Popular Science Monthly/Volume 15/October 1879/Protoplasm and Life

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OCTOBER, 1879.

By Professor G. J. ALLMAN, LL. D., F. R. S.,

MORE than forty years have now passed away since the French naturalist, Dujardin, drew attention to the fact that the bodies of some of the lowest members of the animal kingdom consist of a structureless, semi-fluid, contractile substance, to which he gave the name of Sarcode. A similar substance occurring in the cells of plants was afterward studied by Hugo von Mohl, and named by him Protoplasm. It remained for Max Schultze to demonstrate that the sarcode of animals and the protoplasm of plants were identical.

The conclusions of Max Schultze have been in all respects confirmed by subsequent research, and it has further been rendered certain that this same protoplasm lies at the base of all the phenomena of life, whether in the animal or the vegetable kingdom. Thus has arisen the most important and significant generalization in the whole domain of biological science.

Within the last few years protoplasm has again been made a subject of special study; unexpected and often startling facts have been brought to light, and a voluminous literature has gathered round this new center of research. I believe, therefore, that I can not do better than call your attention to some of the more important results of these inquiries, and endeavor to give you some knowledge of the properties of protoplasm, and of the part it plays in the two great kingdoms of organic nature.

As has just been said, protoplasm lies at the base of every vital phenomenon. It is, as Huxley has well expressed it, "the physical basis of life." Wherever there is life, from its lowest to its highest manifestations, there is protoplasm; wherever there is protoplasm, there, too, is life. Thus, coextensive with the whole of organic nature —every vital act being referable to some mode or property of protoplasm—it becomes to the biologist what the ether is to the physicist; only that instead of being an hypothetical conception, accepted as a reality from its adequacy in the explanation of phenomena, it is a tangible and visible reality, which the chemist may analyze in his laboratory, the biologist scrutinize beneath his microscope and his dissecting needle.

The chemical composition of protoplasm is very complex, and has not been exactly determined. It may, however, be stated that protoplasm is essentially a combination of albuminoid bodies, and that its principal elements are, therefore, oxygen, carbon, hydrogen, and nitrogen. In its typical state it presents the condition of a semi-fluid substance—a tenacious, glairy liquid, with a consistence somewhat like that of the white of an unboiled egg[2] While we watch it beneath the microscope, movements are set up in it: waves traverse its surface, or it may be seen to flow away in streams, either broad and attaining but a slight distance from the main mass, or else stretching away far from their source, as narrow liquid threads, which may continue simple, or may divide into branches, each following its own independent course; or the streams may flow one into the other, as streamlets would flow into rivulets and rivulets into rivers, and this not only where gravity would carry them, but in a direction diametrically opposed to gravitation: now we see it spreading itself out on all sides into a thin liquid stratum, and again drawing itself together within the narrow limits which had at first confined it, and all this without any obvious impulse from without which would send the ripples over its surface or set the streams flowing from its margin. Though it is certain that all these phenomena are in response to some stimulus exerted on it by the outer world, they are such as we never meet with in a simply physical fluid—they are spontaneous movements resulting from its proper irritability, from its essential constitution as living matter.

Examine it closer, bring to bear on it the highest powers of your microscope—you will probably find disseminated through it countless multitudes of exceedingly minute granules; but you may also find it absolutely homogeneous, and, whether containing granules or not, it is certain that you will find nothing to which the term organization can be applied. You have before you a glairy, tenacious fluid, which, if not absolutely homogeneous, is yet totally destitute of structure.

And yet no one who contemplates this spontaneously moving matter can deny that it is alive. Liquid as it is, it is a living liquid; organless and structureless as it is, it manifests the essential phenomena of life.

The picture which I have thus endeavored to trace for you in a few leading outlines is that of protoplasm in its most generalized aspect. Such generalizations, however, are in themselves unable to satisfy the conditions demanded by an exact scientific inquiry, and I propose now, before passing to the further consideration of the place and purport of protoplasm in nature, to bring before you some definite examples of protoplasm, such as are actually met with in the organic world.

A quantity of a peculiar slimy matter was dredged in the North Atlantic by the naturalists of the exploring-ship Porcupine from a depth of from 5,000 to 25,000 feet. It is described as exhibiting, when examined on the spot, spontaneous movements, and as being obviously endowed with life. Specimens of this, preserved in spirits, were examined by Professor Huxley, and declared by him to consist of protoplasm, vast masses of which must thus in a living state extend over wide areas of sea-bottom. To this wonderful slime Huxley gave the name of Bathybius Haeckelii.

Bathybius has since been subjected to an exhaustive examination by Professor Haeckel, who believes that he is able to confirm in all points the conclusions of Huxley, and arrives at the conviction that the bottom of the open ocean at depths below 5,000 feet is covered with an enormous mass of living protoplasm, which lingers there in the simplest and most primitive condition, having as yet acquired no definite form. He suggests that it may have originated by spontaneous generation, but leaves this question for future investigations to decide.

The reality of Bathybius, however, has not been universally accepted. In the more recent investigations of the Challenger the explorers have failed in their attempts to bring further evidence of the existence of masses of amorphous protoplasm spreading over the bed of the ocean. They have met with no trace of Bathybius in any of the regions explored by them, and they believe that they are justified in the conclusion that the matter found in the dredgings of the Porcupine and preserved in spirits for further examination, was only an inorganic precipitate due to the action of the alcohol.

It is not easy to believe, however, that the very elaborate investigations of Huxley and Haeckel can be thus disposed of. These, moreover, have received strong confirmation from the still more recent observations of the Arctic voyager Bessels, who was one of the explorers of the ill-fated Polaris, and who states that he dredged from the Greenland seas masses of living undifferentiated protoplasm. Bessels assigns to these the name of Protobathyhius, but they are apparently indistinguishable from the Bathybius of the Porcupine. Further arguments against the reality of Bathybius will therefore be needed before a doctrine founded on observations so carefully conducted shall be relegated to the region of confuted hypotheses.

Assuming, then, that Bathybius, however much its supposed wide distribution may have been limited by more recent researches, has a real existence, it presents us with a condition of living matter the most rudimental it is possible to conceive. No law of morphology has as yet exerted itself in this formless slime. Even the simplest individualization is absent. We have a living mass, but we. know not where to draw its boundary-lines; it is living matter, but we can scarcely call it a living being.

We are not, however, confined to Bathybius for examples of protoplasm in a condition of extreme simplicity. Haeckel has found, inhabiting the fresh waters in the neighborhood of Jena, minute lumps of protoplasm which, when placed under the microscope, were seen to have no constant shape, their outline being in a state of perpetual change, caused by the protrusion from various parts of their surface of broad lobes and thick, finger-like projections, which, after remaining visible for a time, would be withdrawn, to make their appearance again on some other part of the surface.

These changeable protrusions of its substance, without fixed position or definite form, are eminently characteristic of protoplasm in some of its simplest conditions. They have been termed "Pseudopodia," and will frequently come before you in what I have yet to say.

To the little protoplasmic lumps thus constituted Haeckel bas given the name of Protamœba primitiva. They may be compared to minute detached pieces of Bathybius. He has seen them multiplying themselves by spontaneous division into two pieces, which, on becoming independent, increase in size, and acquire all the characters of the parent.

Several other beings as simple as Protamœba have been described by various observers, and especially by Haeckel, who brings the whole together into a group to which he gives the name of Monera, suggested by the extreme simplicity of the beings included in it.

But we must now pass to a stage a little higher in the development of protoplasmic beings. Widely distributed in the fresh and salt waters of Britain, and probably of almost all parts of the world, are small particles of protoplasm closely resembling the Protamœba just described. Like it, they have no definite shape, and are perpetually changing their form, throwing out and drawing in thick lobes and finger-like pseudopodia, in which their body seems to flow away over the field of the microscope. They are no longer, however, the homogeneous particle of protoplasm which forms the body of Protamœba. Toward the center a small globular mass of firmer protoplasm has become differentiated off from the remainder, and forms what is known as a nucleus, while the protoplasm forming the extreme outer boundary differs slightly from the rest, being more transparent, destitute of granules, and apparently somewhat firmer than the interior. We may also notice that at one spot a clear spherical space has made its appearance, but that while we watch it has suddenly contracted and vanished, and after a few seconds has begun to dilate so as again to come into view, once more to disappear, then again to return, and all this in regular rhythmical sequence. This little rhythmically pulsating cavity is called the "contractile vacuole." It is of very frequent occurrence among those beings which lie low down in the scale of life.

