On Growth and Form

“Cum formarum naturalium et corporalium esse non consistat nisi in unione ad materiam, ejusdem agentis esse videtur eas producere cujus est materiam transmutare. Secundo, quia cum hujusmodi formae non excedant virtutem et ordinem et facultatem principiorum agentium in natura, nulla videtur necessitas eorum originem in principia reducere altiora.” Aquinas, De Pot. Q. iii, a, 11. (Quoted in Brit. Assoc. Address, Section D, 1911.)

“...I would that all other natural phenomena might similarly be deduced from mechanical principles. For many things move me to suspect that everything depends upon certain forces, in virtue of which the particles of bodies, through forces not yet understood, are either impelled together so as to cohere in regular figures, or are repelled and recede from one another.” Newton, in Preface to the Principia. (Quoted by Mr W. Spottiswoode, Brit. Assoc. Presidential Address, 1878.)

“When Science shall have subjected all natural phenomena to the laws of Theoretical Mechanics, when she shall be able to predict the result of every combination as unerringly as Hamilton predicted conical refraction, or Adams revealed to us the existence of Neptune,—that we cannot say. That day may never come, and it is certainly far in the dim future. We may not anticipate it, we may not even call it possible. But none the less are we bound to look to that day, and to labour for it as the crowning triumph of Science:—when Theoretical Mechanics shall be recognised as the key to every physical enigma, the chart for every traveller through the dark Infinite of Nature.” J. H. Jellett, in Brit. Assoc. Address, Section A, 1874.

Of the chemistry of his day and generation, Kant declared that it was “a science, but not science,”—“eine Wissenschaft, aber nicht Wissenschaft”; for that the criterion of physical science lay in its relation to mathematics. And a hundred years later Du Bois Reymond, profound student of the many sciences on which physiology is based, recalled and reiterated the old saying, declaring that chemistry would only reach the rank of science, in the high and strict sense, when it should be found possible to explain chemical reactions in the light of their causal relation to the velocities, tensions and conditions of equi­lib­rium of the component molecules; that, in short, the chemistry of the future must deal with molecular mechanics, by the methods and in the strict language of mathematics, as the astronomy of Newton and Laplace dealt with the stars in their courses. We know how great a step has been made towards this distant and once hopeless goal, as Kant defined it, since van’t Hoff laid the firm foundations of a math­e­mat­i­cal chemistry, and earned his proud epitaph, Physicam chemiae adiunxit1.

We need not wait for the full realisation of Kant’s desire, in order to apply to the natural sciences the principle which he urged. Though chemistry fall short of its ultimate goal in math­e­mat­i­cal mechanics, nevertheless physiology is vastly strengthened and enlarged by making use of the chemistry, as of the physics, of the age. Little by little it draws nearer to our conception of a true science, with each branch of physical science which it {2} brings into relation with itself: with every physical law and every math­e­mat­i­cal theorem which it learns to take into its employ. Between the physiology of Haller, fine as it was, and that of Helmholtz, Ludwig, Claude Bernard, there was all the difference in the world.

As soon as we adventure on the paths of the physicist, we learn to weigh and to measure, to deal with time and space and mass and their related concepts, and to find more and more our knowledge expressed and our needs satisfied through the concept of number, as in the dreams and visions of Plato and Pythagoras; for modern chemistry would have gladdened the hearts of those great philosophic dreamers.

But the zoologist or morphologist has been slow, where the physiologist has long been eager, to invoke the aid of the physical or math­e­mat­i­cal sciences; and the reasons for this difference lie deep, and in part are rooted in old traditions. The zoologist has scarce begun to dream of defining, in math­e­mat­i­cal language, even the simpler organic forms. When he finds a simple geometrical construction, for instance in the honey-comb, he would fain refer it to psychical instinct or design rather than to the operation of physical forces; when he sees in snail, or nautilus, or tiny foraminiferal or radiolarian shell, a close approach to the perfect sphere or spiral, he is prone, of old habit, to believe that it is after all something more than a spiral or a sphere, and that in this “something more” there lies what neither physics nor mathematics can explain. In short he is deeply reluctant to compare the living with the dead, or to explain by geometry or by dynamics the things which have their part in the mystery of life. Moreover he is little inclined to feel the need of such explanations or of such extension of his field of thought. He is not without some justification if he feels that in admiration of nature’s handiwork he has an horizon open before his eyes as wide as any man requires. He has the help of many fascinating theories within the bounds of his own science, which, though a little lacking in precision, serve the purpose of ordering his thoughts and of suggesting new objects of enquiry. His art of clas­si­fi­ca­tion becomes a ceaseless and an endless search after the blood-relationships of things living, and the pedigrees of things {3} dead and gone. The facts of embryology become for him, as Wolff, von Baer and Fritz Müller proclaimed, a record not only of the life-history of the individual but of the annals of its race. The facts of geographical distribution or even of the migration of birds lead on and on to speculations regarding lost continents, sunken islands, or bridges across ancient seas. Every nesting bird, every ant-hill or spider’s web displays its psychological problems of instinct or intelligence. Above all, in things both great and small, the naturalist is rightfully impressed, and finally engrossed, by the peculiar beauty which is manifested in apparent fitness or “adaptation,”—the flower for the bee, the berry for the bird.

Time out of mind, it has been by way of the “final cause,” by the teleological concept of “end,” of “purpose,” or of “design,” in one or another of its many forms (for its moods are many), that men have been chiefly wont to explain the phenomena of the living world; and it will be so while men have eyes to see and ears to hear withal. With Galen, as with Aristotle, it was the physician’s way; with John Ray, as with Aristotle, it was the naturalist’s way; with Kant, as with Aristotle, it was the philosopher’s way. It was the old Hebrew way, and has its splendid setting in the story that God made “every plant of the field before it was in the earth, and every herb of the field before it grew.” It is a common way, and a great way; for it brings with it a glimpse of a great vision, and it lies deep as the love of nature in the hearts of men.

Half overshadowing the “efficient” or physical cause, the argument of the final cause appears in eighteenth century physics, in the hands of such men as Euler2 and Maupertuis, to whom Leibniz3 had passed it on. Half overshadowed by the mechanical concept, it runs through Claude Bernard’s Leçons sur les {4} phénomènes de la Vie4, and abides in much of modern physiology5. Inherited from Hegel, it dominated Oken’s Naturphilosophie and lingered among his later disciples, who were wont to liken the course of organic evolution not to the straggling branches of a tree, but to the building of a temple, divinely planned, and the crowning of it with its polished minarets6.

It is retained, somewhat crudely, in modern embryology, by those who see in the early processes of growth a significance “rather prospective than retrospective,” such that the embryonic phenomena must be “referred directly to their usefulness in building the body of the future animal7”:—which is no more, and no less, than to say, with Aristotle, that the organism is the τέλος, or final cause, of its own processes of generation and development. It is writ large in that Entelechy8 which Driesch rediscovered, and which he made known to many who had neither learned of it from Aristotle, nor studied it with Leibniz, nor laughed at it with Voltaire. And, though it is in a very curious way, we are told that teleology was “refounded, reformed or rehabilitated9” by Darwin’s theory of natural selection, whereby “every variety of form and colour was urgently and absolutely called upon to produce its title to existence either as an active useful agent, or as a survival” of such active usefulness in the past. But in this last, and very important case, we have reached a “teleology” without a τέλος, {5} as men like Butler and Janet have been prompt to shew: a teleology in which the final cause becomes little more, if anything, than the mere expression or resultant of a process of sifting out of the good from the bad, or of the better from the worse, in short of a process of mechanism10. The apparent manifestations of “purpose” or adaptation become part of a mechanical philosophy, according to which “chaque chose finit toujours par s’accommoder à son milieu11.” In short, by a road which resembles but is not the same as Maupertuis’s road, we find our way to the very world in which we are living, and find that if it be not, it is ever tending to become, “the best of all possible worlds12.”

But the use of the teleological principle is but one way, not the whole or the only way, by which we may seek to learn how things came to be, and to take their places in the harmonious complexity of the world. To seek not for ends but for “antecedents” is the way of the physicist, who finds “causes” in what he has learned to recognise as fundamental properties, or inseparable concomitants, or unchanging laws, of matter and of energy. In Aristotle’s parable, the house is there that men may live in it; but it is also there because the builders have laid one stone upon another: and it is as a mechanism, or a mechanical construction, that the physicist looks upon the world. Like warp and woof, mechanism and teleology are interwoven together, and we must not cleave to the one and despise the other; for their union is “rooted in the very nature of totality13.”

Nevertheless, when philosophy bids us hearken and obey the lessons both of mechanical and of teleological interpretation, the precept is hard to follow: so that oftentimes it has come to pass, just as in Bacon’s day, that a leaning to the side of the final cause “hath intercepted the severe and diligent inquiry of all {6} real and physical causes,” and has brought it about that “the search of the physical cause hath been neglected and passed in silence.” So long and so far as “fortuitous variation14” and the “survival of the fittest” remain engrained as fundamental and satisfactory hypotheses in the philosophy of biology, so long will these “satisfactory and specious causes” tend to stay “severe and diligent inquiry,” “to the great arrest and prejudice of future discovery.”

The difficulties which surround the concept of active or “real” causation, in Bacon’s sense of the word, difficulties of which Hume and Locke and Aristotle were little aware, need scarcely hinder us in our physical enquiry. As students of math­e­mat­i­cal and of empirical physics, we are content to deal with those antecedents, or concomitants, of our phenomena, without which the phenomenon does not occur,—with causes, in short, which, aliae ex aliis aptae et necessitate nexae, are no more, and no less, than conditions sine quâ non. Our purpose is still adequately fulfilled: inasmuch as we are still enabled to correlate, and to equate, our particular phenomena with more and ever more of the physical phenomena around, and so to weave a web of connection and interdependence which shall serve our turn, though the metaphysician withhold from that interdependence the title of causality. We come in touch with what the schoolmen called a ratio cognoscendi, though the true ratio efficiendi is still enwrapped in many mysteries. And so handled, the quest of physical causes merges with another great Aristotelian theme,—the search for relations between things apparently disconnected, and for “similitude in things to common view unlike.” Newton did not shew the cause of the apple falling, but he shewed a similitude between the apple and the stars.

Moreover, the naturalist and the physicist will continue to speak of “causes,” just as of old, though it may be with some mental reservations: for, as a French philosopher said, in a kindred difficulty: “ce sont là des manières de s’exprimer, {7} et si elles sont interdites il faut renoncer à parler de ces choses.”

The search for differences or essential contrasts between the phenomena of organic and inorganic, of animate and inanimate things has occupied many mens’ minds, while the search for community of principles, or essential similitudes, has been followed by few; and the contrasts are apt to loom too large, great as they may be. M. Dunan, discussing the “Problème de la Vie15” in an essay which M. Bergson greatly commends, declares: “Les lois physico-chimiques sont aveugles et brutales; là où elles règnent seules, au lieu d’un ordre et d’un concert, il ne peut y avoir qu’incohérence et chaos.” But the physicist proclaims aloud that the physical phenomena which meet us by the way have their manifestations of form, not less beautiful and scarce less varied than those which move us to admiration among living things. The waves of the sea, the little ripples on the shore, the sweeping curve of the sandy bay between its headlands, the outline of the hills, the shape of the clouds, all these are so many riddles of form, so many problems of morphology, and all of them the physicist can more or less easily read and adequately solve: solving them by reference to their antecedent phenomena, in the material system of mechanical forces to which they belong, and to which we interpret them as being due. They have also, doubtless, their immanent teleological significance; but it is on another plane of thought from the physicist’s that we contemplate their intrinsic harmony and perfection, and “see that they are good.”

Nor is it otherwise with the material forms of living things. Cell and tissue, shell and bone, leaf and flower, are so many portions of matter, and it is in obedience to the laws of physics that their particles have been moved, moulded and conformed16. {8} They are no exception to the rule that Θεὸς ἀεὶ γεωμετρεῖ. Their problems of form are in the first instance math­e­mat­i­cal problems, and their problems of growth are essentially physical problems; and the morphologist is, ipso facto, a student of physical science.

Apart from the physico-chemical problems of modern physiology, the road of physico-math­e­mat­i­cal or dynamical in­ves­ti­ga­tion in morphology has had few to follow it; but the pathway is old. The way of the old Ionian physicians, of Anaxagoras17, of Empedocles and his disciples in the days before Aristotle, lay just by that highwayside. It was Galileo’s and Borelli’s way. It was little trodden for long afterwards, but once in a while Swammerdam and Réaumur looked that way. And of later years, Moseley and Meyer, Berthold, Errera and Roux have been among the little band of travellers. We need not wonder if the way be hard to follow, and if these wayfarers have yet gathered little. A harvest has been reaped by others, and the gleaning of the grapes is slow.

It behoves us always to remember that in physics it has taken great men to discover simple things. They are very great names indeed that we couple with the explanation of the path of a stone, the droop of a chain, the tints of a bubble, the shadows in a cup. It is but the slightest adumbration of a dynamical morphology that we can hope to have, until the physicist and the mathematician shall have made these problems of ours their own, or till a new Boscovich shall have written for the naturalist the new Theoria Philosophiae Naturalis.

How far, even then, mathematics will suffice to describe, and physics to explain, the fabric of the body no man can foresee. It may be that all the laws of energy, and all the properties of matter, and all the chemistry of all the colloids are as powerless to explain the body as they are impotent to comprehend the soul. For my part, I think it is not so. Of how it is that the soul informs the body, physical science teaches me nothing: consciousness is not explained to my comprehension by all the nerve-paths and “neurones” of the physiologist; nor do I ask of physics how goodness shines in one man’s face, and evil betrays itself in another. But of the construction and growth and working {9} of the body, as of all that is of the earth earthy, physical science is, in my humble opinion, our only teacher and guide18.

Often and often it happens that our physical knowledge is inadequate to explain the mechanical working of the organism; the phenomena are superlatively complex, the procedure is involved and entangled, and the in­ves­ti­ga­tion has occupied but a few short lives of men. When physical science falls short of explaining the order which reigns throughout these manifold phenomena,—an order more char­ac­ter­is­tic in its totality than any of its phenomena in themselves,—men hasten to invoke a guiding principle, an entelechy, or call it what you will. But all the while, so far as I am aware, no physical law, any more than that of gravity itself, not even among the puzzles of chemical “stereometry,” or of physiological “surface-action” or “osmosis,” is known to be transgressed by the bodily mechanism.

Some physicists declare, as Maxwell did, that atoms or molecules more complicated by far than the chemist’s hypotheses demand are requisite to explain the phenomena of life. If what is implied be an explanation of psychical phenomena, let the point be granted at once; we may go yet further, and decline, with Maxwell, to believe that anything of the nature of physical complexity, however exalted, could ever suffice. Other physicists, like Auerbach19, or Larmor20, or Joly21, assure us that our laws of thermodynamics do not suffice, or are “inappropriate,” to explain the maintenance or (in Joly’s phrase) the “accelerative absorption” {10} of the bodily energies, and the long battle against the cold and darkness which is death. With these weighty problems I am not for the moment concerned. My sole purpose is to correlate with math­e­mat­i­cal statement and physical law certain of the simpler outward phenomena of organic growth and structure or form: while all the while regarding, ex hypothesi, for the purposes of this correlation, the fabric of the organism as a material and mechanical configuration.

Physical science and philosophy stand side by side, and one upholds the other. Without something of the strength of physics, philosophy would be weak; and without something of philosophy’s wealth, physical science would be poor. “Rien ne retirera du tissu de la science les fils d’or que la main du philosophe y a introduits22.” But there are fields where each, for a while at least, must work alone; and where physical science reaches its limitations, physical science itself must help us to discover. Meanwhile the appropriate and legitimate postulate of the physicist, in approaching the physical problems of the body, is that with these physical phenomena no alien influence interferes. But the postulate, though it is certainly legitimate, and though it is the proper and necessary prelude to scientific enquiry, may some day be proven to be untrue; and its disproof will not be to the physicist’s confusion, but will come as his reward. In dealing with forms which are so concomitant with life that they are seemingly controlled by life, it is in no spirit of arrogant assertiveness that the physicist begins his argument, after the fashion of a most illustrious exemplar, with the old formulary of scholastic challenge,—An Vita sit? Dico quod non.

The terms Form and Growth, which make up the title of this little book, are to be understood, as I need hardly say, in their relation to the science of organisms. We want to see how, in some cases at least, the forms of living things, and of the parts of living things, can be explained by physical con­si­de­ra­tions, and to realise that, in general, no organic forms exist save such as are in conformity with ordinary physical laws. And while growth is a somewhat vague word for a complex matter, which may {11} depend on various things, from simple imbibition of water to the complicated results of the chemistry of nutrition, it deserves to be studied in relation to form, whether it proceed by simple increase of size without obvious alteration of form, or whether it so proceed as to bring about a gradual change of form and the slow development of a more or less complicated structure.

In the Newtonian language of elementary physics, force is recognised by its action in producing or in changing motion, or in preventing change of motion or in maintaining rest. When we deal with matter in the concrete, force does not, strictly speaking, enter into the question, for force, unlike matter, has no independent objective existence. It is energy in its various forms, known or unknown, that acts upon matter. But when we abstract our thoughts from the material to its form, or from the thing moved to its motions, when we deal with the subjective conceptions of form, or movement, or the movements that change of form implies, then force is the appropriate term for our conception of the causes by which these forms and changes of form are brought about. When we use the term force, we use it, as the physicist always does, for the sake of brevity, using a symbol for the magnitude and direction of an action in reference to the symbol or diagram of a material thing. It is a term as subjective and symbolic as form itself, and so is appropriately to be used in connection therewith.

The form, then, of any portion of matter, whether it be living or dead, and the changes of form that are apparent in its movements and in its growth, may in all cases alike be described as due to the action of force. In short, the form of an object is a “diagram of forces,” in this sense, at least, that from it we can judge of or deduce the forces that are acting or have acted upon it: in this strict and particular sense, it is a diagram,—in the case of a solid, of the forces that have been impressed upon it when its conformation was produced, together with those that enable it to retain its conformation; in the case of a liquid (or of a gas) of the forces that are for the moment acting on it to restrain or balance its own inherent mobility. In an organism, great or small, it is not merely the nature of the motions of the living substance that we must interpret in terms of force (according to kinetics), but also {12} the conformation of the organism itself, whose permanence or equi­lib­rium is explained by the interaction or balance of forces, as described in statics.

If we look at the living cell of an Amoeba or a Spirogyra, we see a something which exhibits certain active movements, and a certain fluctuating, or more or less lasting, form; and its form at a given moment, just like its motions, is to be investigated by the help of physical methods, and explained by the invocation of the math­e­mat­i­cal conception of force.

Now the state, including the shape or form, of a portion of matter, is the resultant of a number of forces, which represent or symbolise the manifestations of various kinds of energy; and it is obvious, accordingly, that a great part of physical science must be understood or taken for granted as the necessary preliminary to the discussion on which we are engaged. But we may at least try to indicate, very briefly, the nature of the principal forces and the principal properties of matter with which our subject obliges us to deal. Let us imagine, for instance, the case of a so-called “simple” organism, such as Amoeba; and if our short list of its physical properties and conditions be helpful to our further discussion, we need not consider how far it be complete or adequate from the wider physical point of view23.

This portion of matter, then, is kept together by the intermolecular force of cohesion; in the movements of its particles relatively to one another, and in its own movements relative to adjacent matter, it meets with the opposing force of friction. It is acted on by gravity, and this force tends (though slightly, owing to the Amoeba’s small mass, and to the small difference between its density and that of the surrounding fluid), to flatten it down upon the solid substance on which it may be creeping. Our Amoeba tends, in the next place, to be deformed by any pressure from outside, even though slight, which may be applied to it, and this circumstance shews it to consist of matter in a fluid, or at least semi-fluid, state: which state is further indicated when we observe streaming or current motions in its interior. {13} Like other fluid bodies, its surface, whatsoever other substance, gas, liquid or solid, it be in contact with, and in varying degree according to the nature of that adjacent substance, is the seat of molecular force exhibiting itself as a surface-tension, from the action of which many important consequences follow, which greatly affect the form of the fluid surface.

While the protoplasm of the Amoeba reacts to the slightest pressure, and tends to “flow,” and while we therefore speak of it as a fluid, it is evidently far less mobile than such a fluid, for instance, as water, but is rather like treacle in its slow creeping movements as it changes its shape in response to force. Such fluids are said to have a high viscosity, and this viscosity obviously acts in the way of retarding change of form, or in other words of retarding the effects of any disturbing action of force. When the viscous fluid is capable of being drawn out into fine threads, a property in which we know that the material of some Amoebae differs greatly from that of others, we say that the fluid is also viscid, or exhibits viscidity. Again, not by virtue of our Amoeba being liquid, but at the same time in vastly greater measure than if it were a solid (though far less rapidly than if it were a gas), a process of molecular diffusion is constantly going on within its substance, by which its particles interchange their places within the mass, while surrounding fluids, gases and solids in solution diffuse into and out of it. In so far as the outer wall of the cell is different in character from the interior, whether it be a mere pellicle as in Amoeba or a firm cell-wall as in Protococcus, the diffusion which takes place through this wall is sometimes distinguished under the term osmosis.

Within the cell, chemical forces are at work, and so also in all probability (to judge by analogy) are electrical forces; and the organism reacts also to forces from without, that have their origin in chemical, electrical and thermal influences. The processes of diffusion and of chemical activity within the cell result, by the drawing in of water, salts, and food-material with or without chemical transformation into protoplasm, in growth, and this complex phenomenon we shall usually, without discussing its nature and origin, describe and picture as a force. Indeed we shall manifestly be inclined to use the term growth in two senses, {14} just indeed as we do in the case of attraction or gravitation, on the one hand as a process, and on the other hand as a force.

In the phenomena of cell-division, in the attractions or repulsions of the parts of the dividing nucleus and in the “caryokinetic” figures that appear in connection with it, we seem to see in operation forces and the effects of forces, that have, to say the least of it, a close analogy with known physical phenomena; and to this matter we shall afterwards recur. But though they resemble known physical phenomena, their nature is still the subject of much discussion, and neither the forms produced nor the forces at work can yet be satisfactorily and simply explained. We may readily admit, then, that besides phenomena which are obviously physical in their nature, there are actions visible as well as invisible taking place within living cells which our knowledge does not permit us to ascribe with certainty to any known physical force; and it may or may not be that these phenomena will yield in time to the methods of physical in­ves­ti­ga­tion. Whether or no, it is plain that we have no clear rule or guide as to what is “vital” and what is not; the whole assemblage of so-called vital phenomena, or properties of the organism, cannot be clearly classified into those that are physical in origin and those that are sui generis and peculiar to living things. All we can do meanwhile is to analyse, bit by bit, those parts of the whole to which the ordinary laws of the physical forces more or less obviously and clearly and indubitably apply.

Morphology then is not only a study of material things and of the forms of material things, but has its dynamical aspect, under which we deal with the interpretation, in terms of force, of the operations of Energy. And here it is well worth while to remark that, in dealing with the facts of embryology or the phenomena of inheritance, the common language of the books seems to deal too much with the material elements concerned, as the causes of development, of variation or of hereditary transmission. Matter as such produces nothing, changes nothing, does nothing; and however convenient it may afterwards be to abbreviate our nomenclature and our descriptions, we must most carefully realise in the outset that the spermatozoon, the nucleus, {15} the chromosomes or the germ-plasm can never act as matter alone, but only as seats of energy and as centres of force. And this is but an adaptation (in the light, or rather in the conventional symbolism, of modern physical science) of the old saying of the philosopher: ἀρχὴ γὰρ ἡ φύσις μᾶλλον τῆς ὕλης.

To terms of magnitude, and of direction, must we refer all our conceptions of form. For the form of an object is defined when we know its magnitude, actual or relative, in various directions; and growth involves the same conceptions of magnitude and direction, with this addition, that they are supposed to alter in time. Before we proceed to the consideration of specific form, it will be worth our while to consider, for a little while, certain phenomena of spatial magnitude, or of the extension of a body in the several dimensions of space24.

We are taught by elementary mathematics that, in similar solid figures, the surface increases as the square, and the volume as the cube, of the linear dimensions. If we take the simple case of a sphere, with radius r, the area of its surface is equal to 4πr² , and its volume to (⁴⁄₃)πr³ ; from which it follows that the ratio of volume to surface, or ^V⁄_S , is (¹⁄₃)r. In other words, the greater the radius (or the larger the sphere) the greater will be its volume, or its mass (if it be uniformly dense throughout), in comparison with its superficial area. And, taking L to represent any linear dimension, we may write the general equations in the form

From these elementary principles a great number of consequences follow, all more or less interesting, and some of them of great importance. In the first place, though growth in length (let {17} us say) and growth in volume (which is usually tantamount to mass or weight) are parts of one and the same process or phenomenon, the one attracts our attention by its increase, very much more than the other. For instance a fish, in doubling its length, multiplies its weight by no less than eight times; and it all but doubles its weight in growing from four inches long to five.

In the second place we see that a knowledge of the correlation between length and weight in any particular species of animal, in other words a determination of k in the formula W = k · L³ , enables us at any time to translate the one magnitude into the other, and (so to speak) to weigh the animal with a measuring-rod; this however being always subject to the condition that the animal shall in no way have altered its form, nor its specific gravity. That its specific gravity or density should materially or rapidly alter is not very likely; but as long as growth lasts, changes of form, even though inappreciable to the eye, are likely to go on. Now weighing is a far easier and far more accurate operation than measuring; and the measurements which would reveal slight and otherwise imperceptible changes in the form of a fish—slight relative differences between length, breadth and depth, for instance,—would need to be very delicate indeed. But if we can make fairly accurate determinations of the length, which is very much the easiest dimension to measure, and then correlate it with the weight, then the value of k, according to whether it varies or remains constant, will tell us at once whether there has or has not been a tendency to gradual alteration in the general form. To this subject we shall return, when we come to consider more particularly the rate of growth.

But a much deeper interest arises out of this changing ratio of dimensions when we come to consider the inevitable changes of physical relations with which it is bound up. We are apt, and even accustomed, to think that magnitude is so purely relative that differences of magnitude make no other or more essential difference; that Lilliput and Brobdingnag are all alike, according as we look at them through one end of the glass or the other. But this is by no means so; for scale has a very marked effect upon physical phenomena, and the effect of scale constitutes what is known as the principle of similitude, or of dynamical similarity. {18}

This effect of scale is simply due to the fact that, of the physical forces, some act either directly at the surface of a body, or otherwise in proportion to the area of surface; and others, such as gravity, act on all particles, internal and external alike, and exert a force which is proportional to the mass, and so usually to the volume, of the body.

The strength of an iron girder obviously varies with the cross-section of its members, and each cross-section varies as the square of a linear dimension; but the weight of the whole structure varies as the cube of its linear dimensions. And it follows at once that, if we build two bridges geometrically similar, the larger is the weaker of the two25. It was elementary engineering experience such as this that led Herbert Spencer26 to apply the principle of similitude to biology.

The same principle had been admirably applied, in a few clear instances, by Lesage27, a celebrated eighteenth century physician of Geneva, in an unfinished and unpublished work28. Lesage argued, for instance, that the larger ratio of surface to mass would lead in a small animal to excessive transpiration, were the skin as “porous” as our own; and that we may hence account for the hardened or thickened skins of insects and other small terrestrial animals. Again, since the weight of a fruit increases as the cube of its dimensions, while the strength of the stalk increases as the square, it follows that the stalk should grow out of apparent due proportion to the fruit; or alternatively, that tall trees should not bear large fruit on slender branches, and that melons and pumpkins must lie upon the ground. And again, that in quadrupeds a large head must be supported on a neck which is either {19} excessively thick and strong, like a bull’s, or very short like the neck of an elephant.

But it was Galileo who, wellnigh 300 years ago, had first laid down this general principle which we now know by the name of the principle of similitude; and he did so with the utmost possible clearness, and with a great wealth of illustration, drawn from structures living and dead29. He showed that neither can man build a house nor can nature construct an animal beyond a certain size, while retaining the same proportions and employing the same materials as sufficed in the case of a smaller structure30. The thing will fall to pieces of its own weight unless we either change its relative proportions, which will at length cause it to become clumsy, monstrous and inefficient, or else we must find a new material, harder and stronger than was used before. Both processes are familiar to us in nature and in art, and practical applications, undreamed of by Galileo, meet us at every turn in this modern age of steel.

Again, as Galileo was also careful to explain, besides the questions of pure stress and strain, of the strength of muscles to lift an increasing weight or of bones to resist its crushing stress, we have the very important question of bending moments. This question enters, more or less, into our whole range of problems; it affects, as we shall afterwards see, or even determines the whole form of the skeleton, and is very important in such a case as that of a tall tree31.

Here we have to determine the point at which the tree will curve under its own weight, if it be ever so little displaced from the perpendicular32. In such an in­ves­ti­ga­tion we have to make {20} some assumptions,—for instance, with regard to the trunk, that it tapers uniformly, and with regard to the branches that their sectional area varies according to some definite law, or (as Ruskin assumed33) tends to be constant in any horizontal plane; and the math­e­mat­i­cal treatment is apt to be somewhat difficult. But Greenhill has shewn that (on such assumptions as the above), a certain British Columbian pine-tree, which yielded the Kew flagstaff measuring 221 ft. in height with a diameter at the base of 21 inches, could not possibly, by theory, have grown to more than about 300 ft. It is very curious that Galileo suggested precisely the same height (dugento braccia alta) as the utmost limit of the growth of a tree. In general, as Greenhill shews, the diameter of a homogeneous body must increase as the power 3 ⁄ 2 of the height, which accounts for the slender proportions of young trees, compared with the stunted appearance of old and large ones34. In short, as Goethe says in Wahrheit und Dichtung, “Es ist dafür gesorgt dass die Bäume nicht in den Himmel wachsen.” But Eiffel’s great tree of steel (1000 feet high) is built to a very different plan; for here the profile of the tower follows the logarithmic curve, giving equal strength throughout, according to a principle which we shall have occasion to discuss when we come to treat of “form and mechanical efficiency” in connection with the skeletons of animals.

Among animals, we may see in a general way, without the help of mathematics or of physics, that exaggerated bulk brings with it a certain clumsiness, a certain inefficiency, a new element of risk and hazard, a vague preponderance of disadvantage. The case was well put by Owen, in a passage which has an interest of its own as a premonition (somewhat like De Candolle’s) of the “struggle for existence.” Owen wrote as follows35: “In proportion to the bulk of a species is the difficulty of the contest which, as a living organised whole, the individual of such species {21} has to maintain against the surrounding agencies that are ever tending to dissolve the vital bond, and subjugate the living matter to the ordinary chemical and physical forces. Any changes, therefore, in such external conditions as a species may have been originally adapted to exist in, will militate against that existence in a degree proportionate, perhaps in a geometrical ratio, to the bulk of the species. If a dry season be greatly prolonged, the large mammal will suffer from the drought sooner than the small one; if any alteration of climate affect the quantity of vegetable food, the bulky Herbivore will first feel the effects of stinted nourishment.”

But the principle of Galileo carries us much further and along more certain lines.

The tensile strength of a muscle, like that of a rope or of our girder, varies with its cross-section; and the resistance of a bone to a crushing stress varies, again like our girder, with its cross-section. But in a terrestrial animal the weight which tends to crush its limbs or which its muscles have to move, varies as the cube of its linear dimensions; and so, to the possible magnitude of an animal, living under the direct action of gravity, there is a definite limit set. The elephant, in the dimensions of its limb-bones, is already shewing signs of a tendency to disproportionate thickness as compared with the smaller mammals; its movements are in many ways hampered and its agility diminished: it is already tending towards the maximal limit of size which the physical forces permit. But, as Galileo also saw, if the animal be wholly immersed in water, like the whale, (or if it be partly so, as was in all probability the case with the giant reptiles of our secondary rocks), then the weight is counterpoised to the extent of an equivalent volume of water, and is completely counterpoised if the density of the animal’s body, with the included air, be identical (as in a whale it very nearly is) with the water around. Under these circumstances there is no longer a physical barrier to the indefinite growth in magnitude of the animal36. Indeed, {22} in the case of the aquatic animal there is, as Spencer pointed out, a distinct advantage, in that the larger it grows the greater is its velocity. For its available energy depends on the mass of its muscles; while its motion through the water is opposed, not by gravity, but by “skin-friction,” which increases only as the square of its dimensions; all other things being equal, the bigger the ship, or the bigger the fish, the faster it tends to go, but only in the ratio of the square root of the increasing length. For the mechanical work (W) of which the fish is capable being proportional to the mass of its muscles, or the cube of its linear dimensions: and again this work being wholly done in producing a velocity (V) against a resistance (R) which increases as the square of the said linear dimensions; we have at once

But there is often another side to these questions, which makes them too complicated to answer in a word. For instance, the work (per stroke) of which two similar engines are capable should obviously vary as the cubes of their linear dimensions, for it varies on the one hand with the surface of the piston, and on the other, with the length of the stroke; so is it likewise in the animal, where the cor­re­spon­ding variation depends on the cross-section of the muscle, and on the space through which it contracts. But in two precisely similar engines, the actual available horse-power varies as the square of the linear dimensions, and not as the cube; and this for the obvious reason that the actual energy developed depends upon the heating-surface of the boiler37. So likewise must there be a similar tendency, among animals, for the rate of supply of kinetic energy to vary with the surface of the {23} lung, that is to say (other things being equal) with the square of the linear dimensions of the animal. We may of course (departing from the condition of similarity) increase the heating-surface of the boiler, by means of an internal system of tubes, without increasing its outward dimensions, and in this very way nature increases the respiratory surface of a lung by a complex system of branching tubes and minute air-cells; but nevertheless in two similar and closely related animals, as also in two steam-engines of precisely the same make, the law is bound to hold that the rate of working must tend to vary with the square of the linear dimensions, according to Froude’s law of steamship comparison. In the case of a very large ship, built for speed, the difficulty is got over by increasing the size and number of the boilers, till the ratio between boiler-room and engine-room is far beyond what is required in an ordinary small vessel38; but though we find lung-space increased among animals where greater rate of working is required, as in general among birds, I do not know that it can be shewn to increase, as in the “over-boilered” ship, with the size of the animal, and in a ratio which outstrips that of the other bodily dimensions. If it be the case then, that the working mechanism of the muscles should be able to exert a force proportionate to the cube of the linear bodily dimensions, while the respiratory mechanism can only supply a store of energy at a rate proportional to the square of the said dimensions, the singular result ought to follow that, in swimming for instance, the larger fish ought to be able to put on a spurt of speed far in excess of the smaller one; but the distance travelled by the year’s end should be very much alike for both of them. And it should also follow that the curve of fatigue {24} should be a steeper one, and the staying power should be less, in the smaller than in the larger individual. This is the case of long-distance racing, where the big winner puts on his big spurt at the end. And for an analogous reason, wise men know that in the ’Varsity boat-race it is judicious and prudent to bet on the heavier crew.

Leaving aside the question of the supply of energy, and keeping to that of the mechanical efficiency of the machine, we may find endless biological illustrations of the principle of similitude.

In the case of the flying bird (apart from the initial difficulty of raising itself into the air, which involves another problem) it may be shewn that the bigger it gets (all its proportions remaining the same) the more difficult it is for it to maintain itself aloft in flight. The argument is as follows:

In order to keep aloft, the bird must communicate to the air a downward momentum equivalent to its own weight, and therefore proportional to the cube of its own linear dimensions. But the momentum so communicated is proportional to the mass of air driven downwards, and to the rate at which it is driven: the mass being proportional to the bird’s wing-area, and also (with any given slope of wing) to the speed of the bird, and the rate being again proportional to the bird’s speed; accordingly the whole momentum varies as the wing-area, i.e. as the square of the linear dimensions, and also as the square of the speed. Therefore, in order that the bird may maintain level flight, its speed must be proportional to the square root of its linear dimensions.

Now the rate at which the bird, in steady flight, has to work in order to drive itself forward, is the rate at which it communicates energy to the air; and this is proportional to mV² , i.e. to the mass and to the square of the velocity of the air displaced. But the mass of air displaced per second is proportional to the wing-area and to the speed of the bird’s motion, and therefore to the power 2½ of the linear dimensions; and the speed at which it is displaced is proportional to the bird’s speed, and therefore to the square root of the linear dimensions. Therefore the energy communicated per second (being proportional to the mass and to the square of the speed) is jointly proportional to the power 2½ of the linear dimensions, as above, and to the first power thereof: {25} that is to say, it increases in proportion to the power 3½ of the linear dimensions, and therefore faster than the weight of the bird increases.

The work requiring to be done, then, varies as the power 3½ of the bird’s linear dimensions, while the work of which the bird is capable depends on the mass of its muscles, and therefore varies as the cube of its linear dimensions39. The disproportion does not seem at first sight very great, but it is quite enough to tell. It is as much as to say that, every time we double the linear dimensions of the bird, the difficulty of flight is increased in the ratio of 2³ : 2^3½ , or 8 : 11·3, or, say, 1 : 1·4. If we take the ostrich to exceed the sparrow in linear dimensions as 25 : 1, which seems well within the mark, we have the ratio between 25^3½ and 25³ , or between 5⁷ : 5⁶ ; in other words, flight is just five times more difficult for the larger than for the smaller bird40.

The above in­ves­ti­ga­tion includes, besides the final result, a number of others, explicit or implied, which are of not less importance. Of these the simplest and also the most important is {26} contained in the equation V = √l, a result which happens to be identical with one we had also arrived at in the case of the fish. In the bird’s case it has a deeper significance than in the other; because it implies here not merely that the velocity will tend to increase in a certain ratio with the length, but that it must do so as an essential and primary condition of the bird’s remaining aloft. It is accordingly of great practical importance in aeronautics, for it shews how a provision of increasing speed must accompany every enlargement of our aeroplanes. If a given machine weighing, say, 500 lbs. be stable at 40 miles an hour, then one geometrically similar which weighs, say, a couple of tons must have its speed determined as follows:

That is to say, the larger machine must be capable of a speed equal to 1·414 × 40, or about 56½ miles per hour.

It is highly probable, as Lanchester41 remarks, that Lilienthal met his untimely death not so much from any intrinsic fault in the design or construction of his machine, but simply because his engine fell somewhat short of the power required to give the speed which was necessary for stability. An arrow is a very imperfectly designed aeroplane, but nevertheless it is evidently capable, to a certain extent and at a high velocity, of acquiring “stability” and hence of actual “flight”: the duration and consequent range of its trajectory, as compared with a bullet of similar initial velocity, being correspondingly benefited. When we return to our birds, and again compare the ostrich with the sparrow, we know little or nothing about the speed in flight of the latter, but that of the swift is estimated42 to vary from a minimum of 20 to 50 feet or more per second,—say from 14 to 35 miles per hour. Let us take the same lower limit as not far from the minimal velocity of the sparrow’s flight also; and it {27} would follow that the ostrich, of 25 times the sparrow’s linear dimensions, would be compelled to fly (if it flew at all) with a minimum velocity of 5 × 14, or 70 miles an hour.

The same principle of necessary speed, or the indispensable relation between the dimensions of a flying object and the minimum velocity at which it is stable, accounts for a great number of observed phenomena. It tells us why the larger birds have a marked difficulty in rising from the ground, that is to say, in acquiring to begin with the horizontal velocity necessary for their support; and why accordingly, as Mouillard43 and others have observed, the heavier birds, even those weighing no more than a pound or two, can be effectively “caged” in a small enclosure open to the sky. It tells us why very small birds, especially those as small as humming-birds, and à fortiori the still smaller insects, are capable of “stationary flight,” a very slight and scarcely perceptible velocity relatively to the air being sufficient for their support and stability. And again, since it is in all cases velocity relative to the air that we are speaking of, we comprehend the reason why one may always tell which way the wind blows by watching the direction in which a bird starts to fly.

