaning than the Roman poet attached to the melancholy line. Under whatever disguise it takes refuge, whether fungus or oak, worm or man, the living protoplasm not only ultimately dies and is resolved into its mineral and lifeless constituents, but is always dying, and, strange as the paradox may sound, could not live unless it died.

In the wonderful story of the Peau de Chagrin, the hero becomes possessed of a magical wild ass' skin, which yields him the means of gratifying all his wishes. But its surface represents the duration of the proprietor's life; and for every satisfied desire the skin shrinks in proportion to the intensity of fruition, until at length life and the last handbreadth of the peau de chagrin, disappear with the gratification of a last wish.

Balzac's studies had led him over a wide range of thought and speculation, and his shadowing forth of physiological truth in this strange story may have been intentional. At any rate, the matter of life is a veritable peau de chagrin, and for every vital act it is somewhat the smaller. All work im1 We and ours must die.