We have now before us a being which has arrested the attention of naturalists almost from the commencement of microscopical observation. It is the famous Amœba, for which ponds and pools and gutters on the house-roof have for the last two hundred years been ransacked by the microscopist, who has many a time stood in amazement at the undefinable form and Protean changes of this particle of living matter. It is only the science of our own days, however, which has revealed its biological importance, and shown that in this little soft, nucleated particle we have a body whose significance for the morphology and physiology of living beings can not be over-estimated, for in Amœba we have the essential characters of a cell, the morphological unit of organization, the physiological source of specialized function.

The term "cell" has been so Ions' in use that it can not now be displaced from our terminology; and yet it tends to convey an incorrect notion, suggesting as it does the idea of a hollow body or vesicle, this having been the form under which it was first studied. The cell, however, is essentially a definite mass of protoplasm having a nucleus imbedded in it. It may or may not assume the form of a vesicle; it may or may not be protected by an enveloping membrane; it may or may not contain a contractile vacuole; and the nucleus may or may not contain within it one or more minute secondary nuclei or "nucleoli."

Haeckel has done good service to biology in insisting on the necessity of distinguishing such non-nucleated forms as are presented by Protamœba and the other Monera from the nucleated forms as seen in Amœba. To the latter he would restrict the word cell, while he would assign that of "cytode" to the former.

Let us observe our Amœba a little closer. Like all living beings, it must be nourished. It can not grow as a crystal would grow by accumulating on its surface molecule after molecule of matter. It must feed. It must take into its substance the necessary nutriment; it must assimilate this nutriment, and convert it into the material of which it is itself composed.

If we seek, however, for a mouth by which the nutriment can enter into its body, or a stomach by which this nutriment can be digested, we seek in vain. Yet watch it for a moment as it lies in a drop of water beneath our microscope. Some living denizen of the same drop is in its neighborhood, and its presence exerts on the protoplasm of the Amœba a special stimulus which gives rise to the movements necessary for the prehension of nutriment. A stream of protoplasm instantly runs away from the body of the Amœba toward the destined prey, envelops it in its current, and then flows back with it to the central protoplasm, where it sinks deeper and deeper into the soft, yielding mass, and becomes dissolved, digested, and assimilated in. order that it may increase the size and restore the energy of its captor.

But again, like all living things, Amœba must multiply itself, and so after attaining a certain size its nucleus divides into two halves, and then the surrounding protoplasm becomes similarly cleft, each half retaining one half of the original nucleus. The two new nucleated masses which thus arise now lead an independent life, assimilate nutriment, and attain the size and characters of the parent.

We have just seen that in the body of an Amœba we have the type of a cell. Now, both the fresh waters and the sea contain many living beings besides Amœba which never pass beyond the condition of a simple cell. Many of these, instead of emitting the broad, lobelike pseudopodia of Amoeba, have the faculty of sending out long, thin threads of protoplasm, which they can again retract, and by the aid of which they capture their prey or move from place to place. Simple structureless protoplasm as they are, many of them fashion for themselves an outer membranous or calcareous case, often of symmetrical form and elaborate ornamentation, or construct a silicious skeleton of radiating spicula, or crystal clear concentric spheres of exquisite symmetry and beauty.

Some move about by the aid of a flagellum, or long whip-like projection of their bodies, by which they lash the surrounding waters, and which, unlike the pseudopodia of Amœba, can not, during active life, be withdrawn into the general protoplasm of the body; while among many others locomotion is effected by means of cilia—microscopic vibratile hairs, which are distributed in various ways over the surface, and which, like the pseudopodia and flagella, are simple prolongations of their protoplasm.

In every one of these cases the entire body has the morphological value' of a cell, and in this simple cell reside the whole of the properties which manifest themselves in the vital phenomena of the organism.

The part fulfilled by these simple unicellular beings in the economy of nature has at all times been very great, and many geological formations, largely built up of their calcareous or silicious skeletons, bear testimony to the multitudes in which they must have swarmed in the waters of the ancient earth.

Those which have thus come down to us from ancient times owe their preservation to the presence of the hard, persistent structures secreted by their protoplasm, and must, after all, have formed but a very small proportion of the unicellular organisms which peopled the ancient world, and there fulfilled the duties allotted to them in nature, but whose soft, perishable bodies have left no trace behind.

In our own days similar unicellular organisms are at work, taking their part silently and unobtrusively in the great scheme of creation, and mostly destined, like their predecessors, to leave behind them no record of their existence. The Red-Snow plant, to which is mainly due the beautiful phenomenon by which tracts of Arctic and Alpine snow become tinged of a delicate crimson, is a microscopic organism whose whole body consists of a simple spherical cell. In the protoplasm of this little cell must reside all the essential attributes of life; it must grow by the reception of nutriment; it must repeat by multiplication that form which it has itself inherited from its parent; it must be able to respond to the stimulus of the physical conditions by which it is surrounded. And there it is, with its structure almost on the bounds of extremest simplification, taking its allotted part in the economy of nature, combining into living matter the lifeless elements which lie around it, redeeming from sterility the regions of never-thawing ice, and peopling with its countless millions the wastes of the snow-land.[3]

But organization does not long rest on this low stage of unicellular simplicity, for, as we pass from these lowest forms into higher, we find cell added to cell, until many millions of such units become associated in a single organism, where each cell, or each group of cells, has its own special work, while all combine for the welfare and unity of the whole.

In the most complex animals, however, even in man himself, the component cells, notwithstanding their frequent modification and the usual intimacy of their union, are far from losing their individuality. Examine under the microscope a drop of blood freshly taken from the human subject, or from any of the higher animals. It is seen to be composed of a multitude of red corpuscles, swimming in a nearly colorless liquid, and along with these, but in much smaller numbers, somewhat larger colorless corpuscles. The red corpuscles are modified cells, while the colorless corpuscles are cells still retaining their typical form and properties. These last are little masses of protoplasm, each enveloping a central nucleus. Watch them. They will be seen to change their shape; they will project and withdraw pseudopodia, and creep about like an Amœba. But, more than this, like an Amœba, they will take in solid matter as nutriment. They may be fed with colored food, which will then be seen to have accumulated in the interior of their soft, transparent protoplasm; and in some cases the colorless blood-corpuscles have actually been seen to devour their more diminutive companions, the red ones.

Again, there are certain cells filled with peculiar colored matters, and called pigment-cells, which are especially abundant, as constituents of the skin, in fishes, frogs, and other low vertebrate, as well as many invertebrate, animals. Under certain stimuli, such as that of light, or of emotion, these pigment-cells change their form, protrude or retract pseudopodial prolongations of their protoplasm, and assume the form of stars or of irregularly lobed figures, or again draw themselves together into little globular masses. To this change of form in the pigment-cell the rapid change of color, so frequently noticed in the animals provided with them, is to be attributed.

The animal egg, which in its young state forms an element in the structure of the parent organism, possesses in the relations now under consideration a peculiar interest. The egg is a true cell, consisting essentially of a lump of protoplasm inclosing a nucleus, and having a nucleolus included in the interior of the nucleus. While still very young it has no constant form, and is perpetually changing its shape. Indeed, it is often impossible to distinguish it from the Amœba; and it may, like an Amœba, wander from place to place by the aid of its pseudopodial projections. I have shown elsewhere[4] that the primitive egg of the remarkable hydroid Myriothela manifests amœboid motions; while Haeckel has shown[5] that in the sponges certain Amœba-like organisms, which are seen wandering about in the various canals and cavities of their bodies, and had been until lately regarded as parasites which had gained access from without, are really the eggs of the sponge; and a similar amŒboid condition is presented by the very young eggs of even the highest animals.

Again, Reichenbach has proved[6] that during the development of the crawfish the cells of the embryo throw out pseudopodia by which, exactly as in an Amœba, the yolk-spheres, which serve as nutriment for the embryo, are surrounded and ingulfed in the protoplasm of the cells.