It is not improbable that the ostrich has already reached a magnitude, and we may take it for certain that the moa did so, at which flight by muscular action, according to the normal anatomy of a bird, has become physiologically impossible. The same reasoning applies to the case of man. It would be very difficult, and probably absolutely impossible, for a bird to fly were it the bigness of a man. But Borelli, in discussing this question, laid even greater stress on the obvious fact that a man’s pectoral muscles are so immensely less in proportion than those of a bird, that however we may fit ourselves with wings we can never expect to move them by any power of our own relatively weaker muscles; so it is that artificial flight only became possible when an engine was devised whose efficiency was extraordinarily great in comparison with its weight and size.

Had Leonardo da Vinci known what Galileo knew, he would not have spent a great part of his life on vain efforts to make to himself wings. Borelli had learned the lesson thoroughly, and {28} in one of his chapters he deals with the proposition, “Est impossible, ut homines propriis viribus artificiose volare possint44.”

But just as it is easier to swim than to fly, so is it obvious that, in a denser atmosphere, the conditions of flight would be altered, and flight facilitated. We know that in the carboniferous epoch there lived giant dragon-flies, with wings of a span far greater than nowadays they ever attain; and the small bodies and huge extended wings of the fossil pterodactyles would seem in like manner to be quite abnormal according to our present standards, and to be beyond the limits of mechanical efficiency under present conditions. But as Harlé suggests45, following upon a suggestion of Arrhenius, we have only to suppose that in carboniferous and jurassic days the terrestrial atmosphere was notably denser than it is at present, by reason, for instance, of its containing a much larger proportion of carbonic acid, and we have at once a means of reconciling the apparent mechanical discrepancy.

Very similar problems, involving in various ways the principle of dynamical similitude, occur all through the physiology of locomotion: as, for instance, when we see that a cockchafer can carry a plate, many times his own weight, upon his back, or that a flea can jump many inches high.

Problems of this latter class have been admirably treated both by Galileo and by Borelli, but many later writers have remained ignorant of their work. Linnaeus, for instance, remarked that, if an elephant were as strong in proportion as a stag-beetle, it would be able to pull up rocks by the root, and to level mountains. And Kirby and Spence have a well-known passage directed to shew that such powers as have been conferred upon the insect have been withheld from the higher animals, for the reason that had these latter been endued therewith they would have “caused the early desolation of the world46.” {29}

Such problems as that which is presented by the flea’s jumping powers, though essentially physiological in their nature, have their interest for us here: because a steady, progressive diminution of activity with increasing size would tend to set limits to the possible growth in magnitude of an animal just as surely as those factors which tend to break and crush the living fabric under its own weight. In the case of a leap, we have to do rather with a sudden impulse than with a continued strain, and this impulse should be measured in terms of the velocity imparted. The velocity is proportional to the impulse (x), and inversely proportional to the mass (M) moved: V = x ⁄ M. But, according to what we still speak of as “Borelli’s law,” the impulse (i.e. the work of the impulse) is proportional to the volume of the muscle by which it is produced47, that is to say (in similarly constructed animals) to the mass of the whole body; for the impulse is proportional on the one hand to the cross-section of the muscle, and on the other to the distance through which it contracts. It follows at once from this that the velocity is constant, whatever be the size of the animals: in other words, that all animals, provided always that they are similarly fashioned, with their various levers etc., in like proportion, ought to jump, not to the same relative, but to the same actual height48. According to this, then, the flea is not a better, but rather a worse jumper than a horse or a man. As a matter of fact, Borelli is careful to point out that in the act of leaping the impulse is not actually instantaneous, as in the blow of a hammer, but takes some little time, during which the levers are being extended by which the centre of gravity of the animal is being propelled forwards; and this interval of time will be longer in the case of the longer levers of the larger animal. To some extent, then, this principle acts as a corrective to the more general one, {30} and tends to leave a certain balance of advantage, in regard to leaping power, on the side of the larger animal49.

But on the other hand, the question of strength of materials comes in once more, and the factors of stress and strain and bending moment make it, so to speak, more and more difficult for nature to endow the larger animal with the length of lever with which she has provided the flea or the grasshopper.

To Kirby and Spence it seemed that “This wonderful strength of insects is doubtless the result of something peculiar in the structure and arrangement of their muscles, and principally their extraordinary power of contraction.” This hypothesis, which is so easily seen, on physical grounds, to be unnecessary, has been amply disproved in a series of excellent papers by F. Plateau50.

A somewhat simple problem is presented to us by the act of walking. It is obvious that there will be a great economy of work, if the leg swing at its normal pendulum-rate; and, though this rate is hard to calculate, owing to the shape and the jointing of the limb, we may easily convince ourselves, by counting our steps, that the leg does actually swing, or tend to swing, just as a pendulum does, at a certain definite rate51. When we walk quicker, we cause the leg-pendulum to describe a greater arc, but we do not appreciably cause it to swing, or vibrate, quicker, until we shorten the pendulum and begin to run. Now let two individuals, A and B, walk in a similar fashion, that is to say, with a similar angle of swing. The arc through which the leg swings, or the amplitude of each step, will therefore vary as the length of leg, or say as a ⁄ b; but the time of swing will vary as the square {31} root of the pendulum-length, or √a ⁄ √b. Therefore the velocity, which is measured by amplitude ⁄ time, will also vary as the square-roots of the length of leg: that is to say, the average velocities of A and B are in the ratio of √a : √b.

The smaller man, or smaller animal, is so far at a disadvantage compared with the larger in speed, but only to the extent of the ratio between the square roots of their linear dimensions: whereas, if the rate of movement of the limb were identical, irrespective of the size of the animal,—if the limbs of the mouse for instance swung at the same rate as those of the horse,—then, as F. Plateau said, the mouse would be as slow or slower in its gait than the tortoise. M. Delisle52 observed a “minute fly” walk three inches in half-a-second. This was good steady walking. When we walk five miles an hour we go about 88 inches in a second, or 88 ⁄ 6 = 14·7 times the pace of M. Delisle’s fly. We should walk at just about the fly’s pace if our stature were 1 ⁄ (14·7)² , or 1 ⁄ 216 of our present height,—say 72 ⁄ 216 inches, or one-third of an inch high.

But the leg comprises a complicated system of levers, by whose various exercise we shall obtain very different results. For instance, by being careful to rise upon our instep, we considerably increase the length or amplitude of our stride, and very considerably increase our speed accordingly. On the other hand, in running, we bend and so shorten the leg, in order to accommodate it to a quicker rate of pendulum-swing53. In short, the jointed structure of the leg permits us to use it as the shortest possible pendulum when it is swinging, and as the longest possible lever when it is exerting its propulsive force.

Apart from such modifications as that described in the last paragraph,—apart, that is to say, from differences in mechanical construction or in the manner in which the mechanism is used,—we have now arrived at a curiously simple and uniform result. For in all the three forms of locomotion which we have attempted {32} to study, alike in swimming, in flight and in walking, the general result, attained under very different conditions and arrived at by very different modes of reasoning, is in every case that the velocity tends to vary as the square root of the linear dimensions of the organism.

From all the foregoing discussion we learn that, as Crookes once upon a time remarked54, the form as well as the actions of our bodies are entirely conditioned (save for certain exceptions in the case of aquatic animals, nicely balanced with the density of the surrounding medium) by the strength of gravity upon this globe. Were the force of gravity to be doubled, our bipedal form would be a failure, and the majority of terrestrial animals would resemble short-legged saurians, or else serpents. Birds and insects would also suffer, though there would be some compensation for them in the increased density of the air. While on the other hand if gravity were halved, we should get a lighter, more graceful, more active type, requiring less energy and less heat, less heart, less lungs, less blood.

Throughout the whole field of morphology we may find examples of a tendency (referable doubtless in each case to some definite physical cause) for surface to keep pace with volume, through some alteration of its form. The development of “villi” on the inner surface of the stomach and intestine (which enlarge its surface much as we enlarge the effective surface of a bath-towel), the various valvular folds of the intestinal lining, including the remarkable “spiral fold” of the shark’s gut, the convolutions of the brain, whose complexity is evidently correlated (in part at least) with the magnitude of the animal,—all these and many more are cases in which a more or less constant ratio tends to be maintained between mass and surface, which ratio would have been more and more departed from had it not been for the alterations of surface-form55. {33}

In the case of very small animals, and of individual cells, the principle becomes especially important, in consequence of the molecular forces whose action is strictly limited to the superficial layer. In the cases just mentioned, action is facilitated by increase of surface: diffusion, for instance, of nutrient liquids or respiratory gases is rendered more rapid by the greater area of surface; but there are other cases in which the ratio of surface to mass may make an essential change in the whole condition of the system. We know, for instance, that iron rusts when exposed to moist air, but that it rusts ever so much faster, and is soon eaten away, if the iron be first reduced to a heap of small filings; this is a mere difference of degree. But the spherical surface of the raindrop and the spherical surface of the ocean (though both happen to be alike in math­e­mat­i­cal form) are two totally different phenomena, the one due to surface-energy, and the other to that form of mass-energy which we ascribe to gravity. The contrast is still more clearly seen in the case of waves: for the little ripple, whose form and manner of propagation are governed by surface-tension, is found to travel with a velocity which is inversely as the square root of its length; while the ordinary big waves, controlled by gravitation, have a velocity directly proportional to the square root of their wave-length. In like manner we shall find that the form of all small organisms is largely independent of gravity, and largely if not mainly due to the force of surface-tension: either as the direct result of the continued action of surface tension on the semi-fluid body, or else as the result of its action at a prior stage of development, in bringing about a form which subsequent chemical changes have rendered rigid and lasting. In either case, we shall find a very great tendency in small organisms to assume either the spherical form or other simple forms related to ordinary inanimate surface-tension phenomena; which forms do not recur in the external morphology of large animals, or if they in part recur it is for other reasons. {34}

Now this is a very important matter, and is a notable illustration of that principle of similitude which we have already discussed in regard to several of its manifestations. We are coming easily to a conclusion which will affect the whole course of our argument throughout this book, namely that there is an essential difference in kind between the phenomena of form in the larger and the smaller organisms. I have called this book a study of Growth and Form, because in the most familiar illustrations of organic form, as in our own bodies for example, these two factors are inseparably associated, and because we are here justified in thinking of form as the direct resultant and consequence of growth: of growth, whose varying rate in one direction or another has produced, by its gradual and unequal increments, the successive stages of development and the final configuration of the whole material structure. But it is by no means true that form and growth are in this direct and simple fashion correlative or complementary in the case of minute portions of living matter. For in the smaller organisms, and in the individual cells of the larger, we have reached an order of magnitude in which the intermolecular forces strive under favourable conditions with, and at length altogether outweigh, the force of gravity, and also those other forces leading to movements of convection which are the prevailing factors in the larger material aggregate.

However we shall require to deal more fully with this matter in our discussion of the rate of growth, and we may leave it meanwhile, in order to deal with other matters more or less directly concerned with the magnitude of the cell.

The living cell is a very complex field of energy, and of energy of many kinds, surface-energy included. Now the whole surface-energy of the cell is by no means restricted to its outer surface; for the cell is a very heterogeneous structure, and all its protoplasmic alveoli and other visible (as well as invisible) heterogeneities make up a great system of internal surfaces, at every part of which one “phase” comes in contact with another “phase,” and surface-energy is accordingly manifested. But still, the external surface is a definite portion of the system, with a definite “phase” of its own, and however little we may know of the distribution of the total energy of the system, it is at least plain that {35} the conditions which favour equi­lib­rium will be greatly altered by the changed ratio of external surface to mass which a change of magnitude, unaccompanied by change of form, produces in the cell. In short, however it may be brought about, the phenomenon of division of the cell will be precisely what is required to keep ap­prox­i­mate­ly constant the ratio between surface and mass, and to restore the balance between the surface-energy and the other energies of the system. When a germ-cell, for instance, divides or “segments” into two, it does not increase in mass; at least if there be some slight alleged tendency for the egg to increase in mass or volume during segmentation, it is very slight indeed, generally imperceptible, and wholly denied by some56. The development or growth of the egg from a one-celled stage to stages of two or many cells, is thus a somewhat peculiar kind of growth; it is growth which is limited to increase of surface, unaccompanied by growth in volume or in mass.

In the case of a soap-bubble, by the way, if it divide into two bubbles, the volume is actually diminished57 while the surface-area is greatly increased. This is due to a cause which we shall have to study later, namely to the increased pressure due to the greater curvature of the smaller bubbles.

An immediate and remarkable result of the principles just described is a tendency on the part of all cells, according to their kind, to vary but little about a certain mean size, and to have, in fact, certain absolute limitations of magnitude.

Sachs58 pointed out, in 1895, that there is a tendency for each nucleus to be only able to gather around itself a certain definite amount of protoplasm. Driesch59, a little later, found that, by artificial subdivision of the egg, it was possible to rear dwarf sea-urchin larvae, one-half, one-quarter, or even one-eighth of their {36} normal size; and that these dwarf bodies were composed of only a half, a quarter or an eighth of the normal number of cells. Similar observations have been often repeated and amply confirmed. For instance, in the development of Crepidula (a little American “slipper-limpet,” now much at home on our own oyster-beds), Conklin60 has succeeded in rearing dwarf and giant individuals, of which the latter may be as much as twenty-five times as big as the former. But nevertheless, the individual cells, of skin, gut, liver, muscle, and of all the other tissues, are just the same size in one as in the other,—in dwarf and in giant61. Driesch has laid particular stress upon this principle of a “fixed cell-size.”

We get an excellent, and more familiar illustration of the same principle in comparing the large brain-cells or ganglion-cells, both of the lower and of the higher animals62.

Fig. 1. Motor ganglion-cells, from the cervical spinal cord.
(From Minot, after Irving Hardesty.)

In Fig. 1 we have certain identical nerve-cells taken from various mammals, from the mouse to the elephant, all represented on the same scale of magnification; and we see at once that they are all of much the same order of magnitude. The nerve-cell of the elephant is about twice that of the mouse in linear dimensions, and therefore about eight times greater in volume, or mass. But making some allowance for difference of shape, the linear dimensions of the elephant are to those of the mouse in a ratio certainly not less than one to fifty; from which it would follow that the bulk of the larger animal is something like 125,000 times that of the less. And it also follows, the size of the nerve-cells being {37} about as eight to one, that, in cor­re­spon­ding parts of the nervous system of the two animals, there are more than 15,000 times as many individual cells in one as in the other. In short we may (with Enriques) lay it down as a general law that among animals, whether large or small, the ganglion-cells vary in size within narrow limits; and that, amidst all the great variety of structural type of ganglion observed in different classes of animals, it is always found that the smaller species have simpler ganglia than the larger, that is to say ganglia containing a smaller number of cellular elements63. The bearing of such simple facts as this upon the cell-theory in general is not to be disregarded; and the warning is especially clear against exaggerated attempts to correlate physiological processes with the visible mechanism of associated cells, rather than with the system of energies, or the field of force, which is associated with them. For the life of {38} the body is more than the sum of the properties of the cells of which it is composed: as Goethe said, “Das Lebendige ist zwar in Elemente zerlegt, aber man kann es aus diesen nicht wieder zusammenstellen und beleben.”

Among certain lower and microscopic organisms, such for instance as the Rotifera, we are still more palpably struck by the small number of cells which go to constitute a usually complex organ, such as kidney, stomach, ovary, etc. We can sometimes number them in a few units, in place of the thousands that make up such an organ in larger, if not always higher, animals. These facts constitute one among many arguments which combine to teach us that, however important and advantageous the subdivision of organisms into cells may be from the constructional, or from the dynamical point of view, the phenomenon has less essential importance in theoretical biology than was once, and is often still, assigned to it.

Again, just as Sachs shewed that there was a limit to the amount of cytoplasm which could gather round a single nucleus, so Boveri has demonstrated that the nucleus itself has definite limitations of size, and that, in cell-division after fertilisation, each new nucleus has the same size as its parent-nucleus64.

In all these cases, then, there are reasons, partly no doubt physiological, but in very large part purely physical, which set limits to the normal magnitude of the organism or of the cell. But as we have already discussed the existence of absolute and definite limitations, of a physical kind, to the possible increase in magnitude of an organism, let us now enquire whether there be not also a lower limit, below which the very existence of an organism is impossible, or at least where, under changed conditions, its very nature must be profoundly modified.

Among the smallest of known organisms we have, for instance, Micromonas mesnili, Bonel, a flagellate infusorian, which measures about ·34 µ, or ·00034 mm., by ·00025 mm.; smaller even than this we have a pathogenic micrococcus of the rabbit, M. progrediens, Schröter, the diameter of which is said to be only ·00015 mm. or ·15 µ, or 1·5 × 10⁻⁵ cm.,—about equal to the thickness of {39} the thinnest gold-leaf; and as small if not smaller still are a few bacteria and their spores. But here we have reached, or all but reached the utmost limits of ordinary microscopic vision; and there remain still smaller organisms, the so-called “filter-passers,” which the ultra-microscope reveals, but which are mainly brought within our ken only by the maladies, such as hydrophobia, foot-and-mouth disease, or the “mosaic” disease of the tobacco-plant, to which these invisible micro-organisms give rise65. Accordingly, since it is only by the diseases which they occasion that these tiny bodies are made known to us, we might be tempted to suppose that innumerable other invisible organisms, smaller and yet smaller, exist unseen and unrecognised by man.

Fig. 2. Relative magnitudes of: A, human blood-corpuscle (7·5 µ in diameter); B, Bacillus anthracis (4 – 15 µ × 1 µ); C, various Micrococci (diam. 0·5 – 1 µ, rarely 2 µ); D, Micromonas progrediens, Schröter (diam. 0·15 µ).

To illustrate some of these small magnitudes I have adapted the preceding diagram from one given by Zsigmondy66. Upon the {40} same scale the minute ultramicroscopic particles of colloid gold would be represented by the finest dots which we could make visible to the naked eye upon the paper.

A bacillus of ordinary, typical size is, say, 1 µ in length. The length (or height) of a man is about a million and three-quarter times as great, i.e. 1·75 metres, or 1·75 × 10⁶ µ; and the mass of the man is in the neighbourhood of five million, million, million (5 × 10¹⁸) times greater than that of the bacillus. If we ask whether there may not exist organisms as much less than the bacillus as the bacillus is less than the dimensions of a man, it is very easy to see that this is quite impossible, for we are rapidly approaching a point where the question of molecular dimensions, and of the ultimate divisibility of matter, begins to call for our attention, and to obtrude itself as a crucial factor in the case.

According to this estimate, then, our little Micrococcus progrediens should contain only about 10,000 atoms of sulphur, an element indispensable to its protoplasmic constitution; and it follows that an organism of one-tenth the diameter of our micrococcus would only contain 10 sulphur-atoms, and therefore only ten chemical “molecules” or units of protoplasm!

According to the most recent estimates, the weight of the hydrogen molecule is somewhat less than that on which Errera based his calculations, namely about 16 × 10⁻²² mgms. and according to this value, our micrococcus would contain just about 27,000 albumin molecules. In other words, whichever determination we accept, we see that an organism one-tenth as large as our micrococcus, in linear dimensions, would only contain some thirty molecules of albumin; or, in other words, our micrococcus is only about thirty times as large, in linear dimensions, as a single albumin molecule71.

We must doubtless make large allowances for uncertainty in the assumptions and estimates upon which these calculations are based; and we must also remember that the data with which the physicist provides us in regard to molecular magnitudes are, to a very great extent, maximal values, above which the molecular magnitude (or rather the sphere of the molecule’s range of motion) is not likely to lie: but below which there is a greater element of uncertainty as to its possibly greater minuteness. But nevertheless, when we shall have made all reasonable allowances for uncertainty upon the physical side, it will still be clear that the smallest known bodies which are described as organisms draw nigh towards molecular magnitudes, and we must recognise that the subdivision of the organism cannot proceed to an indefinite extent, and in all probability cannot go very much further than it appears to have done in these already discovered forms. For, even, after giving all due regard to the complexity of our unit (that is to say the albumin-molecule), with all the increased possibilities of interrelation with its neighbours which this complexity implies, we cannot but see that physiologically, and comparatively speaking, we have come down to a very simple thing.

While such con­si­de­ra­tions as these, based on the chemical composition of the organism, teach us that there must be a definite lower limit to its magnitude, other con­si­de­ra­tions of a purely physical kind lead us to the same conclusion. For our discussion of the principle of similitude has already taught us that, long before we reach these almost infinitesimal magnitudes, the {43} diminishing organism will have greatly changed in all its physical relations, and must at length arrive under conditions which must surely be incompatible with anything such as we understand by life, at least in its full and ordinary development and manifestation.

We are told, for instance, that the powerful force of surface-tension, or capillarity, begins to act within a range of about 1 ⁄ 500,000 of an inch, or say 0·05 µ. A soap-film, or a film of oil upon water, may be attenuated to far less magnitudes than this; the black spots upon a soap-bubble are known, by various concordant methods of measurement, to be only about 6 × 10⁻⁷ cm., or about ·006 µ thick, and Lord Rayleigh and M. Devaux72 have obtained films of oil of ·002 µ, or even ·001 µ in thickness.

But while it is possible for a fluid film to exist in these almost molecular dimensions, it is certain that, long before we reach them, there must arise new conditions of which we have little knowledge and which it is not easy even to imagine.

It would seem that, in an organism of ·1 µ in diameter, or even rather more, there can be no essential distinction between the interior and the surface layers. No hollow vesicle, I take it, can exist of these dimensions, or at least, if it be possible for it to do so, the contained gas or fluid must be under pressures of a formidable kind73, and of which we have no knowledge or experience. Nor, I imagine, can there be any real complexity, or heterogeneity, of its fluid or semi-fluid contents; there can be no vacuoles within such a cell, nor any layers defined within its fluid substance, for something of the nature of a boundary-film is the necessary condition of the existence of such layers. Moreover, the whole organism, provided that it be fluid or semi-fluid, can only be spherical in form. What, then, can we attribute, in the way of properties, to an organism of a size as small as, or smaller than, say ·05 µ? It must, in all probability, be a homogeneous, structureless sphere, composed of a very small number of albuminoid or other molecules. Its vital properties and functions must be extraordinarily limited; its specific outward characters, even if we could see it, must be nil; and its specific properties must be little more than those of an ion-laden corpuscle, enabling it to perform {44} this or that chemical reaction, or to produce this or that pathogenic effect. Even among inorganic, non-living bodies, there must be a certain grade of minuteness at which the ordinary properties become modified. For instance, while under ordinary circumstances cry­stal­li­sa­tion starts in a solution about a minute solid fragment or crystal of the salt, Ostwald has shewn that we may have particles so minute that they fail to serve as a nucleus for cry­stal­li­sa­tion,—which is as much as to say that they are too minute to have the form and properties of a “crystal”; and again, in his thin oil-films, Lord Rayleigh has noted the striking change of physical properties which ensues when the film becomes attenuated to something less than one close-packed layer of molecules74.

Thus, as Clerk Maxwell put it, “molecular science sets us face to face with physiological theories. It forbids the physiologist from imagining that structural details of infinitely small dimensions [such as Leibniz assumed, one within another, ad infinitum] can furnish an explanation of the infinite variety which exists in the properties and functions of the most minute organisms.” And for this reason he reprobates, with not undue severity, those advocates of pangenesis and similar theories of heredity, who would place “a whole world of wonders within a body so small and so devoid of visible structure as a germ.” But indeed it scarcely needed Maxwell’s criticism to shew forth the immense physical difficulties of Darwin’s theory of Pangenesis: which, after all, is as old as Democritus, and is no other than that Promethean particulam undique desectam of which we have read, and at which we have smiled, in our Horace.

There are many other ways in which, when we “make a long excursion into space,” we find our ordinary rules of physical behaviour entirely upset. A very familiar case, analysed by Stokes, is that the viscosity of the surrounding medium has a relatively powerful effect upon bodies below a certain size. A droplet of water, a thousandth of an inch (25 µ) in diameter, cannot fall in still air quicker than about an inch and a half per second; and as its size decreases, its resistance varies as the diameter, and not (as with larger bodies) as the surface of the {45} drop. Thus a drop one-tenth of that size (2·5 µ), the size, apparently, of the drops of water in a light cloud, will fall a hundred times slower, or say an inch a minute; and one again a tenth of this diameter (say ·25 µ, or about twice as big, in linear dimensions, as our micrococcus), will scarcely fall an inch in two hours. By reason of this principle, not only do the smaller bacteria fall very slowly through the air, but all minute bodies meet with great proportionate resistance to their movements in a fluid. Even such comparatively large organisms as the diatoms and the foraminifera, laden though they are with a heavy shell of flint or lime, seem to be poised in the water of the ocean, and fall in it with exceeding slowness.

The Brownian movement has also to be reckoned with,—that remarkable phenomenon studied nearly a century ago (1827) by Robert Brown, facile princeps botanicorum. It is one more of those fundamental physical phenomena which the biologists have contributed, or helped to contribute, to the science of physics.

The quivering motion, accompanied by rotation, and even by translation, manifested by the fine granular particles issuing from a crushed pollen-grain, and which Robert Brown proved to have no vital significance but to be manifested also by all minute particles whatsoever, organic and inorganic, was for many years unexplained. Nearly fifty years after Brown wrote, it was said to be “due, either directly to some calorical changes continually taking place in the fluid, or to some obscure chemical action between the solid particles and the fluid which is indirectly promoted by heat75.” Very shortly after these last words were written, it was ascribed by Wiener to molecular action, and we now know that it is indeed due to the impact or bombardment of molecules upon a body so small that these impacts do not for the moment, as it were, “average out” to ap­prox­i­mate equality on all sides. The movement becomes manifest with particles of somewhere about 20 µ in diameter, it is admirably displayed by particles of about 12 µ in diameter, and becomes more marked the smaller the particles are. The bombardment causes our particles to behave just like molecules of uncommon size, and this {46} behaviour is manifested in several ways76. Firstly, we have the quivering movement of the particles; secondly, their movement backwards and forwards, in short, straight, disjointed paths; thirdly, the particles rotate, and do so the more rapidly the smaller they are, and by theory, confirmed by observation, it is found that particles of 1 µ in diameter rotate on an average through 100° per second, while particles of 13 µ in diameter turn through only 14° per minute. Lastly, the very curious result appears, that in a layer of fluid the particles are not equally distributed, nor do they all ever fall, under the influence of gravity, to the bottom. But just as the molecules of the atmosphere are so distributed, under the influence of gravity, that the density (and therefore the number of molecules per unit volume) falls off in geometrical progression as we ascend to higher and higher layers, so is it with our particles, even within the narrow limits of the little portion of fluid under our microscope. It is only in regard to particles of the simplest form that these phenomena have been theoretically investigated77, and we may take it as certain that more complex particles, such as the twisted body of a Spirillum, would show other and still more complicated manifestations. It is at least clear that, just as the early microscopists in the days before Robert Brown never doubted but that these phenomena were purely vital, so we also may still be apt to confuse, in certain cases, the one phenomenon with the other. We cannot, indeed, without the most careful scrutiny, decide whether the movements of our minutest organisms are intrinsically “vital” (in the sense of being beyond a physical mechanism, or working model) or not. For example, Schaudinn has suggested that the undulating movements of Spirochaete pallida must be due to the presence of a minute, unseen, “undulating membrane”; and Doflein says of the same species that “sie verharrt oft mit eigenthümlich zitternden Bewegungen zu einem Orte.” Both movements, the trembling or quivering {47} movement described by Doflein, and the undulating or rotating movement described by Schaudinn, are just such as may be easily and naturally interpreted as part and parcel of the Brownian phenomenon.

While the Brownian movement may thus simulate in a deceptive way the active movements of an organism, the reverse statement also to a certain extent holds good. One sometimes lies awake of a summer’s morning watching the flies as they dance under the ceiling. It is a very remarkable dance. The dancers do not whirl or gyrate, either in company or alone; but they advance and retire; they seem to jostle and rebound; between the rebounds they dart hither or thither in short straight snatches of hurried flight; and turn again sharply in a new rebound at the end of each little rush. Their motions are wholly “erratic,” independent of one another, and devoid of common purpose. This is nothing else than a vastly magnified picture, or simulacrum, of the Brownian movement; the parallel between the two cases lies in their complete irregularity, but this in itself implies a close resemblance. One might see the same thing in a crowded market-place, always provided that the bustling crowd had no business whatsoever. In like manner Lucretius, and Epicurus before him, watched the dust-motes quivering in the beam, and saw in them a mimic representation, rei simulacrum et imago, of the eternal motions of the atoms. Again the same phenomenon may be witnessed under the microscope, in a drop of water swarming with Paramoecia or suchlike Infusoria; and here the analogy has been put to a numerical test. Following with a pencil the track of each little swimmer, and dotting its place every few seconds (to the beat of a metronome), Karl Przibram found that the mean successive distances from a common base-line obeyed with great exactitude the “Einstein formula,” that is to say the particular form of the “law of chance” which is applicable to the case of the Brownian movement78. The phenomenon is (of course) merely analogous, and by no means identical with the Brownian movement; for the range of motion of the little active organisms, whether they be gnats or infusoria, is vastly greater than that of the minute particles which are {48} passive under bombardment; but nevertheless Przibram is inclined to think that even his comparatively large infusoria are small enough for the molecular bombardment to be a stimulus, though not the actual cause, of their irregular and interrupted movements.

There is yet another very remarkable phenomenon which may come into play in the case of the minutest of organisms; and this is their relation to the rays of light, as Arrhenius has told us. On the waves of a beam of light, a very minute particle (in vacuo) should be actually caught up, and carried along with an immense velocity; and this “radiant pressure” exercises its most powerful influence on bodies which (if they be of spherical form) are just about ·00016 mm., or ·16 µ in diameter. This is just about the size, as we have seen, of some of our smallest known protozoa and bacteria, while we have some reason to believe that others yet unseen, and perhaps the spores of many, are smaller still. Now we have seen that such minute particles fall with extreme slowness in air, even at ordinary atmospheric pressures: our organism measuring ·16 µ would fall but 83 metres in a year, which is as much as to say that its weight offers practically no impediment to its transference, by the slightest current, to the very highest regions of the atmosphere. Beyond the atmosphere, however, it cannot go, until some new force enable it to resist the attraction of terrestrial gravity, which the viscosity of an atmosphere is no longer at hand to oppose. But it is conceivable that our particle may go yet farther, and actually break loose from the bonds of earth. For in the upper regions of the atmosphere, say fifty miles high, it will come in contact with the rays and flashes of the Northern Lights, which consist (as Arrhenius maintains) of a fine dust, or cloud of vapour-drops, laden with a charge of negative electricity, and projected outwards from the sun. As soon as our particle acquires a charge of negative electricity it will begin to be repelled by the similarly laden auroral particles, and the amount of charge necessary to enable a particle of given size (such as our little monad of ·16 µ) to resist the attraction of gravity may be calculated, and is found to be such as the actual conditions can easily supply. Finally, when once set free from the entanglement of the earth’s {49} atmosphere, the particle may be propelled by the “radiant pressure” of light, with a velocity which will carry it.—like Uriel gliding on a sunbeam,—as far as the orbit of Mars in twenty days, of Jupiter in eighty days, and as far as the nearest fixed star in three thousand years! This, and much more, is Arrhenius’s contribution towards the acceptance of Lord Kelvin’s hypothesis that life may be, and may have been, disseminated across the bounds of space, throughout the solar system and the whole universe!

It may well be that we need attach no great practical importance to this bold conception; for even though stellar space be shewn to be mare liberum to minute material travellers, we may be sure that those which reach a stellar or even a planetary bourne are infinitely, or all but infinitely, few. But whether or no, the remote possibilities of the case serve to illustrate in a very vivid way the profound differences of physical property and potentiality which are associated in the scale of magnitude with simple differences of degree.

When we study magnitude by itself, apart, that is to say, from the gradual changes to which it may be subject, we are dealing with a something which may be adequately represented by a number, or by means of a line of definite length; it is what mathematicians call a scalar phenomenon. When we introduce the conception of change of magnitude, of magnitude which varies as we pass from one direction to another in space, or from one instant to another in time, our phenomenon becomes capable of representation by means of a line of which we define both the length and the direction; it is (in this particular aspect) what is called a vector phenomenon.

When we deal with magnitude in relation to the dimensions of space, the vector diagram which we draw plots magnitude in one direction against magnitude in another,—length against height, for instance, or against breadth; and the result is simply what we call a picture or drawing of an object, or (more correctly) a “plane projection” of the object. In other words, what we call Form is a ratio of magnitudes, referred to direction in space.

When in dealing with magnitude we refer its variations to successive intervals of time (or when, as it is said, we equate it with time), we are then dealing with the phenomenon of growth; and it is evident, therefore, that this term growth has wide meanings. For growth may obviously be positive or negative; that is to say, a thing may grow larger or smaller, greater or less; and by extension of the primitive concrete signification of the word, we easily and legitimately apply it to non-material things, such as temperature, and say, for instance, that a body “grows” hot or cold. When in a two-dimensional diagram, we represent a magnitude (for instance length) in relation to time (or “plot” {51} length against time, as the phrase is), we get that kind of vector diagram which is commonly known as a “curve of growth.” We perceive, accordingly, that the phenomenon which we are now studying is a velocity (whose “dimensions” are ^Space⁄_Time or ^L⁄_T); and this phenomenon we shall speak of, simply, as a rate of growth.

In various conventional ways we can convert a two-dimensional into a three-dimensional diagram. We do so, for example, by means of the geometrical method of “perspective” when we represent upon a sheet of paper the length, breadth and depth of an object in three-dimensional space; but we do it more simply, as a rule, by means of “contour-lines,” and always when time is one of the dimensions to be represented. If we superimpose upon one another (or even set side by side) pictures, or plane projections, of an organism, drawn at successive intervals of time, we have such a three-dimensional diagram, which is a partial representation (limited to two dimensions of space) of the organism’s gradual change of form, or course of development; and in such a case our contour-lines may, for the purposes of the embryologist, be separated by intervals representing a few hours or days, or, for the purposes of the palaeontologist, by interspaces of unnumbered and innumerable years79.

Such a diagram represents in two of its three dimensions form, and in two, or three, of its dimensions growth; and so we see how intimately the two conceptions are correlated or inter-related to one another. In short, it is obvious that the form of an animal is determined by its specific rate of growth in various directions; accordingly, the phenomenon of rate of growth deserves to be studied as a necessary preliminary to the theoretical study of form, and, math­e­mat­i­cally speaking, organic form itself appears to us as a function of time80. {52}

At the same time, we need only consider this part of our subject somewhat briefly. Though it has an essential bearing on the problems of morphology, it is in greater degree involved with physiological problems; and furthermore, the statistical or numerical aspect of the question is peculiarly adapted for the math­e­mat­i­cal study of variation and correlation. On these important subjects we shall scarcely touch; for our main purpose will be sufficiently served if we consider the char­ac­teris­tics of a rate of growth in a few illustrative cases, and recognise that this rate of growth is a very important specific property, with its own char­ac­ter­is­tic value in this organism or that, in this or that part of each organism, and in this or that phase of its existence.

The statement which we have just made that “the form of an organism is determined by its rate of growth in various directions,” is one which calls (as we have partly seen in the foregoing chapter) for further explanation and for some measure of qualification. Among organic forms we shall have frequent occasion to see that form is in many cases due to the immediate or direct action of certain molecular forces, of which surface-tension is that which plays the greatest part. Now when surface-tension (for instance) causes a minute semi-fluid organism to assume a spherical form, or gives the form of a catenary or an elastic curve to a film of protoplasm in contact with some solid skeletal rod, or when it acts in various other ways which are productive of definite contours, this is a process of conformation that, both in appearance and reality, is very different from the process by which an ordinary plant or animal grows into its specific form. In both cases, change of form is brought about by the movement of portions of matter, and in both cases it is ultimately due to the action of molecular forces; but in the one case the movements of the particles of matter lie for the most part within molecular range, while in the other we have to deal chiefly with the transference of portions of matter into the system from without, and from one widely distant part of the organism to another. It is to this latter class of phenomena that we usually restrict the term growth; and it is in regard to them that we are in a position to study the rate of action in different directions, and to see that it is merely on a difference of velocities that the modification of form essentially depends. {53} The difference between the two classes of phenomena is somewhat akin to the difference between the forces which determine the form of a rain-drop and those which, by the flowing of the waters and the sculpturing of the solid earth, have brought about the complex configuration of a river; molecular forces are paramount in the conformation of the one, and molar forces are dominant in the other.

At the same time it is perfectly true that all changes of form, inasmuch as they necessarily involve changes of actual and relative magnitude, may, in a sense, be properly looked upon as phenomena of growth; and it is also true, since the movement of matter must always involve an element of time81, that in all cases the rate of growth is a phenomenon to be considered. Even though the molecular forces which play their part in modifying the form of an organism exert an action which is, theoretically, all but instantaneous, that action is apt to be dragged out to an appreciable interval of time by reason of viscosity or some other form of resistance in the material. From the physical or physiological point of view the rate of action even in such cases may be well worth studying; for example, a study of the rate of cell-division in a segmenting egg may teach us something about the work done, and about the various energies concerned. But in such cases the action is, as a rule, so homogeneous, and the form finally attained is so definite and so little dependent on the time taken to effect it, that the specific rate of change, or rate of growth, does not enter into the morphological problem.

To sum up, we may lay down the following general statements. The form of organisms is a phenomenon to be referred in part to the direct action of molecular forces, in part to a more complex and slower process, indirectly resulting from chemical, osmotic and other forces, by which material is introduced into the organism and transferred from one part of it to another. It is this latter complex phenomenon which we usually speak of as “growth.” {54}

Every growing organism, and every part of such a growing organism, has its own specific rate of growth, referred to a particular direction. It is the ratio between the rates of growth in various directions by which we must account for the external forms of all, save certain very minute, organisms. This ratio between rates of growth in various directions may sometimes be of a simple kind, as when it results in the math­e­mat­i­cally definable outline of a shell, or in the smooth curve of the margin of a leaf. It may sometimes be a very constant one, in which case the organism, while growing in bulk, suffers little or no perceptible change in form; but such equi­lib­rium seldom endures for more than a season, and when the ratio tends to alter, then we have the phenomenon of morphological “development,” or steady and persistent change of form.

This elementary concept of Form, as determined by varying rates of Growth, was clearly apprehended by the math­e­mat­i­cal mind of Haller,—who had learned his mathematics of the great John Bernoulli, as the latter in turn had learned his physiology from the writings of Borelli. Indeed it was this very point, the apparently unlimited extent to which, in the development of the chick, inequalities of growth could and did produce changes of form and changes of anatomical “structure,” that led Haller to surmise that the process was actually without limits, and that all development was but an unfolding, or “evolutio,” in which no part came into being which had not essentially existed before82. In short the celebrated doctrine of “preformation” implied on the one hand a clear recognition of what, throughout the later stages of development, growth can do, by hastening the increase in size of one part, hindering that of another, changing their relative magnitudes and positions, and altering their forms; while on the other hand it betrayed a failure (inevitable in those days) to recognise the essential difference between these movements of masses and the molecular processes which precede and accompany {55} them, and which are char­ac­ter­is­tic of another order of magnitude.

By other writers besides Haller the very general, though not strictly universal connection between form and rate of growth has been clearly recognised. Such a connection is implicit in those “proportional diagrams” by which Dürer and some of his brother artists were wont to illustrate the successive changes of form, or of relative dimensions, which attend the growth of the child, to boyhood and to manhood. The same connection was recognised, more explicitly, by some of the older embryologists, for instance by Pander83, and appears, as a survival of the doctrine of preformation, in his study of the development of the chick. And long afterwards, the embryological aspect of the case was emphasised by His, who pointed out, for instance, that the various foldings of the blastoderm, by which the neural and amniotic folds were brought into being, were essentially and obviously the resultant of unequal rates of growth,—of local accelerations or retardations of growth,—in what to begin with was an even and uniform layer of embryonic tissue. If we imagine a flat sheet of paper, parts of which are caused (as by moisture or evaporation) to expand or to contract, the plane surface is at once dimpled, or “buckled,” or folded, by the resultant forces of expansion or contraction: and the various distortions to which the plane surface of the “germinal disc” is subject, as His shewed once and for all, are precisely analogous. An experimental demonstration still more closely comparable to the actual case of the blastoderm, is obtained by making an “artificial blastoderm,” of little pills or pellets of dough, which are caused to grow, with varying velocities, by the addition of varying quantities of yeast. Here, as Roux is careful to point out84, we observe that it is not only the growth of the individual cells, but the traction exercised through their mutual interconnections, which brings about the foldings and other distortions of the entire structure. {56}

But this again was clearly present to Haller’s mind, and formed an essential part of his embryological doctrine. For he has no sooner treated of incrementum, or celeritas incrementi, than he proceeds to deal with the contributory and complementary phenomena of expansion, traction (adtractio)85, and pressure, and the more subtle influences which he denominates vis derivationis et revulsionis86: these latter being the secondary and correlated effects on growth in one part, brought about, through such changes as are produced (for instance) in the circulation, by the growth of another.

Let us admit that, on the physiological side, Haller’s or His’s methods of explanation carry us back but a little way; yet even this little way is something gained. Nevertheless, I can well remember the harsh criticism, and even contempt, which His’s doctrine met with, not merely on the ground that it was inadequate, but because such an explanation was deemed wholly inappropriate, and was utterly disavowed87. Hertwig, for instance, asserted that, in embryology, when we found one embryonic stage preceding another, the existence of the former was, for the embryologist, an all-sufficient “causal explanation” of the latter. “We consider (he says), that we are studying and explaining a causal relation when we have demonstrated that the gastrula arises by invagination of a blastosphere, or the neural canal by the infolding of a cell plate so as to constitute a tube88.” For Hertwig, therefore, as {57} Roux remarks, the task of investigating a physical mechanism in embryology,—“der Ziel das Wirken zu erforschen,”—has no existence at all. For Balfour also, as for Hertwig, the mechanical or physical aspect of organic development had little or no attraction. In one notable instance, Balfour himself adduced a physical, or quasi-physical, explanation of an organic process, when he referred the various modes of segmentation of an ovum, complete or partial, equal or unequal and so forth, to the varying amount or the varying distribution of food yolk in association with the germinal protoplasm of the egg89. But in the main, Balfour, like all the other embryologists of his day, was engrossed by the problems of phylogeny, and he expressly defined the aims of comparative embryology (as exemplified in his own textbook) as being “twofold: (1) to form a basis for Phylogeny. and (2) to form a basis for Organogeny or the origin and evolution of organs90.”

It has been the great service of Roux and his fellow-workers of the school of “Ent­wicke­lungs­me­cha­nik,” and of many other students to whose work we shall refer, to try, as His tried91 to import into embryology, wherever possible, the simpler concepts of physics, to introduce along with them the method of experiment, and to refuse to be bound by the narrow limitations which such teaching as that of Hertwig would of necessity impose on the work and the thought and on the whole philosophy of the biologist.

Before we pass from this general discussion to study some of the particular phenomena of growth, let me give a single illustration, from Darwin, of a point of view which is in marked contrast to Haller’s simple but essentially math­e­mat­i­cal conception of Form.

There is a curious passage in the Origin of Species92, where Darwin is discussing the leading facts of embryology, and in particular Von Baer’s “law of embryonic resemblance.” Here Darwin says “We are so much accustomed to see a difference in {58} structure between the embryo and the adult, that we are tempted to look at this difference as in some necessary manner contingent on growth. But there is no reason why, for instance, the wing of a bat, or the fin of a porpoise, should not have been sketched out with all their parts in proper proportion, as soon as any part became visible.” After pointing out with his habitual care various exceptions, Darwin proceeds to lay down two general principles, viz. “that slight variations generally appear at a not very early period of life,” and secondly, that “at whatever age a variation first appears in the parent, it tends to reappear at a cor­re­spon­ding age in the offspring.” He then argues that it is with nature as with the fancier, who does not care what his pigeons look like in the embryo, so long as the full-grown bird possesses the desired qualities; and that the process of selection takes place when the birds or other animals are nearly grown up,—at least on the part of the breeder, and presumably in nature as a general rule. The illustration of these principles is set forth as follows; “Let us take a group of birds, descended from some ancient form and modified through natural selection for different habits. Then, from the many successive variations having supervened in the several species at a not very early age, and having been inherited at a cor­re­spon­ding age, the young will still resemble each other much more closely than do the adults,—just as we have seen with the breeds of the pigeon .... Whatever influence long-continued use or disuse may have had in modifying the limbs or other parts of any species, this will chiefly or solely have affected it when nearly mature, when it was compelled to use its full powers to gain its own living; and the effects thus produced will have been transmitted to the offspring at a cor­re­spon­ding nearly mature age. Thus the young will not be modified, or will be modified only in a slight degree, through the effects of the increased use or disuse of parts.” This whole argument is remarkable, in more ways than we need try to deal with here; but it is especially remarkable that Darwin should begin by casting doubt upon the broad fact that a “difference in structure between the embryo and the adult” is “in some necessary manner contingent on growth”; and that he should see no reason why complicated structures of the adult “should not have been sketched out {59} with all their parts in proper proportion, as soon as any part became visible.” It would seem to me that even the most elementary attention to form in its relation to growth would have removed most of Darwin’s difficulties in regard to the particular phenomena which he is here considering. For these phenomena are phenomena of form, and therefore of relative magnitude; and the magnitudes in question are attained by growth, proceeding with certain specific velocities, and lasting for certain long periods of time. And it is accordingly obvious that in any two related individuals (whether specifically identical or not) the differences between them must manifest themselves gradually, and be but little apparent in the young. It is for the same simple reason that animals which are of very different sizes when adult, differ less and less in size (as well as in form) as we trace them backwards through the foetal stages.

Though we study the visible effects of varying rates of growth throughout wellnigh all the problems of morphology, it is not very often that we can directly measure the velocities concerned. But owing to the obvious underlying importance which the phenomenon has to the morphologist we must make shift to study it where we can, even though our illustrative cases may seem to have little immediate bearing on the morphological problem93.

In a very simple organism, of spherical symmetry, such as the single spherical cell of Protococcus or of Orbulina, growth is reduced to its simplest terms, and indeed it becomes so simple in its outward manifestations that it is no longer of special interest to the morphologist. The rate of growth is measured by the rate of change in length of a radius, i.e. V = (R′ − R) ⁄ T, and from this we may calculate, as already indicated, the rate of growth in terms of surface and of volume. The growing body remains of constant form, owing to the symmetry of the system; because, that is to say, on the one hand the pressure exerted by the growing protoplasm is exerted equally in all directions, after the manner of a hydrostatic pressure, which indeed it actually is: while on the other hand, the “skin” or surface layer of the cell is sufficiently {60} homogeneous to exert at every point an ap­prox­i­mate­ly uniform resistance. Under these conditions then, the rate of growth is uniform in all directions, and does not affect the form of the organism.

But in a larger or a more complex organism the study of growth, and of the rate of growth, presents us with a variety of problems, and the whole phenomenon becomes a factor of great morphological importance. We no longer find that it tends to be uniform in all directions, nor have we any right to expect that it should. The resistances which it meets with will no longer be uniform. In one direction but not in others it will be opposed by the important resistance of gravity; and within the growing system itself all manner of structural differences will come into play, setting up unequal resistances to growth by the varying rigidity or viscosity of the material substance in one direction or another. At the same time, the actual sources of growth, the chemical and osmotic forces which lead to the intussusception of new matter, are not uniformly distributed; one tissue or one organ may well manifest a tendency to increase while another does not; a series of bones, their intervening cartilages, and their surrounding muscles, may all be capable of very different rates of increment. The differences of form which are the resultants of these differences in rate of growth are especially manifested during that part of life when growth itself is rapid: when the organism, as we say, is undergoing its development. When growth in general has become slow, the relative differences in rate between different parts of the organism may still exist, and may be made manifest by careful observation, but in many, or perhaps in most cases, the resultant change of form does not strike the eye. Great as are the differences between the rates of growth in different parts of an organism, the marvel is that the ratios between them are so nicely balanced as they actually are, and so capable, accordingly, of keeping for long periods of time the form of the growing organism all but unchanged. There is the nicest possible balance of forces and resistances in every part of the complex body; and when this normal equi­lib­rium is disturbed, then we get abnormal growth, in the shape of tumours, exostoses, and malformations of every kind. {61}

Fig. 3. Curve of Growth in Man, from birth to 20 yrs ();) from Quetelet’s Belgian data. The upper curve of stature from Bowditch’s Boston data.

If the child be some 20 inches, or say 50 cm. tall at birth, and the man some six feet high, or say 180 cm., at twenty, we may say that his average rate of growth has been (180 − 50) ⁄ 20 cm., or 6·5 centimetres per annum. But we know very well that this is {62} but a very rough preliminary statement, and that the boy grew quickly during some, and slowly during other, of his twenty years. It becomes necessary therefore to study the phenomenon of growth in successive small portions; to study, that is to say, the successive lengths, or the successive small differences, or increments, of length (or of weight, etc.), attained in successive short increments of time. This we do in the first instance in the usual way, by the “graphic method” of plotting length against time, and so constructing our “curve of growth.” Our curve of growth, whether of weight or length (Fig. 3), has always a certain char­ac­ter­is­tic form, or char­ac­ter­is­tic curvature. This is our immediate proof of the fact that the rate of growth changes as time goes on; for had it not been so, had an equal increment of length been added in each equal interval of time, our “curve” would have appeared as a straight line. Such as it is, it tells us not only that the rate of growth tends to alter, but that it alters in a definite and orderly way; for, subject to various minor interruptions, due to secondary causes, our curves of growth are, on the whole, “smooth” curves.

The curve of growth for length or stature in man indicates a rapid increase at the outset, that is to say during the quick growth of babyhood; a long period of slower, but still rapid and almost steady growth in early boyhood; as a rule a marked quickening soon after the boy is in his teens, when he comes to “the growing age”; and finally a gradual arrest of growth as the boy “comes to his full height,” and reaches manhood.

If we carried the curve further, we should see a very curious thing. We should see that a man’s full stature endures but for a spell; long before fifty95 it has begun to abate, by sixty it is notably lessened, in extreme old age the old man’s frame is shrunken and it is but a memory that “he once was tall.” We have already seen, and here we see again, that growth may have a “negative value.” The phenomenon of negative growth in old age extends to weight also, and is evidently largely chemical in origin: the organism can no longer add new material to its fabric fast enough to keep pace with the wastage of time. Our curve {63} of growth is in fact a diagram of activity, or “time-energy” diagram96. As the organism grows it is absorbing energy beyond its daily needs, and accumulating it at a rate depicted in our

curve; but the time comes when it accumulates no longer, and at last it is constrained to draw upon its dwindling store. But in part, the slow decline in stature is an expression of an unequal contest between our bodily powers and the unchanging force of gravity, {64} which draws us down when we would fain rise up97. For against gravity we fight all our days, in every movement of our limbs, in every beat of our hearts; it is the indomitable force that defeats us in the end, that lays us on our deathbed, that lowers us to the grave98.

Side by side with the curve which represents growth in length, or stature, our diagram shows the curve of weight99. That this curve is of a very different shape from the former one, is accounted for in the main (though not wholly) by the fact which we have already dealt with, that, whatever be the law of increment in a linear dimension, the law of increase in volume, and therefore in weight, will be that these latter magnitudes tend to vary as the cubes of the linear dimensions. This however does not account for the change of direction, or “point of inflection” which we observe in the curve of weight at about one or two years old, nor for certain other differences between our two curves which the scale of our diagram does not yet make clear. These differences are due to the fact that the form of the child is altering with growth, that other linear dimensions are varying somewhat differently from length or stature, and that consequently the growth in bulk or weight is following a more complicated law.

Our curve of growth, whether for weight or length, is a direct picture of velocity, for it represents, as a connected series, the successive epochs of time at which successive weights or lengths are attained. But, as we have already in part seen, a great part of the interest of our curve lies in the fact that we can see from it, not only that length (or some other magnitude) is changing, but that the rate of change of magnitude, or rate of growth, is itself changing. We have, in short, to study the phenomenon of acceleration: we have begun by studying a velocity, or rate of {65} change of magnitude; we must now study an acceleration, or rate of change of velocity. The rate, or velocity, of growth is measured by the slope of the curve; where the curve is steep, it means that growth is rapid, and when growth ceases the curve appears as a horizontal line. If we can find a means, then, of representing at successive epochs the cor­re­spon­ding slope, or steepness, of the curve, we shall have obtained a picture of the rate of change of velocity, or the acceleration of growth. The measure of the steepness of a curve is given by the tangent to the curve, or we may estimate it by taking for equal intervals of time (strictly speaking, for each infinitesimal interval of time) the actual increment added during that interval of time: and in practice this simply amounts to taking the successive differences between the values of length (or of weight) for the successive ages which we have begun by studying. If we then plot these successive differences against time, we obtain a curve each point upon which represents a velocity, and the whole curve indicates the rate of change of velocity, and we call it an acceleration-curve. It contains, in truth, nothing whatsoever that was not implicit in our former curve; but it makes clear to our eye, and brings within the reach of further in­ves­ti­ga­tion, phenomena that were hard to see in the other mode of representation.

The acceleration-curve of height, which we here illustrate, in Fig. 4, is very different in form from the curve of growth which we have just been looking at; and it happens that, in this case, there is a very marked difference between the curve which we obtain from Quetelet’s data of growth in height and that which we may draw from any other series of observations known to me from British, French, American or German writers. It begins (as will be seen from our next table) at a very high level, such as it never afterwards attains; and still stands too high, during the first three or four years of life, to be represented on the scale of the accompanying diagram. From these high velocities it falls away, on the whole, until the age when growth itself ceases, and when the rate of growth, accordingly, has, for some years together, the constant value of nil; but the rate of fall, or rate of change of velocity, is subject to several changes or interruptions. During the first three or four years of life the fall is continuous and rapid, {66} but it is somewhat arrested for a while in childhood, from about five years old to eight. According to Quetelet’s data, there is another slight interruption in the falling rate between the ages of about fourteen and sixteen; but in place of this almost insignificant interruption, the English and other statistics indicate a sudden

Fig. 4. Mean annual increments of stature (), Belgian and American.

and very marked acceleration of growth beginning at about twelve years of age, and lasting for three or four years; when this period of acceleration is over, the rate begins to fall again, and does so with great rapidity. We do not know how far the absence of this striking feature in the Belgian curve is due to the imperfections of Quetelet’s data, or whether it is a real and significant feature in the small-statured race which he investigated.

Even apart from these data of Quetelet’s (which seem to constitute an extreme case), it is evident that there are very {68} marked differences between different races, as we shall presently see there are between the two sexes, in regard to the epochs of acceleration of growth, in other words, in the “phase” of the curve.

It is evident that, if we pleased, we might represent the rate of change of acceleration on yet another curve, by constructing a table of “second differences”; this would bring out certain very interesting phenomena, which here however we must not stay to discuss.

The acceleration-curve for man’s weight (Fig. 5), whether we draw it from Quetelet’s data, or from the British, American and other statistics of later writers, is on the whole similar to that which we deduce from the statistics of these latter writers in regard to height or stature; that is to say, it is not a curve which continually descends, but it indicates a rate of growth which is subject to important fluctuations at certain epochs of life. We see that it begins at a high level, and falls continuously and rapidly100 {69} during the first two or three years of life. After a slight recovery, it runs nearly level during boyhood from about five to twelve years old; it then rapidly rises, in the “growing period” of the early teens, and slowly and steadily falls from about the age of sixteen onwards. It does not reach the base-line till the man is about seven or eight and twenty, for normal increase of weight continues during the years when the man is “filling out,” long after growth in height has ceased; but at last, somewhere about thirty, the velocity reaches zero, and even falls below it, for then the man usually begins to lose weight a little. The subsequent slow changes in this acceleration-curve we need not stop to deal with.

Fig. 5. Mean annual increments of weight, in man and woman; from Quetelet’s data.

In the same diagram (Fig. 5) I have set forth the acceleration-curves in respect of increment of weight for both man and woman, according to Quetelet. That growth in boyhood and growth in girlhood follow a very different course is a matter of common knowledge; but if we simply plot the ordinary curve of growth, or velocity-curve, the difference, on the small scale of our diagrams, {70} is not very apparent. It is admirably brought out, however, in the acceleration-curves. Here we see that, after infancy, say from three years old to eight, the velocity in the girl is steady, just as in the boy, but it stands on a lower level in her case than in his: the little maid at this age is growing slower than the boy. But very soon, and while his acceleration-curve is still represented by a straight line, hers has begun to ascend, and until the girl is about thirteen or fourteen it continues to ascend rapidly. After that age, as after sixteen or seventeen in the boy’s case, it begins to descend. In short, throughout all this period, it is a very similar curve in the two sexes; but it has its notable differences, in amplitude and especially in phase. Last of all, we may notice that while the acceleration-curve falls to a negative value in the male about or even a little before the age of thirty years, this does not happen among women. They continue to grow in weight, though slowly, till very much later in life; until there comes a final period, in both sexes alike, during which weight, and height and strength all alike diminish.

The result of these differences (which are essentially phase-differences) between the two sexes in regard to the velocity of growth and to the rate of change of that velocity, is to cause the ratio between the weights of the two sexes to fluctuate in a somewhat complicated manner. At birth the baby-girl weighs on the average nearly 10 per cent. less than the boy. Till about two years old she tends to gain upon him, but she then loses again until the age of about five; from five she gains for a few years somewhat rapidly, and the girl of ten to twelve is only some 3 per cent. less in weight than the boy. The boy in his teens gains {71} steadily, and the young woman of twenty is nearly 15 per cent. lighter than the man. This ratio of difference again slowly diminishes, and between fifty and sixty stands at about 12 per cent., or not far from the mean for all ages; but once more as old age advances, the difference tends, though very slowly, to increase (Fig. 6).

Fig. 6. Percentage ratio, throughout life, of female weight to male; from Quetelet’s data.

While careful observations on the rate of growth in other animals are somewhat scanty, they tend to show so far as they go that the general features of the phenomenon are always much the same. Whether the animal be long-lived, as man or the elephant, or short-lived, like horse or dog, it passes through the same phases of growth101. In all cases growth begins slowly; it attains a maximum velocity early in its course, and afterwards slows down (subject to temporary accelerations) towards a point where growth ceases altogether. But especially in the cold-blooded animals, such as fishes, the slowing-down period is very greatly protracted, and the size of the creature would seem never actually to reach, but only to approach asymptotically, to a maximal limit.

The size ultimately attained is a resultant of the rate, and of {72} the duration, of growth. It is in the main true, as Minot has said, that the rabbit is bigger than the guinea-pig because he grows the faster; but that man is bigger than the rabbit because he goes on growing for a longer time.

In ordinary physical investigations dealing with velocities, as for instance with the course of a projectile, we pass at once from the study of acceleration to that of momentum and so to that of force; for change of momentum, which is proportional to force, is the product of the mass of a body into its acceleration or change of velocity. But we can take no such easy road of kinematical in­ves­ti­ga­tion in this case. The “velocity” of growth is a very different thing from the “velocity” of the projectile. The forces at work in our case are not susceptible of direct and easy treatment; they are too varied in their nature and too indirect in their action for us to be justified in equating them directly with the mass of the growing structure.

In the acceleration-curves which we have shown above (Figs. 2, 3), it will be seen that the curve starts at a considerable interval from the actual date of birth; for the first two increments which we can as yet compare with one another are those attained during the first and second complete years of life. Now we can in many cases “interpolate” with safety between known points upon a curve, but it is very much less safe, and is not very often justifiable (at least until we understand the physical principle involved, and its math­e­mat­i­cal expression), to “extrapolate” beyond the limits of our observations. In short, we do not yet know whether our curve continued to ascend as we go backwards to the date of birth, or whether it may not have changed its direction, and descended, perhaps, to zero-value. In regard to length, or stature, however, we can obtain the requisite information from certain tables of Rüssow’s103, who gives the stature of the infant month by month during the first year of its life, as follows:

If we multiply these monthly differences, or mean monthly velocities, by 12, to bring them into a form comparable with the {74} annual velocities already represented on our acceleration-curves, we shall see that the one series of observations joins on very well with the other; and in short we see at once that our acceleration-curve rises steadily and rapidly as we pass back towards the date of birth.

Fig. 7. Curve of growth (in length or stature) of child, before and after birth. (From His and Rüssow’s data.)

But birth itself, in the case of a viviparous animal, is but an unimportant epoch in the history of growth. It is an epoch whose relative date varies according to the particular animal: the foal and the lamb are born relatively later, that is to say when development has advanced much farther, than in the case of man; the kitten and the puppy are born earlier and therefore more helpless than we are; and the mouse comes into the world still earlier and more inchoate, so much so that even the little marsupial is scarcely more unformed and embryonic. In all these cases alike, we must, in order to study the curve of growth in its entirety, take full account of prenatal or intra-uterine growth. {75}

According to His104, the following are the mean lengths of the unborn human embryo, from month to month.

Fig. 8. Mean monthly increments of length or stature of child (in cms.).

These data link on very well to those of Rüssow, which we have just considered, and (though His’s measurements for the pre-natal months are more detailed than are those of Rüssow for the first year of post-natal life) we may draw a continuous curve of growth (Fig. 7) and curve of acceleration of growth (Fig. 8) for the combined periods. It will at once be seen that there is a “point of inflection” somewhere about the fifth month of intra-uterine life105: up to that date growth proceeds with a continually increasing {76} velocity; but after that date, though growth is still rapid, its velocity tends to fall away. There is a slight break between our two separate sets of statistics at the date of birth, while this is the very epoch regarding which we should particularly like to have precise and continuous information. Undoubtedly there is a certain slight arrest of growth, or diminution of the rate of growth, about the epoch of birth: the sudden change in the {77} method of nutrition has its inevitable effect; but this slight temporary set-back is immediately followed by a secondary, and temporary, acceleration.

Fig. 9. Curve of pre-natal growth (length or stature) of child; and cor­re­spon­ding curve of mean monthly increments (mm.).

It is worth our while to draw a separate curve to illustrate on a larger scale His’s careful data for the ten months of pre-natal life (Fig. 9). We see that this curve of growth is a beautifully regular one, and is nearly symmetrical on either side of that point of inflection of which we have already spoken; it is a curve for which we might well hope to find a simple math­e­mat­i­cal expression. The acceleration-curve shown in Fig. 9 together with the pre-natal curve of growth, is not taken directly from His’s recorded data, but is derived from the tangents drawn to a smoothed curve, cor­re­spon­ding as nearly as possible to the actual curve of growth: the rise to a maximal velocity about the fifth month and the subsequent gradual fall are now demonstrated even more clearly than before. In Fig. 10, which is a curve of growth of the bamboo106, we see (so far as it goes) the same essential features, {78} the slow beginning, the rapid increase of velocity, the point of inflection, and the subsequent slow negative acceleration107.

Fig. 10. Curve of growth of bamboo (from Ostwald, after Kraus).

The magnitudes and velocities which we are here dealing with are, of course, mean values derived from a certain number, sometimes a large number, of individual cases. But no statistical account of mean values is complete unless we also take account of the amount of variability among the individual cases from which the mean value is drawn. To do this throughout would lead us into detailed investigations which lie far beyond the scope of this elementary book; but we may very briefly illustrate the nature of the process, in connection with the phenomena of growth which we have just been studying.

It was in connection with these phenomena, in the case of man, that Quetelet first conceived the statistical study of variation, on lines which were afterwards expounded and developed by Galton, and which have grown, in the hands of Karl Pearson and others, into the modern science of Biometrics.

When Quetelet tells us, for instance, that the mean stature of the ten-year old boy is 1·273 metres, this implies, according to the law of error, or law of probabilities, that all the individual measurements of ten-year-old boys group themselves in an orderly way, that is to say according to a certain definite law, about this mean value of 1·273. When these individual measurements are grouped and plotted as a curve, so as to show the number of individual cases at each individual length, we obtain a char­ac­ter­is­tic curve of error or curve of frequency; and the “spread” of this curve is a measure of the amount of variability in this particular case. A certain math­e­mat­i­cal measure of this “spread,” as described in works upon statistics, is called the Index of Variability, or Standard Deviation, and is usually denominated by the letter σ. It is practically equivalent to a determination of the point upon the frequency curve where it changes its curvature on either side of the mean, and where, from being concave towards the middle line, it spreads out to be convex thereto. When we divide this {79} value by the mean, we get a figure which is independent of any particular units, and which is called the Coefficient of Variability. (It is usually multiplied by 100, to make it of a more convenient amount; and we may then define this coefficient, C, as = (σ ⁄ M) × 100.)

In regard to the growth of man, Pearson has determined this coefficient of variability as follows: in male new-born infants, the coefficient in regard to weight is 15·66, and in regard to stature, 6·50; in male adults, for weight 10·83, and for stature, 3·66. The amount of variability tends, therefore, to decrease with growth or age.

Similar determinations have been elaborated by Bowditch, by Boas and Wissler, and by other writers for intermediate ages, especially from about five years old to eighteen, so covering a great part of the whole period of growth in man108.

The result is very curious indeed. We see, from Fig. 11, that the curve of variability is very similar to what we have called the acceleration-curve (Fig. 4): that is to say, it descends when the rate of growth diminishes, and rises very markedly again when, in late boyhood, the rate of growth is temporarily accelerated. We {80} see, in short, that the amount of variability in stature or in weight is a function of the rate of growth in these magnitudes, though we are not yet in a position to equate the terms precisely, one with another.

Fig. 11. Coefficients of variability of stature in Man (). from Boas and Wissler’s data.

The whole phenomenon of variability in regard to magnitude and to rate of increment is in the highest degree suggestive: inasmuch as it helps further to remind and to impress upon us that specific rate of growth is the real physiological factor which we want to get at, of which specific magnitude, dimensions and form, and all the variations of these, are merely the concrete and visible resultant. But the problems of variability, though they are intimately related to the general problem of growth, carry us very soon beyond our present limitations.

Just as the human curve of growth has its slight but well-marked interruptions, or variations in rate, coinciding with such epochs as birth and puberty, so is it with other animals, and this phenomenon is particularly striking in the case of animals which undergo a regular metamorphosis.

In the accompanying curve of growth in weight of the mouse (Fig. 12), based on W. Ostwald’s observations111, we see a distinct slackening of the rate when the mouse is about a fortnight old, at which period it opens its eyes and very soon afterwards is weaned. At about six weeks old there is another well-marked retardation of growth, following on a very rapid period, and coinciding with the epoch of puberty. {83}

Fig. 13 shews the curve of growth of the silkworm112, during its whole larval life, up to the time of its entering the chrysalis stage.

The silkworm moults four times, at intervals of about a week, the first moult being on the sixth or seventh day after hatching. A distinct retardation of growth is exhibited on our curve in the case of the third and fourth moults; while a similar retardation accompanies the first and second moults also, but the scale of our diagram does not render it visible. When the worm is about seven weeks old, a remarkable process of “purgation” takes place, as a preliminary to entering on the pupal, or chrysalis, stage; and the great and sudden loss of weight which accompanies this process is the most marked feature of our curve.

Fig. 12. Growth in weight of Mouse. (After W. Ostwald.)

The rate of growth in the tadpole113 (Fig. 14) is likewise marked by epochs of retardation, and finally by a sudden and drastic change. There is a slight diminution in weight immediately after {84} the little larva frees itself from the egg; there is a retardation of growth about ten days later, when the external gills disappear; and finally, the complete metamorphosis, with the loss of the tail, the growth of the legs and the cessation of branchial respiration, is accompanied by a loss of weight amounting to wellnigh half the weight of the full-grown larva. {85}

Fig. 13. Growth in weight of Silkworm. (From Ostwald, after Luciani and Lo Monaco.)

While as a general rule, the better the animals be fed the quicker they grow and the sooner they metamorphose, Barfürth has pointed out the curious fact that a short spell of starvation, just before metamorphosis is due, appears to hasten the change.

Fig. 14. Growth in weight of Tadpole. (From Ostwald, after Schaper.)

The negative growth, or actual loss of bulk and weight which often, and perhaps always, accompanies metamorphosis, is well shewn in the case of the eel114. The contrast of size is great between {87} the flattened, lancet-shaped Leptocephalus larva and the little black cylindrical, almost thread-like elver, whose magnitude is less than that of the Leptocephalus in every dimension, even, at first, in length (Fig. 15).

Fig. 15. Development of Eel; from Leptocephalus larvae to young Elver. (From Ostwald after Joh. Schmidt.)

Fig. 16. Growth in length of Spirogyra. (From Ostwald, after Hofmeister.)

From the higher study of the physiology of growth we learn that such fluctuations as we have described are but special interruptions in a process which is never actually continuous, but is perpetually interrupted in a rhythmic manner115. Hofmeister shewed, for instance, that the growth of Spirogyra proceeds by fits and starts, by periods of activity and rest, which alternate with one another at intervals of so many minutes (Fig. 16). And Bose, by very refined methods of experiment, has shewn that plant-growth really proceeds by tiny and perfectly rhythmical pulsations recurring at regular intervals of a few seconds of time. Fig. 17 shews, according to Bose’s observations116, the growth of a crocus, under a very high magnification. The stalk grows by little jerks, each with an amplitude of about ·002 mm., every {88} twenty seconds or so, and after each little increment there is a partial recoil.

Fig. 17. Pulsations of growth in Crocus, in micro-millimetres. (After Bose.)

The differences in regard to rate of growth between various parts or organs of the body, internal and external, can be amply illustrated in the case of man, and also, but chiefly in regard to external form, in some few other creatures118. It is obvious that there lies herein an endless field for the math­e­mat­i­cal study of correlation and of variability, but with this aspect of the case we cannot deal.

In the accompanying table, I shew, from some of Vierordt’s data, the relative weights, at various ages, compared with the weight at birth, of the entire body, of the brain, heart and liver; {89} and also the percentage relation which each of these organs bears, at the several ages, to the weight of the whole body.

From the first portion of the table, it will be seen that none of these organs by any means keep pace with the body as a whole in regard to growth in weight; in other words, there must be some other part of the fabric, doubtless the muscles and the bones, which increase more rapidly than the average increase of the body. Heart and liver both grow nearly at the same rate, and by the {90} age of twenty-five they have multiplied their weight at birth by about thirteen times, while the weight of the entire body has been multiplied by about twenty-one; but the weight of the brain has meanwhile been multiplied only about three and a quarter times. In the next place, we see the very remarkable phenomenon that the brain, growing rapidly till the child is about four years old, then grows more much slowly till about eight or nine years old, and after that time there is scarcely any further perceptible increase. These phenomena are dia­gram­ma­ti­cally illustrated in Fig. 18.

Fig. 18. Relative growth in weight (in Man) of Brain, Heart, and whole Body.

The second part of the table shews the steadily decreasing weights of the organs in question as compared with the body; the brain falling from over 12 per cent. at birth to little over 2 per cent. at five and twenty; the heart from ·75 to ·46 per cent.; and the liver from 4·57 to 2·75 per cent. of the whole bodily weight.

It is plain, then, that there is no simple and direct relation, holding good throughout life, between the size of the body as a whole and that of the organs we have just discussed; and the changing ratio of magnitude is especially marked in the case of the brain, which, as we have just seen, constitutes about one-eighth of the whole bodily weight at birth, and but one-fiftieth at five and twenty. The same change of ratio is observed in other animals, in equal or even greater degree. For instance, Max Weber121 tells us that in the lion, at five weeks, four months, eleven months, and lastly when full-grown, the brain-weight represents the following fractions of the weight of the whole body, viz. 1 ⁄ 18, 1 ⁄ 80, 1 ⁄ 184, and 1 ⁄ 546. And Kellicott has, in like manner, shewn that in the dogfish, while some organs (e.g. rectal gland, pancreas, etc.) increase steadily and very nearly proportionately to the body as a whole, the brain, and some other organs also, grow in a diminishing ratio, which is capable of representation, ap­prox­i­mate­ly, by a logarithmic curve122.

But if we confine ourselves to the adult, then, as Raymond Pearl has shewn in the case of man, the relation of brain-weight to age, to stature, or to weight, becomes a comparatively simple one, and may be sensibly expressed by a straight line, or simple equation.

As regards external form, very similar differences exist, which however we must express in terms not of weight but of length. Thus the annexed table shews the changing ratios of the vertical length of the head to the entire stature; and while this ratio constantly diminishes, it will be seen that the rate of change is greatest (or the coefficient of acceleration highest) between the ages of about two and five years.

In one of Quetelet’s tables (supra, p. 63), he gives measurements of the total span of the outstretched arms in man, from year to year, compared with the vertical stature. The two measurements are so nearly identical in actual magnitude that a direct comparison by means of curves becomes unsatisfactory; but I have reduced Quetelet’s data to percentages, and it will be seen from Fig. 19 that the percentage proportion of span to height undergoes a remarkable and steady change from birth to the age of twenty years; the man grows more rapidly in stretch of arms than he does in height, and the span which was less than {94} the stature at birth by about 1 per cent. exceeds it at the age of twenty by about 4 per cent. After the age of twenty, Quetelet’s data are few and irregular, but it is clear that the span goes on for a long while increasing in proportion to the stature. How far the phenomenon is due to actual growth of the arms and how far to the increasing breadth of the chest is not yet ascertained.

Fig. 19. Ratio of stature in Man, to span of outstretched arms.
(From Quetelet’s data.)

The differences of rate of growth in different parts of the body are very simply brought out by the following table, which shews the relative growth of certain parts and organs of a young trout, at intervals of a few days during the period of most rapid development. It would not be difficult, from a picture of the little trout at any one of these stages, to draw its ap­prox­i­mate form at any other, by the help of the numerical data here set forth126. {95}

While it is inequality of growth in different directions that we can most easily comprehend as a phenomenon leading to gradual change of outward form, we shall see in another chapter127 that differences of rate at different parts of a longitudinal system, though always in the same direction, also lead to very notable and regular trans­for­ma­tions. Of this phenomenon, the difference in rate of longitudinal growth between head and body is a simple case, and the difference which accompanies and results from it in the bodily form of the child and the man is easy to see. A like phenomenon has been studied in much greater detail in the case of plants, by Sachs and certain other botanists, after a method in use by Stephen Hales a hundred and fifty years before128.

On the growing root of a bean, ten narrow zones were marked off, starting from the apex, each zone a millimetre in breadth. After twenty-four hours’ growth, at a certain constant temperature, the whole marked portion had grown from 10 mm. to 33 mm. in length; but the individual zones had grown at very unequal rates, as shewn in the annexed table129.

Fig. 20. Rate of growth in successive zones near the tip of the bean-root.

The several values in this table lie very nearly (as we see by Fig. 20) in a smooth curve; in other words a definite law, or principle of continuity, connects the rates of growth at successive points along the growing axis of the root. Moreover this curve, in its general features, is singularly like those acceleration-curves which we have already studied, in which we plotted the rate of growth against successive intervals of time, as here we have plotted it against successive spatial intervals of an actual growing structure. If we suppose for a moment that the velocities of growth had been transverse to the axis, instead of, as in this case, longitudinal and parallel with it, it is obvious that these same velocities would have given us a leaf-shaped structure, of which our curve in Fig. 20 (if drawn to a suitable scale) would represent the actual outline on either side of the median axis; or, again, if growth had been not confined to one plane but symmetrical about the axis, we should have had a sort of turnip-shaped root, {97} having the form of a surface of revolution generated by the same curve. This then is a simple and not unimportant illustration of the direct and easy passage from velocity to form.

We have seen that a slow but definite change of form is a common accompaniment of increasing age, and is brought about as the simple and natural result of an altered ratio between the rates of growth in different dimensions: or rather by the progressive change necessarily brought about by the difference in their accelerations. There are many cases however in which the change is all but imperceptible to ordinary measurement, and many others in which some one dimension is easily measured, but others are hard to measure with cor­re­spon­ding accuracy. {98} For instance, in any ordinary fish, such as a plaice or a haddock, the length is not difficult to measure, but measurements of breadth or depth are very much more uncertain. In cases such as these, while it remains difficult to define the precise nature of the change of form, it is easy to shew that such a change is taking place if we make use of that ratio of length to weight which we have spoken of in the preceding chapter. Assuming, as we may fairly do, that weight is directly proportional to bulk or volume, we may express this relation in the form W ⁄ L³ = k, where k is a constant, to be determined for each particular case. (W and L are expressed in grammes and centimetres, and it is usual to multiply the result by some figure, such as 1000, so as to give the constant k a value near to unity.)

Now while this k may be spoken of as a “constant,” having a certain mean value specific to each species of organism, and depending on the form of the organism, any change to which it may be subject will be a very delicate index of progressive changes of form; for we know that our measurements of length are, on the average, very accurate, and weighing is a still more delicate method of comparison than any linear measurement.

Fig. 21. Changes in the weight-length ratio of Plaice, with increasing size.

Thus, in the case of plaice, when we deal with the mean values for a large number of specimens, and when we are careful to deal only with such as are caught in a particular locality and at a particular time, we see that k is by no means constant, but steadily increases to a maximum, and afterwards slowly declines with the increasing size of the fish (Fig. 21). To begin with, therefore, the weight is increasing more rapidly than the cube of the length, and it follows that the length itself is increasing less rapidly than some other linear dimension; while in later life this condition is reversed. The maximum is reached when the length of the fish is somewhere near to 30 cm., and it is tempting to suppose that with this “point of inflection” there is associated some well-marked epoch in the fish’s life. As a matter of fact, the size of 30 cm. is ap­prox­i­mate­ly that at which sexual maturity may be said to begin, or is at least near enough to suggest a close connection between the two phenomena. The first step towards further in­ves­ti­ga­tion of the {100} apparent coincidence would be to determine the coefficient k of the two sexes separately, and to discover whether or not the point of inflection is reached (or sexual maturity is reached) at a smaller size in the male than in the female plaice; but the material for this in­ves­ti­ga­tion is at present scanty.

Fig. 22. Periodic annual change in the weight-length ratio of Plaice.

A still more curious and more unexpected result appears when we compare the values of k for the same fish at different seasons of the year131. When for simplicity’s sake (as in the accompanying table and Fig. 22) we restrict ourselves to fish of one particular size, it is not necessary to determine the value of k, because a change in the ratio of length to weight is obvious enough; but when we have small numbers, and various sizes, to deal with, the determination of k may help us very much. It will be seen, then, that in the case of plaice the ratio of weight to length exhibits a regular periodic variation with the course of the seasons. {101}

With unchanging length, the weight and therefore the bulk of the fish falls off from about November to March or April, and again between May or June and November the bulk and weight are gradually restored. The explanation is simple, and depends wholly on the process of spawning, and on the subsequent building up again of the tissues and the reproductive organs. It follows that, by this method, without ever seeing a fish spawn, and without ever dissecting one to see the state of its reproductive system, we can ascertain its spawning season, and determine the beginning and end thereof, with great accuracy.