I had shown some years ago[7] that in Myriothela pseudopodial processes are being constantly projected from the walls of the alimentary canal into its cavity. They appear as direct extensions of a layer of clear, soft, homogeneous protoplasm, which lies over the surface of the naked cells lining the cavity, and which I now regard as the "Hautschicht," or cortical layer, of these cells. I then suggested that the function of these pseudopodia lay in seizing, in the manner of an Amœba, such alimentary matter as may be found in the contents of the canal, and applying it to the nutrition of the hydroid.

What I had thus suggested with regard to Myriothela has been since proved in certain planarian worms by Metschnikoff,[8] who has seen the cells which line the alimentary canal in these animals act like independent Amœbæ, and ingulf in their protoplasm such solid nutriment as may be contained in the canal. When the planaria was fed with coloring matter these amœboid cells became gorged with the colored particles just as would have happened in an Amœba when similarly fed.

But it is not alone in such loosely aggregated cells as those of the blood, or in the amœboid cells of the alimentary canal, or in such scattered constituents of the tissues as the pigment-cells, or in cells destined for an ultimate state of freedom, as the egg, that there exists an independence. The whole complex organism is a society of cells, in which every individual cell possesses an independence, an autonomy, not at once so obvious as in the blood-cells, but not the least real. With this autonomy of each element there is at the same time a subordination of each to the whole, thus establishing a unity in the entire organism, and a concert and harmony between all the phenomena of its life.

In this society of cells each has its own work to perform, and the life of the organism is made up of the lives of its component cells. Here it is that we find most distinctly expressed the great law of the physiological division of labor. In the lowest organisms, where the whole being consists of a single cell, the performance of all the processes which constitute its life must devolve on the protoplasm of this one cell; but as we pass to more highly organized beings, the work becomes distributed among a multitude of workers. These workers are the cells which now make up the complex organism. The distribution of labor, however, is not a uniform one, and we are not to suppose that the work performed by each cell is but a repetition of that of every other. For the life-processes, which are accumulated in the single cell of the unicellular organism become in the more complex organism differentiated, some being intensified and otherwise modified and allocated to special cells, or to special groups of cells, which we call organs, and whose proper duty is now to take charge of the special processes which have been assigned to them. In all this we have a true division of labor—a division of labor, however, by no means absolute; for the processes which are essential to the life of the cell must still continue common to all the cells of the organism. No cell, however great may be the differentiation of function in the organism, can dispense with its irritability, the one constant and essential property of every living cell. There thus devolves on each cell or group of cells some special work which contributes to the well-being of all, and their combined labors secure the necessary conditions of life for every cell in the community, and result in those complex and wonderful phenomena which constitute the life of the higher organisms.

We have hitherto considered the cell only as a mass of active nucleated protoplasm, either absolutely naked, or partially inclosed in a protective case, which still permits free contact of the protoplasm with the surrounding medium. In very many instances, however, the protoplasm becomes confined within resisting walls, which entirely shut it in from all direct contact with the medium which surrounds it. With the plant this is almost always so after the earliest stages of its life. Here the protoplasm of the cells is endowed with the faculty of secreting over its surface a firm, resisting membrane, composed of cellulose, a substance destitute of nitrogen, thus totally different from the contained protoplasm, and incapable of manifesting any of the phenomena of life. .

Within the walls of cellulose the protoplasm is now closely imprisoned, but we are not on that account to suppose that it has lost its activity, or has abandoned its work as a living being. Though it is now no longer in direct contact with the surrounding medium, it is not the less dependent on it, and the reaction between the imprisoned protoplasm and the outer world is still permitted by the permeability of the surrounding wall of cellulose.

When the protoplasm thus becomes surrounded by a cellulose wall it seldom retains the uniform arrangement of its parts which is often found in the naked cells. Minute cavities or vacuoles make their appearance in it; these increase in size and run one into the other, and may finally form one large cavity in the center, which becomes filled with a watery fluid, known as the cell-sap. This condition of the cell was the first observed, and it was it which suggested the often inapplicable term "cell." By the formation of this central sap cavity the surrounding protoplasm is pushed aside, and pressed against the cellulose wall, over which it now extends as a continuous layer. The nucleus either continues near the center, enveloped by a layer of protoplasm, which is connected by radiating bands of protoplasm with that of the walls, or it accompanies the displaced protoplasm, and lies imbedded in this on the walls of the cell.

We have abundant evidence to show that the imprisoned protoplasm loses none of its activity. The Characæ constitute an exceedingly interesting group of simple plants, common in the clear water of ponds and of slowly running streams. The cells of which they are built up are comparatively large, and, like almost all vegetable cells, are each inclosed in a wall of cellulose. The cellulose is perfectly transparent, and if the microscope, even with a low power, be brought to bear on one of these cells, a portion of its protoplasm may be seen in active rotation, flowing up one side of the long, tubular cell and down the other, and sweeping on with it such more solid particles as may become enveloped in its current. In another water-plant, the Vallsnerla spiralis, a similar active rotation of the protoplasm may be seen in the cells of the leaf, where the continuous stream of liquid protoplasm sweeping along the green granules of chlorophyl, and even carrying the globular nucleus with it in its current, presents one of the most beautiful of the many beautiful phenomena which the microscope has revealed to us.

In many other cells with large sap-cavities, such as those which form the stinging hairs of nettles and other kinds of vegetable hairs, the protoplasmic lining of the wall may send off into the sap-cavity projecting ridges and strings, forming an irregular network, along which, under a high power of the microscope, a slow streaming of granules may be witnessed. The form and position of this protoplasmic network undergo constant changes, and the analogy with the changes of form in an Amœba becomes obvious. The external wall of cellulose renders it impossible for the confined protoplasm to emit, like a naked Amœba, pseudopodia from its outer side; but on the inner side there is no obstacle to the extension of the protoplasm, and here the cavity of the cell becomes more or less completely traversed by protoplasmic projections from the wall. These often stretch themselves out in the form of thin filaments, which, meeting with a neighboring one, become fused into it; they show currents of granules streaming along their length, and after a time become withdrawn and disappear. The vegetable cell, in short, with its surrounding wall of cellulose, is in all essential points a closely imprisoned rhizopod.

Further proof that the imprisoned protoplasm has lost by its imprisonment none of its essential irritability, is afforded by the fact that if the transparent cell of a Nitella, one of the simple water-plants just referred to, be touched under the microscope with the point of a blunt needle, its green protoplasm will be seen to recede, under the irritation of the needle, from the cellulose wall. If the cellulose wall of the comparatively large cell which forms the entire plant in a Vaucheria, a unicellular alga very common in shallow ditches, be ruptured under the microscope, its protoplasm will escape, and may then be often seen to throw out pseudopodial projections and exhibit amœboid movements.

Even in the higher plants, without adducing such obvious and well-known instances as those of the sensitive-plant and Venus's flytrap, the irritability of the protoplasm may be easily rendered manifest. There are many herbaceous plants, in which, if the young, succulent stem of a vigorously growing specimen receive a sharp blow, of such a nature, however, as not to bruise its tissues, or in any way wound it, the blow will sometimes be immediately followed by a drooping of the stem commencing at some distance above the point to which the stroke had been applied; its strength appears to have here suddenly left it; it is no longer able to bear its own weight, and seems to be dying. The protoplasm, however, of its cells, is in this instance not killed, it is only stunned by the violence of the blow, and needs time for its restoration. After remaining, it may be for some hours, in this drooping and flaccid state, the stem begins to raise itself, and soon regains its original vigor. This experiment will generally succeed well in plants with a rather large terminal spike or raceme, when the stroke is applied some little distance below the inflorescence shortly before the expansion of the flower.

In the several instances now adduced, the protoplasm is in the mature state of the plant entirely included within a wall of cellulose. Some recent beautiful observations, however, of Mr. Francis Darwin have shown that even in the higher plants truly naked protoplasm may occur. From the cells of certain glandular hairs contained within the cup-like receptacles formed by the united bases of two opposite leaves in the teasel (Dipsacus), he has seen emitted long, pseudopodia-like projections of the protoplasm. What may be the significance of this very exceptional phenomenon is still undetermined. It is probably, as Mr. Darwin supposes, connected with the absorption of nitrogenous matter.