As a final illustration of the rate of growth, and of unequal growth in various directions, I give the following table of data regarding the ox, extending over the first three years, or nearly so, of the animal’s life. The observed data are (1) the weight of the animal, month by month, (2) the length of the back, from the occiput to the root of the tail, and (3) the height to the withers. To these data I have added (1) the ratio of length to height, (2) the coefficient (k) expressing the ratio of weight to the cube of the length, and (3) a similar coefficient (k′) for the height of the animal. It will be seen that, while all these ratios tend to alter continuously, shewing that the animal’s form is steadily altering as it approaches maturity, the ratio between length and weight {102} changes comparatively little. The simple ratio between length and height increases considerably, as indeed we should expect; for we know that in all Ungulate animals the legs are remarkably

long at birth in comparison with other dimensions of the body. It is somewhat curious, however, that this ratio seems to fall off a little in the third year of growth, the animal continuing to grow in height to a marked degree after growth in length has become very slow. The ratio between height and weight is by much the most variable of our three ratios; the coefficient W ⁄ H³ steadily increases, and is more than twice as great at three years old as it was at birth. This illustrates the important, but obvious fact, that the coefficient k is most variable in the case of that dimension which grows most uniformly, that is to say most nearly in proportion to the general bulk of the animal. In short, the successive values of k, as determined (at successive epochs) for one dimension, are a measure of the variability of the others.

From the whole of the foregoing discussion we see that a certain definite rate of growth is a char­ac­ter­is­tic or specific phenomenon, deep-seated in the physiology of the organism; and that a very large part of the specific morphology of the organism depends upon the fact that there is not only an average, or aggregate, rate of growth common to the whole, but also a variation of rate in different parts of the organism, tending towards a specific rate char­ac­ter­is­tic of each different part or organ. The smallest change in the relative magnitudes of these partial or localised velocities of growth will be soon manifested in more and more striking differences of form. This is as much as to say that the time-element, which is implicit in the idea of growth, can never (or very seldom) be wholly neglected in our consideration of form132. It is scarcely necessary to enlarge here upon our statement, for not only is the truth of it self-evident, but it will find illustration again and again throughout this book. Nevertheless, let us go out of our way for a moment to consider it in reference to a particular case, and to enquire whether it helps to remove any of the difficulties which that case appears to present. {104}

Fig. 23. Variability of length of tail-forceps in a sample of Earwigs. (After Bateson, P. Z. S. 1892, p. 588.)

In a very well-known paper, Bateson shewed that, among a large number of earwigs, collected in a particular locality, the males fell into two groups, characterised by large or by small tail-forceps, with very few instances of intermediate magnitude. This distribution into two groups, according to magnitude, is illustrated in the accompanying diagram (Fig. 23); and the phenomenon was described, and has been often quoted, as one of dimorphism, or discontinuous variation. In this diagram the time-element does not appear; but it is certain, and evident, that it lies close behind. Suppose we take some organism which is born not at all times of the year (as man is) but at some one particular season (for instance a fish), then any random sample will consist of individuals whose ages, and therefore whose magnitudes, will form a discontinuous series; and by plotting these magnitudes on a curve in relation to the number of individuals of each particular magnitude, we obtain a curve such as that shewn in Fig. 24, the first practical use of which is to enable us to analyse our sample into its constituent “age-groups,” or in other words to determine ap­prox­i­mate­ly the age, or ages of the fish. And if, instead of measuring the whole length of our fish, we had confined ourselves to particular parts, such as head, or {105} tail or fin, we should have obtained discontinuous curves of distribution, precisely analogous to those for the entire animal. Now we know that the differences with which Bateson was dealing were entirely a question of magnitude, and we cannot help seeing that the discontinuous distributions of magnitude represented by his earwigs’ tails are just such as are illustrated by the magnitudes of the older and younger fish; we may indeed go so far as to say that the curves are precisely comparable, for in both cases we see a char­ac­ter­is­tic feature of detail, namely that the “spread” of the curve is greater in the second wave than in the first, that is to say (in the case of the fish) in the older as well as larger series. Over the reason for this phenomenon, which is simple and all but obvious, we need not pause.

Fig. 24. Variability of length of body in a sample of Plaice.

It is evident, then, that in this case of “dimorphism,” the tails of the one group of earwigs (which Bateson calls the “high males”) have either grown faster, or have been growing for a longer period of time, than those of the “low males.” If we could be certain that the whole random sample of earwigs were of one and the same age, then we should have to refer the phenomenon of dimorphism to a physiological phenomenon, simple in kind (however remarkable and unexpected); viz. that there were two alternative {106} values, very different from one another, for the mean velocity of growth, and that the individual earwigs varied around one or other of these mean values, in each case according to the law of probabilities. But on the other hand, if we could believe that the two groups of earwigs were of different ages, then the phenomenon would be simplicity itself, and there would be no more to be said about it133.

Before we pass from the subject of the relative rate of growth of different parts or organs, we may take brief note of the fact that various experiments have been made to determine whether the normal ratios are maintained under altered circumstances of nutrition, and especially in the case of partial starvation. For instance, it has been found possible to keep young rats alive for many weeks on a diet such as is just sufficient to maintain life without permitting any increase of weight. The rat of three weeks old weighs about 25 gms., and under a normal diet should weigh at ten weeks old about 150 gms., in the male, or 115 gms. in the female; but the underfed rat is still kept at ten weeks old to the weight of 25 gms. Under normal diet the proportions of the body change very considerably between the ages of three and ten weeks. For instance the tail gets relatively longer; and even when the total growth of the rat is prevented by underfeeding, the form continues to alter so that this increasing length of the tail is still manifest134. {107}

Again as physiologists have long been aware, there is a marked difference in the variation of weight of the different organs, according to whether the animal’s total weight remain constant, or be caused to diminish by actual starvation; and further striking differences appear when the diet is not only scanty, but ill-balanced. But these phenomena of abnormal growth, however interesting from the physiological view, are of little practical importance to the morphologist.

The rates of growth which we have hitherto dealt with are based on special investigations, conducted under particular local conditions. For instance, Quetelet’s data, so far as we have used them to illustrate the rate of growth in man, are drawn from his study of the population of Belgium. But apart from that “fortuitous” individual variation which we have already considered, it is obvious that the normal rate of growth will be found to vary, in man and in other animals, just as the average stature varies, in different localities, and in different “races.” This phenomenon is a very complex one, and is doubtless a resultant of many undefined contributory causes; but we at least gain something in regard to it, when we discover that the rate of growth is directly affected by temperature, and probably by other physical {108} conditions. Réaumur was the first to shew, and the observation was repeated by Bonnet136, that the rate of growth or development of the chick was dependent on temperature, being retarded at temperatures below and somewhat accelerated at temperatures above the normal temperature of incubation, that is to say the temperature of the sitting hen. In the case of plants the fact that growth is greatly affected by temperature is a matter of familiar knowledge; the subject was first carefully studied by Alphonse De Candolle, and his results and those of his followers are discussed in the textbooks of Botany137.

The annexed diagram (Fig. 25), shewing the growth in length of the roots of some common plants during an identical period of forty-eight hours, at temperatures varying from about 14° to 37° C., is a sufficient illustration of the phenomenon. We see that in all cases there is a certain optimum temperature at which the rate of growth is a maximum, and we can also see that on either side of this optimum temperature the acceleration of growth, positive or negative, with increase of temperature is rapid, while at a distance from the optimum it is very slow. From the data given by Sachs and others, we see further that this optimum temperature is very much the same for all the common plants of our own climate which have as yet been studied; in them it is {109} somewhere about 26° C. (or say 77° F.), or about the temperature of a warm summer’s day; while it is found, very naturally, to be considerably higher in the case of plants such as the melon or the maize, which are at home in warmer regions that our own.

Fig. 25. Relation of rate of growth to temperature in certain plants. (From Sachs’s data.)

In a large number of physical phenomena, and in a very marked degree in all chemical reactions, it is found that rate of action is affected, and for the most part accelerated, by rise of temperature; and this effect of temperature tends to follow a definite “exponential” law, which holds good within a considerable range of temperature, but is altered or departed from when we pass beyond certain normal limits. The law, as laid down by van’t Hoff for chemical reactions, is, that for an interval of n degrees the velocity varies as xⁿ , x being called the “temperature coefficient”139 for the reaction in question. {110}

Van’t Hoff’s law, which has become a fundamental principle of chemical mechanics, is likewise applicable (with certain qualifications) to the phenomena of vital chemistry; and it follows that, on very much the same lines, we may speak of the “temperature coefficient” of growth. At the same time we must remember that there is a very important difference (though we can scarcely call it a fundamental one) between the purely physical and the physiological phenomenon, in that in the former we study (or seek and profess to study) one thing at a time, while in the latter we have always to do with various factors which intersect and interfere; increase in the one case (or change of any kind) tends to be continuous, in the other case it tends to be brought to arrest. This is the simple meaning of that Law of Optimum, laid down by Errera and by Sachs as a general principle of physiology: namely that every physiological process which varies (like growth itself) with the amount or intensity of some external influence, does so according to a law in which progressive increase is followed by progressive decrease; in other words the function has its optimum condition, and its curve shews a definite maximum. In the case of temperature, as Jost puts it, it has on the one hand its accelerating effect which tends to follow van’t Hoff’s law. But it has also another and a cumulative effect upon the organism: “Sie schädigt oder sie ermüdet ihn, und je höher sie steigt, desto rascher macht sie die Schädigung geltend und desto schneller schreitet sie voran.” It would seem to be this double effect of temperature in the case of the organism which gives us our “optimum” curves, which are the expression, accordingly, not of a primary phenomenon, but of a more or less complex resultant. Moreover, as Blackman and others have pointed out, our “optimum” temperature is very ill-defined until we take account also of the duration of our experiment; for obviously, a high temperature may lead to a short, but exhausting, spell of rapid growth, while the slower rate manifested at a lower temperature may be the best in the end. {111} The mile and the hundred yards are won by different runners; and maximum rate of working, and maximum amount of work done, are two very different things140.

In the case of maize, a certain series of experiments shewed that the growth in length of the roots varied with the temperature as follows141:

This first approximation might be considerably improved by taking account of all the experimental values, two only of which we have as yet made use of; but even as it is, we see by Fig. 26 that it is in very fair accordance with the actual results of observation, within those particular limits of temperature to which the experiment is confined. {112}

For an experiment on Lupinus albus, quoted by Asa Gray142, I have worked out the cor­re­spon­ding coefficient, but a little more carefully. Its value I find to be 1·16, or very nearly identical with that we have just found for the maize; and the cor­re­spon­dence between the calculated curve and the actual observations is now a close one.

Fig. 26. Relation of rate of growth to temperature in Maize. Observed values (after Köppen), and calculated curve.

Fig. 27. Relation of rate of growth to temperature in rootlets of Pea. (From Miss I. Leitch’s data.)

While the above results conform fairly well to the law of the temperature coefficient, it is evident that the imbibition of water plays so large a part in the process of elongation of the root or stem that the phenomenon is rather a physical than a chemical one: and on this account, as Blackman has remarked, the data commonly given for the rate of growth in plants are apt to be {114} irregular, and sometimes (we might even say) misleading144. The fact also, which we have already learned, that the elongation of a shoot tends to proceed by jerks, rather than smoothly, is another indication that the phenomenon is not purely and simply a chemical one. We have abundant illustrations, however, among animals, in which we may study the temperature coefficient under circumstances where, though the phenomenon is always complicated by osmotic factors, true metabolic growth or chemical combination plays a larger role. Thus Mlle. Maltaux and Professor Massart145 have studied the rate of division in a certain flagellate, Chilomonas paramoecium, and found the process to take 29 minutes at 15° C., 12 at 25°, and only 5 minutes at 35° C. These velocities are in the ratio of 1 : 2·4 : 5·76, which ratio corresponds precisely to a temperature coefficient of 2·4 for each rise of 10°, or about 1·092 for each degree centigrade.

By means of this principle we may throw light on the apparently complicated results of many experiments. For instance, Fig. 28 is an illustration, which has been often copied, of O. Hertwig’s work on the effect of temperature on the rate of development of the tadpole146.

From inspection of this diagram, we see that the time taken to attain certain stages of development (denoted by the numbers III–VII) was as follows, at 20° and at 10° C., respectively.

That is to say, the time taken to produce a given result at {115} 10° was (on the average) somewhere about 55·6 ⁄ 16·7, or 3·33, times as long as was required at 20°.

Fig. 28. Diagram shewing time taken (in days), at various temperatures (°C.), to reach certain stages of development in the Frog: viz. I, gastrula; II, medullary plate; III, closure of medullary folds; IV, tail-bud; V, tail and gills; VI, tail-fin; VII, operculum beginning; VIII, do. closing; IX, first appearance of hind-legs. (From Jenkinson, after O. Hertwig, 1898.)

That is to say, between the intervals of 10° and 20° C., if it take m days, at a certain given temperature, for a certain stage of development to be attained, it will take m × 1·128ⁿ days, when the temperature is n degrees less, for the same stage to be arrived at.

Fig. 29. Calculated values, cor­re­spon­ding to preceding figure.

Fig. 29 is calculated throughout from this value; and it will be seen that it is extremely concordant with the original diagram, as regards all the stages of development and the whole range of temperatures shewn: in spite of the fact that the coefficient on which it is based was derived by an easy method from a very few points in the original curves. {117}

Karl Peter147, experimenting chiefly on echinoderm eggs, and also making use of Hertwig’s experiments on young tadpoles, gives the normal temperature coefficients for intervals of 10° C. (commonly written Q₁₀) as follows.

These values are not only concordant, but are evidently of the same order of magnitude as the temperature-coefficient in ordinary chemical reactions. Peter has also discovered the very interesting fact that the temperature-coefficient alters with age, usually but not always becoming smaller as age increases.

Furthermore, the temperature coefficient varies with the temperature, diminishing as the temperature rises,—a rule which van’t Hoff has shewn to hold in ordinary chemical operations. Thus, in Rana the temperature coefficient at low temperatures may be as high as 5·6: which is just another way of saying that at low temperatures development is exceptionally retarded.

In certain fish, such as plaice and haddock, I and others have found clear evidence that the ascending curve of growth is subject to seasonal interruptions, the rate during the winter months being always slower than in the months of summer: it is as though we superimposed a periodic, annual, sine-curve upon the continuous curve of growth. And further, as growth itself grows less and less from year to year, so will the difference between the winter and the summer rate also grow less and less. The fluctuation in rate {118} will represent a vibration which is gradually dying out; the amplitude of the sine-curve will gradually diminish till it disappears; in short, our phenomenon is simply expressed by what is known as a “damped sine-curve.” Exactly the same thing occurs in man, though neither in his case nor in that of the fish have we sufficient data for its complete illustration.

We can demonstrate the fact, however, in the case of man by the help of certain very interesting measurements which have been recorded by Daffner148, of the height of German cadets, measured at half-yearly intervals.

In the accompanying diagram (Fig. 30) the half-yearly increments are set forth, from the above table, and it will be seen that they form two even and entirely separate series. The curve joining up each series of points is an acceleration-curve; and the comparison of the two curves gives a clear view of the relative rates of growth during winter and summer, and the fluctuation which these velocities undergo during the years in question. The dotted line represents, ap­prox­i­mate­ly, the acceleration-curve in its continuous fluctuation of alternate seasonal decrease and increase.

In the case of trees, the seasonal fluctuations of growth149 admit {119} of easy determination, and it is a point of considerable interest to compare the phenomenon in evergreen and in deciduous trees. I happen to have no measurements at hand with which to make this comparison in the case of our native trees, but from a paper by Mr Charles E. Hall150 I have compiled certain mean values for growth in the climate of Uruguay.

Fig. 30. Half-yearly increments of growth, in cadets of various ages. (From Daffner’s data.)

The measurements taken were those of the girth of the tree, in mm., at three feet from the ground. The evergreens included species of Pinus, Eucalyptus and Acacia; the deciduous trees included Quercus, Populus, Robinia and Melia. I have merely taken mean values for these two groups, and expressed the monthly values as percentages of the mean annual increase. The result (as shewn by Fig. 31) is very much what we might have expected. The growth of the deciduous trees is completely arrested in winter-time, and the arrest is all but complete over {120} a considerable period of time; moreover, during the warm season, the monthly values are regularly graded (ap­prox­i­mate­ly in a sine-curve) with a clear maximum (in the southern hemisphere) about the month of December. In the evergreen trees, on the other hand, the amplitude of the periodic wave is very much less; there is a notable amount of growth all the year round, and, while there is a marked diminution in rate during the coldest months, there is a tendency towards equality over a considerable part of the warmer season. It is probable that some of the species examined, and especially the pines, were definitely retarded in growth, either by a temperature above their optimum, or by deficiency of moisture, during the hottest period of the year; with the result that the seasonal curve in our diagram has (as it were) its region of maximum cut off.

Fig. 31. Periodic annual fluctuation in rate of growth of trees (in the southern hemisphere).

In the case of trees, the seasonal periodicity of growth is so well marked that we are entitled to make use of the phenomenon in a converse way, and to draw deductions as to variations in {121} climate during past years from the record of varying rates of growth which the tree, by the thickness of its annual rings, has preserved for us. Mr. A. E. Douglass, of the University of Arizona, has made a careful study of this question151, and I have received (through Professor H. H. Turner of Oxford) some measurements of the average width of the successive annual rings in “yellow pine,” 500 years old, from Arizona, in which trees the annual rings are very clearly distinguished. From the year 1391 to 1518, the mean of two trees was used; from 1519 to 1912, the mean of five; and the means of these, and sometimes of larger numbers, were found to be very concordant. A correction was applied by drawing a long, nearly straight line through the curve for the whole period, which line was assumed to represent the slowly diminishing mean width of ring accompanying the increase of size, or age, of the tree; and the actual growth as measured was equated with this diminishing mean. The figures used give, accordingly, the ratio of the actual growth in each year to the mean growth cor­re­spon­ding to the age or magnitude of the tree at that epoch.

It was at once manifest that the rate of growth so determined shewed a tendency to fluctuate in a long period of between 100 and 200 years. I then smoothed in groups of 100 (according to Gauss’s method) the yearly values, so that each number thus found represented the mean annual increase during a century: that is to say, the value ascribed to the year 1500 represented the average annual growth during the whole period between 1450 and 1550, and so on. These values give us a curve of beautiful and surprising smoothness, from which we seem compelled to draw the direct conclusion that the climate of Arizona, during the last 500 years, has fluctuated with a regular periodicity of almost precisely 150 years. Here again we should be left in doubt (so far as these {123} observations go) whether the essential factor be a fluctuation of temperature or an alternation of moisture and aridity; but the character of the Arizona climate, and the known facts of recent years, encourage the belief that the latter is the more direct and more important factor.

Fig. 32. Long-period fluctuation in rate of growth of Arizona trees (smoothed in 100-year periods), from A.D. 1390–1490 to A.D. 1810–1910.

It has been often remarked that our common European trees, such for instance as the elm or the cherry, tend to have larger leaves the further north we go; but in this case the phenomenon is to be ascribed rather to the longer hours of daylight than to any difference of temperature152. The point is a physiological one, and consequently of little importance to us here153; the main point for the morphologist is the very simple one that physical or climatic conditions have greatly influenced the rate of growth. The case is analogous to the direct influence of temperature in modifying the colouration of organisms, such as certain butterflies. Now if temperature affects the rate of growth in strict uniformity, alike in all directions and in all parts or organs, its direct effect must be limited to the production of local races or varieties differing from one another in actual magnitude, as the Siberian goldfinch or bullfinch, for instance, differ from our own. But if there be even ever so little of a discriminating action in the enhancement of growth by temperature, such that it accelerates the growth of one tissue or one organ more than another, then it is evident that it must at once lead to an actual difference of racial, or even “specific” form.

It is not to be doubted that the various factors of climate have some such discriminating influence. The leaves of our northern trees may themselves be an instance of it; and we have, {124} probably, a still better instance of it in the case of Alpine plants154, whose general habit is dwarfed, though their floral organs suffer little or no reduction. The subject, however, has been little investigated, and great as its theoretic importance would be to us, we must meanwhile leave it alone.

The curves of growth which we have now been studying represent phenomena which have at least a two-fold interest, morphological and physiological. To the morphologist, who recognises that form is a “function” of growth, the important facts are mainly these: (1) that the rate of growth is an orderly phenomenon, with general features common to very various organisms, while each particular organism has its own char­ac­ter­is­tic phenomena, or “specific constants”; (2) that rate of growth varies with temperature, that is to say with season and with climate, and with various other physical factors, external and internal; (3) that it varies in different parts of the body, and according to various directions or axes; such variations being definitely correlated with one another, and thus giving rise to the char­ac­ter­is­tic proportions, or form, of the organism, and to the changes in form which it undergoes in the course of its development. But to the physiologist, the phenomenon suggests many other important con­si­de­ra­tions, and throws much light on the very nature of growth itself, as a manifestation of chemical and physical energies.

To be content to shew that a certain rate of growth occurs in a certain organism under certain conditions, or to speak of the phenomenon as a “reaction” of the living organism to its environment or to certain stimuli, would be but an example of that “lack of particularity155” in regard to the actual mechanism of physical cause and effect with which we are apt in biology to be too easily satisfied. But in the case of rate of growth we pass somewhat {125} beyond these limitations; for the affinity with certain types of chemical reaction is plain, and has been recognised by a great number of physiologists.

A large part of the phenomenon of growth, both in animals and still more conspicuously in plants, is associated with “turgor,” that is to say, is dependent on osmotic conditions; in other words, the velocity of growth depends in great measure (as we have already seen, p. 113) on the amount of water taken up into the living cells, as well as on the actual amount of chemical metabolism performed by them156. Of the chemical phenomena which result in the actual increase of protoplasm we shall speak presently, but the rôle of water in growth deserves also a passing word, even in our morphological enquiry.

It has been shewn by Loeb that in Cerianthus or Tubularia, for instance, the cells in order to grow must be turgescent; and this turgescence is only possible so long as the salt water in which the cells lie does not overstep a certain limit of concentration. The limit, in the case of Tubularia, is passed when the salt amounts to about 5·4 per cent. Sea-water contains some 3·0 to 3·5 p.c. of salts; but it is when the salinity falls much below this normal, to about 2·2 p.c., that Tubularia exhibits its maximal turgescence, and maximal growth. A further dilution is said to act as a poison to the animal. Loeb has also shewn157 that in certain eggs (e.g. those of the little fish Fundulus) an increasing concentration of the sea-water (leading to a diminishing “water-content” of the egg) retards the rate of segmentation and at length renders segmentation impossible; though nuclear division, by the way, goes on for some time longer.

Among many other observations of the same kind, those of Bialaszewicz158, on the early growth of the frog, are notable. He shews that the growth of the embryo while still within the {126} vitelline membrane depends wholly on the absorption of water; that whether rate of growth be fast or slow (in accordance with temperature) the quantity of water absorbed is constant; and that successive changes of form correspond to definite quantities of water absorbed. The solid residue, as Davenport has also shewn, may actually and notably diminish, while the embryo organism is increasing rapidly in bulk and weight.

On the other hand, in later stages and especially in the higher animals, the percentage of water tends to diminish. This has been shewn by Davenport in the frog, by Potts in the chick, and particularly by Fehling in the case of man159. Fehling’s results are epitomised as follows:

To such phenomena of osmotic balance as the above, or in other words to the dependence of growth on the uptake of water, Höber160 and also Loeb are inclined to refer the modifications of form which certain phyllopod crustacea undergo, when the highly saline waters which they inhabit are further concentrated, or are abnormally diluted. Their growth, according to Schmankewitsch, is retarded by increase of concentration, so that the individuals from the more saline waters appear stunted and dwarfish; and they become altered or transformed in other ways, which for the most part suggest “degeneration,” or a failure to attain full and perfect development161. Important physiological changes also ensue. The rate of multiplication is increased, and parthenogenetic reproduction is encouraged. Male individuals become plentiful in the less saline waters, and here the females bring forth {127} their young alive; males disappear altogether in the more concentrated brines, and then the females lay eggs, which, however, only begin to develop when the salinity is somewhat reduced.

The best-known case is the little “brine-shrimp,” Artemia salina, found, in one form or another, all the world over, and first discovered more than a century and a half ago in the salt-pans at Lymington. Among many allied forms, one, A. milhausenii, inhabits the natron-lakes of Egypt and Arabia, where, under the name of “loul,” or “Fezzan-worm,” it is eaten by the Arabs162. This fact is interesting, because it indicates (and in­ves­ti­ga­tion has apparently confirmed) that the tissues of the creature are not impregnated with salt, as is the medium in which it lives. The fluids of the body, the milieu interne (as Claude Bernard called them163), are no more salt than are those of any ordinary crustacean or other animal, but contain only some 0·8 per cent. of NaCl164, while the milieu externe may contain 10, 20, or more per cent. of this and other salts; which is as much as to say that the skin, or body-wall, of the creature acts as a “semi-permeable membrane,” through which the dissolved salts are not permitted to diffuse, though water passes through freely: until a statical equi­lib­rium (doubtless of a complex kind) is at length attained.

Among the structural changes which result from increased concentration of the brine (partly during the life-time of the individual, but more markedly during the short season which suffices for the development of three or four, or perhaps more, successive generations), it is found that the tail comes to bear fewer and fewer bristles, and the tail-fins themselves tend at last to disappear; these changes cor­re­spon­ding to what have been {128} described as the specific characters of A. milhausenii, and of a still more extreme form, A. köppeniana; while on the other hand, progressive dilution of the water tends to precisely opposite conditions, resulting in forms which have also been described as separate species, and even referred to a separate genus, Callaonella, closely akin to Branchipus (Fig. 33). Pari passu with these changes, there is a marked change in the relative lengths of the fore and hind portions of the body, that is to say, of the “cephalothorax” and abdomen: the latter growing relatively longer, the salter the water. In other words, not only is the rate of growth of the whole

Fig. 33. Brine-shrimps (Artemia), from more or less saline water. Upper figures shew tail-segment and tail-fins; lower figures, relative length of cephalothorax and abdomen. (After Abonyi.)

animal lessened by the saline concentration, but the specific rates of growth in the parts of its body are relatively changed. This latter phenomenon lends itself to numerical statement, and Abonyi has lately shewn that we may construct a very regular curve, by plotting the proportionate length of the creature’s abdomen against the salinity, or density, of the water; and the several species of Artemia, with all their other correlated specific characters, are then found to occupy successive, more or less well-defined, and more or less extended, regions of the curve (Fig. 33). In short, the density of the water is so clearly a “function” of the specific {129} character, that we may briefly define the species Artemia (Callaonella) Jelskii, for instance, as the Artemia of density 1000–1010 (NaCl), or the typical A. salina, or principalis, as the Artemia of density 1018–1025, and so forth. It is a most interesting fact that these Artemiae, under the protection of their semi-permeable skins, are capable of living in waters not only of great density, but of very varied chemical composition. The natron-lakes, for instance, contain large quantities of magnesium

Fig. 34. Percentage ratio of length of abdomen to cephalothorax in brine-shrimps, at various salinities. (After Abonyi.)

sulphate; and the Artemiae continue to live equally well in artificial solutions where this salt, or where calcium chloride, has largely taken the place of sodium chloride in the more common habitat. Furthermore, such waters as those of the natron-lakes are subject to very great changes of chemical composition as concentration proceeds, owing to the different solubilities of the constituent salts. It appears that the forms which the Artemiae assume, and the changes which they undergo, are identical or {130} in­dis­tin­guish­able, whichever of the above salts happen to exist, or to predominate, in their saline habitat. At the same time we still lack (so far as I know) the simple, but crucial experiments which shall tell us whether, in solutions of different chemical composition, it is at equal densities, or at “isotonic” concentrations (that is to say, under conditions where the osmotic pressure, and consequently the rate of diffusion, is identical), that the same structural changes are produced, or cor­re­spon­ding phases of equi­lib­rium attained.

While Höber and others165 have referred all these phenomena to osmosis, Abonyi is inclined to believe that the viscosity, or mechanical resistance, of the fluid also reacts upon the organism; and other possible modes of operation have been suggested. But we may take it for certain that the phenomenon as a whole is not a simple one; and that it includes besides the passive phenomena of intermolecular diffusion, some other form of activity which plays the part of a regulatory mechanism166.

In ordinary chemical reactions we have to deal (1) with a specific velocity proper to the particular reaction, (2) with variations due to temperature and other physical conditions, (3) according to van’t Hoff’s “Law of Mass,” with variations due to the actual quantities present of the reacting substances, and (4) in certain cases, with variations due to the presence of “catalysing agents.” In the simpler reactions, the law of mass involves a steady, gradual slowing-down of the process, according to a logarithmic ratio, as the reaction proceeds and as the initial amount of substance diminishes; a phenomenon, however, which need not necessarily {131} occur in the organism, part of whose energies are devoted to the continual bringing-up of fresh supplies.

Catalytic action occurs when some substance, often in very minute quantity, is present, and by its presence produces or accelerates an action, by opening “a way round,” without the catalytic agent itself being diminished or used up167. Here the velocity curve, though quickened, is not necessarily altered in form, for gradually the law of mass exerts its effect and the rate of the reaction gradually diminishes. But in certain cases we have the very remarkable phenomenon that a body acting as a catalyser is necessarily formed as a product, or bye-product, of the main reaction, and in such a case as this the reaction-velocity will tend to be steadily accelerated. Instead of dwindling away, the reaction will continue with an ever-increasing velocity: always subject to the reservation that limiting conditions will in time make themselves felt, such as a failure of some necessary ingredient, or a development of some substance which shall antagonise or finally destroy the original reaction. Such an action as this we have learned, from Ostwald, to describe as “autocatalysis.” Now we know that certain products of protoplasmic metabolism, such as the enzymes, are very powerful catalysers, and we are entitled to speak of an autocatalytic action on the part of protoplasm itself. This catalytic activity of protoplasm is a very important phenomenon. As Blackman says, in the address already quoted, the botanists (or the zoologists) “call it growth, attribute it to a specific power of protoplasm for assimilation, and leave it alone as a fundamental phenomenon; but they are much concerned as to the distribution of new growth in innumerable specifically distinct forms.” While the chemist, on the other hand, recognises it as a familiar phenomenon, and refers it to the same category as his other known examples of autocatalysis. {132}

This very important, and perhaps even fundamental phenomenon of growth would seem to have been first recognised by Professor Chodat of Geneva, as we are told by his pupil Monnier168. “On peut bien, ainsi que M. Chodat l’a proposé, considérer l’accroissement comme une réaction chimique complexe, dans laquelle le catalysateur est la cellule vivante, et les corps en présence sont l’eau, les sels, et l’acide carbonique.”

Very soon afterwards a similar suggestion was made by Loeb169, in connection with the synthesis of nuclein or nuclear protoplasm; for he remarked that, as in an autocatalysed chemical reaction, the velocity of the synthesis increases during the initial stage of cell-division in proportion to the amount of nuclear matter already synthesised. In other words, one of the products of the reaction, i.e. one of the constituents of the nucleus, accelerates the production of nuclear from cytoplasmic material.

The phenomenon of autocatalysis is by no means confined to living or protoplasmic chemistry, but at the same time it is char­ac­teris­ti­cally, and apparently constantly, associated therewith. And it would seem that to it we may ascribe a considerable part of the difference between the growth of the organism and the simpler growth of the crystal170: the fact, for instance, that the cell can grow in a very low concentration of its nutritive solution, while the crystal grows only in a supersaturated one; and the fundamental fact that the nutritive solution need only contain the more or less raw materials of the complex constituents of the cell, while the crystal grows only in a solution of its own actual substance171.

As F. F. Blackman has pointed out, the multiplication of an organism, for instance the prodigiously rapid increase of a bacterium, {133} which tends to double its numbers every few minutes, till (were it not for limiting factors) its numbers would be all but incalculable in a day172, is a simple but most striking illustration of the potentialities of protoplasmic catalysis; and (apart from the large share taken by mere “turgescence” or imbibition of water) the same is true of the growth, or cell-multiplication, of a multicellular organism in its first stage of rapid acceleration.

It is not necessary for us to pursue this subject much further, for it is sufficiently clear that the normal “curve of growth” of an organism, in all its general features, very closely resembles the velocity-curve of chemical autocatalysis. We see in it the first and most typical phase of greater and greater acceleration; this is followed by a phase in which limiting conditions (whose details are practically unknown) lead to a falling off of the former acceleration; and in most cases we come at length to a third phase, in which retardation of growth is succeeded by actual diminution of mass. Here we may recognise the influence of processes, or of products, which have become actually deleterious; their deleterious influence is staved off for a while, as the organism draws on its accumulated reserves, but they lead ere long to the stoppage of all activity, and to the physical phenomenon of death. But when we have once admitted that the limiting conditions of growth, which cause a phase of retardation to follow a phase of acceleration, are very imperfectly known, it is plain that, ipso facto, we must admit that a resemblance rather than an identity between this phenomenon and that of chemical autocatalysis is all that we can safely assert meanwhile. Indeed, as Enriques has shewn, points of contrast between the two phenomena are not lacking; for instance, as the chemical reaction draws to a close, it is by the gradual attainment of chemical equi­lib­rium: but when organic growth draws to a close, it is by reason of a very different kind of equi­lib­rium, due in the main to the gradual differentiation of the organism into parts, among whose peculiar {134} and specialised functions that of cell-multiplication tends to fall into abeyance173.

It would seem to follow, as a natural consequence, from what has been said, that we could without much difficulty reduce our curves of growth to logarithmic formulae174 akin to those which the physical chemist finds applicable to his autocatalytic reactions. This has been diligently attempted by various writers175; but the results, while not destructive of the hypothesis itself, are only partially successful. The difficulty arises mainly from the fact that, in the life-history of an organism, we have usually to deal (as indeed we have seen) with several recurrent periods of relative acceleration and retardation. It is easy to find a formula which shall satisfy the conditions during any one of these periodic phases, but it is very difficult to frame a comprehensive formula which shall apply to the entire period of growth, or to the whole duration of life.

But if it be meanwhile impossible to formulate or to solve in precise math­e­mat­i­cal terms the equation to the growth of an organism, we have yet gone a very long way towards the solution of such problems when we have found a “qualitative expression,” as Poincaré puts it; that is to say, when we have gained a fair ap­prox­i­mate knowledge of the general curve which represents the unknown function.

As soon as we have touched on such matters as the chemical phenomenon of catalysis, we are on the threshold of a subject which, if we were able to pursue it, would soon lead us far into the special domain of physiology; and there it would be necessary to follow it if we were dealing with growth as a phenomenon in itself, instead of merely as a help to our study and comprehension of form. For instance the whole question of diet, of overfeeding and underfeeding, would present itself for discussion176. But without attempting to open up this large subject, we may say a {135} further passing word upon the essential fact that certain chemical substances have the power of accelerating or of retarding, or in some way regulating, growth, and of so influencing directly the morphological features of the organism.

Thus lecithin has been shewn by Hatai177, Danilewsky178 and others to have a remarkable power of stimulating growth in various animals; and the so-called “auximones,” which Professor Bottomley prepares by the action of bacteria upon peat appear to be, after a somewhat similar fashion, potent accelerators of the growth of plants. But by much the most interesting cases, from our point of view, are those where a particular substance appears to exert a differential effect, stimulating the growth of one part or organ of the body more than another.

It has been known for a number of years that a diseased condition of the pituitary body accompanies the phenomenon known as “acromegaly,” in which the bones are variously enlarged or elongated, and which is more or less exemplified in every skeleton of a “giant”; while on the other hand, disease or extirpation of the thyroid causes an arrest of skeletal development, and, if it take place early, the subject remains a dwarf. These, then, are well-known illustrations of the regulation of function by some internal glandular secretion, some enzyme or “hormone” (as Bayliss and Starling call it), or “harmozone,” as Gley calls it in the particular case where the function regulated is that of growth, with its consequent influence on form.

Among other illustrations (which are plentiful) we have, for instance the growth of the placental decidua, which Loeb has shewn to be due to a substance given off by the corpus luteum of the ovary, giving to the uterine tissues an abnormal capacity for growth, which in turn is called into action by the contact of the ovum, or even of any foreign body. And various sexual characters, such as the plumage, comb and spurs of the cock, are believed in like manner to arise in response to some particular internal secretion. When the source of such a secretion is removed by castration, well-known morphological changes take place in various animals; and when a converse change takes place, the female acquires, in greater or less degree, characters which are {136} proper to the male, as in certain extreme cases, known from time immemorial, when late in life a hen assumes the plumage of the cock.

There are some very remarkable experiments by Gudernatsch, in which he has shewn that by feeding tadpoles (whether of frogs or toads) on thyroid gland substance, their legs may be made to grow out at any time, days or weeks before the normal date of their appearance179. No other organic food was found to produce the same effect; but since the thyroid gland is known to contain iodine180, Morse experimented with this latter substance, and found that if the tadpoles were fed with iodised amino-acids the legs developed precociously, just as when the thyroid gland itself was used. We may take it, then, as an established fact, whose full extent and bearings are still awaiting in­ves­ti­ga­tion, that there exist substances both within and without the organism which have a marvellous power of accelerating growth, and of doing so in such a way as to affect not only the size but the form or proportions of the organism.

If we once admit, as we are now bound to do, the existence of such factors as these, which, by their physiological activity and apart from any direct action of the nervous system, tend towards the acceleration of growth and consequent modification of form, we are led into wide fields of speculation by an easy and a legitimate pathway. Professor Gley carries such speculations a long, long way: for he says181 that by these chemical influences “Toute une partie de la construction des êtres parait s’expliquer d’une façon toute mécanique. La forteresse, si longtemps inaccessible, du vitalisme est entamée. Car la notion morphogénique était, suivant le mot de Dastre182, comme ‘le dernier réduit de la force vitale.’ ”

The physiological speculations we need not discuss: but, to take a single example from morphology, we begin to understand the possibility, and to comprehend the probable meaning, of the {137} all but sudden appearance on the earth of such exaggerated and almost monstrous forms as those of the great secondary reptiles and the great tertiary mammals183. We begin to see that it is in order to account, not for the appearance, but for the disappearance of such forms as these that natural selection must be invoked. And we then, I think, draw near to the conclusion that what is true of these is universally true, and that the great function of natural selection is not to originate, but to remove: donec ad interitum genus id natura redegit184.

The world of things living, like the world of things inanimate, grows of itself, and pursues its ceaseless course of creative evolution. It has room, wide but not unbounded, for variety of living form and structure, as these tend towards their seemingly endless, but yet strictly limited, possibilities of permutation and degree: it has room for the great and for the small, room for the weak and for the strong. Environment and circumstance do not always make a prison, wherein perforce the organism must either live or die; for the ways of life may be changed, and many a refuge found, before the sentence of unfitness is pronounced and the penalty of extermination paid. But there comes a time when “variation,” in form, dimensions, or other qualities of the organism, goes farther than is compatible with all the means at hand of health and welfare for the individual and the stock; when, under the active and creative stimulus of forces from within and from without, the active and creative energies of growth pass the bounds of physical and physiological equi­lib­rium: and so reach the limits which, as again Lucretius tells us, natural law has set between what may and what may not be,

Then, at last, we are entitled to use the customary metaphor, and to see in natural selection an inexorable force, whose function {138} is not to create but to destroy,—to weed, to prune, to cut down and to cast into the fire185.

The phenomenon of regeneration, or the restoration of lost or amputated parts, is a particular case of growth which deserves separate consideration. As we are all aware, this property is manifested in a high degree among invertebrates and many cold-blooded vertebrates, diminishing as we ascend the scale, until at length, in the warm-blooded animals, it lessens down to no more than that vis medicatrix which heals a wound. Ever since the days of Aristotle, and especially since the experiments of Trembley, Réaumur and Spallanzani in the middle of the eighteenth century, the physiologist and the psychologist have alike recognised that the phenomenon is both perplexing and important. The general phenomenon is amply discussed elsewhere, and we need only deal with it in its immediate relation to growth186.

Regeneration, like growth in other cases, proceeds with a velocity which varies according to a definite law; the rate varies with the time, and we may study it as velocity and as acceleration.

Let us take, as an instance, Miss M. L. Durbin’s measurements of the rate of regeneration of tadpoles’ tails: the rate being here measured in terms, not of mass, but of length, or longitudinal increment187.

From a number of tadpoles, whose average length was 34·2 mm., their tails being on an average 21·2 mm. long, about half the tail {139} (11·5 mm.) was cut off, and the amounts regenerated in successive periods are shewn as follows:

The first line of numbers in this table, if plotted as a curve against the number of days, will give us a very satisfactory view of the “curve of growth” within the period of the observations: that is to say, of the successive relations of length to time, or the velocity of the process. But the third line is not so satisfactory, and must not be plotted directly as an acceleration curve. For it is evident that the “rates” here determined do not correspond to velocities at the dates to which they are referred, but are the mean velocities over a preceding period; and moreover the periods over which these means are taken are here of very unequal length. But we may draw a good deal more information from this experiment, if we begin by drawing a smooth curve, as nearly as possible through the points cor­re­spon­ding to the amounts regenerated (according to the first line of the table); and if we then interpolate from this smooth curve the actual lengths attained, day by day, and derive from these, by subtraction, the successive daily increments, which are the measure of the daily mean velocities (Table, p. 141). (The more accurate and strictly correct method would be to draw successive tangents to the curve.)

In our curve of growth (Fig. 35) we cannot safely interpolate values for the first three days, that is to say for the dates between amputation and the first actual measurement of the regenerated part. What goes on in these three days is very important; but we know nothing about it, save that our curve descended to zero somewhere or other within that period. As we have already learned, we can more or less safely interpolate between known points, or actual observations; but here we have no known starting-point. In short, for all that the observations tell us, and for all that the appearance of the curve can suggest, the curve of growth may have descended evenly to the base-line, which it would then have reached about the end of the second {140} day; or it may have had within the first three days a change of direction, or “point of inflection,” and may then have sprung at once from the base-line at zero. That is to say, there may

Fig. 35. Curve of regenerative growth in tadpoles’ tails. (From M. L. Durbin’s data.)

have been an intervening “latent period,” during which no growth occurred, between the time of injury and the first measurement of regenerative growth;

Fig. 36. Mean daily increments, cor­re­spon­ding to Fig. 35.

or, for all we yet know, regeneration may have begun at once, but with a velocity much less than that which it afterwards attained. This apparently trifling difference would correspond to a very great difference in the nature of the phenomenon, and would lead to a very striking difference in the curve which we have next to draw.

The curve already drawn (Fig. 35) illustrates, as we have seen, the relation of length to time, i.e. L ⁄ T = V. The second (Fig. 36) represents the rate of change of velocity; it sets V against T;

and V ⁄ T or L ⁄ T² , represents (as we have learned) the acceleration of growth, this being simply the “differential coefficient,” the first derivative of the former curve.

Fig. 37. Logarithms of values shewn in Fig. 36.

Now, plotting this acceleration curve from the date of the first measurement made three days after the amputation of the tail (Fig. 36), we see that it has no point of inflection, but falls steadily, only more and more slowly, till at last it comes down nearly to the base-line. The velocities of growth are continually diminishing. As regards the missing portion at the beginning of the curve, we cannot be sure whether it bent round and came down to zero, or whether, as in our ordinary acceleration curves of growth from birth onwards, it started from a maximum. The former is, in this case, obviously the more probable, but we cannot be sure.

As regards that large portion of the curve which we are acquainted with, we see that it resembles the curve known as a rectangular hyperbola, which is the form assumed when two variables (in this case V and T) vary inversely as one another. If we take the logarithms of the velocities (as given in the table) and plot them against time (Fig. 37), we see that they fall, ap­prox­i­mate­ly, into a straight line; and if this curve be plotted on the {143} proper scale we shall find that the angle which it makes with the base is about 25°, of which the tangent is ·46, or in round numbers ½.

Had the angle been 45° (tan 45° = 1), the curve would have been actually a rectangular hyperbola, with V T = constant. As it is, we may assume, provisionally, that it belongs to the same family of curves, so that V^m Tⁿ , or V^m ⁄ n T, or V T^n ⁄ m , are all severally constant. In other words, the velocity varies inversely as some power of the time, or vice versa. And in this particular case, the equation V T² = constant, holds very nearly true; that is to say the velocity varies, or tends to vary, inversely as the square of the time. If some general law akin to this could be established as a general law, or even as a common rule, it would be of great importance.

Fig. 38. Rate of regenerative growth in larger tadpoles.

But though neither in this case nor in any other can the minute increments of growth during the first few hours, or the first couple of days, after injury, be directly measured, yet the most important point is quite capable of solution. What the foregoing curve leaves us in ignorance of, is simply whether growth starts at zero, with zero velocity, and works up quickly to a maximum velocity from which it afterwards gradually falls away; or whether after a latent period, it begins, so to speak, in full force. The answer {144} to this question-depends on whether, in the days following the first actual measurement, we can or cannot detect a daily increment in velocity, before that velocity begins its normal course of diminution. Now this preliminary ascent to a maximum, or point of inflection of the curve, though not shewn in the above-quoted experiment, has been often observed: as for instance, in another similar experiment by the author of the former, the tadpoles being in this case of larger size (average 49·1 mm.)188.

Or, by graphic interpolation,

The acceleration curve is drawn in Fig. 39.

Here we have just what we lacked in the former case, namely a visible point of inflection in the curve about the seventh day (Figs. 38, 39), whose existence is confirmed by successive observations on the 3rd, 5th, 7th and 10th days, and which justifies to some extent our extrapolation for the otherwise unknown period up to and ending with the third day; but even here there is a short space near the very beginning during which we are not quite sure of the precise slope of the curve.

We have now learned that, according to these experiments, with which many others are in substantial agreement, the rate of growth in the regenerative process is as follows. After a very short latent period, not yet actually proved but whose existence is highly probable, growth commences with a velocity which very {145} rapidly increases to a maximum. The curve quickly,—almost suddenly,—changes its direction, as the velocity begins to fall; and the rate of fall, that is, the negative acceleration, proceeds at a slower and slower rate, which rate varies inversely as some power of the time, and is found in both of the above-quoted experiments to be very ap­prox­i­mate­ly as 1 ⁄ T² . But it is obvious that the value which we have found for the latter portion of the curve (however closely it be conformed to) is only an empirical value; it has only a temporary usefulness, and must in time give place to a formula which shall represent the entire phenomenon, from start to finish.

Fig. 39. Daily increment, or amount regenerated, cor­re­spon­ding to Fig. 38.

While the curve of regenerative growth is apparently different from the curve of ordinary growth as usually drawn (and while this apparent difference has been commented on and treated as valid by certain writers) we are now in a position to see that it only looks different because we are able to study it, if not from the beginning, at least very nearly so: while an ordinary curve of growth, as it is usually presented to us, is one which dates, not {146} from the beginning of growth, but from the comparatively late, and unimportant, and even fallacious epoch of birth. A complete curve of growth, starting from zero, has the same essential char­ac­teris­tics as the regeneration curve.

Indeed the more we consider the phenomenon of regeneration, the more plainly does it shew itself to us as but a particular case of the general phenomenon of growth189, following the same lines, obeying the same laws, and merely started into activity by the special stimulus, direct or indirect, caused by the infliction of a wound. Neither more nor less than in other problems of physiology are we called upon, in the case of regeneration, to indulge in metaphysical speculation, or to dwell upon the beneficent purpose which seemingly underlies this process of healing and restoration.

It is a very general rule, though apparently not a universal one, that regeneration tends to fall somewhat short of a complete restoration of the lost part; a certain percentage only of the lost tissues is restored. This fact was well known to some of those old investigators, who, like the Abbé Trembley and like Voltaire, found a fascination in the study of artificial injury and the regeneration which followed it. Sir John Graham Dalyell, for instance, says, in the course of an admirable paragraph on regeneration190: “The reproductive faculty ... is not confined to one portion, but may extend over many; and it may ensue even in relation to the regenerated portion more than once. Nevertheless, the faculty gradually weakens, so that in general every successive regeneration is smaller and more imperfect than the organisation preceding it; and at length it is exhausted.”

In certain minute animals, such as the Infusoria, in which the capacity for “regeneration” is so great that the entire animal may be restored from the merest fragment, it becomes of great interest to discover whether there be some definite size at which the fragment ceases to display this power. This question has {147} been studied by Lillie191, who found that in Stentor, while still smaller fragments were capable of surviving for days, the smallest portions capable of regeneration were of a size equal to a sphere of about 80 µ in diameter, that is to say of a volume equal to about one twenty-seventh of the average entire animal. He arrives at the remarkable conclusion that for this, and for all other species of animals, there is a “minimal organisation mass,” that is to say a “minimal mass of definite size consisting of nucleus and cytoplasm within which the organisation of the species can just find its latent expression.” And in like manner, Boveri192 has shewn that the fragment of a sea-urchin’s egg capable of growing up into a new embryo, and so discharging the complete functions of an entire and uninjured ovum, reaches its limit at about one-twentieth of the original egg,—other writers having found a limit at about one-fourth. These magnitudes, small as they are, represent objects easily visible under a low power of the microscope, and so stand in a very different category to the minimal magnitudes in which life itself can be manifested, and which we have discussed in chapter II.

A number of phenomena connected with the linear rate of regeneration are illustrated and epitomised in the accompanying diagram (Fig. 40), which I have constructed from certain data given by Ellis in a paper on the relation of the amount of tail regenerated to the amount removed, in Tadpoles. These data are summarised in the next Table. The tadpoles were all very much of a size, about 40 mm.; the average length of tail was very near to 26 mm., or 65 per cent. of the whole body-length; and in four series of experiments about 10, 20, 40 and 60 per cent. of the tail were severally removed. The amount regenerated in successive intervals of three days is shewn in our table. By plotting the actual amounts regenerated against these three-day intervals of time, we may interpolate values for the time taken to regenerate definite percentage amounts, 5 per cent., 10 per cent., etc. of the {148}

Fig. 40. Relation between the percentage amount of tail removed, the percentage restored, and the time required for its restoration. (From M. M. Ellis’s data.)

amount removed; and my diagram is constructed from the four sets of values thus obtained, that is to say from the four sets of experiments which differed from one another in the amount of tail amputated. To these we have to add the general result of a fifth series of experiments, which shewed that when as much as 75 per cent. of the tail was cut off, no regeneration took place at all, but the animal presently died. In our diagram, then, each {149} curve indicates the time taken to regenerate n per cent. of the amount removed. All the curves converge towards infinity, when the amount removed (as shewn by the ordinate) approaches 75 per cent.; and all of the curves start from zero, for nothing is regenerated where nothing had been removed. Each curve approximates in form to a cubic parabola.

The amount regenerated varies also with the age of the tadpole and with other factors, such as temperature; in other words, for any given age, or size, of tadpole and also for various specific temperatures, a similar diagram might be constructed.

The power of reproducing, or regenerating, a lost limb is particularly well developed in arthropod animals, and is sometimes accompanied by remarkable modification of the form of the regenerated limb. A case in point, which has attracted much attention, occurs in connection with the claws of certain Crustacea193.

In many Crustacea we have an asymmetry of the great claws, one being larger than the other and also more or less different in form. For instance, in the common lobster, one claw, the larger of the two, is provided with a few great “crushing” teeth, while the smaller claw has more numerous teeth, small and serrated. Though Aristotle thought otherwise, it appears that the crushing-claw may be on the right or left side, indifferently; whether it be on one or the other is a problem of “chance.” It is otherwise in many other Crustacea, where the larger and more powerful claw is always left or right, as the case may be, according to the species: where, in other words, the “probability” of the large or the small claw being left or being right is tantamount to certainty194.

The one claw is the larger because it has grown the faster; {150} it has a higher “coefficient of growth,” and accordingly, as age advances, the disproportion between the two claws becomes more and more evident. Moreover, we must assume that the char­ac­ter­is­tic form of the claw is a “function” of its magnitude; the knobbiness is a phenomenon coincident with growth, and we never, under any circumstances, find the smaller claw with big crushing teeth and the big claw with little serrated ones. There are many other somewhat similar cases where size and form are manifestly correlated, and we have already seen, to some extent, that the phenomenon of growth is accompanied by certain ratios of velocity that lead inevitably to changes of form. Meanwhile, then, we must simply assume that the essential difference between the two claws is one of magnitude, with which a certain differentiation of form is inseparably associated.

If we amputate a claw, or if, as often happens, the crab “casts it off,” it undergoes a process of regeneration,—it grows anew, and evidently does so with an accelerated velocity, which acceleration will cease when equi­lib­rium of the parts is once more attained: the accelerated velocity being a case in point to illustrate that vis revulsionis of Haller, to which we have already referred.

With the help of this principle, Przibram accounts for certain curious phenomena which accompany the process of regeneration. As his experiments and those of Morgan shew, if the large or knobby claw (A) be removed, there are certain cases, e.g. the common lobster, where it is directly regenerated. In other cases, e.g. Alpheus195, the other claw (B) assumes the size and form of that which was amputated, while the latter regenerates itself in the form of the other and weaker one; A and B have apparently changed places. In a third case, as in the crabs, the A-claw regenerates itself as a small or B-claw, but the B-claw remains for a time unaltered, though slowly and in the course of repeated moults it later on assumes the large and heavily toothed A-form.

Much has been written on this phenomenon, but in essence it is very simple. It depends upon the respective rates of growth, upon a ratio between the rate of regeneration and the rate of growth of the uninjured limb: complicated a little, however, by {151} the possibility of the uninjured limb growing all the faster for a time after the animal has been relieved of the other. From the time of amputation, say of A, A begins to grow from zero, with a high “regenerative” velocity; while B, starting from a definite magnitude, continues to increase, with its normal or perhaps somewhat accelerated velocity. The ratio between the two velocities of growth will determine whether, by a given time, A has equalled, outstripped, or still fallen short of the magnitude of B.

That this is the gist of the whole problem is confirmed (if confirmation be necessary) by certain experiments of Wilson’s. It is known that by section of the nerve to a crab’s claw, its growth is retarded, and as the general growth of the animal proceeds the claw comes to appear stunted or dwarfed. Now in such a case as that of Alpheus, we have seen that the rate of regenerative growth in an amputated large claw fails to let it reach or overtake the magnitude of the growing little claw: which latter, in short, now appears as the big one. But if at the same time as we amputate the big claw we also sever the nerve to the lesser one, we so far slow down the latter’s growth that the other is able to make up to it, and in this case the two claws continue to grow at ap­prox­i­mate­ly equal rates, or in other words continue of coequal size.

The phenomenon of regeneration goes some way towards helping us to comprehend the phenomenon of “multiplication by fission,” as it is exemplified at least in its simpler cases in many worms and worm-like animals. For physical reasons which we shall have to study in another chapter, there is a natural tendency for any tube, if it have the properties of a fluid or semi-fluid substance, to break up into segments after it comes to a certain length; and nothing can prevent its doing so, except the presence of some controlling force, such for instance as may be due to the pressure of some external support, or some superficial thickening or other intrinsic rigidity of its own substance. If we add to this natural tendency towards fission of a cylindrical or tubular worm, the ordinary phenomenon of regeneration, we have all that is essentially implied in “reproduction by fission.” And in so far {152} as the process rests upon a physical principle, or natural tendency, we may account for its occurrence in a great variety of animals, zoologically dissimilar; and also for its presence here and absence there, in forms which, though materially different in a physical sense, are zoologically speaking very closely allied.

But the phenomena of regeneration, like all the other phenomena of growth, soon carry us far afield, and we must draw this brief discussion to a close.

For the main features which appear to be common to all curves of growth we may hope to have, some day, a physical explanation. In particular we should like to know the meaning of that point of inflection, or abrupt change from an increasing to a decreasing velocity of growth which all our curves, and especially our acceleration curves, demonstrate the existence of, provided only that they include the initial stages of the whole phenomenon: just as we should also like to have a full physical or physiological explanation of the gradually diminishing velocity of growth which follows, and which (though subject to temporary interruption or abeyance) is on the whole char­ac­ter­is­tic of growth in all cases whatsoever. In short, the char­ac­ter­is­tic form of the curve of growth in length (or any other linear dimension) is a phenomenon which we are at present unable to explain, but which presents us with a definite and attractive problem for future solution. It would seem evident that the abrupt change in velocity must be due, either to a change in that pressure outwards from within, by which the “forces of growth” make themselves manifest, or to a change in the resistances against which they act, that is to say the tension of the surface; and this latter force we do not by any means limit to “surface-tension” proper, but may extend to the development of a more or less resistant membrane or “skin,” or even to the resistance of fibres or other histological elements, binding the boundary layers to the parts within. I take it that the sudden arrest of velocity is much more likely to be due to a sudden increase of resistance than to a sudden diminution of internal energies: in other words, I suspect that it is coincident with some notable event of histological differentiation, such as {153} the rapid formation of a comparatively firm skin; and that the dwindling of velocities, or the negative acceleration, which follows, is the resultant or composite effect of waning forces of growth on the one hand, and increasing superficial resistance on the other. This is as much as to say that growth, while its own energy tends to increase, leads also, after a while, to the establishment of resistances which check its own further increase.

Our knowledge of the whole complex phenomenon of growth is so scanty that it may seem rash to advance even this tentative suggestion. But yet there are one or two known facts which seem to bear upon the question, and to indicate at least the manner in which a varying resistance to expansion may affect the velocity of growth. For instance, it has been shewn by Frazee196 that electrical stimulation of tadpoles, with small current density and low voltage, increases the rate of regenerative growth. As just such an electrification would tend to lower the surface-tension, and accordingly decrease the external resistance, the experiment would seem to support, in some slight degree, the suggestion which I have made.

We may summarise, as follows, the main results of the foregoing discussion:

• (1) Except in certain minute organisms and minute parts of organisms, whose form is due to the direct action of molecular forces, we may look upon the form of the organism as a “function of growth,” or a direct expression of a rate of growth which varies according to its different directions.

• (2) Rate of growth is subject to definite laws, and the velocities in different directions tend to maintain a ratio which is more or less constant for each specific organism; and to this regularity is due the fact that the form of the organism is in general regular and constant.

• (3) Nevertheless, the ratio of velocities in different directions is not absolutely constant, but tends to alter or fluctuate in a regular way; and to these progressive changes are due the changes of form which accompany “development,” and the slower changes of form which continue perceptibly in after life.

• (4) The rate of growth is a function of the age of the organism, it has a maximum somewhat early in life, after which epoch of maximum it slowly declines.

• (5) The rate of growth is directly affected by temperature, and by other physical conditions.

• (6) It is markedly affected, in the way of acceleration or retardation, at certain physiological epochs of life, such as birth, puberty, or metamorphosis.

• (7) Under certain circumstances, growth may be negative, the organism growing smaller: and such negative growth is a common accompaniment of metamorphosis, and a frequent accompaniment of old age.

• (8) The phenomenon of regeneration is associated with a large temporary increase in the rate of growth (or “acceleration” of growth) of the injured surface; in other respects, regenerative growth is similar to ordinary growth in all its essential phenomena.

In this discussion of growth, we have left out of account a vast number of processes, or phenomena, by which, in the physiological mechanism of the body, growth is effected and controlled. We have dealt with growth in its relation to magnitude, and to {155} that relativity of magnitudes which constitutes form; and so we have studied it as a phenomenon which stands at the beginning of a morphological, rather than at the end of a physiological enquiry. Under these restrictions, we have treated it as far as possible, or in such fashion as our present knowledge permits, on strictly physical lines.

In all its aspects, and not least in its relation to form, the growth of organisms has many analogies, some close and some perhaps more remote, among inanimate things. As the waves grow when the winds strive with the other forces which govern the movements of the surface of the sea, as the heap grows when we pour corn out of a sack, as the crystal grows when from the surrounding solution the proper molecules fall into their appropriate places: so in all these cases, very much as in the organism itself, is growth accompanied by change of form, and by a development of definite shapes and contours. And in these cases (as in all other mechanical phenomena), we are led to equate our various magnitudes with time, and so to recognise that growth is essentially a question of rate, or of velocity.

The differences of form, and changes of form, which are brought about by varying rates (or “laws”) of growth, are essentially the same phenomenon whether they be, so to speak, episodes in the life-history of the individual, or manifest themselves as the normal and distinctive char­ac­teris­tics of what we call separate species of the race. From one form, or ratio of magnitude, to another there is but one straight and direct road of transformation, be the journey taken fast or slow; and if the transformation take place at all, it will in all likelihood proceed in the self-same way, whether it occur within the life-time of an individual or during the long ancestral history of a race. No small part of what is known as Wolff’s or von Baer’s law, that the individual organism tends to pass through the phases char­ac­ter­is­tic of its ancestors, or that the life-history of the individual tends to recapitulate the ancestral history of its race, lies wrapped up in this simple account of the relation between rate of growth and form.

But enough of this discussion. Let us leave for a while the subject of the growth of the organism, and attempt to study the conformation, within and without, of the individual cell.

In the early days of the cell-theory, more than seventy years ago, Goodsir was wont to speak of cells as “centres of growth” or “centres of nutrition,” and to consider them as essentially “centres of force.” He looked forward to a time when the forces connected with the cell should be particularly investigated: when, that is to say, minute anatomy should be studied in its dynamical aspect. “When this branch of enquiry,” he says “shall have been opened up, we shall expect to have a science of organic forces, having direct relation to anatomy, the science of organic forms198.” And likewise, long afterwards, Giard contemplated a science of morphodynamique,—but still looked upon it as forming so guarded and hidden a “territoire scientifique, que la plupart des naturalistes de nos jours ne le verront que comme Moïse vit la terre promise, seulement de loin et sans pouvoir y entrer199.”

To the external forms of cells, and to the forces which produce and modify these forms, we shall pay attention in a later chapter. But there are forms and con­fi­gur­a­tions of matter within the cell, which also deserve to be studied with due regard to the forces, known or unknown, of whose resultant they are the visible expression.

In the long interval since Goodsir’s day, the visible structure, the conformation and configuration, of the cell, has been studied far more abundantly than the purely dynamic problems that are associated therewith. The overwhelming progress of microscopic observation has multiplied our knowledge of cellular and intracellular structure; and to the multitude of visible structures it {157} has been often easier to attribute virtues than to ascribe intelligible functions or modes of action. But here and there nevertheless, throughout the whole literature of the subject, we find recognition of the inevitable fact that dynamical problems lie behind the morphological problems of the cell.

Bütschli pointed out forty years ago, with emphatic clearness, the failure of morphological methods, and the need for physical methods, if we were to penetrate deeper into the essential nature of the cell200. And such men as Loeb and Whitman, Driesch and Roux, and not a few besides, have pursued the same train of thought and similar methods of enquiry.

Whitman201, for instance, puts the case in a nutshell when, in speaking of the so-called “caryokinetic” phenomena of nuclear division, he reminds us that the leading idea in the term “caryokinesis” is motion,—“motion viewed as an exponent of forces residing in, or acting upon, the nucleus. It regards the nucleus as a seat of energy, which displays itself in phenomena of motion202.”

In short it would seem evident that, except in relation to a dynamical in­ves­ti­ga­tion, the mere study of cell structure has but little value of its own. That a given cell, an ovum for instance, contains this or that visible substance or structure, germinal vesicle or germinal spot, chromatin or achromatin, chromosomes or centrosomes, obviously gives no explanation of the activities of the cell. And in all such hypotheses as that of “pangenesis,” in all the theories which attribute specific properties to micellae, {158} idioplasts, ids, or other constituent particles of protoplasm or of the cell, we are apt to fall into the error of attributing to matter what is due to energy and is manifested in force: or, more strictly speaking, of attributing to material particles individually what is due to the energy of their collocation.

The tendency is a very natural one, as knowledge of structure increases, to ascribe particular virtues to the material structures themselves, and the error is one into which the disciple is likely to fall, but of which we need not suspect the master-mind. The dynamical aspect of the case was in all probability kept well in view by those who, like Goodsir himself, first attacked the problem of the cell and originated our conceptions of its nature and functions.

But if we speak, as Weismann and others speak, of an “hereditary substance,” a substance which is split off from the parent-body, and which hands on to the new generation the char­ac­teris­tics of the old, we can only justify our mode of speech by the assumption that that particular portion of matter is the essential vehicle of a particular charge or distribution of energy, in which is involved the capability of producing motion, or of doing “work.”

For, as Newton said, to tell us that a thing “is endowed with an occult specific quality, by which it acts and produces manifest effects, is to tell us nothing; but to derive two or three general principles of motion203 from phenomena would be a very great step in philosophy, though the causes of these principles were not yet discovered.” The things which we see in the cell are less important than the actions which we recognise in the cell; and these latter we must especially scrutinize, in the hope of discovering how far they may be attributed to the simple and well-known physical forces, and how far they be relevant or irrelevant to the phenomena which we associate with, and deem essential to, the manifestation of life. It may be that in this way we shall in time draw nigh to the recognition of a specific and ultimate residuum. {159}

And lacking, as we still do lack, direct knowledge of the actual forces inherent in the cell, we may yet learn something of their distribution, if not also of their nature, from the outward and inward configuration of the cell, and from the changes taking place in this configuration; that is to say from the movements of matter, the kinetic phenomena, which the forces in action set up.

The fact that the germ-cell develops into a very complex structure, is no absolute proof that the cell itself is structurally a very complicated mechanism: nor yet, though this is somewhat less obvious, is it sufficient to prove that the forces at work, or latent, within it are especially numerous and complex. If we blow into a bowl of soapsuds and raise a great mass of many-hued and variously shaped bubbles, if we explode a rocket and watch the regular and beautiful configuration of its falling streamers, if we consider the wonders of a limestone cavern which a filtering stream has filled with stalactites, we soon perceive that in all these cases we have begun with an initial system of very slight complexity, whose structure in no way foreshadowed the result, and whose comparatively simple intrinsic forces only play their part by complex interaction with the equally simple forces of the surrounding medium. In an earlier age, men sought for the visible embryo, even for the homunculus, within the reproductive cells; and to this day, we scrutinize these cells for visible structure, unable to free ourselves from that old doctrine of “pre-formation204.”

Moreover, the microscope seemed to substantiate the idea (which we may trace back to Leibniz205 and to Hobbes206), that there is no limit to the mechanical complexity which we may postulate in an organism, and no limit, therefore, to the hypotheses which we may rest thereon.

But no microscopical examination of a stick of sealing-wax, no study of the material of which it is composed, can enlighten {160} us as to its electrical manifestations or properties. Matter of itself has no power to do, to make, or to become: it is in energy that all these potentialities reside, energy invisibly associated with the material system, and in interaction with the energies of the surrounding universe.

That “function presupposes structure” has been declared an accepted axiom of biology. Who it was that so formulated the aphorism I do not know; but as regards the structure of the cell it harks back to Brücke, with whose demand for a mechanism, or organisation, within the cell histologists have ever since been attempting to comply207. But unless we mean to include thereby invisible, and merely chemical or molecular, structure, we come at once on dangerous ground. For we have seen, in a former chapter, that some minute “organisms” are already known of such all but infinitesimal magnitudes that everything which the morphologist is accustomed to conceive as “structure” has become physically impossible; and moreover recent research tends generally to reduce, rather than to extend, our conceptions of the visible structure necessarily inherent in living protoplasm. The microscopic structure which, in the last resort or in the simplest cases, it seems to shew, is that of a more or less viscous colloid, or rather mixture of colloids, and nothing more. Now, as Clerk Maxwell puts it, in discussing this very problem, “one material system can differ from another only in the configuration and motion which it has at a given instant208.” If we cannot assume differences in structure, we must assume differences in motion, that is to say, in energy. And if we cannot do this, then indeed we are thrown back upon modes of reasoning unauthorised in physical science, and shall find ourselves constrained to assume, or to “admit, that the properties of a germ are not those of a purely material system.” {161}

But we are by no means necessarily in this dilemma. For though we come perilously near to it when we contemplate the lowest orders of magnitude to which life has been attributed, yet in the case of the ordinary cell, or ordinary egg or germ which is going to develop into a complex organism, if we have no reason to assume or to believe that it comprises an intricate “mechanism,” we may be quite sure, both on direct and indirect evidence, that, like the powder in our rocket, it is very heterogeneous in its structure. It is a mixture of substances of various kinds, more or less fluid, more or less mobile, influenced in various ways by chemical, electrical, osmotic, and other forces, and in their admixture separated by a multitude of surfaces, or boundaries, at which these, or certain of these forces are made manifest.

Indeed, such an arrangement as this is already enough to constitute a “mechanism”; for we must be very careful not to let our physical or physiological concept of mechanism be narrowed to an interpretation of the term derived from the delicate and complicated contrivances of human skill. From the physical point of view, we understand by a “mechanism” whatsoever checks or controls, and guides into determinate paths, the workings of energy; in other words, whatsoever leads in the degradation of energy to its manifestation in some determinate form of work, at a stage short of that ultimate degradation which lapses in uniformly diffused heat. This, as Warburg has well explained, is the general effect or function of the physiological machine, and in particular of that part of it which we call “cell-structure209.” The normal muscle-cell is something which turns energy, derived from oxidation, into work; it is a mechanism which arrests and utilises the chemical energy of oxidation in its downward course; but the same cell when injured or disintegrated, loses its “usefulness,” and sets free a greatly increased proportion of its energy in the form of heat.

But very great and wonderful things are done after this manner by means of a mechanism (whether natural or artificial) of extreme simplicity. A pool of water, by virtue of its surface, {162} is an admirable mechanism for the making of waves; with a lump of ice in it, it becomes an efficient and self-contained mechanism for the making of currents. The great cosmic mechanisms are stupendous in their simplicity; and, in point of fact, every great or little aggregate of heterogeneous matter (not identical in “phase”) involves, ipso facto, the essentials of a mechanism. Even a non-living colloid, from its intrinsic heterogeneity, is in this sense a mechanism, and one in which energy is manifested in the movement and ceaseless rearrangement of the constituent particles. For this reason Graham (if I remember rightly) speaks somewhere or other of the colloid state as “the dynamic state of matter”; or in the same philosopher’s phrase (of which Mr Hardy210 has lately reminded us), it possesses “energia211.”

Let us turn then to consider, briefly and dia­gram­ma­ti­cally, the structure of the cell, a fertilised germ-cell or ovum for instance, not in any vain attempt to correlate this structure with the structure or properties of the resulting and yet distant organism; but merely to see how far, by the study of its form and its changing internal configuration, we may throw light on certain forces which are for the time being at work within it.

We may say at once that we can scarcely hope to learn more of these forces, in the first instance, than a few facts regarding their direction and magnitude; the nature and specific identity of the force or forces is a very different matter. This latter problem is likely to be very difficult of elucidation, for the reason, among others, that very different forces are often very much alike in their outward and visible manifestations. So it has come to pass that we have a multitude of discordant hypotheses as to the nature of the forces acting within the cell, and producing, in cell division, the “caryokinetic” figures of which we are about to speak. One student may, like Rhumbler, choose to account for them by an hypothesis of mechanical traction, acting on a reticular web of protoplasm212; another, like Leduc, may shew us how in {163} many of their most striking features they may be admirably simulated by the diffusion of salts in a colloid medium; others again, like Gallardo213 and Hartog, and Rhumbler (in his earlier papers)214, insist on their resemblance to the phenomena of electricity and magnetism215; while Hartog believes that the force in question is only analogous to these, and has a specific identity of its own216. All these conflicting views are of secondary importance, so long as we seek only to account for certain con­fi­gur­a­tions which reveal the direction, rather than the nature, of a force. One and the same system of lines of force may appear in a field of magnetic or of electrical energy, of the osmotic energy of diffusion, of the gravitational energy of a flowing stream. In short, we may expect to learn something of the pure or abstract dynamics, long before we can deal with the special physics of the cell. For indeed (as Maillard has suggested), just as uniform expansion about a single centre, to whatsoever physical cause it may be due will lead to the configuration of a sphere, so will any two centres or foci of potential (of whatsoever kind) lead to the con­fi­gur­a­tions with which Faraday made us familiar under the name of “lines of force217”; and this is as much as to say that the phenomenon, {164} though physical in the concrete, is in the abstract purely math­e­mat­i­cal, and in its very essence is neither more nor less than a property of three-dimensional space.

But as a matter of fact, in this instance, that is to say in trying to explain the leading phenomena of the caryokinetic division of the cell, we shall soon perceive that any explanation which is based, like Rhumbler’s, on mere mechanical traction, is obviously inadequate, and we shall find ourselves limited to the hypothesis of some polarised and polarising force, such as we deal with, for instance, in the phenomena of magnetism or electricity.

Let us speak first of the cell itself, as it appears in a state of rest, and let us proceed afterwards to study the more active phenomena which accompany its division.

Our typical cell is a spherical body; that is to say, the uniform surface-tension at its boundary is balanced by the outward resistance of uniform forces within. But at times the surface-tension may be a fluctuating quantity, as when it produces the rhythmical contractions or “Ransom’s waves” on the surface of a trout’s egg; or again, while the egg is in contact with other bodies, the surface-tension may be locally unequal and variable, giving rise to an amoeboid figure, as in the egg of Hydra218.

Within the ovum is a nucleus or germinal vesicle, also spherical, and consisting as a rule of portions of “chromatin,” aggregated together within a more fluid drop. The fact has often been commented upon that, in cells generally, there is no correlation of form (though there apparently is of size) between the nucleus and the “cytoplasm,” or main body of the cell. So Whitman219 remarks that “except during the process of division the nucleus seldom departs from its typical spherical form. It divides and sub-divides, ever returning to the same round or oval form .... How different with the cell. It preserves the spherical form as rarely as the nucleus departs from it. Variation in form marks the beginning and the end of every important chapter in its {165} history.” On simple dynamical grounds, the contrast is easily explained. So long as the fluid substance of the nucleus is qualitatively different from, and incapable of mixing with, the fluid or semi-fluid protoplasm which surrounds it, we shall expect it to be, as it almost always is, of spherical form. For, on the one hand, it is bounded by a liquid film, whose surface-tension is uniform; and on the other, it is immersed in a medium which transmits on all sides a uniform fluid pressure220. For a similar reason the contractile vacuole of a Protozoon is spherical in form: it is just a “drop” of fluid, bounded by a uniform surface-tension and through whose boundary-film diffusion is taking place. But here, owning to the small difference between the fluid constituting, and that surrounding, the drop, the surface-tension equi­lib­rium is unstable; it is apt to vanish, and the rounded outline of the drop, like a burst bubble, disappears in a moment221. The case of the spherical nucleus is closely akin to the spherical form of the yolk within the bird’s egg222. But if the substance of the cell acquire a greater solidity, as for instance in a muscle {166} cell, or by reason of mucous accumulations in an epithelium cell, then the laws of fluid pressure no longer apply, the external pressure on the nucleus tends to become unsymmetrical, and its shape is modified accordingly. “Amoeboid” movements may be set up in the nucleus by anything which disturbs the symmetry of its own surface-tension. And the cases, as in many Rhizopods, where “nuclear material” is scattered in small portions throughout the cell instead of being aggregated in a single nucleus, are probably capable of very simple explanation by supposing that the “phase difference” (as the chemists say) between the nuclear and the protoplasmic substance is comparatively slight, and the surface-tension which tends to keep them separate is correspondingly small223.

It has been shewn that ordinary nuclei, isolated in a living or fresh state, easily flow together; and this fact is enough to suggest that they are aggregations of a particular substance rather than bodies deserving the name of particular organs. It is by reason of the same tendency to confluence or aggregation of particles that the ordinary nucleus is itself formed, until the imposition of a new force leads to its disruption.

Apart from that invisible or ultra-microscopic heterogeneity which is inseparable from our notion of a “colloid,” there is a visible heterogeneity of structure within both the nucleus and the outer protoplasm. The former, for instance, contains a rounded nucleolus or “germinal spot,” certain conspicuous granules or strands of the peculiar substance called chromatin, and a coarse meshwork of a protoplasmic material known as “linin” or achromatin; the outer protoplasm, or cytoplasm, is generally believed to consist throughout of a sponge-work, or rather alveolar meshwork, of more and less fluid substances; and lastly, there are generally to be detected one or more very minute bodies, usually in the cytoplasm, sometimes within the nucleus, known as the centrosome or centrosomes.

The morphologist is accustomed to speak of a “polarity” of {167} the cell, meaning thereby a symmetry of visible structure about a particular axis. For instance, whenever we can recognise in a cell both a nucleus and a centrosome, we may consider a line drawn through the two as the morphological axis of polarity; in an epithelium cell, it is obvious that the cell is morphologically symmetrical about a median axis passing from its free surface to its attached base. Again, by an extension of the term “polarity,” as is customary in dynamics, we may have a “radial” polarity, between centre and periphery; and lastly, we may have several apparently independent centres of polarity within the single cell. Only in cells of quite irregular, or amoeboid form, do we fail to recognise a definite and symmetrical “polarity.” The morphological “polarity” is accompanied by, and is but the outward expression (or part of it) of a true dynamical polarity, or distribution of forces; and the “lines of force” are rendered visible by concatenation of particles of matter, such as come under the influence of the forces in action.

When the lines of force stream inwards from the periphery towards a point in the interior of the cell, the particles susceptible of attraction either crowd towards the surface of the cell, or, when retarded by friction, are seen forming lines or “fibrillae” which radiate outwards from the centre and constitute a so-called “aster.” In the cells of columnar or ciliated epithelium, where the sides of the cell are symmetrically disposed to their neighbours but the free and attached surfaces are very diverse from one another in their external relations, it is these latter surfaces which constitute the opposite poles; and in accordance with the parallel lines of force so set up, we very frequently see parallel lines of granules which have ranged themselves perpendicularly to the free surface of the cell (cf. fig. 97).