That there is no essential difference between the protoplasm of plants and that of animals is rendered further evident by other motor phenomena, which we are in the habit of regarding as the exclusive attribute of animals. Many of the more simply organized plants give origin to peculiar cells called spores, which separate from the parent, and, like the seeds of the higher plants, are destined to repeat its form. In many cases these spores are eminently locomotive. They are then termed "swarm-spores," and their movements are brought about, sometimes by changes of shape, when they move about in the manner of an Amœba, but more frequently by minute vibratile cilia, or by more strongly developed flagella or whip-like projections of their protoplasm. These cilia and flagella are absolutely indistinguishable from similar structures widely distributed among animals, and by their vibratory or lashing strokes upon the surrounding water the swarm spores are rapidly carried from place to place. In these motions they often present a curious semblance of volition, for if the swarm-spore meet with an obstacle in its course, it will, as if to avoid it, change the direction of its motion, and retreat by a reversion of the stroke of its cilia. They are usually attracted by light, and congregate at the light side of the vessel which contains them, though in some cases light has the opposite effect on them, and they recede from it.

Another fact may here be adduced to show the uniform character of protoplasm, and how very different are its properties from those of lifeless matter, namely, the faculty which all living protoplasm possesses of resisting the entrance of coloring matter into its substance. As many here present are aware, microscopists are in the habit of using in their investigations various coloring matters, such as solutions of carmine. These act differently on the different tissues, staining some, for example, more deeply than others, and thus enabling the histologist to detect certain elements of structure, which would otherwise remain unknown. Now, if a solution of carmine be brought into contact with living protoplasm, this will remain, so long as it continues alive, unaffected by the coloring matter. But if the protoplasm be killed, the carmine will at once pervade its whole substance, and stain it throughout with a color more intense than even that of the coloring solution itself.

But no more illustrative example can be offered of the properties of protoplasm as living matter, independently of any part it may take in organization, than that presented by the Myxomycetæ.

The Myxomycetæ constitute a group of remarkable organisms, which, from their comparatively large size and their consisting, during a great part of their lives, of naked protoplasm, have afforded a fine field for research, and have become one of the chief sources from which our knowledge of the nature and phenomena of protoplasm has been derived.

They have generally been associated by botanists with the fungi, but though their affinities with these are perhaps closer than with any other plants, they differ from them in so many points, especially in their development, as to render this association untenable. They are found in moist situations, growing on old tan or on moss, or decaying leaves or rotten wood, over which they spread in the form of a network of naked protoplasmic filaments, of a soft, creamy consistence, and usually of a yellowish color.

Under the microscope the filaments of the network exhibit active spontaneous movements, which, in the larger branches, are visible under an ordinary lens, or even by the naked eye. A succession of undulations may then be noticed passing along the course of the threads. Under higher magnifying powers a constant movement of granules may be seen flowing along the threads, and streaming from branch to branch of this wonderful network. Here and there offshoots of the protoplasm are projected, and again withdrawn in the manner of the pseudopodia of an Amœba, while the whole organism may be occasionally seen to abandon the support over which it had grown, and to creep over neighboring surfaces, thus far resembling in all respects a colossal ramified Amœba. It is also curiously sensitive to light, and may be sometimes found to have retreated during the day to the dark side of the leaves, or into the recesses of the tan over which it had been growing, and again to creep out on the approach of night.

After a time there arise from the surface of this protoplasmic net oval capsules or spore-cases, in which are contained the spores or reproductive bodies of the Myxomycetæ. When the spore-case has arrived at maturity, it bursts and allows the spores to escape. These are in the form of spherical cells, each included in a delicate membranous wall, and when they fall into water the wall becomes ruptured, and the little cell creeps out. This consists of a little mass of protoplasm with a round central nucleus, inclosing a nucleolus, and with a clear vacuole, which exhibits a rhythmically pulsating movement. The little naked spore thus set at liberty is soon seen to be drawn out at one point into a long, vibratile, whip-like flagellum, which by its lashing action carries the spore from place to place. After a time the flagellum disappears, and the spore may now be seen emitting and withdrawing finger-like pseudopodia, by means of which it creeps about like an Amœba, and like an Amœba devours solid particles by ingulfing them in its soft protoplasm.

So far these young Amœba-like Myxomycetæ have enjoyed each an independent existence. Now, however, a singular and significant phenomenon is presented. Two or more of these Myxamœbæ, as they have been called, approach one another, come into contact, and finally become completely fused together into a single mass of protoplasm, in which the components are no longer to be distinguished. To the body thus formed by the fusion of the Myxamœbæ the name of "Plasmodium" has been given.

The plasmodium continues, like the simple amœbiform bodies of which it is composed, to grow by the ingestion and assimilation of solid nutriment, which it envelops in its substance; it throws out ramifying and mosculating processes, and finally becomes converted into a protoplasmic network, which in its turn gives rise to spore-cases with their contained spores, and thus completes the cycle of its development.

Under certain external conditions, the Myxomycetæ have been observed to pass from an active mobile state into a resting state, and this may occur both in the amœbiform spores and in the plasmodium. When the plasmodium is about to pass into a resting state, it usually withdraws its finer branches, and expels such solid ingesta as may be included in it. Its motions then gradually cease, it breaks up into a multitude of polyhedral cells, which, however, remain connected, and the whole body dries into a horny brittle mass, known by the name of "sclerotium."

In this condition, without giving the slightest sign of life, the sclerotium may remain for many months. Life, however, is not destroyed; its manifestations are only suspended; and if after an indefinite time the apparently dead sclerotium be placed in water, it immediately begins to swell up, the membranous covering of its component cells becomes dissolved and disappears, and the cells themselves flow together into an active amœboid plasmodium.

We have already seen that every cell possesses an autonomy or independent individuality, and from this we should expect that, like all living beings, it had the faculty of multiplying itself, and of becoming the parent of other cells. This is truly the case, and the process of cell-multiplication has of late years been studied, with the result of adding largely to our knowledge of the phenomena of life.

The labors of Strasburger, of Auerbach, of Oscar Hertwig, of Eduard van Beneden, Bütschli, Fol, and others, here come prominently before us, but neither the time at my disposal nor the purport of this address will allow me to do more than call your attention to some of the more striking results of their investigations.

By far the most frequent mode of multiplication among cells shows itself in a spontaneous division of the protoplasm into two separate portions, which then become independent of one another, so that instead of the single parent cell two new ones have made their appearance. In this process the nucleus usually takes an important part. Strasburger has studied it with great care in certain plant-cells, such as the so-called "corpuscula" or "secondary embryo-sacs" of the Coniferæ and the cells of Spirogyra; and has further shown a close correspondence between cell division in animals and that in plants.

It may be generally stated as the results of his observations on the corpuscula of the Coniferæ, that the nucleus of the cell about to divide assumes a spindle-shape, and at the same time presents a peculiar striated differentiation, as if it were composed of parallel filaments reaching from end to end of the spindle. These filaments become thickened in the middle, and there form by the approximation of the thickened portions a transverse plate of protoplasm (the "nucleusplate"). This soon splits into two halves, which recede from one another toward the poles of the spindle, traveling in this course along the filaments, which remain continuous from end to end. When arrived near the poles they form there two new nuclei, still connected with one another by the intervening portion of the spindle.

In the equator of this intervening portion there is now formed in a similar way a second plate of protoplasm (the "cell-plate"), which, extending to the walls of the dividing cell, cuts the whole protoplasm into two halves, each half containing one of the newly-formed nuclei. This partition plate is at first single, but it soon splits into two lamina?, which become the apposed bounding surfaces of the two protoplasm masses into which the mother cell has been divided. A wall of cellulose is then all at once secerted between them, and the two daughter cells are complete.