A simple manifestation of “polarity” may be well illustrated by the phenomenon of diffusion, where we may conceive, and may automatically reproduce, a “field of force,” with its poles and visible lines of equipotential, very much as in Faraday’s conception of the field of force of a magnetic system. Thus, in one of Leduc’s experiments224, if we spread a layer of salt solution over a level {168} plate of glass, and let fall into the middle of it a drop of indian ink, or of blood, we shall find the coloured particles travelling outwards from the central “pole of concentration” along the lines of diffusive force, and so mapping out for us a “monopolar field” of diffusion: and if we set two such drops side by side, their lines of diffusion will oppose, and repel, one another. Or, instead of the uniform layer of salt solution, we may place at a little distance from one another a grain of salt and a drop of blood, representing two opposite poles: and so obtain a picture of a “bipolar field” of diffusion. In either case, we obtain results closely analogous to the “morphological,” but really dynamical, polarity of the organic cell. But in all probability, the dynamical polarity, or asymmetry of the cell is a very complicated phenomenon: for the obvious reason that, in any system, one asymmetry will tend to beget another. A chemical asymmetry will induce an inequality of surface-tension, which will lead directly to a modification of form; the chemical asymmetry may in turn be due to a process of electrolysis in a polarised electrical field; and again the chemical heterogeneity may be intensified into a chemical “polarity,” by the tendency of certain substances to seek a locus of greater or less surface-energy. We need not attempt to grapple with a subject so complicated, and leading to so many problems which lie beyond the sphere of interest of the morphologist. But yet the morphologist, in his study of the cell, cannot quite evade these important issues; and we shall return to them again when we have dealt somewhat with the form of the cell, and have taken account of some of the simpler phenomena of surface-tension.

We are now ready, and in some measure prepared, to study the numerous and complex phenomena which usually accompany the division of the cell, for instance of the fertilised egg.

Division of the cell is essentially accompanied, and preceded, by a change from radial or monopolar to a definitely bipolar polarity.

In the hitherto quiescent, or apparently quiescent cell, we perceive certain movements, which correspond precisely to what must accompany and result from a “polarisation” of forces within the {169} cell: of forces which, whatever may be their specific nature, at least are capable of polarisation, and of producing consequent attraction or repulsion between charged particles of matter. The opposing forces which were distributed in equi­lib­rium throughout the substance of the cell become focussed at two “centrosomes,” which may or may not be already distinguished as visible portions of matter; in the egg, one of these is always near to, and the other remote from, the “animal pole” of the egg, which pole is visibly as well as chemically different from the other, and is the region in which the more rapid and conspicuous developmental changes will presently begin. Between the two centrosomes, a spindle-shaped

Fig. 41. Caryokinetic figure in a dividing cell (or blastomere) of the Trout’s egg. (After Prenant, from a preparation by Prof. P. Bouin.)

figure appears, whose striking resemblance to the lines of force made visible by iron-filings between the poles of a magnet, was at once recognised by Hermann Fol, when in 1873 he witnessed for the first time the phenomenon in question. On the farther side of the centrosomes are seen star-like figures, or “asters,” in which we can without difficulty recognise the broken lines of force which run externally to those stronger lines which lie nearer to the polar axis and which constitute the “spindle.” The lines of force are rendered visible or “material,” just as in the experiment of the iron-filings, by the fact that, in the heterogeneous substance of the cell, certain portions of matter are more “permeable” to the acting force than the rest, become themselves polarised after the {170} fashion of a magnetic or “paramagnetic” body, arrange themselves in an orderly way between the two poles of the field of force, cling to one another as it were in threads225, and are only prevented by the friction of the surrounding medium from approaching and congregating around the adjacent poles.

As the field of force strengthens, the more will the lines of force be drawn in towards the interpolar axis, and the less evident will be those remoter lines which constitute the terminal, or extrapolar, asters: a clear space, free from materialised lines of force, may thus tend to be set up on either side of the spindle, the so-called “Bütschli space” of the histologists226. On the other hand, the lines of force constituting the spindle will be less concentrated if they find a path of less resistance at the periphery of the cell: as happens, in our experiment of the iron-filings, when we encircle the field of force with an iron ring. On this principle, the differences observed between cells in which the spindle is well developed and the asters small, and others in which the spindle is weak and the asters enormously developed, can be easily explained by variations in the potential of the field, the large, conspicuous asters being probably correlated with a marked permeability of the surface of the cell.

The visible field of force, though often called the “nuclear spindle,” is formed outside of, but usually near to, the nucleus. Let us look a little more closely into the structure of this body, and into the changes which it presently undergoes.

Within its spherical outline (Fig. 42), it contains an “alveolar” {171} meshwork (often described, from its appearance in optical section, as a “reticulum”), consisting of more solid substances, with more fluid matter filling up the interalveolar meshes. This phenomenon is nothing else than what we call in ordinary language, a “froth” or a “foam.” It is a surface-tension phenomenon, due to the interacting surface-tensions of two intermixed fluids, not very different in density, as they strive to separate. Of precisely the same kind (as Bütschli was the first to shew) are the minute alveolar networks which are to be discerned in the cytoplasm of the cell227, and which we now know to be not inherent in the nature of protoplasm, or of living matter in general, but to be due to various causes, natural as well as artificial. The microscopic honeycomb structure of cast metal under various conditions of cooling, even on a grand scale the columnar structure of basaltic rock, is an example of the same surface-tension phenomenon. {172}

Fig. 42.

Fig. 43.

The following is a brief epitome of the visible changes undergone by a typical cell, leading up to the act of segmentation, and constituting the phenomenon of mitosis or caryokinetic division. In the egg of a sea-urchin, we see with almost diagrammatic completeness what is set forth here231.

Fig. 44.

Fig. 45.

• 1. The chromatin, which to begin with was distributed in granules on the otherwise achromatic reticulum (Fig. 42), concentrates to form a skein or spireme, which may be a continuous thread from the first (Figs. 43, 44), or from the first segmented. In any case it divides transversely sooner or later into a number of chromosomes (Fig. 45), which as a rule have the shape of little rods, straight or curved, often bent into a V, but which may also be ovoid, or round, or even annular. Certain deeply staining masses, the nucleoli, which may be present in the resting nucleus, do not take part in the process of chromosome formation; they are either cast out of the nucleus and are dissolved in the cytoplasm, or fade away in situ.

• 2. Meanwhile, the deeply staining granule (here extra-nuclear), known as the centrosome, has divided in two. The two resulting granules travel to opposite poles of the nucleus, and {174} there each becomes surrounded by a system of radiating lines, the asters; immediately around the centrosome is a clear space, the centrosphere (Figs. 43–45). Between the two centrosomes with their asters stretches a bundle of achromatic fibres, the spindle.

• 3. The surface-film bounding the nucleus has broken down, the definite nuclear boundaries are lost, and the spindle now stretches through the nuclear material, in which lie the chromosomes (Figs. 45, 46). These chromosomes now arrange themselves midway between the poles of the spindle, where they form what is called the equatorial plate (Fig. 47).

  

  Fig. 46.

  Fig. 47.

• 4. Each chromosome splits longitudinally into two: usually at this stage,—but it is to be noticed that the splitting may have taken place so early as the spireme stage (Fig. 48).

• 5. The halves of the split chromosomes now separate from one another, and travel in opposite directions towards the two poles (Fig. 49). As they move, it becomes apparent that the spindle consists of a median bundle of “fibres,” the central spindle, running from pole to pole, and a more superficial sheath of “mantle-fibres,” to which the chromosomes seem to be attached, and by which they seem to be drawn towards the asters.

• 6. The daughter chromosomes, arranged now in two groups, become closely crowded in a mass near the centre of each aster {175} (Fig. 50). They fuse together and form once more an alveolar reticulum and may occasionally at this stage form another spireme.

  

  Fig. 48.

  Fig. 49.

  A boundary or surface wall is now developed round each reconstructed nuclear mass, and the spindle-fibres disappear (Fig. 51). The centrosome remains, as a rule, outside the nucleus.

  

  Fig. 50.

  Fig. 51.

• 7. On the central spindle, in the position of the equatorial plate, there has appeared during the migration of the chromosomes, a “cell-plate” of deeply staining thickenings (Figs. 50, 51). This is more conspicuous in plant-cells. {176}

• 8. A constriction has meanwhile appeared in the cytoplasm, and the cell divides through the equatorial plane. In plant-cells the line of this division is foreshadowed by the “cell-plate,” which extends from the spindle across the entire cell, and splits into two layers, between which appears the membrane by which the daughter cells are cleft asunder. In animal cells the cell-plate does not attain such dimensions, and no cell-wall is formed.

The whole, or very nearly the whole of these nuclear phenomena may be brought into relation with that polarisation of forces, in the cell as a whole, whose field is made manifest by the “spindle” and “asters” of which we have already spoken: certain particular phenomena, directly attributable to surface-tension and diffusion, taking place in more or less obvious and inevitable dependence upon the polar system*.

* The reference numbers in the following account refer to the paragraphs and figures of the preceding summary of visible nuclear phenomena.

At the same time, in attempting to explain the phenomena, we cannot say too clearly, or too often, that all that we are meanwhile justified in doing is to try to shew that such and such actions lie within the range of known physical actions and phenomena, or that known physical phenomena produce effects similar to them. We want to feel sure that the whole phenomenon is not sui generis, but is somehow or other capable of being referred to dynamical laws, and to the general principles of physical science. But when we speak of some particular force or mode of action, using it as an illustrative hypothesis, we must stop far short of the implication that this or that force is necessarily the very one which is actually at work within the living cell; and certainly we need not attempt the formidable task of trying to reconcile, or to choose between, the various hypotheses which have already been enunciated, or the several assumptions on which they depend.

Any region of space within which action is manifested is a field of force; and a simple example is a bipolar field, in which the action is symmetrical with reference to the line joining two points, or poles, and also with reference to the “equatorial” plane equidistant from both. We have such a “field of force” in {177} the neighbourhood of the centrosome of the ripe cell or ovum, when it is about to divide; and by the time the centrosome has divided, the field is definitely a bipolar one.

The quality of a medium filling the field of force may be uniform, or it may vary from point to point. In particular, it may depend upon the magnitude of the field; and the quality of one medium may differ from that of another. Such variation of quality, within one medium, or from one medium to another, is capable of diagrammatic representation by a variation of the direction or the strength of the field (other conditions being the same) from the state manifested in some uniform medium taken as a standard. The medium is said to be permeable to the force, in greater or less degree than the standard medium, according as the variation of the density of the lines of force from the standard case, under otherwise identical conditions, is in excess or defect. A body placed in the medium will tend to move towards regions of greater or less force according as its permeability is greater or less than that of the surrounding medium232. In the common experiment of placing iron-filings between the two poles of a magnetic field, the filings have a very high permeability; and not only do they themselves become polarised so as to attract one another, but they tend to be attracted from the weaker to the stronger parts of the field, and as we have seen, were it not for friction or some other resistance, they would soon gather together around the nearest pole. But if we repeat the same experiment with such a metal as bismuth, which is very little permeable to the magnetic force, then the conditions are reversed, and the particles, being repelled from the stronger to the weaker parts of the field, tend to take up their position as far from the poles as possible. The particles have become polarised, but in a sense opposite to that of the surrounding, or adjacent, field.

Now, in the field of force whose opposite poles are marked by {178} the centrosomes the nucleus appears to act as a more or less permeable body, as a body more permeable than the surrounding medium, that is to say the “cytoplasm” of the cell. It is accordingly attracted by, and drawn into, the field of force, and tries, as it were, to set itself between the poles and as far as possible from both of them. In other words, the centrosome-foci will be apparently drawn over its surface, until the nucleus as a whole is involved within the field of force, which is visibly marked out by the “spindle” (par. 3, Figs. 44, 45).

If the field of force be electrical, or act in a fashion analogous to an electrical field, the charged nucleus will have its surface-tensions diminished233: with the double result that the inner alveolar meshwork will be broken up (par. 1), and that the spherical boundary of the whole nucleus will disappear (par. 2). The break-up of the alveoli (by thinning and rupture of their partition walls) leads to the formation of a net, and the further break-up of the net may lead to the unravelling of a thread or “spireme” (Figs. 43, 44).

Here there comes into play a fundamental principle which, in so far as we require to understand it, can be explained in simple words. The effect (and we might even say the object) of drawing the more permeable body in between the poles, is to obtain an “easier path” by which the lines of force may travel; but it is obvious that a longer route through the more permeable body may at length be found less advantageous than a shorter route through the less permeable medium. That is to say, the more permeable body will only tend to be drawn in to the field of force until a point is reached where (so to speak) the way round and the way through are equally advantageous. We should accordingly expect that (on our hypothesis) there would be found cases in which the nucleus was wholly, and others in which it was only partially, and in greater or less degree, drawn in to the field between the centrosomes. This is precisely what is found to occur in actual fact. Figs. 44 and 45 represent two so-called “types,” of a phase which follows that represented in Fig. 43. According to the usual descriptions (and in particular to Professor {179} E. B. Wilson’s234), we are told that, in such a case as Fig. 44, the “primary spindle” disappears and the centrosomes diverge to opposite poles of the nucleus; such a condition being found in many plant-cells, and in the cleavage-stages of many eggs. In Fig. 45, on the other hand, the primary spindle persists, and subsequently comes to form the main or “central” spindle; while at the same time we see the fading away of the nuclear membrane, the breaking up of the spireme into separate chromosomes, and an ingrowth into the nuclear area of the “astral rays,”—all as in Fig. 46, which represents the next succeeding phase of Fig. 45. This condition, of Fig. 46, occurs in a variety of cases; it is well seen in the epidermal cells of the salamander, and is also on the whole char­ac­ter­is­tic of the mode of formation of the “polar bodies.” It is clear and obvious that the two “types” correspond to mere differences of degree, and are such as would naturally be brought about by differences in the relative permeabilities of the nuclear mass and of the surrounding cytoplasm, or even by differences in the magnitude of the former body.

But now an important change takes place, or rather an important difference appears; for, whereas the nucleus as a whole tended to be drawn in to the stronger parts of the field, when it comes to break up we find, on the contrary, that its contained spireme-thread or separate chromosomes tend to be repelled to the weaker parts. Whatever this difference may be due to,—whether, for instance, to actual differences of permeability, or possibly to differences in “surface-charge,”—the fact is that the chromatin substance now behaves after the fashion of a “diamagnetic” body, and is repelled from the stronger to the weaker parts of the field. In other words, its particles, lying in the inter-polar field, tend to travel towards the equatorial plane thereof (Figs. 47, 48), and further tend to move outwards towards the periphery of that plane, towards what the histologist calls the “mantle-fibres,” or outermost of the lines of force of which the spindle is made up (par. 5, Fig. 47). And if this comparatively non-permeable chromatin substance come to consist of separate portions, more or less elongated in form, these portions, or separate “chromosomes,” will adjust themselves longitudinally, {180} in a peripheral equatorial circle (Figs. 48, 49). This is precisely what actually takes place. Moreover, before the breaking up of the nucleus, long before the chromatin material has broken up into separate chromosomes, and at the very time when it is being fashioned into a “spireme,” this body already lies in a polar field, and must already have a tendency to set itself in the equatorial plane thereof. But the long, continuous spireme thread is unable, so long as the nucleus retains its spherical boundary wall, to adjust itself in a simple equatorial annulus; in striving to do so, it must tend to coil and “kink” itself, and in so doing (if all this be so), it must tend to assume the char­ac­ter­is­tic convolutions of the “spireme.”

Fig. 52. Chromosomes, undergoing splitting and separation.
(After Hatschek and Flemming, diagrammatised.)

After the spireme has broken up into separate chromosomes, these particles come into a position of temporary, and unstable, equi­lib­rium near the periphery of the equatorial plane, and here they tend to place themselves in a symmetrical arrangement (Fig. 52). The particles are rounded, linear, sometimes annular, similar in form and size to one another; and lying as they do in a fluid, and subject to a symmetrical system of forces, it is not surprising that they arrange themselves in a symmetrical manner, the precise arrangement depending on the form of the particles themselves. This symmetry may perhaps be due, as has already been suggested, to induced electrical charges. In discussing Brauer’s observations on the splitting of the chromatic filament, and the symmetrical arrangement of the separate granules, in Ascaris megalocephala, Lillie235 {181} remarks: “This behaviour is strongly suggestive of the division of a colloidal particle under the influence of its surface electrical charge, and of the effects of mutual repulsion in keeping the products of division apart.” It is also probable that surface-tensions between the particles and the surrounding protoplasm would bring about an identical result, and would sufficiently account for the obvious, and at first sight, very curious, symmetry. We know that if we float a couple of matches in water they tend to approach one another, till they lie close together, side by side; and, if we lay upon a smooth wet plate four matches, half broken across, a precisely similar attraction brings the four matches together in the form of a symmetrical cross. Whether one of these, or some other, be the actual explanation of the phenomenon, it is at least plain that by some physical cause, some mutual and symmetrical attraction or repulsion of the particles, we must seek

Fig. 53. Annular chromosomes, formed in the spermatogenesis of the Mole-cricket. (From Wilson, after Vom Rath.)

to account for the curious symmetry of these so-called “tetrads.” The remarkable annular chromosomes, shewn in Fig. 53, can also be easily imitated by means of loops of thread upon a soapy film when the film within the annulus is broken or its tension reduced.

So far as we have now gone, there is no great difficulty in pointing to simple and familiar phenomena of a field of force which are similar, or comparable, to the phenomena which we witness within the cell. But among these latter phenomena there are others for which it is not so easy to suggest, in accordance with known laws, a simple mode of physical causation. It is not at once obvious how, in any simple system of symmetrical forces, {182} the chromosomes, which had at first been apparently repelled from the poles towards the equatorial plane, should then be split asunder, and should presently be attracted in opposite directions, some to one pole and some to the other. Remembering that it is not our purpose to assert that some one particular mode of action is at work, but merely to shew that there do exist physical forces, or distributions of force, which are capable of producing the required result, I give the following suggestive hypothesis, which I owe to my colleague Professor W. Peddie.

As we have begun by supposing that the nuclear, or chromosomal matter differs in permeability from the medium, that is to say the cytoplasm, in which it lies, let us now make the further assumption that its permeability is variable, and depends upon the strength of the field.

Fig. 54.

In Fig. 54, we have a field of force (representing our cell), consisting of a homogeneous medium, and including two opposite poles: lines of force are indicated by full lines, and loci of constant magnitude of force are shewn by dotted lines.

Let us now consider a body whose permeability (µ) depends on the strength of the field F. At two field-strengths, such as F_a, F_b, let the permeability of the body be equal to that of the {183} medium, and let the curved line in Fig. 55 represent generally its permeability at other field-strengths; and let the outer and inner dotted curves in Fig. 54 represent respectively the loci of the field-strengths F_b and F_a. The body if it be placed in the medium within either branch of the inner curve, or outside the outer curve, will tend to move into the neighbourhood of the adjacent pole. If it be placed in the region intermediate to the two dotted curves, it will tend to move towards regions of weaker field-strength.

Fig. 55.

The locus F_b is therefore a locus of stable position, towards which the body tends to move; the locus F_a is a locus of unstable position, from which it tends to move. If the body were placed across F_a, it might be torn asunder into two portions, the split coinciding with the locus F_a.

Suppose a number of such bodies to be scattered throughout the medium. Let at first the regions F_a and F_b be entirely outside the space where the bodies are situated: and, in making this supposition we may, if we please, suppose that the loci which we are calling F_a and F_b are meanwhile situated somewhat farther from the axis than in our figure, that (for instance) F_a is situated where we have drawn F_b, and that F_b is still further out. The bodies then tend towards the poles; but the tendency may be very small if, in Fig. 55, the curve and its intersecting straight line do not diverge very far from one another beyond F_a; in other {184} words, if, when situated in this region, the permeability of the bodies is not very much in excess of that of the medium.

Let the poles now tend to separate farther and farther from one another, the strength of each pole remaining unaltered; in other words, let the centrosome-foci recede from one another, as they actually do, drawing out the spindle-threads between them. The loci F_a, F_b, will close in to nearer relative distances from the poles. In doing so, when the locus F_a crosses one of the bodies, the body may be torn asunder; if the body be of elongated shape, and be crossed at more points than one, the forces at work will tend to exaggerate its foldings, and the tendency to rupture is greatest when F_a is in some median position (Fig. 56).

Fig. 56.

When the locus F_a has passed entirely over the body, the body tends to move towards regions of weaker force; but when, in turn, the locus F_b has crossed it, then the body again moves towards regions of stronger force, that is to say, towards the nearest pole. And, in thus moving towards the pole, it will do so, as appears actually to be the case in the dividing cell, along the course of the outer lines of force, the so-called “mantle-fibres” of the histologist236.

Such con­si­de­ra­tions as these give general results, easily open to modification in detail by a change of any of the arbitrary postulates which have been made for the sake of simplicity. Doubtless there are many other assumptions which would more or less meet the case; for instance, that of Ida H. Hyde that, {185} during the active phase of the chromatin molecule (during which it decomposes and sets free nucleic acid) it carries a charge opposite to that which it bears during its resting, or alkaline phase; and that it would accordingly move towards different poles under the influence of a current, wandering with its negative charge in an alkaline fluid during its acid phase to the anode, and to the kathode during its alkaline phase. A whole field of speculation is opened up when we begin to consider the cell not merely as a polarised electrical field, but also as an electrolytic field, full of wandering ions. Indeed it is high time we reminded ourselves that we have perhaps been dealing too much with ordinary physical analogies: and that our whole field of force within the cell is of an order of magnitude where these grosser analogies may fail to serve us, and might even play us false, or lead us astray. But our sole object meanwhile, as I have said more than once, is to demonstrate, by such illustrations as these, that, whatever be the actual and as yet unknown modus operandi, there are physical conditions and distributions of force which could produce just such phenomena of movement as we see taking place within the living cell. This, and no more, is precisely what Descartes is said to have claimed for his description of the human body as a “mechanism237.”

The foregoing account is based on the provisional assumption that the phenomena of caryokinesis are analogous to, if not identical with those of a bipolar electrical field; and this comparison, in my opinion, offers without doubt the best available series of analogies. But we must on no account omit to mention the fact that some of Leduc’s diffusion-experiments offer very remarkable analogies to the diagrammatic phenomena of caryokinesis, as shewn in the annexed figure238. Here we have two identical (not opposite) poles of osmotic concentration, formed by placing a drop of indian ink in salt water, and then on either side of this central drop, a hypertonic drop of salt solution more lightly coloured. On either side the pigment of the central drop has been drawn towards the focus nearest to it; but in the middle line, the pigment {186} is drawn in opposite directions by equal forces, and so tends to remain undisturbed, in the form of an “equatorial plate.”

Nor should we omit to take account (however briefly and inadequately) of a novel and elegant hypothesis put forward by A. B. Lamb. This hypothesis makes use of a theorem of Bjerknes, to the effect that synchronously vibrating or pulsating bodies in a liquid field attract or repel one another according as their oscillations are identical or opposite in phase. Under such circumstances, true currents, or hydrodynamic lines of force, are produced, identical in form with the lines of force of a magnetic field; and other particles floating, though not necessarily pulsating, in the liquid field, tend to be attracted or repelled by the pulsating bodies according as they are lighter or heavier than the surrounding fluid. Moreover (and this is the most remarkable point of all), the lines of force set up by the oppositely pulsating bodies are the same as those which are produced by opposite magnetic poles: though in the former case repulsion, and in the latter case attraction, takes place between the two poles239.

Fig. 57. Artificial caryokinesis (after Leduc), for comparison with Fig. 41, p. 169.

But to return to our general discussion.

While it can scarcely be too often repeated that our enquiry is not directed towards the solution of physiological problems, save {187} only in so far as they are inseparable from the problems presented by the visible con­fi­gur­a­tions of form and structure, and while we try, as far as possible, to evade the difficult question of what particular forces are at work when the mere visible forms produced are such as to leave this an open question, yet in this particular case we have been drawn into the use of electrical analogies, and we are bound to justify, if possible, our resort to this particular mode of physical action. There is an important paper by R. S. Lillie, on the “Electrical Convection of certain Free Cells and Nuclei240,” which, while I cannot quote it in direct support of the suggestions which I have made, yet gives just the evidence we need in order to shew that electrical forces act upon the constituents of the cell, and that their action discriminates between the two species of colloids represented by the cytoplasm and the nuclear chromatin. And the difference is such that, in the presence of an electrical current, the cell substance and the nuclei (including sperm-cells) tend to migrate, the former on the whole with the positive, the latter with the negative stream: a difference of electrical potential being thus indicated between the particle and the surrounding medium, just as in the case of minute suspended particles of various kinds in various feebly conducing media241. And the electrical difference is doubtless greatest, in the case of the cell constituents, just at the period of mitosis: when the chromatin is invariably in its most deeply staining, most strongly acid, and therefore, presumably, in its most electrically negative phase. In short, {188} Lillie comes easily to the conclusion that “electrical theories of mitosis are entitled to more careful consideration than they have hitherto received.”

Among other investigations, all leading towards the same general conclusion, namely that differences of electric potential play a great part in the phenomenon of cell division, I would mention a very noteworthy paper by Ida H. Hyde242, in which the writer shews (among other important observations) that not only is there a measurable difference of potential between the animal and vegetative poles of a fertilised egg (Fundulus, toad, turtle, etc.), but that this difference is not constant, but fluctuates, or actually reverses its direction, periodically, at epochs coinciding with successive acts of segmentation or other important phases in the development of the egg243; just as other physical rhythms, for instance in the production of CO₂ , had already been shewn to do. Hence we shall be by no means surprised to find that the “materialised” lines of force, which in the earlier stages form the convergent curves of the spindle, are replaced in the later phases of caryokinesis by divergent curves, indicating that the two foci, which are marked out within the field by the divided and reconstituted nuclei, are now alike in their polarity (Figs. 58, 59).

It is certain, to my mind, that these observations of Miss Hyde’s, and of Lillie’s, taken together with those of many writers on the behaviour of colloid particles generally in their relation to an electrical field, have a close bearing upon the physiological side of our problem, the full discussion of which lies outside our present field.

The break-up of the nucleus, already referred to and ascribed to a diminution of its surface-tension, is accompanied by certain diffusion phenomena which are sometimes visible to the eye; and we are reminded of Lord Kelvin’s view that diffusion is implicitly {189} associated with surface-tension changes, of which the first step is a minute puckering of the surface-skin, a sort of interdigitation with the surrounding medium. For instance, Schewiakoff has observed in Euglypha244 that, just before the break-up of the nucleus, a system of rays appears, concentred about it, but having nothing to do with the polar asters: and during the existence of this striation, the nucleus enlarges very considerably, evidently by imbibition of fluid from the surrounding protoplasm. In short, diffusion is at work, hand in hand with, and as it were in opposition to, the surface-tensions which define the nucleus. By diffusion, hand in hand with surface-tension, the alveoli of the nuclear meshwork are formed, enlarged, and finally ruptured: diffusion sets up the movements which give rise to the appearance of rays, or striae, around the nucleus: and through increasing diffusion, and weakening surface-tension, the rounded outline of the nucleus finally disappears. {190}

Fig. 58. Final stage in the first seg­men­ta­tion of the egg of Cere­brat­u­lus. (From Pre­nant, after Coe.)245

Fig. 59. Diagram of field of force with two similar poles.

As we study these manifold phenomena, in the individual cases of particular plants and animals, we recognise a close identity of type, coupled with almost endless variation of specific detail; and in particular, the order of succession in which certain of the phenomena occur is variable and irregular. The precise order of the phenomena, the time of longitudinal and of transverse fission of the chromatin thread, of the break-up of the nuclear wall, and so forth, will depend upon various minor contingencies and “interferences.” And it is worthy of particular note that these variations, in the order of events and in other subordinate details, while doubtless attributable to specific physical conditions, would seem to be without any obvious clas­si­fi­ca­tory value or other biological significance246.

As regards the actual mechanical division of the cell into two halves, we shall see presently that, in certain cases, such as that of a long cylindrical filament, surface-tension, and what is known as the principle of “minimal area,” go a long way to explain the mechanical process of division; and in all cells whatsoever, the process of division must somehow be explained as the result of a conflict between surface-tension and its opposing forces. But in such a case as our spherical cell, it is not very easy to see what physical cause is at work to disturb its equi­lib­rium and its integrity.

The fact that, when actual division of the cell takes place, it does so at right angles to the polar axis and precisely in the direction of the equatorial plane, would lead us to suspect that the new surface formed in the equatorial plane sets up an annular tension, directed inwards, where it meets the outer surface layer of the cell itself. But at this point, the problem becomes more complicated. Before we could hope to comprehend it, we should have not only to enquire into the potential distribution at the surface of the cell in relation to that which we have seen to exist in its interior, but we should probably also have to take account of the differences of potential which the material arrangements along the lines of force must themselves tend to produce. Only {191} thus could we approach a comprehension of the balance of forces which cohesion, friction, capillarity and electrical distribution combine to set up.

The manner in which we regard the phenomenon would seem to turn, in great measure, upon whether or no we are justified in assuming that, in the liquid surface-film of a minute spherical cell, local, and symmetrically localised, differences of surface-tension are likely to occur. If not, then changes in the conformation of the cell such as lead immediately to its division must be ascribed not to local changes in its surface-tension, but rather to direct changes in internal pressure, or to mechanical forces due to an induced surface-distribution of electrical potential.

It has seemed otherwise to many writers, and we have a number of theories of cell division which are all based directly on inequalities or asymmetry of surface-tension. For instance, Bütschli suggested, some forty years ago247, that cell division is brought about by an increase of surface-tension in the equatorial region of the cell. This explanation, however, can scarcely hold; for it would seem that such an increase of surface-tension in the equatorial plane would lead to the cell becoming flattened out into a disc, with a sharply curved equatorial edge, and to a streaming of material towards the equator. In 1895, Loeb shewed that the streaming went on from the equator towards the divided nuclei, and he supposed that the violence of these streaming movements brought about actual division of the cell: a hypothesis which was adopted by many other physiologists248. This streaming movement would suggest, as Robertson has pointed out, a diminution of surface-tension in the region of the equator. Now Quincke has shewn that the formation of soaps at the surface of an oil-droplet results in a diminution of the surface-tension of the latter; and that if the saponification be local, that part of the surface tends to spread. By laying a thread moistened with a dilute solution of caustic alkali, or even merely smeared with soap, across a drop of oil, Robertson has further shewn that the drop at once divides into two: the edges of the drop, that is to say the ends of the {192} diameter across which the thread lies, recede from the thread, so forming a notch at each end of the diameter, while violent streaming motions are set up at the surface, away from the thread in the direction of the two opposite poles. Robertson249 suggests, accordingly, that the division of the cell is actually brought about by a lowering of the equatorial surface-tension, and that this in turn is due to a chemical action, such as a liberation of cholin, or of soaps of cholin, through the splitting of lecithin in nuclear synthesis.

But purely chemical changes are not of necessity the fundamental cause of alteration in the surface-tension of the egg, for the action of electrolytes on surface-tension is now well known and easily demonstrated. So, according to other views than those with which we have been dealing, electrical charges are sufficient in themselves to account for alterations of surface-tension; while these in turn account for that protoplasmic streaming which, as so many investigators agree, initiates the segmentation of the egg250. A great part of our difficulty arises from the fact that in such a case as this the various phenomena are so entangled and apparently concurrent that it is hard to say which initiates another, and to which this or that secondary phenomenon may be considered due. Of recent years the phenomenon of adsorption has been adduced (as we have already briefly said) in order to account for many of the events and appearances which are associated with the asymmetry, and lead towards the division, of the cell. But our short discussion of this phenomenon may be reserved for another chapter.

However, we are not directly concerned here with the phenomena of segmentation or cell division in themselves, except only in so far as visible changes of form are capable of easy and obvious correlation with the play of force. The very fact of “development” indicates that, while it lasts, the equi­lib­rium of the egg is never complete251. And we may simply conclude the {193} matter by saying that, if you have caryokinetic figures developing inside the cell, that of itself indicates that the dynamic system and the localised forces arising from it are in continual alteration; and, consequently, changes in the outward configuration of the system are bound to take place.

As regards the phenomena of fertilisation,—of the union of the spermatozoon with the “pronucleus” of the egg,—we might study these also in illustration, up to a certain point, of the polarised forces which are manifestly at work. But we shall merely take, as a single illustration, the paths of the male and female pronuclei, as they travel to their ultimate meeting place.

The spermatozoon, when within a very short distance of the egg-cell, is attracted by it. Of the nature of this attractive force we have no certain knowledge, though we would seem to have a pregnant hint in Loeb’s discovery that, in the neighbourhood of other substances, such even as a fragment, or bead, of glass, the spermatozoon undergoes a similar attraction. But, whatever the force may be, it is one acting normally to the surface of the ovum, and accordingly, after entry, the sperm-nucleus points straight towards the centre of the egg; from the fact that other spermatozoa, subsequent to the first, fail to effect an entry, we may safely conclude that an immediate consequence of the entry of the spermatozoon is an increase in the surface-tension of the egg252. Somewhere or other, near or far away, within the egg, lies its own nuclear body, the so-called female pronucleus, and we find after a while that this has fused with the head of the spermatozoon (or male pronucleus), and that the body resulting from their fusion has come to occupy the centre of the egg. This must be due (as Whitman pointed out long ago) to a force of attraction acting between the two bodies, and another force acting upon one or other or both in the direction of the centre of the cell. Did we know the magnitude of these several forces, it would be a very easy task to calculate the precise path which the two pronuclei would follow, leading to conjugation and the central {194} position. As we do not know the magnitude, but only the direction, of these forces we can only make a general statement: (1) the paths of both moving bodies will lie wholly within a plane triangle drawn between the two bodies and the centre of the cell; (2) unless the two bodies happen to lie, to begin with, precisely on a diameter of the cell, their paths until they meet one another will be curved paths, the convexity of the curve being towards the straight line joining the two bodies; (3) the two bodies will meet a little before they reach the centre; and, having met and fused, will travel on to reach the centre in a straight line. The actual study and observation of the path followed is not very easy, owing to the fact that what we usually see is not the path itself, but only a projection of the path upon the plane of the microscope; but the curved path is particularly well seen in the frog’s egg, where the path of the spermatozoon is marked by a little streak of brown pigment, and the fact of the meeting of the pronuclei before reaching the centre has been repeatedly seen by many observers.

The problem is nothing else than a particular case of the famous problem of three bodies, which has so occupied the astronomers; and it is obvious that the foregoing brief description is very far from including all possible cases. Many of these are particularly described in the works of Fol, Roux, Whitman and others253.

The intracellular phenomena of which we have now spoken have assumed immense importance in biological literature and discussion during the last forty years; but it is open to us to doubt whether they will be found in the end to possess more than a remote and secondary biological significance. Most, if not all of them, would seem to follow immediately and inevitably from very simple assumptions as to the physical constitution of the cell, and from an extremely simple distribution of polarised forces within it. We have already seen that how a thing grows, and what it grows into, is a dynamic and not a merely material problem; so far as the material substance is concerned, it is so only by reason {195} of the chemical, electrical or other forces which are associated with it. But there is another consideration which would lead us to suspect that many features in the structure and configuration of the cell are of very secondary biological importance; and that is, the great variation to which these phenomena are subject in similar or closely related organisms, and the apparent impossibility of correlating them with the peculiarities of the organism as a whole. “Comparative study has shewn that almost every detail of the processes (of mitosis) described above is subject to variation in different forms of cells254.” A multitude of cells divide to the accompaniment of caryokinetic phenomena; but others do so without any visible caryokinesis at all. Sometimes the polarised field of force is within, sometimes it is adjacent to, and at other times it lies remote from the nucleus. The distribution of potential is very often symmetrical and bipolar, as in the case described; but a less symmetrical distribution often occurs, with the result that we have, for a time at least, numerous centres of force, instead of the two main correlated poles: this is the simple explanation of the numerous stellate figures, or “Strahlungen,” which have been described in certain eggs, such as those of Chaetopterus. In one and the same species of worm (Ascaris megalocephala), one group or two groups of chromosomes may be present. And remarkably constant, in general, as the number of chromosomes in any one species undoubtedly is, yet we must not forget that, in plants and animals alike, the whole range of observed numbers is but a small one; for (as regards the germ-nuclei) few organisms have less than six chromosomes, and fewer still have more than sixteen255. In closely related animals, such as various species of Copepods, and even in the same species of worm or insect, the form of the chromosomes, and their arrangement in relation to the nuclear spindle, have been found to differ in the various ways alluded to above. In short, there seem to be strong grounds for believing that these and many similar phenomena are in no way specifically related to the particular organism in which they have {196} been observed, and are not even specially and indisputably connected with the organism as such. They include such manifestations of the physical forces, in their various permutations and combinations, as may also be witnessed, under appropriate conditions, in non-living things.

When we attempt to separate our purely morphological or “purely embryological” studies from physiological and physical investigations, we tend ipso facto to regard each particular structure and configuration as an attribute, or a particular “character,” of this or that particular organism. From this assumption we are apt to go on to the drawing of new conclusions or the framing of new theories as to the ancestral history, the clas­si­fi­ca­tory position, the natural affinities of the several organisms: in fact, to apply our embryological knowledge mainly, and at times exclusively, to the study of phylogeny. When we find, as we are not long of finding, that our phylogenetic hypotheses, as drawn from embryology, become complex and unwieldy, we are nevertheless reluctant to admit that the whole method, with its fundamental postulates, is at fault. And yet nothing short of this would seem to be the case, in regard to the earlier phases at least of embryonic development. All the evidence at hand goes, as it seems to me, to shew that embryological data, prior to and even long after the epoch of segmentation, are essentially a subject for physiological and physical in­ves­ti­ga­tion and have but the very slightest link with the problems of systematic or zoological clas­si­fi­ca­tion. Comparative embryology has its own facts to classify, and its own methods and principles of clas­si­fi­ca­tion. Thus we may classify eggs according to the presence or absence, the paucity or abundance, of their associated food-yolk, the chromosomes according to their form and their number, the segmentation according to its various “types,” radial, bilateral, spiral, and so forth. But we have little right to expect, and in point of fact we shall very seldom and (as it were) only accidentally find, that these embryological categories coincide with the lines of “natural” or “phylogenetic” clas­si­fi­ca­tion which have been arrived at by the systematic zoologist.

The cell, which Goodsir spoke of as a “centre of force,” is in {197} reality a “sphere of action” of certain more or less localised forces; and of these, surface-tension is the particular force which is especially responsible for giving to the cell its outline and its morphological individuality. The partially segmented differs from the totally segmented egg, the unicellular Infusorian from the minute multicellular Turbellarian, in the intensity and the range of those surface-tensions which in the one case succeed and in the other fail to form a visible separation between the “cells.” Adam Sedgwick used to call attention to the fact that very often, even in eggs that appear to be totally segmented, it is yet impossible to discover an actual separation or cleavage, through and through between the cells which on the surface of the egg are so clearly delimited; so far and no farther have the physical forces effectuated a visible “cleavage.” The vacuolation of the protoplasm in Actinophrys or Actinosphaerium is due to localised surface-tensions, quite irrespective of the multinuclear nature of the latter organism. In short, the boundary walls due to surface-tension may be present or may be absent with or without the delimination of the other specific fields of force which are usually correlated with these boundaries and with the independent individuality of the cells. What we may safely admit, however, is that one effect of these circumscribed fields of force is usually such a separation or segregation of the protoplasmic constituents, the more fluid from the less fluid and so forth, as to give a field where surface-tension may do its work and bring a visible boundary into being. When the formation of a “surface” is once effected, its physical condition, or phase, will be bound to differ notably from that of the interior of the cell, and under appropriate chemical conditions the formation of an actual cell-wall, cellulose or other, is easily intelligible. To this subject we shall return again, in another chapter.

From the moment that we enter on a dynamical conception of the cell, we perceive that the old debates were in vain as to what visible portions of the cell were active or passive, living or non-living. For the manifestations of force can only be due to the interaction of the various parts, to the transference of energy from one to another. Certain properties may be manifested, certain functions may be carried on, by the protoplasm apart {198} from the nucleus; but the interaction of the two is necessary, that other and more important properties or functions may be manifested. We know, for instance, that portions of an Infusorian are incapable of regenerating lost parts in the absence of a nucleus, while nucleated pieces soon regain the specific form of the organism: and we are told that reproduction by fission cannot be initiated, though apparently all its later steps can be carried on, independently of nuclear action. Nor, as Verworn pointed out, can the nucleus possibly be regarded as the “sole vehicle of inheritance,” since only in the conjunction of cell and nucleus do we find the essentials of cell-life. “Kern und Protoplasma sind nur vereint lebensfähig,” as Nussbaum said. Indeed we may, with E. B. Wilson, go further, and say that “the terms ‘nucleus’ and ‘cell-body’ should probably be regarded as only topographical expressions denoting two differentiated areas in a common structural basis.”

Endless discussion has taken place regarding the centrosome, some holding that it is a specific and essential structure, a permanent corpuscle derived from a similar pre-existing corpuscle, a “fertilising element” in the spermatozoon, a special “organ of cell-division,” a material “dynamic centre” of the cell (as Van Beneden and Boveri call it); while on the other hand, it is pointed out that many cells live and multiply without any visible centrosomes, that a centrosome may disappear and be created anew, and even that under artificial conditions abnormal chemical stimuli may lead to the formation of new centrosomes. We may safely take it that the centrosome, or the “attraction sphere,” is essentially a “centre of force,” and that this dynamic centre may or may not be constituted by (but will be very apt to produce) a concrete and visible concentration of matter.

It is far from correct to say, as is often done, that the cell-wall, or cell-membrane, belongs “to the passive products of protoplasm rather than to the living cell itself”; or to say that in the animal cell, the cell-wall, because it is “slightly developed,” is relatively unimportant compared with the important role which it assumes in plants. On the contrary, it is quite certain that, whether visibly differentiated into a semi-permeable membrane, or merely constituted by a liquid film, the surface of the cell is the seat of {199} important forces, capillary and electrical, which play an essential part in the dynamics of the cell. Even in the thickened, largely solidified cellulose wall of the plant-cell, apart from the mechanical resistances which it affords, the osmotic forces developed in connection with it are of essential importance.

But if the cell acts, after this fashion, as a whole, each part interacting of necessity with the rest, the same is certainly true of the entire multicellular organism: as Schwann said of old, in very precise and adequate words, “the whole organism subsists only by means of the reciprocal action of the single elementary parts256.”

As Wilson says again, “the physiological autonomy of the individual cell falls into the background ... and the apparently composite character which the multicellular organism may exhibit is owing to a secondary distribution of its energies among local centres of action257.”

It is here that the homology breaks down which is so often drawn, and overdrawn, between the unicellular organism and the individual cell of the metazoon258.

Whitman, Adam Sedgwick259, and others have lost no opportunity of warning us against a too literal acceptation of the cell-theory, against the view that the multicellular organism is a colony (or, as Haeckel called it (in the case of the plant), a “republic”) of independent units of life260. As Goethe said long ago, “Das lebendige ist zwar in Elemente {200} zerlegt, aber man kann es aus diesen nicht wieder zusammenstellen und beleben;” the dictum of the Cellularpathologie being just the opposite, “Jedes Thier erscheint als eine Summe vitaler Einheiten, von denen jede den vollen Charakter des Lebens an sich trägt.”

Hofmeister and Sachs have taught us that in the plant the growth of the mass, the growth of the organ, is the primary fact, that “cell formation is a phenomenon very general in organic life, but still only of secondary significance.” “Comparative embryology” says Whitman, “reminds us at every turn that the organism dominates cell-formation, using for the same purpose one, several, or many cells, massing its material and directing its movements and shaping its organs, as if cells did not exist261.” So Rauber declared that, in the whole world of organisms, “das Ganze liefert die Theile, nicht die Theile das Ganze: letzteres setzt die Theile zusammen, nicht diese jenes262.” And on the botanical side De Bary has summed up the matter in an aphorism, “Die Pflanze bildet Zellen, nicht die Zelle bildet Pflanzen.”

Discussed almost wholly from the concrete, or morphological point of view, the question has for the most part been made to turn on whether actual protoplasmic continuity can be demonstrated between one cell and another, whether the organism be an actual reticulum, or syncytium. But from the dynamical point of view the question is much simpler. We then deal not with material continuity, not with little bridges of connecting protoplasm, but with a continuity of forces, a comprehensive field of force, which runs through and through the entire organism and is by no means restricted in its passage to a protoplasmic continuum. And such a continuous field of force, somehow shaping the whole organism, independently of the number, magnitude and form of the individual cells, which enter, like a froth, into its fabric, seems to me certainly and obviously to exist. As Whitman says, “the fact that physiological unity is not broken by cell-boundaries is confirmed in so many ways that it must be accepted as one of the fundamental truths of biology263.”

Protoplasm, as we have already said, is a fluid or rather a semifluid substance, and we need not pause here to attempt to describe the particular properties of the semifluid, colloid, or jelly-like substances to which it is allied; we should find it no easy matter. Nor need we appeal to precise theoretical definitions of fluidity, lest we come into a debateable land. It is in the most general sense that protoplasm is “fluid.” As Graham said (of colloid matter in general), “its softness partakes of fluidity, and enables the colloid to become a vehicle for liquid diffusion, like water itself264.” When we can deal with protoplasm in sufficient quantity we see it flow; particles move freely through it, air-bubbles and liquid droplets shew round or spherical within it; and we shall have much to say about other phenomena manifested by its own surface, which are those especially char­ac­ter­is­tic of liquids. It may encompass and contain solid bodies, and it may “secrete” within or around itself solid substances; and very often in the complex living organism these solid substances formed by the living protoplasm, like shell or nail or horn or feather, may remain when the protoplasm which formed them is dead and gone; but the protoplasm itself is fluid or semifluid, and accordingly permits of free (though not necessarily rapid) diffusion and easy convection of particles within itself. This simple fact is of elementary importance in connection with form, and with what appear at first sight to be common char­ac­teris­tics or peculiarities of the forms of living things.

The older naturalists, in discussing the differences between inorganic and organic bodies, laid stress upon the fact or statement that the former grow by “agglutination,” and the latter by {202} what they termed “intussusception.” The contrast is true, rather, of solid as compared with jelly-like bodies of all kinds, living or dead, the great majority of which as it so happens, but by no means all, are of organic origin.

A crystal “grows” by deposition of new molecules, one by one and layer by layer, superimposed or aggregated upon the solid substratum already formed. Each particle would seem to be influenced, practically speaking, only by the particles in its immediate neighbourhood, and to be in a state of freedom and independence from the influence, either direct or indirect, of its remoter neighbours. As Lord Kelvin and others have explained the formation and the resulting forms of crystals, so we believe that each added particle takes up its position in relation to its immediate neighbours already arranged, generally in the holes and corners that their arrangement leaves, and in closest contact with the greatest number265. And hence we may repeat or imitate this process of arrangement, with great or apparently even with precise accuracy (in the case of the simpler crystalline systems), by piling up spherical pills or grains of shot. In so doing, we must have regard to the fact that each particle must drop into the place where it can go most easily, or where no easier place offers. In more technical language, each particle is free to take up, and does take up, its position of least potential energy relative to those already deposited; in other words, for each particle motion is induced until the energy of the system is so distributed that no tendency or resultant force remains to move it more. The application of this principle has been shewn to lead to the production of planes266 (in all cases where by the limitation of material, surfaces must occur); and where we have planes, straight edges and solid angles must obviously also occur; and, if equi­lib­rium is {203} to follow, must occur symmetrically. Our piling up of shot, or manufacture of mimic crystals, gives us visible demonstration that the result is actually to obtain, as in the natural crystal, plane surfaces and sharp angles, symmetrically disposed.

But the living cell grows in a totally different way, very much as a piece of glue swells up in water, by “imbibition,” or by interpenetration into and throughout its entire substance. The semifluid colloid mass takes up water, partly to combine chemically with its individual molecules267, partly by physical diffusion into the interstices between these molecules, and partly, as it would seem, in other ways; so that the entire phenomenon is a very complex and even an obscure one. But, so far as we are concerned, the net result is a very simple one. For the equi­lib­rium or tendency to equi­lib­rium of fluid pressure in all parts of its interior while the process of imbibition is going on, the constant rearrangement of its fluid mass, the contrast in short with the crystalline method of growth where each particle comes to rest to move (relatively to the whole) no more, lead the mass of jelly to swell up, very much as a bladder into which we blow air, and so, by a graded and harmonious distribution of forces, to assume everywhere a rounded and more or less bubble-like external form268. So, when the same school of older naturalists called attention to a new distinction or contrast of form between the organic and inorganic objects, in that the contours of the former tended to roundness and curvature, and those of the latter to be bounded by straight lines, planes and sharp angles, we see that this contrast was not a new and different one, but only another aspect of their former statement, and an immediate consequence of the difference between the processes of agglutination and intussusception.

This common and general contrast between the form of the crystal on the one hand, and of the colloid or of the organism on the other, must by no means be pressed too far. For Lehmann, {204} in his great work on so-called Fluid Crystals269, to which we shall afterwards return, has shewn how, under certain circumstances, surface-tension phenomena may coexist with cry­stal­li­sa­tion, and produce a form of minimal potential which is a resultant of both: the fact being that the bonds maintaining the crystalline arrangement are now so much looser than in the solid condition that the tendency to least total surface-area is capable of being satisfied. Thus the phenomenon of “liquid cry­stal­li­sa­tion” does not destroy the distinction between crystalline and colloidal forms, but gives added unity and continuity to the whole series of phenomena270. Lehmann has also demonstrated phenomena within the crystal, known for instance as transcry­stal­li­sa­tion, which shew us that we must not speak unguardedly of the growth of crystals as limited to deposition upon a surface, and Bütschli has already pointed out the possible great importance to the biologist of the various phenomena which Lehmann has described271.

So far then, as growth goes on, unaffected by pressure or other external force, the fluidity of protoplasm, its mobility internal and external, and the manner in which particles move with comparative freedom from place to place within, all manifestly tend to the production of swelling, rounded surfaces, and to their great predominance over plane surfaces in the contour of the organism. These rounded contours will tend to be preserved, for a while, in the case of naked protoplasm by its viscosity, and in the presence of a cell-wall by its very lack of fluidity. In a general way, the presence of curved boundary surfaces will be especially obvious in the unicellular organisms, and still more generally in the external forms of all organisms; and wherever mutual pressure between adjacent cells, or other adjacent parts, has not come into play to flatten the rounded surfaces into planes.

But the rounded contours that are assumed and exhibited by {205} a piece of hard glue, when we throw it into water and see it expand as it sucks the water up, are not nearly so regular or so beautiful as are those which appear when we blow a bubble, or form a drop, or pour water into a more or less elastic bag. For these curving contours depend upon the properties of the bag itself, of the film or membrane that contains the mobile gas, or that contains or bounds the mobile liquid mass. And hereby, in the case of the fluid or semifluid mass, we are introduced to the subject of surface tension: of which indeed we have spoken in the preceding chapter, but which we must now examine with greater care.

Among the forces which determine the forms of cells, whether they be solitary or arranged in contact with one another, this force of surface-tension is certainly of great, and is probably of paramount importance. But while we shall try to separate out the phenomena which are directly due to it, we must not forget that, in each particular case, the actual conformation which we study may be, and usually is, the more or less complex resultant of surface tension acting together with gravity, mechanical pressure, osmosis, or other physical forces.

Surface tension is that force by which we explain the form of a drop or of a bubble, of the surfaces external and internal of a “froth” or collocation of bubbles, and of many other things of like nature and in like circumstances272. It is a property of liquids (in the sense at least with which our subject is concerned), and it is manifested at or very near the surface, where the liquid comes into contact with another liquid, a solid or a gas. We note here that the term surface is to be interpreted in a wide sense; for wherever we have solid particles imbedded in a fluid, wherever we have a non-homogeneous fluid or semi-fluid such as a particle {206} of protoplasm, wherever we have the presence of “impurities,” as in a mass of molten metal, there we have always to bear in mind the existence of “surfaces” and of surface tensions, not only on the exterior of the mass but also throughout its interstices, wherever like meets unlike.

Surface tension is due to molecular force, to force that is to say arising from the action of one molecule upon another, and it is accordingly exerted throughout a small thickness of material, comparable to the range of the molecular forces. We imagine that within the interior of the liquid mass such molecular interactions negative one another: but that at and near the free surface, within a layer or film ap­prox­i­mate­ly equal to the range of the molecular force, there must be a lack of such equi­lib­rium and consequently a manifestation of force.

The action of the molecular forces has been variously explained. But one simple explanation (or mode of statement) is that the molecules of the surface layer (whose thickness is definite and constant) are being constantly attracted into the interior by those which are more deeply situated, and that consequently, as molecules keep quitting the surface for the interior, the bulk of the latter increases while the surface diminishes; and the process continues till the surface itself has become a minimum, the surface-shrinkage exhibiting itself as a surface-tension. This is a sufficient description of the phenomenon in cases where a portion of liquid is subject to no other than its own molecular forces, and (since the sphere has, of all solids, the smallest surface for a given volume) it accounts for the spherical form of the raindrop, of the grain of shot, or of the living cell in many simple organisms. It accounts also, as we shall presently see, for a great number of much more complicated forms, manifested under less simple conditions.

Let us here briefly note that surface tension is, in itself, a comparatively small force, and easily measurable: for instance that of water is equivalent to but a few grains per linear inch, or a few grammes per metre. But this small tension, when it exists in a curved surface of very great curvature, gives rise to a very great pressure directed towards the centre of curvature. We can easily calculate this pressure, and so satisfy ourselves that, when the radius of curvature is of molecular dimensions, the {207} pressure is of the magnitude of thousands of atmospheres,—a conclusion which is supported by other physical con­si­de­ra­tions.

The contraction of a liquid surface and other phenomena of surface tension involve the doing of work, and the power to do work is what we call energy. It is obvious, in such a simple case as we have just considered, that the whole energy of the system is diffused throughout its molecules; but of this whole stock of energy it is only that part which comes into play at or very near to the surface which normally manifests itself in work, and hence we may speak (though the term is open to some objections) of a specific surface energy. The consideration of surface energy, and of the manner in which its amount is increased and multiplied by the multiplication of surfaces due to the subdivision of the organism into cells, is of the highest importance to the physiologist; and even the morphologist cannot wholly pass it by, if he desires to study the form of the cell in its relation to the phenomena of surface tension or “capillarity.” The case has been set forth with the utmost possible lucidity by Tait and by Clerk Maxwell, on whose teaching the following paragraphs are based: they having based their teaching upon that of Gauss,—who rested on Laplace.

Let E be the whole potential energy of a mass M of liquid; let e₀ be the energy per unit mass of the interior liquid (we may call it the internal energy); and let e be the energy per unit mass for a layer of the skin, of surface S, of thickness t, and density ρ (e being what we call the surface energy). It is obvious that the total energy consists of the internal plus the surface energy, and that the former is distributed through the whole mass, minus its surface layers. That is to say, in math­e­mat­i­cal language,

and this is as much as to say that the total energy of the system may be taken to consist of two portions, one uniform throughout the whole mass, and another, which is proportional on the one hand to the amount of surface, and on the other hand is proportional to the difference between e and e₀ , that is to say to the difference between the unit values of the internal and the surface energy. {208}

It was Gauss who first shewed after this fashion how, from the mutual attractions between all the particles, we are led to an expression which is what we now call the potential energy of the system; and we know, as a fundamental theorem of dynamics, that the potential energy of the system tends to a minimum, and in that minimum finds, as a matter of course, its stable equi­lib­rium.

We see in our last equation that the term M e₀ is irreducible, save by a reduction of the mass itself. But the other term may be diminished (1) by a reduction in the area of surface, S, or (2) by a tendency towards equality of e and e₀ , that is to say by a diminution of the specific surface energy, e.

These then are the two methods by which the energy of the system will manifest itself in work. The one, which is much the more important for our purposes, leads always to a diminution of surface, to the so-called “principle of minimal areas”; the other, which leads to the lowering (under certain circumstances) of surface tension, is the basis of the theory of Adsorption, to which we shall have some occasion to refer as the modus operandi in the development of a cell-wall, and in a variety of other histological phenomena. In the technical phraseology of the day, the “capacity factor” is involved in the one case, and the “intensity factor” in the other.

Inasmuch as we are concerned with the form of the cell it is the former which becomes our main postulate: telling us that the energy equations of the surface of a cell, or of the free surfaces of cells partly in contact, or of the partition-surfaces of cells in contact with one another or with an adjacent solid, all indicate a minimum of potential energy in the system, by which the system is brought, ipso facto, into equi­lib­rium. And we shall not fail to observe, with something more than mere historical interest and curiosity, how deeply and intrinsically there enter into this whole class of problems the “principle of least action” of Maupertuis, the “lineae curvae maximi minimive proprietate gaudentes” of Euler, by which principles these old natural philosophers explained correctly a multitude of phenomena, and drew the lines whereon the foundations of great part of modern physics are well and truly laid. {209}

In all cases where the principle of maxima and minima comes into play, as it conspicuously does in the systems of liquid films which are governed by the laws of surface-tension, the figures and conformations produced are characterised by obvious and remarkable symmetry. Such symmetry is in a high degree char­ac­ter­is­tic of organic forms, and is rarely absent in living things,—save in such cases as amoeba, where the equi­lib­rium on which symmetry depends is likewise lacking. And if we ask what physical equi­lib­rium has to do with formal symmetry and regularity, the reason is not far to seek; nor can it be put better than in the following words of Mach’s273. “In every symmetrical system every deformation that tends to destroy the symmetry is complemented by an equal and opposite deformation that tends to restore it. In each deformation positive and negative work is done. One condition, therefore, though not an absolutely sufficient one, that a maximum or minimum of work corresponds to the form of equi­lib­rium, is thus supplied by symmetry. Regularity is successive symmetry. There is no reason, therefore, to be astonished that the forms of equi­lib­rium are often symmetrical and regular.”

As we proceed in our enquiry, and especially when we approach the subject of tissues, or agglomerations of cells, we shall have from time to time to call in the help of elementary mathematics. But already, with very little math­e­mat­i­cal help, we find ourselves in a position to deal with some simple examples of organic forms.

When we melt a stick of sealing-wax in the flame, surface tension (which was ineffectively present in the solid but finds play in the now fluid mass), rounds off its sharp edges into curves, so striving towards a surface of minimal area; and in like manner, by melting the tip of a thin rod of glass, Leeuwenhoek made the little spherical beads which served him for a microscope274. When any drop of protoplasm, either over all its surface or at some free end, as at the extremity of the pseudopodium of an amoeba, is {210} seen likewise to “round itself off,” that is not an effect of “vital contractility,” but (as Hofmeister shewed so long ago as 1867) a simple consequence of surface tension; and almost immediately afterwards Engelmann275 argued on the same lines, that the forces which cause the contraction of protoplasm in general may “be just the same as those which tend to make every non-spherical drop of fluid become spherical!” We are not concerned here with the many theories and speculations which would connect the phenomena of surface tension with contractility, muscular movement or other special physiological functions, but we find ample room to trace the operation of the same cause in producing, under conditions of rest and equi­lib­rium, certain definite and inevitable forms of surface.

It is however of great importance to observe that the living cell is one of those cases where the phenomena of surface tension are by no means limited to the outer surface; for within the heterogeneous substance of the cell, between the protoplasm and its nuclear and other contents, and in the alveolar network of the cytoplasm itself (so far as that “alveolar structure” is actually present in life), we have a multitude of interior surfaces; and, especially among plants, we may have a large inner surface of “interfacial” contact, where the protoplasm contains cavities or “vacuoles” filled with a different and more fluid material, the “cell-sap.” Here we have a great field for the development of surface tension phenomena: and so long ago as 1865, Nägeli and Schwendener shewed that the streaming currents of plant cells might be very plausibly explained by this phenomenon. Even ten years earlier, Weber had remarked upon the resemblance between these protoplasmic streamings and the streamings to be observed in certain inanimate drops, for which no cause but surface tension could be assigned276.

The case of amoeba, though it is an elementary case, is at the same time a complicated one. While it remains “amoeboid,” it is never at rest or in equi­lib­rium; it is always moving, from one to another of its protean changes of configuration; its surface tension is constantly varying from point to point. Where the {211} surface tension is greater, that portion of the surface will contract into spherical or spheroidal forms; where it is less the surface will correspondingly extend. While generally speaking the surface energy has a minimal value, it is not necessarily constant. It may be diminished by a rise of temperature; it may be altered by contact with adjacent substances277, by the transport of constituent materials from the interior to the surface, or again by actual chemical and fermentative change. Within the cell, the surface energies developed about its heterogeneous contents will constantly vary as these contents are affected by chemical metabolism. As the colloid materials are broken down and as the particles in suspension are diminished in size the “free surface energy” will be increased, but the osmotic energy will be diminished278. Thus arise the various fluctuations of surface tension and the various phenomena of amoeboid form and motion, which Bütschli and others have reproduced or imitated by means of the fine emulsions which constitute their “artificial amoebae.” A multitude of experiments shew how extraordinarily delicate is the adjustment of the surface tension forces, and how sensitive they are to the least change of temperature or chemical state. Thus, on a plate which we have warmed at one side, a drop of alcohol runs towards the warm area, a drop of oil away from it; and a drop of water on the glass plate exhibits lively movements when {212} we bring into its neighbourhood a heated wire, or a glass rod dipped in ether. When we find that a plasmodium of Aethalium, for instance, creeps towards a damp spot, or towards a warm spot, or towards substances that happen to be nutritious, and again creeps away from solutions of sugar or of salt, we seem to be dealing with phenomena every one of which can be paralleled by ordinary phenomena of surface tension279. Even the soap-bubble itself is imperfectly in equi­lib­rium, for the reason that its film, like the protoplasm of amoeba or Aethalium, is an excessively heterogeneous substance. Its surface tensions vary from point to point, and chemical changes and changes of temperature increase and magnify the variation. The whole surface of the bubble is in constant movement as the concentrated portions of the soapy fluid make their way outwards from the deeper layers; it thins and it thickens, its colours change, currents are set up in it, and little bubbles glide over it; it continues in this state of constant movement, as its parts strive one with another in all their interactions towards equi­lib­rium280.

In the case of the naked protoplasmic cell, as the amoeboid phase is emphatically a phase of freedom and activity, of chemical and physiological change, so, on the other hand, is the spherical form indicative of a phase of rest or comparative inactivity. In the one phase we see unequal surface tensions manifested in the creeping movements of the amoeboid body, in the rounding off of the ends of the pseudopodia, in the flowing out of its substance over a particle of “food,” and in the current-motions in the interior of its mass; till finally, in the other phase, when internal homogeneity and equi­lib­rium have been attained and the potential {213} energy of the system is for the time being at a minimum, the cell assumes a rounded or spherical form, passing into a state of “rest,” and (for a reason which we shall presently see) becoming at the same time “encysted.”

Fig. 60.

In a budding yeast-cell (Fig. 60), we see a more definite and restricted change of surface tension. When a “bud” appears, whether with or without actual growth by osmosis or otherwise of the mass, it does so because at a certain part of the cell-surface the surface tension has more or less suddenly diminished, and the area of that portion expands accordingly; but in turn the surface tension of the expanded area will make itself felt, and the bud will be rounded off into a more or less spherical form.

The yeast-cell with its bud is a simple example of a principle which we shall find to be very important. Our whole treatment of cell-form in relation to surface-tension depends on the fact (which Errera was the first to point out, or to give clear expression to) that the incipient cell-wall retains with but little impairment the properties of a liquid film281, and that the growing cell, in spite of the membrane by which it has already begun to be surrounded, behaves very much like a fluid drop. But even the ordinary yeast-cell shows, by its ovoid and non-spherical form, that it has acquired its shape under the influence of some force other than that uniform and symmetrical surface-tension which would be productive of a sphere; and this or any other asymmetrical form, once acquired, may be retained by virtue of the solidification and consequent rigidity of the membranous wall of the cell. Unless such rigidity ensue, it is plain that such a conformation as that of the cell with its attached bud could not be long retained, amidst the constantly varying conditions, as a figure of even partial equi­lib­rium. But as a matter of fact, the cell in this case is not in equi­lib­rium at all; it is in process of budding, and is slowly altering its shape by rounding off the bud. It is plain that over its surface the surface-energies are unequally distributed, owing to some heterogeneity of the substance; and to this matter we shall afterwards return. In like manner the developing egg {214} through all its successive phases of form is never in complete equi­lib­rium; but is merely responding to constantly changing conditions, by phases of partial, transitory, unstable and conditional equi­lib­rium.

It is obvious that there are innumerable solitary plant-cells, and unicellular organisms in general, which, like the yeast-cell, do not correspond to any of the simple forms that may be generated under the influence of simple and homogeneous surface-tension; and in many cases these forms, which we should expect to be unstable and transitory, have become fixed and stable by reason of the comparatively sudden or rapid solidification of the envelope. This is the case, for instance, in many of the more complicated forms of diatoms or of desmids, where we are dealing, in a less striking but even more curious way than in the budding yeast-cell, not with one simple act of formation, but with a complicated result of successive stages of localised growth, interrupted by phases of partial consolidation. The original cell has acquired or assumed a certain form, and then, under altering conditions and new distributions of energy, has thickened here or weakened there, and has grown out or tended (as it were) to branch, at particular points. We can often, or indeed generally, trace in each particular stage of growth or at each particular temporary growing point, the laws of surface tension manifesting themselves in what is for the time being a fluid surface; nay more, even in the adult and completed structure, we have little difficulty in tracing and recognising (for instance in the outline of such a desmid as Euastrum) the rounded lobes that have successively grown or flowed out from the original rounded and flattened cell. What we see in a many chambered foraminifer, such as Globigerina or Rotalia, is just the same thing, save that it is carried out in greater completeness and perfection. The little organism as a whole is not a figure of equi­lib­rium or of minimal area; but each new bud or separate chamber is such a figure, conditioned by the forces of surface tension, and superposed upon the complex aggregate of similar bubbles after these latter have become consolidated one by one into a rigid system.

Let us now make some enquiry regarding the various forms {215} which, under the influence of surface tension, a surface can possibly assume. In doing so, we are obviously limited to conditions under which other forces are relatively unimportant, that is to say where the “surface energy” is a considerable fraction of the whole energy of the system; and this in general will be the case when we are dealing with portions of liquid so small that their dimensions come within what we have called the molecular range, or, more generally, in which the “specific surface” is large282: in other words it will be small or minute organisms, or the small cellular elements of larger organisms, whose forms will be governed by surface-tension; while the general forms of the larger organisms will be due to other and non-molecular forces. For instance, a large surface of water sets itself level because here gravity is predominant; but the surface of water in a narrow tube is manifestly curved, for the reason that we are here dealing with particles which are mutually within the range of each other’s molecular forces. The same is the case with the cell-surfaces and cell-partitions which we are presently to study, and the effect of gravity will be especially counteracted and concealed when, as in the case of protoplasm in a watery fluid, the object is immersed in a liquid of nearly its own specific gravity.

We have already learned, as a fundamental law of surface-tension phenomena, that a liquid film in equi­lib­rium assumes a form which gives it a minimal area under the conditions to which it is subject. And these conditions include (1) the form of the boundary, if such exist, and (2) the pressure, if any, to which the film is subject; which pressure is closely related to the volume, of air or of liquid, which the film (if it be a closed one) may have to contain. In the simplest of cases, when we take up a soap-film on a plane wire ring, the film is exposed to equal atmospheric pressure on both sides, and it obviously has its minimal area in the form of a plane. So long as our wire ring lies in one plane (however irregular in outline), the film stretched across it will still be in a plane; but if we bend the ring so that it lies no longer in a plane, then our film will become curved into a surface which may be extremely complicated, but is still the smallest possible {216} surface which can be drawn continuously across the uneven boundary.

The question of pressure involves not only external pressures acting on the film, but also that which the film itself is capable of exerting. For we have seen that the film is always contracting to its smallest limits; and when the film is curved, this obviously leads to a pressure directed inwards,—perpendicular, that is to say, to the surface of the film. In the case of the soap-bubble, the uniform contraction of whose surface has led to its spherical form, this pressure is balanced by the pressure of the air within; and if an outlet be given for this air, then the bubble contracts with perceptible force until it stretches across the mouth of the tube, for instance the mouth of the pipe through which we have blown the bubble. A precisely similar pressure, directed inwards, is exercised by the surface layer of a drop of water or a globule of mercury, or by the surface pellicle on a portion or “drop” of protoplasm. Only we must always remember that in the soap-bubble, or the bubble which a glass-blower blows, there is a twofold pressure as compared with that which the surface-film exercises on the drop of liquid of which it is a part; for the bubble consists (unless it be so thin as to consist of a mere layer of molecules283) of a liquid layer, with a free surface within and another without, and each of these two surfaces exercises its own independent and coequal tension, and cor­re­spon­ding pressure284.

If we stretch a tape upon a flat table, whatever be the tension of the tape it obviously exercises no pressure upon the table below. But if we stretch it over a curved surface, a cylinder for instance, it does exercise a downward pressure; and the more curved the surface the greater is this pressure, that is to say the greater is this share of the entire force of tension which is resolved in the downward direction. In math­e­mat­i­cal language, the pressure (p) varies directly as the tension (T), and inversely as the radius of curvature (R): that is to say, p = T ⁄ R, per unit of surface. {217}

If instead of a cylinder, which is curved only in one direction, we take a case where there are curvatures in two dimensions (as for instance a sphere), then the effects of these must be simply added to one another, and the resulting pressure p is equal to T ⁄ R + T ⁄ R′ or p = T(1 ⁄ R + 1 ⁄ R′)*.

And if in addition to the pressure p, which is due to surface tension, we have to take into account other pressures, p′, p″, etc., which are due to gravity or other forces, then we may say that the total pressure, P = p′ + p″ + T(1 ⁄ R + 1 ⁄ R′). While in some cases, for instance in speaking of the shape of a bird’s egg, we shall have to take account of these extraneous pressures, in the present part of our subject we shall for the most part be able to neglect them.

Our equation is an equation of equi­lib­rium. The resistance to compression,—the pressure outwards,—of our fluid mass, is a constant quantity (P); the pressure inwards, T(1 ⁄ R + 1 ⁄ R′), is also constant; and if (unlike the case of the mobile amoeba) the surface be homogeneous, so that T is everywhere equal, it follows that throughout the whole surface 1 ⁄ R + 1 ⁄ R′ = C (a constant).

Now equi­lib­rium is attained after the surface contraction has done its utmost, that is to say when it has reduced the surface to the smallest possible area; and so we arrive, from the physical side, at the conclusion that a surface such that 1 ⁄ R + 1 ⁄ R′ = C, in other words a surface which has the same mean curvature at all points, is equivalent to a surface of minimal area: and to the same conclusion we may also arrive through purely analytical mathematics. It is obvious that the plane and the sphere are two examples of such surfaces, for in both cases the radius of curvature is everywhere constant, being equal to infinity in the case of the plane, and to some definite magnitude in the case of the sphere.

From the fact that we may extend a soap-film across a ring of wire however fantastically the latter may be bent, we realise that there is no limit to the number of surfaces of minimal area which may be constructed or may be imagined; and while some of these are very complicated indeed, some, for instance a spiral helicoid screw, are relatively very simple. But if we limit ourselves to {218} surfaces of revolution (that is to say, to surfaces symmetrical about an axis), we find, as Plateau was the first to shew, that those which meet the case are very few in number. They are six in all, namely the plane, the sphere, the cylinder, the catenoid, the unduloid, and a curious surface which Plateau called the nodoid.

These several surfaces are all closely related, and the passage from one to another is generally easy. Their math­e­mat­i­cal interrelation is expressed by the fact (first shewn by Delaunay286, in 1841) that the plane curves by whose rotation they are generated are themselves generated as “roulettes” of the conic sections.

Let us imagine a straight line upon which a circle, an ellipse or other conic section rolls; the focus of the conic section will describe a line in some relation to the fixed axis, and this line (or roulette), rotating around the axis, will describe in space one or other of the six surfaces of revolution with which we are dealing.

Fig. 61.

If we imagine an ellipse so to roll over a line, either of its foci will describe a sinuous or wavy line (Fig. 61B) at a distance alternately maximal and minimal from the axis; and this wavy line, by rotation about the axis, becomes the meridional line of the surface which we call the unduloid. The more unequal the two axes are of our ellipse, the more pronounced will be the sinuosity of the described roulette. If the two axes be equal, then our ellipse becomes a circle, and the path described by its rolling centre is a straight line parallel to the axis (A); and obviously the solid of revolution generated therefrom will be a cylinder. If one axis of our ellipse vanish, while the other remain of finite length, then the ellipse is reduced to a straight line, and its roulette will appear as a succession of semicircles touching one another upon the axis (C); the solid of revolution will be a series of equal spheres. If as before one axis of the ellipse vanish, but the other be infinitely long, then the curve described by the rotation {219} of this latter will be a circle of infinite radius, i.e. a straight line infinitely distant from the axis; and the surface of rotation is now a plane. If we imagine one focus of our ellipse to remain at a given distance from the axis, but the other to become infinitely remote, that is tantamount to saying that the ellipse becomes transformed into a parabola; and by the rolling of this curve along the axis there is described a catenary (D), whose solid of revolution is the catenoid.

Lastly, but this is a little more difficult to imagine, we have the case of the hyperbola.

We cannot well imagine the hyperbola rolling upon a fixed straight line so that its focus shall describe a continuous curve. But let us suppose that the fixed line is, to begin with, asymptotic to one branch of the hyperbola, and that the rolling proceed until the line is now asymptotic to the other branch, that is to say touching it at an infinite distance; there will then be math­e­mat­i­cal continuity if we recommence rolling with this second branch, and so in turn with the other, when each has run its course. We shall see, on reflection, that the line traced by one and the same focus will be an “elastic curve” describing a succession of kinks or knots (E), and the solid of revolution described by this meridional line about the axis is the so-called nodoid.

The physical transition of one of these surfaces into another can be experimentally illustrated by means of soap-bubbles, or better still, after the method of Plateau, by means of a large globule of oil, supported when necessary by wire rings, within a fluid of specific gravity equal to its own.

To prepare a mixture of alcohol and water of a density precisely equal to that of the oil-globule is a troublesome matter, and a method devised by Mr C. R. Darling is a great improvement on Plateau’s287. Mr Darling uses the oily liquid orthotoluidene, which does not mix with water, has a beautiful and conspicuous red colour, and has precisely the same density as water when both are kept at a temperature of 24° C. We have therefore only to run the liquid into water at this temperature in order to produce beautifully spherical drops of any required size: and by adding {220} a little salt to the lower layers of water, the drop may be made to float or rest upon the denser liquid.

We have already seen that the soap-bubble, spherical to begin with, is transformed into a plane when we relieve its internal pressure and let the film shrink back upon the orifice of the pipe. If we blow a small bubble and then catch it up on a second pipe, so that it stretches between, we may gradually draw the two pipes apart, with the result that the spheroidal surface will be gradually flattened in a longitudinal direction, and the bubble will be transformed into a cylinder. But if we draw the pipes yet farther apart, the cylinder will narrow in the middle into a sort of hourglass form, the increasing curvature of its transverse section being balanced by a gradually increasing negative curvature in the longitudinal section. The cylinder has, in turn, been converted into an unduloid. When we hold a portion of a soft glass tube in the flame, and “draw it out,” we are in the same identical fashion converting a cylinder into an unduloid (Fig. 62A); when on the other hand we stop the end and blow, we again convert the cylinder into an unduloid (B), but into one which is now positively, while the former was negatively curved. The two figures are essentially the same, save that the two halves of the one are reversed in the other.

Fig. 62.

That spheres, cylinders and unduloids are of the commonest occurrence among the forms of small unicellular organism, or of individual cells in the simpler aggregates, and that in the processes of growth, reproduction and development transitions are frequent from one of these forms to another, is obvious to the naturalist, and we shall deal presently with a few illustrations of these phenomena.

But before we go further in this enquiry, it will be necessary to consider, to some small extent at least, the curvatures of the six different surfaces, that is to say, to determine what modification {221} is required, in each case, of the general equation which applies to them all. We shall find that with this question is closely connected the question of the pressures exercised by, or impinging on the film, and also the very important question of the limitations which, from the nature of the case, exist to prevent the extension of certain of the figures beyond certain bounds. The whole subject is math­e­mat­i­cal, and we shall only deal with it in the most elementary way.

We have seen that, in our general formula, the expression 1 ⁄ R + 1 ⁄ R′ = C, a constant; and that this is, in all cases, the condition of our surface being one of minimal area. In other words, it is always true for one and all of the six surfaces which we have to consider. But the constant C may have any value, positive, negative, or nil.

In the case of the plane, where R and R′ are both infinite, it is obvious that 1 ⁄ R + 1 ⁄ R′ = 0. The expression therefore vanishes, and our dynamical equation of equi­lib­rium becomes P = p. In short, we can only have a plane film, or we shall only find a plane surface in our cell, when on either side thereof we have equal pressures or no pressure at all. A simple case is the plane partition between two equal and similar cells, as in a filament of spirogyra.

In the case of the sphere, the radii are all equal, R = R′; they are also positive, and T (1 ⁄ R + 1 ⁄ R′), or 2 T ⁄ R, is a positive quantity, involving a positive pressure P, on the other side of the equation.

In the cylinder, one radius of curvature has the finite and positive value R; but the other is infinite. Our formula becomes T ⁄ R, to which corresponds a positive pressure P, supplied by the surface-tension as in the case of the sphere, but evidently of just half the magnitude developed in the latter case for a given value of the radius R.

In the unduloid, unlike the four surfaces which we have just been considering, it is obvious that the curvatures change from one point to another. At the middle of one of the swollen portions, or “beads,” the two curvatures are both positive; the expression (1 ⁄ R + 1 ⁄ R′) is therefore positive, and it is also finite. The film, accordingly, exercises a positive tension inwards, which must be compensated by a finite and positive outward pressure P. At the middle of one of the narrow necks, between two adjacent beads, there is obviously, in the transverse direction, a much stronger curvature than in the former case, and the curvature which balances it is now a negative one. But the sum of the two must remain positive, as well as constant; and we therefore see that the convex or positive curvature must always be greater than the concave or negative curvature at the same point. This is plainly the case in our figure of the unduloid.

The nodoid is, like the unduloid, a continuous curve which keeps altering its curvature as it alters its distance from the axis; but in this case the resultant pressure inwards is negative instead of positive. But this curve is a complicated one, and a full discussion of it would carry us beyond our scope.

Fig. 63.