It sometimes happens in the generation of cells that a young brood of cells arises from the parent cell by what is called "free-cell formation." In this only a part of the protoplasm of the mother cell is used up in the production of the offspring. It is seen chiefly in the formation of the spores of the lower plants, in the first foundation of the embryo in the higher, and in the formation of the endosperm—a cellular mass which serves as the first nutriment for the embryo—in the seeds of most Phanerogams. The formation of the endosperm has been carefully studied by Strasburger in the embyro-sac of the kidney-bean, and may serve as an example of the process of free-cell formation. The embryo-sac is morphologically a large cell with its protoplasm, nucleus, and cellulose wall, while the endosperm which arises within it is composed of a multitude of minute cells united into a tissue. The formation of the endosperm is preceded by the dissolution and disappearance of the nucleus of the embryo-sac, and then in the midst of the protoplasm of the sac several new nuclei make their appearance. Around each of these as a center the protoplasm of the mother cell is seen to have become differentiated in the form of a clear spherule, and we have thus corresponding to each of the new nuclei a young naked cell, which soon secretes over its surface a membrane of cellulose. The new cells, when once formed, multiply by division, press one on the other, and so combining into a cellular mass, constitute the completed endosperm.

Related to the formation of new cells, whether by division or by free-cell formation, is another very interesting phenomenon of living protoplasm known as "rejuvenescence." In this the whole protoplasm of a cell, by a new arrangement of its parts, assumes a new shape and acquires new properties. It then abandons its cellulose chamber, and enters on a new and independent life in the surrounding medium.

A good example of this is afforded by the formation of swarm spores in Œdogonium, one of the fresh-water algæ. Here the whole of the protoplasm of an adult cell contracts, and by the expulsion of its cell-sap changes from a cylindrical to a globular shape. Then one spot becomes clear, and a pencil of vibratile cilia here shows itself. The cellulose wall which had hitherto confined it now becomes ruptured, and the protoplasmic sphere, endowed with new faculties of development and with powers of active locomotion, escapes as a swarm spore, which, after enjoying for a time the free life of an animal, comes to rest, and develops itself into a new plant.

The beautiful researches which have within the last few years been made by the observers already mentioned, on the division of animal cells, show how close is the agreement between plants and animals in all the leading phenomena of cell-division, and afford one more proof of the essential unity of the two great organic kingdoms.

There is one form of cell which, in its relation to the organic world, possesses a significance beyond that of every other, namely, the egg. As already stated, the egg is, wherever it occurs, a typical cell, consisting essentially of a globule of protoplasm enveloping a nucleus (the "germinal vesicle"), and with one or more nucleoli (the "germinal spots") in the interior of the nucleus. This cell, distinguishable by no tangible characters from thousands of other cells, is nevertheless destined to run through a definite series of developmental changes, which have as their end the building up of an organism like that to which the egg owes its origin.

It is obvious that such complex organisms as thus result—composed, it may be, of countless millions of cells—can be derived from the simple egg-cell only by a process of cell-multiplication. The birth of new cells derived from the primary cell or egg thus lies as the basis of embryonic development. It is here that the phenomena of cell-multiplication in the animal kingdom can in general be most satisfactorily observed, and the greater number of recent researches into the nature of these phenomena, have found their most fertile field in the early periods of the development of the egg.

A discussion of the still earlier changes which the egg undergoes in order to bring it into the condition in which cell-multiplication may be possible, would, however full of interest, be here out of place; and I shall therefore confine myself to the first moments of actual development—to what is called "the cleavage of the egg"—which is nothing more than a multiplication of the egg-cell by repeated division. I shall further confine myself to an account of this phenomenon as presented in typical cases, leaving out of consideration certain modifications which would only complicate and obscure our picture.

The egg, notwithstanding the preliminary changes to which I have alluded, is still at the commencement of development a true cell. It has its protoplasm and its nucleus, and it is, as a rule, enveloped in a delicate membrane. The protoplasm forms what is known as the vitellus, or yolk, and the surrounding membrane is called the "vitellary membrane." The division which is now about to take place in it is introduced by a change of form in the nucleus. This becomes elongated, and assumes the shape of a spindle, similar to what we have already seen in the cell-division of plants. On each pole of the spindle transparent protoplasm collects, forming here a clear spherical area.

At this time a very striking and characteristic phenomenon is witnessed in the egg. Each pole of the spindle has become the center of a system of rays which stream out in all directions into the surrounding protoplasm. The protoplasm thus shows, enveloped in its mass, two sun-like figures, whose centers are connected with one another by the spindle-shaped nucleus. To this, with the sun-like rays streaming from its poles, Auerbach gives the name of "Karyolitic figure," suggested by its connection with the breaking up of the original nucleus, to which our attention must next be directed.

A phenomenon similar to one we have already seen in cell-division among plants now shows itself. The nucleus becomes broken up into a number of filaments, which lie together in a bundle, each filament stretching from pole to pole of the spindle. Exactly in its central point every filament shows a knot-like enlargement, and from the close approximation of the knots there results a thick zone of protoplasm in the equator of the spindle. Each knot soon divides into two halves, and each half recedes from the equator and travels along the filament toward its extremity. When arrived at the poles of the spindle each set of half knots becomes fused together into a globular body, while the intervening portion of the spindle, becoming torn up, and gradually drawn into the substance of the two globular masses, finally disappears. And now, instead of the single fusiform nucleus, whose changes we have been tracing, we have two new globular nuclei, each occupying the place of one of its poles, and formed at its expense.[9] The egg now begins to divide along a plane at right angles to a line connecting the two nuclei. The division takes place without the formation of a cell-plate such as we saw in the division of the plant cell, and is introduced by a constriction of its protoplasm, which commences at the circumference just within the vitelline membrane, and extending toward the center, divides the whole mass of protoplasm into two halves, each including within it one of the new nuclei. Thus the simple cell which constituted the condition of the egg at the commencement of development becomes divided into two similar cells. This forms the first stage of cleavage. Each of these two young cells divides in its turn in a direction at right angles to the first division plane, while by continued repetition of the same act the whole of the protoplasm or yolk becomes broken up into a vast, multitude of cells, and the unicellular organism—the egg, with which we began our history—has become converted into an organism composed of many thousands of cells. This is one of the most widely distributed phenomena of the organic world. It is called "the cleavage of the egg," and consists essentially in the production, by division, of successive broods of cells from a single ancestral cell—the egg.

It is no part of my purpose to carry on the phenomena of development further than this. Such of my hearers as may desire to become acquainted with the further history of the embryo, I would refer to the excellent address delivered two years ago at the Plymouth meeting of the Association by one of my predecessors in this chair—Professor Allen Thompson.

That protoplasm, however, may present a phenomenon the reverse of that in which a simple cell becomes multiplied into many, is shown by a phenomenon already referred to—the production of plasmodia in the Myxomycetæ by the fusion into one another of cells originally distinct.

The genus Myriothela will afford another example in which the formation of plasmodia becomes introduced into the cycle of development. The primitive eggs are here, as elsewhere, true cells with nucleolated nuclei, but without any boundary membrane. They are formed in considerable numbers, but remain only for a short time separate and distinct. After this they begin to exhibit amœboid changes of shape, project pseudopodial prolongations which coalesce with those of others in their vicinity, and, finally, a multitude of these primitive ova become fused together into a common plasmodium, in which, as in the simple egg-cell of other animals, the phenomena of development take place.

In many of the lower plants a very similar coalescence is known to take place between the protoplasmic bodies of separate cells, and constitutes the phenomenon of conjugation. Spirogyra is a genus of algæ, consisting of long, green threads common in ponds. Every thread is composed of a series of cylindrical chambers of transparent cellulose placed end to end, each containing a sac of protoplasm with a large quantity of cell-sap, and with a green band of chlorophyl wound spirally on its walls. When the threads have attained their full growth they approach one another in pairs, and lie in close proximity, parallel one to the other. A communication is then established by means of short connecting-tubes between the chambers of adjacent filaments, and across the channel thus formed the whole of the protoplasm of one of the conjugating chambers passes into the cavity of the other, and then immediately fuses with the protoplasm it finds there. The single mass thus formed shapes itself into a solid oval body, known as a "zygospore." This now frees itself from the filament, secretes over its naked surface a new wall of cellulose, and, when placed in the conditions necessary for its development, attaches itself by one end, and then, by repeated acts of cell-division, grows into a many-celled filament like those in which it originated.

The formation of plasmodia, regarded as a coalescence and absolute fusion into one another of separate, naked masses of protoplasm, is a phenomenon of great significance. It is highly probable that, notwithstanding the complete loss of individuality in the combining elements, such differences as may have been present in these will always find themselves expressed in the properties of the resulting plasmodia—a fact of great importance in its bearing on the phenomena of inheritance. Recent researches, indeed, render it almost certain that fertilization, whether in the animal or the vegetable kingdom, consists essentially in the coalescence and consequent loss of individuality of the protoplasmic contents of two cells.

In by far the greater number of plants the protoplasm of most of the cells which are exposed to the sunlight undergoes a curious and important differentiation, part of it becoming separated from the remainder in the form usually of green granules, known as chlorophyll. The chlorophyl-granules thus consist of true protoplasm, their color being due to the presence of a green coloring matter, which may be extracted, leaving behind the colorless protoplasmic base.

The coloring matter of chlorophyl presents under the spectroscope a very characteristic spectrum. For our knowledge of its optical properties, on which time will not now permit me to dwell, we are mainly indebted to the researches of your townsman, Dr. Sorby, who has made these the subject of a series of elaborate investigations, which have contributed largely to the advancement of an important department of physical science.

That the chlorophyl is a living substance, like the uncolored protoplasm of the cell, is sufficiently obvious. When once formed, the chlorophyl-granule may grow by intussusception of nutriment to many times its original size, and may multiply itself by division.

To the presence of chlorophyl is due one of the most striking aspects of external nature—the green color of the vegetation which clothes the surface of the earth: and with its formation is introduced a function of fundamental importance in the economy of plants, for it is on the cells which contain this substance that devolves the faculty of decomposing carbonic acid. On this depends the assimilation of plants, a process which becomes manifest externally by the exhalation of oxygen. Now, it is under the influence of light on the chlorophyl containing cell that this evolution of oxygen is brought about. The recent observations of Draper and of Pfeffer have shown that in this action the solar spectrum is not equally effective in all its parts; that the yellow and least refrangible rays are those which act with most intensity; that the violent and other highly refrangible rays of the visible spectrum take but a very subordinate part in assimilation; and that the invisible rays which lie beyond the violet are totally inoperative.

In almost every grain of chlorophyl one or more starch-granules may be seen. This starch is chemically isomeric with the cellulose cell-wall, with woody fiber, and other hard parts of plants, and is one of the most important products of assimilation. When plants whose chlorophyl contains starch are left for a sufficient time in darkness, the starch is absorbed and completely disappears; but when they are restored to the light the starch reappears in the chlorophyl of the cells.

With this dependence of assimilation on the presence of chlorophyl a new physiological division of labor is introduced into the life of plants. In the higher plants, while the work of assimilation is allocated to the chlorophyl-containing cells, that of cell division and growth devolves on another set of cells, which, lying deeper in the plant, are removed from the direct action of light, and in which chlorophyl is therefore never produced. In certain lower plants, in consequence of their simplicity of structure and the fact that all the cells are equally exposed to the influence of light, this physiological division of labor shows itself in a somewhat different fashion. Thus in some of the simple green algae, such as Spirogyra and Hydrodictyon, assimilation takes place as in other cases during the day, while their cell division and growth takes place chiefly, if not exclusively, at night. Strasburger, in his remarkable observations on cell-divisions in Spirogyra, was obliged to adopt an artificial device in order to compel the Spirogyra to postpone the division of its cells to the morning.

Here the functions of assimilation and growth devolve on one and the same cell, but, while one of these functions is exercised only during the day, the time for the other is the night. It seems impossible for the same cell at the same time to exercise both functions, and these are here accordingly divided between different periods of the twenty-four hours.

The action of chlorophyl in bringing about the decomposition of carbonic acid is not, as was recently believed, absolutely confined to plants. In some of the lower animals, such as Stentor and other infusoria, the Green Hydra, and certain green planarise and other worms, chlorophyl is differentiated in their protoplasm, and probably always acts here under the influence of light exactly as in plants.

Indeed, it has been proved[10] by some recent researches of Mr. Geddes, that the green planarias when placed in water and exposed to the sunlight give out bubbles of gas which contain from forty-four to fifty-five per cent, of oxygen. Mr. Geddes has further shown that these animals contain granules of starch in their tissues, and in this fact we have another striking point of resemblance between them and plants.

A similar approximation of the two organic kingdoms has been shown by the beautiful researches of Mr. Darwin—confirmed and extended by his son, Mr. Francis Darwin—on Drosera and other so called carnivorous plants. These researches, as is now well known, have shown that in all carnivorous plants there is a mechanism fitted for the capture of living prey, and that the animal matter of the prey is absorbed by the plant after having been digested by a secretion which acts like the gastric juice of animals.

Again, Nägeli has recently shown[11] that the cell of the yeast-fungus contains about two per cent, of peptine, a substance hitherto known only as a product of the digestion of azotized matter by animals.

Indeed, all recent research has been bringing out in a more and more decisive manner the fact that there is no dualism in life—that the life of the animal and the life of the plant are, like their protoplasm, in all essential points identical.

But there is, perhaps, nothing which shows more strikingly the identity of the protoplasm in plants and animals, and the absence of any deep-pervading difference between the life of the animal and that of the plant, than the fact that plants may be placed, just like animals, under the influence of anaesthetics.

When the vapor of chloroform or of ether is inhaled by the human subject, it passes into the lungs, where it is absorbed by the blood, and thence carried by the circulation to all the tissues of the body. The first to be affected by it is the delicate nervous element of the brain, and loss of consciousness is the result. If the action of the anæsthetic be continued, all the other tissues are in their turn attacked by it and their irritability arrested. A set of phenomena entirely parallel to these may be presented by plants.

We owe to Claude Bernard a series of interesting and most instructive experiments on the action of ether and chloroform on plants. He exposed to the vapor of ether a healthy and vigorous sensitive plant, by confining it under a bell-glass into which he introduced a sponge filled with ether. At the end of half an hour the plant was in a state of anæsthesia. All its leaflets remained fully extended, but they showed no tendency to shrink when touched. It was then withdrawn from the influence of the ether, when it gradually recovered its irritability, and finally responded, as before, to the touch.

It is obvious that the irritability of the protoplasm was here arrested by the anæsthetic, so that the plant became unable to give a response to the action of an external stimulus.

It is not, however, the irritability of the protoplasm of only the motor elements of plants that anæsthetics are capable of arresting. These may act also on the protoplasm of those cells whose function lies in chemical synthesis, such as is manifested in the phenomena of the germination of the seed and in nutrition generally, and Claude Bernard has shown that germination is suspended by the action of ether or chloroform.

Seeds of cress, a plant whose germination is very rapid, were placed. in conditions favorable to a speedy germination, and while thus placed were exposed to the vapor of ether. The germination, which would otherwise have shown itself by the next day, was arrested. For five or six days the seeds were kept under the influence of the ether, and showed during this time no disposition to germinate. They were not killed, however, they only slept; for, on the substitution of common air for the etherized air with which they had been surrounded, germination at once set in and proceeded with activity.

Experiments were also made on that function of plants by which they absorb carbonic acid and exhale oxygen, and which, as we have already seen, is carried on through the agency of the green protoplasm or chlorophyl, under the influence of light—a function which is commonly, but erroneously, called the respiration of plants.

Aquatic plants afford the most convenient subjects for such experiments. If one of these be placed in a jar of water holding ether or chloroform in solution, and a bell-glass be placed over the submerged plant, we shall find that the plant no longer absorbs carbonic acid or emits oxygen. It remains, however, quite green and healthy. In order to awaken the plant, it is only necessary to place it in non-etherized water, when it will begin once more to absorb carbonic acid, and exhale oxygen under the influence of sunlight.

The same great physiologist has also investigated the action of anaesthetics on fermentation. It is well known that alcoholic fermentation is due to the presence of a minute fungus, the yeast-fungus, the living protoplasm of whose cells has the property of separating solutions of sugar into alcohol, which remains in the liquid, and carbonic acid, which escapes into the air.

Now, if the yeast-plant be placed along with sugar in etherized water, it will no longer act as a ferment. It is anæsthesiated, and can not respond to the stimulus which, under ordinary circumstances, it would find in the presence of the sugar. If, now, it be placed on a filter, and the ether washed completely away, it will, on restoration to a saccharine liquid, soon resume its duty of separating the sugar into alcohol and carbonic acid.

Claude Bernard has further palled attention to a very significant fact which is observable in this experiment. While the proper alcoholic fermentation is entirely arrested by the etherization of the yeast-plant, there still goes on in the saccharine solution a curious chemical change, the cane-sugar of the solution being converted into grape-sugar, a substance identical in its chemical composition with the cane-sugar, but different in its molecular constitution. Now, it is well known from the researches of Berthelot that this conversion of cane-sugar into grape-sugar is due to a peculiar inversive ferment, which, while it accompanies the living yeast-plant, is itself soluble and destitute of life. Indeed, it has been shown that, in its natural conditions, the yeast-fungus is unable of itself to assimilate cane-sugar, and that, in order that this may be brought into a state fitted for the nutrition of the fungus, it must be first digested and converted into grape-sugar, exactly as happens in our own digestive organs. To quote Claude Bernard's graphic account: "The fungus ferment has thus beside it in the same yeast a sort of servant given by Nature to effect this digestion. The servant is the unorganized inversive ferment. This ferment is soluble, and, as it is not a plant, but an unorganized body destitute of sensibility, it has not gone to sleep under the action of the ether, and thus continues to fulfill its task."

In the experiment already recorded on the germination of seeds the interest is by no means confined to that which attaches itself to the arrest of the organizing functions of the seed, those namely which manifest themselves in the development of the radicle, and plumule, and other organs of the young plant. Another phenomenon of great significance becomes at the same time apparent: the anæsthetic exerts no action on the concomitant chemical phenomena which in germinating seeds show themselves in the transformation of starch into sugar under the influence of diastase (a soluble and non-living ferment which also exists in the seed), and the absorption of oxygen with the exhalation of carbonic acid. These go on as usual, the anæsthesiated seed continuing to respire, as proved by the accumulation of carbonic acid in the surrounding air. The presence of the carbonic acid was rendered evident by placing in the same vessel with the seeds which were the object of the experiment a solution of barytes, when the carbonate became precipitated from the solution in quantity equal to that produced in a similar experiment with seeds germinating in unetherized air.

So, also, in the experiment which proves that the faculty possessed by the chlorophyllian cells of absorbing carbonic acid and exhaling oxygen under the influence of light may be arrested by anæsthetics, it could be seen that the plant, while in a state of anæsthesia, continued to respire in the manner of animals; that is, it continued to absorb oxygen and exhale carbonic acid. This is the true respiratory function which was previously masked by the predominant function of assimilation, which devolves on the green cells of plants, and which manifests itself under the influence of light in the absorption of carbonic acid and the exhalation of oxygen.

It must not, however, be supposed that the respiration of plants is entirely independent of life. The conditions which bring the oxygen of the air and the combustible matter of the respiring plant into such relations as may allow them to act on one another are still under its control, and we must conclude that in Claude Bernard's experiment the anæsthesia had not been carried so far as to arrest such properties of the living tissues as are needed for this.

The quite recent researches of Schützenberger, who has investigated the process of respiration as it takes place in the cell of the yeast-fungus, have shown that vitality is a factor in this process. He has shown that fresh yeast, placed in water, breathes like an aquatic animal, disengaging carbonic acid, and causing the oxygen contained in the water to disappear. That this phenomenon is a function of the living cell is proved by the fact that, if the yeast be first heated to 60° C. and then placed in the oxygenated water, the quantity of oxygen in the water remains unchanged; in other words, the yeast ceases to breathe.

Schützenberger has further shown that light exerts no influence on the respiration of the yeast-cell—that the absorption of oxygen by the cell takes place in the dark exactly as in sunlight. On the other hand, the influence of temperature is well marked. Respiration is almost entirely arrested at temperatures below 10° C, it reaches its maximum at about 40° C, while at 60° C. it again ceases.

All this proves that the respiration of living beings is identical, whether manifested in the plant or in the animal. It is essentially a destructive phenomenon—as much so as the burning of a piece of charcoal in the open air, and, like it, is characterized by the disappearance of oxygen and the formation of carbonic acid.

One of the most valuable results of the recent careful application of the experimental method of research to the life-phenomena of plants is thus the complete demolition of the supposed antagonism between respiration in plants and that in animals.


I have thus endeavored to give you in a few broad outlines a sketch of the nature and properties of one special modification of matter, which will yield to none other in the interest which attaches to its study, and in the importance of the part allocated to it in the economy of nature. Did the occasion permit, I might have entered into many details which I have left untouched; but enough has been said to convince you that in protoplasm we find the only form of matter in which life can manifest itself; and that, though the outer conditions of life—heat, air, water, food—may all be present, protoplasm would still be needed, in order that these conditions may be utilized; in order that the energy of lifeless nature may be converted into that of the countless multitudes of animal and vegetable forms which dwell upon the surface of the earth or people the great depths of its seas.

We are thus led to the conception of an essential unity in the two great kingdoms of organic nature—a structural unity, in the fact that every living being has protoplasm as the essential matter of every living element of its structure; and a physiological unity, in the universal attribute of irritability which has its seat in this same protoplasm, and is the prime mover of every phenomenon of life.

We have seen how little mere form has to do with the essential properties of protoplasm. This may shape itself into cells, and the cells may combine into organs in ever-increasing complexity, and protoplasm-force may be thus intensified, and, by the mechanism of organization, turned to the best possible account; but we must still go back to protoplasm as a naked, formless plasma if we would find—freed from all non-essential complications—the agent to which has been assigned the duty of building up structure and of transforming the energy of lifeless matter into that of living.

To suppose, however, that all protoplasm is identical where no difference cognizable by any means at our disposal can be detected would be an error. Of two particles of protoplasm, between which we may defy all the power of the microscope, all the resources of the laboratory, to detect a difference, one can develop only to a jelly-fish, the other only to a man, and one conclusion alone is here possible—that deep within them there must be a fundamental difference which thus determines their inevitable destiny, but of which we know nothing, and can assert nothing beyond the statement that it must depend on their hidden molecular constitution.

In the molecular condition of protoplasm there is probably as much complexity as in the disposition of organs in the most highly differentiated organisms; and between two masses of protoplasm indistinguishable from one another there may be as much molecular difference as there is between the form and arrangement of organs in the most widely separated animals or plants.

Herein lies the many-sidedness of protoplasm; herein lies its significance as the basis of all morphological expression, as the agent of all physiological work, while in all this there must be an adaptiveness to purpose as great as any claimed for the most complicated organism.

From the facts which have been now brought to your notice there is but one legitimate conclusion—that life is a property of protoplasm. In this assertion there is nothing that need startle us. The essential phenomena of living beings are not so widely separated from the phenomena of lifeless matter as to render it impossible to recognize an analogy between them; for even irritability, the one grand character of all living beings, is not more difficult to be conceived of as a property of matter than the physical phenomena of radial energy.

It is quite true that between lifeless and living matter there is a vast difference, a difference greater far than any which can be found between the most diverse manifestations of lifeless matter. Though the refined synthesis of modern chemistry may have succeeded in forming a few principles which until lately had been deemed the proper product of vitality, the fact still remains that no one has ever yet built up one particle of living matter out of lifeless elements—that every living creature, from the simplest dweller on the confines of organization up to the highest and most complex organism, has its origin in preexistent living matter—that the protoplasm of to-day is but the continuation of the protoplasm of other ages, handed down to us through periods of indefinable and indeterminable time.

Yet with all this, vast as the differences may be, there is nothing which precludes a comparison of the properties of living matter with those of lifeless.

When, however, we say that life is a property of protoplasm, we assert as much as we are justified in doing. Here we stand upon the boundary between life in its proper conception, as a group of phenomena having irritability as their common bond, and that other and higher group of phenomena which we designate as consciousness or thought, and which, however intimately connected with those of life, are yet essentially distinct from them.

When the heart of a recently killed frog is separated from its body and touched with the point of a needle, it begins to beat under the excitation of the stimulus, and we believe ourselves justified in referring the contraction of the cardiac fibers to the irritability of their protoplasm as its proper cause. We see in it a remarkable phenomenon, but one nevertheless in which we can see unmistakable analogies with phenomena purely physical. There is no greater difficulty in conceiving of contractility as a property of protoplasm than there is of conceiving of attraction as a property of the magnet.

When a thought passes through the mind, it is associated, as we have now abundant reason for believing, with some change in the protoplasm of the cerebral cells. Are we, therefore, justified in regarding thought as a property of the protoplasm of these cells, in the sense in which we regard muscular contraction as a property of the protoplasm of muscle, or is it really a property residing in something far different, but which may yet need for its manifestation the activity of cerebral protoplasm?

If we could see any analogy between thought and any one of the admitted phenomena of matter, we should be justified in accepting the first of these conclusions as the simplest, and as affording an hypothesis most in accordance with the comprehensiveness of natural laws; but between thought and the physical phenomena of matter there is not only no analogy, but there is no conceivable analogy; and the obvious and continuous path which we have hitherto followed up in our reasonings from the phenomena of lifeless matter through those of living matter here comes suddenly to an end. The chasm between unconscious life and thought is deep and impassable, and no transitional phenomena can be found by which as by a bridge we may span it over; for even from irritability, to which, on a superficial view, consciousness may seem related, it is as absolutely distinct as it is from any of the ordinary phenomena of matter.

It has been argued that because physiological activity must be a property of every living cell, psychical activity must be equally so, and the language of the metaphysician has been carried into biology, and the "cell-soul" spoken of as a conception inseparable from that of life.

That psychical phenomena, however, characterized as they essentially are by consciousness, are not necessarily coextensive with those of life, there can not be a doubt. How far back in the scale of life consciousness may exist we have as yet no means of determining, nor is it necessary for our argument that we should. Certain it is that many things, to all appearance the result of volition, are capable of being explained as absolutely unconscious acts; and when the swimming swarm-spore of an alga avoids collision, and, by a reversal of the stroke of its cilia, backs from an obstacle lying in its course, there is almost certainly in all this nothing but a purely unconscious act. It is but a case in which we find expressed the great law of the adaptation of living beings to the conditions which surround them. The irritability of the protoplasm of the ciliated spore responding to an external stimulus sets in motion a mechanism derived by inheritance from its ancestors, and whose parts are correlated to a common end—the preservation of the individual.

But even admitting that every living cell were a conscious and thinking being, are we therefore justified in asserting that its consciousness, like its irritability, is a property of the matter of which it is composed? The sole argument on which this view is made to rest is that from analogy. It is argued that because the life-phenomena, which are invariably found in the cell, must be regarded as a property of the cell, the phenomena of consciousness by which they are accompanied must be also so regarded. The weak point in the argument is the absence of all analogy between the things compared, and, as the conclusion rests solely on the argument from analogy, the two must fall to the ground together.

In a lecture[12] to which I once had the pleasure of listening—a lecture characterized no less by lucid exposition than by the fascinating form in which its facts were presented to the hearers—Professor Huxley argues that no difference, however great, between the phenomena of living matter and those of the lifeless elements of which this matter is composed should militate against our attributing to protoplasm the phenomena of life as properties essentially inherent in it; since we know that the result of a chemical combination of physical elements may exhibit physical properties totally different from those of the elements combined; the physical phenomena presented by water, for example, having no resemblance to those of its combining elements, oxygen and hydrogen.

I believe that Professor Huxley intended to apply this argument only to the phenomena of life in the stricter sense of the word. As such it is conclusive. But when it is pushed further, and extended to the phenomena of consciousness, it loses all its force. The analogy, perfectly valid in the former case, here fails. The properties of the chemical compound are like those of its components, still physical properties. They come within the wide category of the universally accepted properties of matter, while those of consciousness belong to a category absolutely distinct—one which presents not a trace of a connection with any of those which physicists have agreed in assigning to matter as its proper characteristics. The argument thus breaks down, for its force depends on analogy alone, and here all analogy vanishes.

That consciousness is never manifested except in the presence of cerebral matter or of something like it, there can not be a question; but this is a very different thing from its being a property of such matter in the sense in which polarity is a property of the magnet, or irritability of protoplasm. The generation of the rays which lie invisible beyond the violet in the spectrum of the sun can not be regarded as a property of the medium which by changing their refrangibility can alone render them apparent.

I know that there is a special charm in those broad generalizations which would refer many very different phenomena to a common source. But in this very charm there is undoubtedly a danger, and we must be all the more careful lest it should exert an influence in arresting the progress of truth, just as at an earlier period traditional beliefs exerted an authority from which the mind but slowly and with difficulty succeeded in emancipating itself.

But have we, it may be asked, made in all this one step forward toward an explanation of the phenomena of consciousness or the discovery of its source? Assuredly not. The power of conceiving of a substance different from that of matter is still beyond the limits of human intelligence, and the physical or objective conditions which are the concomitants of thought are the only ones of which it is possible to know anything, and the only ones whose study is of value.

We are not, however, on that account forced to the conclusion that there is nothing in the universe but matter and force. The simplest physical law is absolutely inconceivable by the highest of the brutes, and no one would be justified in assuming that man had already attained the limit of his powers. Whatever may be that mysterious bond which connects organization with psychical endowments, the one grand fact—a fact of inestimable importance—stands out clear and freed from all obscurity and doubt, that from the first dawn of intelligence there is with every advance in organization a corresponding advance in mind. Mind as well as body is thus traveling onward through higher and still higher phases; the great law of evolution is shaping the destiny of our race; and though now we may at most but indicate some weak point in the generalization which would refer consciousness as well as life to a common material source, who can say that in the far-off future there may not yet be evolved other and higher faculties from which light may stream in upon the darkness, and reveal to man the great mystery of thought?

  1. Inaugural Address at the Sheffield meeting, August 20, 1879.
  2. In speaking of protoplasm as a liquid, it must be borne in mind that this expression refers only to its physical consistence—a condition depending mainly on the amount of water with which it is combined, and subject to considerable variation, from the solid form in which we find it in the dormant embryo of seeds, to the thin, watery state in which it occurs in the leaves of Valisneria. Its distinguishing properties are totally different from those of a purely physical liquid, and are subject to an entirely different set of laws.
  3. The Red-Snow plant (Protococcus nivalis) acts on the atmosphere through the agency of chlorophyl, like the ordinary green plants. As in these, chlorophyl is developed in it, and is only withdrawn from view by the predominant red pigment to which the Protococcus owes one of its most striking characteristics.
  4. "On the Structure and Development of Myriothela" "Philosophical Transactions," vol. clxv., 1875, p. 552.
  5. "Jenaische Zeitschrift," 1871.
  6. "Die Embryonanlage und erste Entwickelung des Flusskrebse," "Zeitschrift für wissenschaftliche Zoologie," 1877.
  7. Loc. cit.
  8. "Ueber die Verdauungsorgane einiger Süsswasser-Turbellarien," "Zoologischer Anzeiger," December, 1878.
  9. Though none of the above-mentioned observers, to whom we owe our knowledge of the phenomena here described, seem to have thought of connecting the fibrous condition assumed by the spindle with any special structure of the quiescent nucleus, it is highly probable that it consists in a rearrangement of fibers already present. That this is really the case is borne out by the observations of Schleicher on the division of cartilage-cells. ("Die Knorpelzelltheilung," "Arch, für mikr. Anat.," Band xvi., Heft 2, 1878.) From these it would appear that, in the division of cartilage-cells, the investing membrane of the nucleus first becomes torn up, and then the filaments, rodlets, and granules, which, according to him, form its body, enter into a state of intense motor activity, and may be seen arranging themselves into star-like, or wreath-like, or irregular figures, while the whole nucleus, now deprived of its membrane, may wander about the cell, traveling toward one of its poles, and then toward the other; or it may at one time contract, and then again dilate, to such an extent as nearly to fill the entire cell. To this nuclear activity Schleicher applies the term "Karyokinesis." It results in a nearly parallel arrangement of the nuclear filaments. Then these converge at their extremities and become more widely separated in the middle, so as to give to the nucleus the form of a spindle. The filaments then become fused together at each pole of the spindle, so as to form the two new nuclei, which are at first nearly homogeneous, but which afterward become broken up into their component filaments, rods, and granules.
  10. "Sur la Fonction de la Chlorophyll dans les Planaires vertes," "Comptes Rendus," December, 1878.
  11. "Ueber die chemische Zusammensetzung der Hefo," "Sitzungsbericht der math, phys. Classe der k.k. Akad. der Wissens. zu München," 1878.
  12. "The Physical Basis of Life" (see "Essays and Reviews," by T. H. Huxley).