In one of Plateau’s experiments, a bubble of oil (protected from gravity by the specific gravity of the surrounding fluid being identical with its own) is balanced between two annuli. It may then be brought to assume the form of Fig. 63, that is to say the form of a cylinder with spherical ends; and there is then everywhere, owing to the convexity of the surface film, a pressure inwards upon the fluid contents of the bubble. If the surrounding liquid be ever so little heavier or lighter than that which constitutes the drop, then the conditions of equi­lib­rium will be accordingly {223} modified, and the cylindrical drop will assume the form of an unduloid (Fig. 64 A, B), with its dilated portion below or above,

Fig. 64.

as the case may be; and our cylinder may also, of course, be converted into an unduloid either by elongating it further, or by abstracting a portion of its oil, until at length rupture ensues and the cylinder breaks up into two new spherical drops. In all cases alike, the unduloid, like the original cylinder, will be capped by spherical ends, which are the sign, and the consequence, of the positive pressure produced by the curved walls of the unduloid. But if our initial cylinder, instead of being tall, be a flat or dumpy one (with certain definite relations of height to breadth), then new phenomena may be exhibited. For now, if a little oil be cautiously withdrawn from the mass by help of a small syringe, the cylinder may be made to flatten down so that its upper and lower surfaces become plane; which is of itself an indication that the pressure inwards is now nil. But at the very moment when the upper and lower surfaces become plane, it will be found that the sides curve inwards, in the fashion shewn in Fig. 65B. This figure is a catenoid, which, as

Fig. 65.

we have already seen, is, like the plane itself, a surface exercising no pressure, and which therefore may coexist with the plane as part of one and the same system. We may continue to withdraw more oil from our bubble, drop by drop, and now the upper and lower surfaces dimple down into concave portions of spheres, as the result of the negative internal pressure; and thereupon the peripheral catenoid surface alters its form (perhaps, on this small scale, imperceptibly), and becomes a portion of a nodoid (Fig. 65A). {224} It represents, in fact, that portion of the nodoid, which in Fig. 66 lies between such points as O, P. While it is easy to

Fig. 66.

draw the outline, or meridional section, of the nodoid (as in Fig. 66), it is obvious that the solid of revolution to be derived from it, can never be realised in its entirety: for one part of the solid figure would cut, or entangle with, another. All that we can ever do, accordingly, is to realise isolated portions of the nodoid.

If, in a sequel to the preceding experiment of Plateau’s, we use solid discs instead of annuli, so as to enable us to exert direct mechanical pressure upon our globule of oil, we again begin by adjusting the pressure of these discs so that the oil assumes the form of a cylinder: our discs, that is to say, are adjusted to exercise a mechanical pressure equal to what in the former case was supplied by the surface-tension of the spherical caps or ends of the bubble. If we now increase the pressure slightly, the peripheral walls will become convexly curved, exercising a precisely cor­re­spon­ding pressure. Under these circumstances the form assumed by the sides of our figure will be that of a portion of an unduloid. If we increase the pressure between the discs, the peripheral surface of oil will bulge out more and more, and will presently constitute a portion of a sphere. But we may continue the process yet further, and within certain limits we shall find that the system remains perfectly stable. What is this new curved surface which has arisen out of the sphere, as the latter was produced from the unduloid? It is no other than a portion of a nodoid, that part which in Fig. 66 lies between such limits as M and N. But this surface, which is concave in both directions towards the surface of the oil within, is exerting a pressure upon the latter, just as did the sphere out of which a moment ago it was transformed; and we had just stated, in considering the previous experiment, that the pressure inwards exerted by the nodoid was a negative one. The explanation of this seeming discrepancy lies in the simple fact that, if we follow the outline {225} of our nodoid curve in Fig. 66 from O, P, the surface concerned in the former case, to M, N, that concerned in the present, we shall see that in the two experiments the surface of the liquid is not homologous, but lies on the positive side of the curve in the one case and on the negative side in the other.

Of all the surfaces which we have been describing, the sphere is the only one which can enclose space; the others can only help to do so, in combination with one another or with the sphere itself. Thus we have seen that, in normal equi­lib­rium, the cylindrical vesicle is closed at either end by a portion of a sphere, and so on. Moreover the sphere is not only the only one of our figures which can enclose a finite space; it is also, of all possible figures, that which encloses the greatest volume with the least area of surface; it is strictly and absolutely the surface of minimal area, and it is therefore the form which will be naturally assumed by a unicellular organism (just as by a raindrop), when it is practically homogeneous and when, like Orbulina floating in the ocean, its surroundings are likewise practically homogeneous and symmetrical. It is only relatively speaking that all the rest are surfaces minimae areae; they are so, that is to say, under the given conditions, which involve various forms of pressure or restraint. Such restraints are imposed, for instance, by the pipes or annuli with the help of which we draw out our cylindrical or unduloid oil-globule or soap-bubble; and in the case of the organic cell, similar restraints are constantly supplied by solidification, partial or complete, local or general, of the cell-wall.

Before we pass to biological illustrations of our surface-tension figures, we have still another preliminary matter to deal with. We have seen from our description of two of Plateau’s classical experiments, that at some particular point one type of surface gives place to another; and again, we know that, when we draw out our soap-bubble into and then beyond a cylinder, there comes a certain definite point at which our bubble breaks in two, and leaves us with two bubbles of which each is a sphere, or a portion of a sphere. In short there are certain definite limits to the dimensions of our figures, within which limits equi­lib­rium is stable but at which it becomes unstable, and above which it {226} breaks down. Moreover in our composite surfaces, when the cylinder for instance is capped by two spherical cups or lenticular discs, there is a well-defined ratio which regulates their respective curvatures, and therefore their respective dimensions. These two matters we may deal with together.

Let us imagine a liquid drop which by appropriate conditions has been made to assume the form of a cylinder; we have already seen that its ends will be terminated by portions of spheres. Since one and the same liquid film covers the sides and ends of the drop (or since one and the same delicate membrane encloses the sides and ends of the cell), we assume the surface-tension (T) to be everywhere identical; and it follows, since the internal fluid-pressure is also everywhere identical, that the expression (1 ⁄ R + 1 ⁄ R′) for the cylinder is equal to the cor­re­spon­ding expression, which we may call (1 ⁄ r + 1 ⁄ r′), in the case of the terminal spheres. But in the cylinder 1 ⁄ R′ = 0, and in the sphere 1 ⁄ r = 1 ⁄ r′. Therefore our relation of equality becomes 1 ⁄ R = 2 ⁄ r, or r = 2 R; that is to say, the sphere in question has just twice the radius of the cylinder of which it forms a cap.

Fig. 67.

Among the many important theoretical discoveries which we owe to Plateau, one to which we have just referred is of peculiar importance: namely that, with the exception of the sphere and the plane, the surfaces with which we have been dealing are only in complete equi­lib­rium within certain dimensional limits, or in other words, have a certain definite limit of stability; only the plane and the sphere, or any portions of a sphere, are perfectly stable, because they are perfectly symmetrical, figures. For experimental demonstration, the case of the cylinder is the simplest. If we produce a liquid film having the form of a cylinder, either by

Fig. 68.

drawing out a bubble or by supporting between two rings a globule of oil, the experiment proceeds easily until the length of the cylinder becomes just about three times as great as its diameter. But somewhere about this limit the cylinder alters its form; it begins to narrow at the waist, so passing into an unduloid, and the deformation progresses quickly until at last our cylinder breaks in two, and its two halves assume a spherical form. It is found, by theoretical con­si­de­ra­tions, that the precise limit of stability is at the point when the length of the cylinder is exactly equal to its circumference, that is to say, when L = 2πR, or when the ratio of length to diameter is represented by π.

In the case of the catenoid, Plateau’s experimental procedure was as follows. To support his globule of oil (in, as usual, a mixture of alcohol and water of its own specific gravity), he used {228} a pair of metal rings, which happened to have a diameter of 71 millimetres; and, in a series of experiments, he set these rings apart at distances of 55, 49, 47, 45, and 43 mm. successively. In each case he began by bringing his oil-globule into a cylindrical form, by sucking superfluous oil out of the drop until this result was attained; and always, for the reason with which we are now acquainted, the cylindrical sides were associated with spherical ends to the cylinder. On continuing to withdraw oil in the hope of converting these spherical ends into planes, he found, naturally, that the sides of the cylinder drew in to form a concave surface; but it was by no means easy to get the extremities actually plane: and unless they were so, thus indicating that the surface-pressure of the drop was nil, the curvature of the sides could not be that of a catenoid. For in the first experiment, when the rings were 55 mm. apart, as soon as the convexity of the ends was to a certain extent diminished, it spontaneously increased again; and the transverse constriction of the globule correspondingly deepened, until at a certain point equi­lib­rium set in anew. Indeed, the more oil he removed, the more convex became the ends, until at last the increasing transverse constriction led to the breaking of the oil-globule into two. In the third experiment, when the rings were 47 mm. apart, it was easy to obtain end-surfaces that were actually plane, and they remained so even though more oil was withdrawn, the transverse constriction deepening accordingly. Only after a considerable amount of oil had been sucked up did the plane terminal surface become gradually convex, and presently the narrow waist, narrowing more and more, broke across in the usual way. Finally in the fifth experiment, where the rings were still nearer together, it was again possible to bring the ends of the oil-globule to a plane surface, as in the third and fourth experiments, and to keep this surface plane in spite of some continued withdrawal of oil. But very soon the ends became gradually concave, and the concavity deepened as more and more oil was withdrawn, until at a certain limit, the whole oil-globule broke up in general disruption.

We learn from this that the limiting size of the catenoid was reached when the distance of the supporting rings was to their diameter as 47 to 71, or, as nearly as possible, as two to three; {229} and as a matter of fact it can be shewn that 2 ⁄ 3 is the true theoretical value. Above this limit of 2 ⁄ 3, the inevitable convexity of the end-surfaces shows that a positive pressure inwards is being exerted by the surface film, and this teaches us that the sides of the figure actually constitute not a catenoid but an unduloid, whose spontaneous changes tend to a form of greater stability. Below the 2 ⁄ 3 limit the catenoid surface is essentially unstable, and the form into which it passes under certain conditions of disturbance such as that of the excessive withdrawal of oil, is that of a nodoid (Fig. 65A).

The unduloid has certain peculiar properties as regards its limitations of stability. But as to these we need mention two facts only: (1) that when the unduloid, which we produce with our soap-bubble or our oil-globule, consists of the figure containing a complete constriction, it has somewhat wide limits of stability; but (2) if it contain the swollen portion, then equi­lib­rium is limited to the condition that the figure consists simply of one complete unduloid, that is to say that its ends are constituted by the narrowest portions, and its middle by the widest portion of the entire curve. The theoretical proof of this latter fact is difficult, but if we take the proof for granted, the fact will serve to throw light on what we have learned regarding the stability of the cylinder. For, when we remember that the meridional section of our unduloid is generated by the rolling of an ellipse upon a straight line in its own plane, we shall easily see that the length of the entire unduloid is equal to the circumference of the generating ellipse. As the unduloid becomes less and less sinuous in outline, it gradually approaches, and in time reaches, the form of a cylinder; and correspondingly, the ellipse which generated it has its foci more and more approximated until it passes into a circle. The cylinder of a length equal to the circumference of its generating circle is therefore precisely homologous to an unduloid whose length is equal to the circumference of its generating ellipse; and this is just what we recognise as constituting one complete segment of the unduloid.

While the figures of equi­lib­rium which are at the same time surfaces of revolution are only six in number, there is an infinite {230} number of figures of equi­lib­rium, that is to say of surfaces of constant mean curvature, which are not surfaces of revolution; and it can be shewn math­e­mat­i­cally that any given contour can be occupied by a finite portion of some one such surface, in stable equi­lib­rium. The experimental verification of this theorem lies in the simple fact (already noted) that however we may bend a wire into a closed curve, plane or not plane, we may always, under appropriate precautions, fill the entire area with an unbroken film.

Of the regular figures of equi­lib­rium, that is to say surfaces of constant mean curvature, apart from the surfaces of revolution which we have discussed, the helicoid spiral is the most interesting to the biologist. This is a helicoid generated by a straight line perpendicular to an axis, about which it turns at a uniform rate while at the same time it slides, also uniformly, along this same axis. At any point in this surface, the curvatures are equal and of opposite sign, and the sum of the curvatures is accordingly nil. Among what are called “ruled surfaces” (which we may describe as surfaces capable of being defined by a system of stretched strings), the plane and the helicoid are the only two whose mean curvature is null, while the cylinder is the only one whose curvature is finite and constant. As this simplest of helicoids corresponds, in three dimensions, to what in two dimensions is merely a plane (the latter being generated by the rotation of a straight line about an axis without the superadded gliding motion which generates the helicoid), so there are other and much more complicated helicoids which correspond to the sphere, the unduloid and the rest of our figures of revolution, the generating planes of these latter being supposed to wind spirally about an axis. In the case of the cylinder it is obvious that the resulting figure is in­dis­tin­guish­able from the cylinder itself. In the case of the unduloid we obtain a grooved spiral, such as we may meet with in nature (for instance in Spirochætes, Bodo gracilis, etc.), and which accordingly it is of interest to us to be able to recognise as a surface of minimal area or constant curvature.

The foregoing con­si­de­ra­tions deal with a small part only of the theory of surface tension, or of capillarity: with that part, namely, which relates to the forms of surface which are {231} capable of subsisting in equi­lib­rium under the action of that force, either of itself or subject to certain simple constraints. And as yet we have limited ourselves to the case of a single surface, or of a single drop or bubble, leaving to another occasion a discussion of the forms assumed when such drops or vesicles meet and combine together. In short, what we have said may help us to understand the form of a cell,—considered, as with certain limitations we may legitimately consider it, as a liquid drop or liquid vesicle; the conformation of a tissue or cell-aggregate must be dealt with in the light of another series of theoretical con­si­de­ra­tions. In both cases, we can do no more than touch upon the fringe of a large and difficult subject. There are many forms capable of realisation under surface tension, and many of them doubtless to be recognised among organisms, which we cannot touch upon in this elementary account. The subject is a very general one; it is, in its essence, more math­e­mat­i­cal than physical; it is part of the mathematics of surfaces, and only comes into relation with surface tension, because this physical phenomenon illustrates and exemplifies, in a concrete way, most of the simple and symmetrical conditions with which the general math­e­mat­i­cal theory is capable of dealing. And before we pass to illustrate by biological examples the physical phenomena which we have described, we must be careful to remember that the physical conditions which we have hitherto presupposed will never be wholly realised in the organic cell. Its substance will never be a perfect fluid, and hence equi­lib­rium will be more or less slowly reached; its surface will seldom be perfectly homogeneous, and therefore equi­lib­rium will (in the fluid condition) seldom be perfectly attained; it will very often, or generally, be the seat of other forces, symmetrical or unsymmetrical; and all these causes will more or less perturb the effects of surface tension acting by itself. But we shall find that, on the whole, these effects of surface tension though modified are not obliterated nor even masked; and accordingly the phenomena to which I have devoted the foregoing pages will be found manifestly recurring and repeating themselves among the phenomena of the organic cell.

In a spider’s web we find exemplified several of the principles {232} of surface tension which we have now explained. The thread is formed out of the fluid secretion of a gland, and issues from the body as a semi-fluid cylinder, that is to say in the form of a surface of equi­lib­rium, the force of expulsion giving it its elongation and that of surface tension giving it its circular section. It is prevented, by almost immediate solidification on exposure to the air, from breaking up into separate drops or spherules, as it would otherwise tend to do as soon as the length of the cylinder had passed its limit of stability. But it is otherwise with the sticky secretion which, coming from another gland, is simultaneously poured over the issuing thread when it is to form the spiral portion of the web. This latter secretion is more fluid than the first, and retains its fluidity for a very much longer time, finally drying up after several hours. By capillarity it “wets” the thread, spreading itself over it in an even film, which film is now itself a cylinder. But this liquid cylinder has its limit of stability when its length equals its own circumference, and therefore just at the points so defined it tends to disrupt into separate segments: or rather, in the actual case, at points somewhat more distant, owing to the imperfect fluidity of the viscous film, and still more to the frictional drag upon it of the inner solid cylinder, or thread, with which it is in contact. The cylinder disrupts in the usual manner, passing first into the wavy outline of an unduloid, whose swollen portions swell more and more till the contracted parts break asunder, and we arrive at a series of spherical drops or beads, of equal size, strung at equal intervals along the thread. If we try to spread varnish over a thin stretched wire, we produce automatically the same identical result288; unless our varnish be such as to dry almost instantaneously, it gathers into beads, and do what we can, we fail to spread it smooth. It follows that, according to the viscidity and drying power of the varnish, the process may stop or seem to stop at any point short of the formation of the perfect spherules; it is quite possible, therefore, that as our final stage we may only obtain half-formed beads, or the wavy outline of an unduloid. The formation of the beads may be facilitated or hastened by jerking the stretched thread, as the spider actually does: the {233} effect of the jerk being to disturb and destroy the unstable equi­lib­rium of the viscid cylinder289. Another very curious phenomenon here presents itself.

In Plateau’s experimental separation of a cylinder of oil into two spherical portions, it was noticed that, when contact was nearly broken, that is to say when the narrow neck of the unduloid had become very thin, the two spherical bullae, instead of absorbing the fluid out of the narrow neck into themselves as they had done with the preceding portion, drew out this small remaining part of the liquid into a thin thread as they completed their spherical form and consequently receded from one another: the reason being that, after the thread or “neck” has reached a certain tenuity, the internal friction of the fluid prevents or retards its rapid exit from the little thread to the adjacent spherule. It is for the same reason that we are able to draw a glass rod or tube, which we have heated in the middle, into a long and uniform cylinder or thread, by quickly separating the two ends. But in the case of the glass rod, the long thin intermediate cylinder quickly cools and solidifies, while in the ordinary separation of a liquid cylinder the cor­re­spon­ding intermediate cylinder remains liquid; and therefore, like any other liquid cylinder, it is liable to break up, provided that its dimensions exceed the normal limit of stability. And its length is generally such that it breaks at two points, thus leaving two terminal portions continuous with the spheres and becoming confluent with these, and one median portion which resolves itself into a comparatively tiny spherical drop, midway between the original and larger two. Occasionally, the same process of formation of a connecting thread repeats itself a second time, between the small intermediate spherule and the large spheres; and in this case we obviously obtain two additional spherules, still smaller in size, and lying one on either side of our first little one. This whole phenomenon, of equal and regularly interspaced beads, often with little beads regularly interspaced between the larger ones, and possibly also even a third series of still smaller beads regularly intercalated, may be easily observed in a spider’s web, such as that of Epeira, very often with beautiful regularity,—which {234} naturally, however, is sometimes interrupted and disturbed owing to a slight want of homogeneity in the secreted fluid; and the same phenomenon is repeated on a grosser scale when the web is bespangled with dew, and every thread bestrung with pearls innumerable. To the older naturalists, these regularly arranged and beautifully formed globules on the spider’s web were a cause of great wonder and admiration. Blackwall, counting some twenty globules in a tenth of an inch, calculated that a large garden-spider’s web comprised about 120,000 globules; the net was spun and finished in about forty minutes, and Blackwall was evidently filled with astonishment at the skill and quickness with which the spider manufactured these little beads. And no wonder, for according to the above estimate they had to be made at the rate of about 50 per second290.

Fig. 69. Hair of Trianea, in glycerine. (After Berthold.)

The little delicate beads which stud the long thin pseudopodia of a foraminifer, such as Gromia, or which in like manner appear upon the cylindrical film of protoplasm which covers the long radiating spicules of Globigerina, represent an identical phenomenon. Indeed there are many cases, in which we may study in a protoplasmic filament the whole process of formation of such beads. If we squeeze out on to a slide the viscid contents of a mistletoe berry, the long sticky threads into which the substance runs shew the whole phenomenon particularly well. Another way to demonstrate it was noticed many years ago by Hofmeister and afterwards explained by Berthold. The hairs of certain water-plants, such as Hydrocharis or Trianea, constitute very long cylindrical cells, the protoplasm being supported, and maintained in equi­lib­rium by its contact with the cell-wall. But if we immerse the filament in some dense fluid, a little sugar-solution for instance, or dilute glycerine, the cell-sap tends to diffuse outwards, the protoplasm parts company with its surrounding and supporting wall, {235} and lies free as a protoplasmic cylinder in the interior of the cell. Thereupon it immediately shews signs of instability, and commences to disrupt. It tends to gather into spheres, which however, as in our illustration, may be prevented by their narrow quarters from assuming the complete spherical form; and in between these spheres, we have more or less regularly alternate ones, of smaller size291. Similar, but less regular, beads or droplets may be caused to appear, under stimulation by an alternating current, in the protoplasmic threads within the living cells of the hairs of Tradescantia. The explanation usually given is, that the viscosity of the protoplasm is reduced, or its fluidity increased; but an increase of the surface tension would seem a more likely reason292.

We may take note here of a remarkable series of phenomena, which, though they seem at first sight to be of a very different order, are closely related to the phenomena which attend and which bring about the breaking-up of a liquid cylinder or thread.

Fig. 70. Phases of a Splash. (From Worthington.)

In some of Mr Worthington’s most beautiful experiments on {236} splashes, it was found that the fall of a round pebble into water from a considerable height, caused the rise of a filmy sheet of water in the form of a cup or cylinder; and the edge of this cylindrical film tended to be cut up into alternate lobes and notches, and the prominent lobes or “jets” tended, in more extreme cases, to break off or to break up into spherical beads (Fig. 70)293. A precisely similar appearance is seen, on a great scale, in the thin edge of a breaking wave: when the smooth cylindrical edge, at a given moment, shoots out an array of tiny jets which break up into the droplets which constitute “spray” (Fig. 71, a, b). We are at once reminded of the beautifully symmetrical notching on the calycles of many hydroids, which little cups before they became stiff and rigid had begun their existence as liquid or semi-liquid films.

Fig. 71. A breaking wave. (From Worthington.)

The phenomenon is two-fold. In the first place, the edge of our tubular or crater-like film forms a liquid ring or annulus, which is closely comparable with the liquid thread or cylinder which we have just been considering, if only we conceive the thread to be bent round into the ring. And accordingly, just as the thread spontaneously segments, first into an unduloid, and then into separate spherical drops, so likewise will the edge of our annulus tend to do. This phase of notching, or beading, of the edge of the film is beautifully seen in many of Worthington’s experiments294. In the second place, the very fact of the rising of the crater means that liquid is flowing up from below towards the rim; and the segmentation of the rim means that channels of easier flow are {237} created, along which the liquid is led, or is driven, into the protuberances: and these are thus exaggerated into the jets or arms which are sometimes so conspicuous at the edge of the crater. In short, any film or film-like cup, fluid or semi-fluid in its consistency, will, like the straight liquid cylinder, be unstable: and its instability will manifest itself (among other ways) in a tendency to segmentation or notching of the edge; and just such a peripheral notching is a conspicuous feature of many minute organic cup-like structures. In the case of the hydroid calycle (Fig. 72), we are led to the conclusion that the two common and conspicuous features of notching or indentation of the cup, and of constriction or annulation of the long cylindrical stem, are phenomena of the same order and are due to surface-tension in both cases alike.

Fig. 72. Calycles of Campanularian zoophytes.  (A) C. integra;  (B) C. groenlandica;  (C) C. bispinosa;  (D) C. raridentata.

Another phenomenon displayed in the same experiments is the formation of a rope-like or cord-like thickening of the edge of the annulus. This is due to the more or less sudden checking at the rim of the flow of liquid rising from below: and a similar peripheral thickening is frequently seen, not only in some of our hydroid cups, but in many Vorticellas (cf. Fig. 75), and other organic cup-like conformations. A perusal of Mr Worthington’s book will soon suggest that these are not the only manifestations of surface-tension in connection with splashes which present curious resemblances and analogies to phenomena of organic form.

The phenomena of an ordinary liquid splash are so swiftly {238} transitory that their study is only rendered possible by “instantaneous” photography: but this excessive rapidity is not an essential part of the phenomenon. For instance, we can repeat and demonstrate many of the simpler phenomena, in a permanent or quasi-permanent form, by splashing water on to a surface of dry sand, or by firing a bullet into a soft metal target. There is nothing, then, to prevent a slow and lasting manifestation, in a viscous medium such as a protoplasmic organism, of phenomena which appear and disappear with prodigious rapidity in a more mobile liquid. Nor is there anything peculiar in the “splash” itself; it is simply a convenient method of setting up certain motions or currents, and producing certain surface-forms, in a liquid medium,—or even in such an extremely imperfect fluid as is represented (in another series of experiments) by a bed of sand. Accordingly, we have a large range of possible conditions under which the organism might conceivably display con­fi­gur­a­tions analogous to, or identical with, those which Mr Worthington has shewn us how to exhibit by one particular experimental method.

To one who has watched the potter at his wheel, it is plain that the potter’s thumb, like the glass-blower’s blast of air, depends for its efficacy upon the physical properties of the medium on which it operates, which for the time being is essentially a fluid. The cup and the saucer, like the tube and the bulb, display (in their simple and primitive forms) beautiful surfaces of equi­lib­rium as manifested under certain limiting conditions. They are neither more nor less than glorified “splashes,” formed slowly, under conditions of restraint which enhance or reveal their math­e­mat­i­cal symmetry. We have seen, and we shall see again before we are done, that the art of the glass-blower is full of lessons for the naturalist as also for the physicist: illustrating as it does the development of a host of math­e­mat­i­cal con­fi­gur­a­tions and organic conformations which depend essentially on the establishment of a constant and uniform pressure within a closed elastic shell or fluid envelope. In like manner the potter’s art illustrates the somewhat obscurer and more complex problems (scarcely less frequent in biology) of a figure of equi­lib­rium which is an open surface, or solid, of revolution. It is clear, at the same time, that the two series of problems are closely akin; for the {239} glass-blower can make most things that the potter makes, by cutting off portions of his hollow ware. And besides, when this fails, and the glass-blower, ceasing to blow, begins to use his rod to trim the sides or turn the edges of wineglass or of beaker, he is merely borrowing a trick from the craft of the potter.

It would be venturesome indeed to extend our comparison with these liquid surface-tension phenomena from the cup or calycle of the hydrozoon to the little hydroid polype within: and yet I feel convinced that there is something to be learned by such a comparison, though not without much detailed consideration and math­e­mat­i­cal study of the surfaces concerned. The cylindrical body of the tiny polype, the jet-like row of tentacles, the beaded annulations which these tentacles exhibit, the web-like film which sometimes (when they stand a little way apart) conjoins their bases, the thin annular film of tissue which surrounds the little organism’s mouth, and the manner in which this annular “peristome” contracts295, like a shrinking soap-bubble, to close the aperture, are every one of them features to which we may find a singular and striking parallel in the surface-tension phenomena which Mr Worthington has illustrated and demonstrated in the case of the splash.

Here however, we may freely confess that we are for the present on the uncertain ground of suggestion and conjecture; and so must we remain, in regard to many other simple and symmetrical organic forms, until their form and dynamical stability shall have been investigated by the mathematician: in other words, until the mathematicians shall have become persuaded that there is an immense unworked field wherein they may labour, in the detailed study of organic form.

According to Plateau, the viscidity of the liquid, while it helps to retard the breaking up of the cylinder and so increases the length of the segments beyond that which theory demands, has nevertheless less influence in this direction than we might have expected. On the other hand, any external support or adhesion, such as contact with a solid body, will be equivalent to a reduction of surface-tension and so will very greatly increase the {240} stability of our cylinder. It is for this reason that the mercury in our thermometer tubes does not as a rule separate into drops, though it occasionally does so, much to our inconvenience. And again it is for this reason that the protoplasm in a long and growing tubular or cylindrical cell does not necessarily divide into separate cells and internodes, until the length of these far exceeds the theoretic limits. Of course however and whenever it does so, we must, without ever excluding the agency of surface tension, remember that there may be other forces affecting the latter, and accelerating or retarding that manifestation of surface tension by which the cell is actually rounded off and divided.

In most liquids, Plateau asserts that, on the average, the influence of viscosity is such as to cause the cylinder to segment when its length is about four times, or at most from four to six times that of its diameter: instead of a fraction over three times as, in a perfect fluid, theory would demand. If we take it at four times, it may then be shewn that the resulting spheres would have a diameter of about 1·8 times, and their distance apart would be equal to about 2·2 times the diameter of the original cylinder. The calculation is not difficult which would shew how these numbers are altered in the case of a cylinder formed around a solid core, as in the case of the spider’s web. Plateau has also made the interesting observation that the time taken in the process of division of the cylinder is directly proportional to the diameter of the cylinder, while varying considerably with the nature of the liquid. This question, of the time occupied in the division of a cell or filament, in relation to the dimensions of the latter, has not so far as I know been enquired into by biologists.

From the simple fact that the sphere is of all surfaces that whose surface-area for a given volume is an absolute minimum, we have already seen it to be plain that it is the one and only figure of equi­lib­rium which will be assumed under surface-tension by a drop or vesicle, when no other disturbing factors are present. One of the most important of these disturbing factors will be introduced, in the form of complicated tensions and pressures, when one drop is in contact with another drop and when a system of intermediate films or partition walls is developed between them. {241} This subject we shall discuss later, in connection with cell-aggregates or tissues, and we shall find that further theoretical con­si­de­ra­tions are needed as a preliminary to any such enquiry. Meanwhile let us consider a few cases of the forms of cells, either solitary, or in such simple aggregates that their individual form is little disturbed thereby.

Let us clearly understand that the cases we are about to consider are those cases where the perfect symmetry of the sphere is replaced by another symmetry, less complete, such as that of an ellipsoidal or cylindrical cell. The cases of asymmetrical deformation or displacement, such as is illustrated in the production of a bud or the development of a lateral branch, are much simpler. For here we need only assume a slight and localised variation of surface-tension, such as may be brought about in various ways through the heterogeneous chemistry of the cell; to this point we shall return in our chapter on Adsorption. But the diffused and graded asymmetry of the system, which brings about for instance the ellipsoidal shape of a yeast-cell, is another matter.

If the sphere be the one surface of complete symmetry and therefore of independent equi­lib­rium, it follows that in every cell which is otherwise conformed there must be some definite force to cause its departure from sphericity; and if this cause be the very simple and obvious one of the resistance offered by a solidified envelope, such as an egg-shell or firm cell-wall, we must still seek for the deforming force which was in action to bring about the given shape, prior to the assumption of rigidity. Such a cause may be either external to, or may lie within, the cell itself. On the one hand it may be due to external pressure or to some form of mechanical restraint: as it is in all our experiments in which we submit our bubble to the partial restraint of discs or rings or more complicated cages of wire; and on the other hand it may be due to intrinsic causes, which must come under the head either of differences of internal pressure, or of lack of homogeneity or isotropy in the surface itself296. {242}

Our full formula of equi­lib­rium, or equation to an elastic surface, is P = p_e + (T ⁄ R + T′ ⁄ R′), where P is the internal pressure, p_e any extraneous pressure normal to the surface, R, R′ the radii of curvature at a point, and T, T′, the cor­re­spon­ding tensions, normal to one another, of the envelope.

Now in any given form which we are seeking to account for, R, R′ are known quantities; but all the other factors of the equation are unknown and subject to enquiry. And somehow or other, by this formula, we must account for the form of any solitary cell whatsoever (provided always that it be not formed by successive stages of solidification), the cylindrical cell of Spirogyra, the ellipsoidal yeast-cell, or (as we shall see in another chapter) the shape of the egg of any bird. In using this formula hitherto, we have taken it in a simplified form, that is to say we have made several limiting assumptions. We have assumed that P was simply the uniform hydrostatic pressure, equal in all directions, of a body of liquid; we have assumed that the tension T was simply due to surface-tension in a homogeneous liquid film, and was therefore equal in all directions, so that T = T′; and we have only dealt with surfaces, or parts of a surface, where extraneous pressure, p_n, was non-existent. Now in the case of a bird’s egg, the external pressure p_n, that is to say the pressure exercised by the walls of the oviduct, will be found to be a very important factor; but in the case of the yeast-cell or the Spirogyra, wholly immersed in water, no such external pressure comes into play. We are accordingly left, in such cases as these last, with two hypotheses, namely that the departure from a spherical form is due to inequalities in the internal pressure P, or else to inequalities in the tension T, that is to say to a difference between T and T′. In other words, it is theoretically possible that the oval form of a yeast-cell is due to a greater internal pressure, a greater “tendency to grow,” in the direction of the longer axis of the ellipse, or alternatively, that with equal and symmetrical tendencies to growth there is associated a difference of external resistance in {243} respect of the tension of the cell-wall. Now the former hypothesis is not impossible; the protoplasm is far from being a perfect fluid; it is the seat of various internal forces, sometimes manifestly polar; and accordingly it is quite possible that the internal forces, osmotic and other, which lead to an increase of the content of the cell and are manifested in pressure outwardly directed upon its wall may be unsymmetrical, and such as to lead to a deformation of what would otherwise be a simple sphere. But while this hypothesis is not impossible, it is not very easy of acceptance. The protoplasm, though not a perfect fluid, has yet on the whole the properties of a fluid; within the small compass of the cell there is little room for the development of unsymmetrical pressures; and, in such a case as Spirogyra, where a large part of the cavity is filled by a fluid and watery cell-sap, the conditions are still more obviously those under which a uniform hydrostatic pressure is to be expected. But in variations of T, that is to say of the specific surface-tension per unit area, we have an ample field for all the various deformations with which we shall have to deal. Our condition now is, that (T ⁄ R + T′ ⁄ R′) = a constant; but it no longer follows, though it may still often be the case, that this will represent a surface of absolute minimal area. As soon as T and T′ become unequal, it is obvious that we are no longer dealing with a perfectly liquid surface film; but its departure from a perfect fluidity may be of all degrees, from that of a slight non-isotropic viscosity to the state of a firm elastic membrane297. And it matters little whether this viscosity or semi-rigidity be manifested in the self-same layer which is still a part of the protoplasm of the cell, or in a layer which is completely differentiated into a distinct and separate membrane. As soon as, by secretion or “adsorption,” the molecular constitution of the surface layer is altered, it is clearly conceivable that the alteration, or the secondary chemical changes which follow it, may be such as to produce an anisotropy, and to render the molecular forces less capable in one direction than another of exerting that contractile force by which they are striving to reduce to an absolute minimum the {244} surface area of the cell. A slight inequality in two opposite directions will produce the ellipsoid cell, and a very great inequality will give rise to the cylindrical cell298.

I take it therefore, that the cylindrical cell of Spirogyra, or any other cylindrical cell which grows in freedom from any manifest external restraint, has assumed that particular form simply by reason of the molecular constitution of its developing surface-membrane; and that this molecular constitution was anisotropous, in such a way as to render extension easier in one direction than another.

Such a lack of homogeneity or of isotropy, in the cell-wall is often rendered visible, especially in plant-cells, in various ways, in the form of concentric lamellae, annular and spiral striations, and the like.

But this phenomenon, while it brings about a certain departure from complete symmetry, is still compatible with, and coexistent with, many of the phenomena which we have seen to be associated with surface-tension. The symmetry of tensions still leaves the cell a solid of revolution, and its surface is still a surface of equi­lib­rium. The fluid pressure within the cylinder still causes the film or membrane which caps its ends to be of a spherical form. And in the young cell, where the surface pellicle is absent or but little differentiated, as for instance in the oögonium of Achlya, or in the young zygospore of Spirogyra, we always see the tendency of the entire structure towards a spherical form reasserting itself: unless, as in the latter case, it be overcome by direct compression within the cylindrical mother-cell. Moreover, in those cases where the adult filament consists of cylindrical cells, we see that the young, germinating spore, at first spherical, very soon assumes with growth an elliptical or ovoid form: the direct result of an incipient anisotropy of its envelope, which when more developed will convert the ovoid into a cylinder. We may also notice that a truly cylindrical cell is comparatively rare; for in most cases, what we call a cylindrical cell shews a distinct bulging of its sides; it is not truly a cylinder, but a portion of a spheroid or ellipsoid. {245}

Unicellular organisms in general, including the protozoa, the unicellular cryptogams, the various bacteria, and the free, isolated cells, spores, ova, etc. of higher organisms, are referable for the most part to a very small number of typical forms; but besides a certain number of others which may be so referable, though obscurely, there are obviously many others in which either no symmetry is to be recognized, or in which the form is clearly not one of equi­lib­rium. Among these latter we have Amoeba itself, and all manner of amoeboid organisms, and also many curiously shaped cells, such as the Trypanosomes and various other aberrant Infusoria. We shall return to the consideration of these; but in the meanwhile it will suffice to say that, as their surfaces are not equi­lib­rium-surfaces, so neither are the living cells themselves in any stable equi­lib­rium. On the contrary, they are in continual flux and movement, each portion of the surface constantly changing its form, and passing from one phase to another of an equi­lib­rium which is never stable for more than a moment. The former class, which rest in stable equi­lib­rium, must fall (as we have seen) into two classes,—those whose equi­lib­rium arises from liquid surface-tension alone, and those in whose conformation some other pressure or restraint has been superimposed upon ordinary surface-tension.

To the fact that these little organisms belong to an order of magnitude in which form is mainly, if not wholly, conditioned and controlled by molecular forces, is due the limited range of forms which they actually exhibit. These forms vary according to varying physical conditions. Sometimes they do so in so regular and orderly a way that we instinctively explain them merely as “phases of a life-history,” and leave physical properties and physical causation alone: but many of their variations of form we treat as exceptional, abnormal, decadent or morbid, and are apt to pass these over in neglect, while we give our attention to what we suppose to be the typical or “char­ac­ter­is­tic” form or attitude. In the case of the smallest organisms, the bacteria, micrococci, and so forth, the range of form is especially limited, owing to their minuteness, the powerful pressure which their highly curved surfaces exert, and the comparatively homogeneous nature of their substance. But within their narrow range of possible diversity {246} these minute organisms are protean in their changes of form. A certain species will not only change its shape from stage to stage of its little “cycle” of life; but it will be remarkably different in outward form according to the circumstances under which we find it, or the histological treatment to which we submit it. Hence the pathological student, commencing the study of bacteriology, is early warned to pay little heed to differences of form, for purposes of recognition or specific identification. Whatever grounds we may have for attributing to these organisms a permanent or stable specific identity (after the fashion of the higher plants and animals), we can seldom safely do so on the ground of definite and always recognisable form: we may

Fig. 73. A flagellate “monad,” Distigma proteus, Ehr. (After Saville Kent.)

Fig. 74. Noctiluca miliaris.

often be inclined, in short, to ascribe to them a physiological (sometimes a “pathogenic”), rather than a morphological specificity.

Among the Infusoria, we have a small number of forms whose symmetry is distinctly spherical, for instance among the small flagellate monads; but even these are seldom actually spherical except when we see them in a non-flagellate and more or less encysted or “resting” stage. In this condition, it need hardly be remarked that the spherical form is common and general among a great variety of unicellular organisms. When our little monad developes a flagellum, that is in itself an indication of “polarity” or symmetrical non-homogeneity of the cell; and accordingly, we {247} usually see signs of an unequal tension of the membrane in the neighbourhood of the base of the flagellum. Here the tension is usually less than elsewhere, and the radius of curvature is accordingly less: in other words that end of the cell is drawn out to a tapering point (Fig. 73). But sometimes it is the other way, as in Noctiluca, where the large flagellum springs from a depression in the otherwise uniformly rounded cell. In this case the explanation seems to lie in the many strands of radiating protoplasm which converge upon this point, and may be supposed to keep it relatively fixed by their viscosity, while the rest of the cell-surface is free to expand (Fig. 74).

Fig. 75. Various species of Vorticella. (Mostly after Saville Kent.)

A very large number of Infusoria represent unduloids, or portions of unduloids, and this type of surface appears and reappears in a great variety of forms. The cups of the various species of Vorticella (Fig. 75) are nothing in the world but a beautiful series of unduloids, or partial unduloids, in every gradation from a form that is all but cylindrical to one that is all but a perfect sphere. These unduloids are not completely symmetrical, but they are such unduloids as develop themselves when we suspend an oil-globule between two unequal rings, or blow a soap-bubble between two unequal pipes; for, just as in these cases, the surface of our Vorticella bell finds its terminal supports, on the one hand in its attachment to its narrow stalk, and on the other in the thickened ring from which spring its circumoral cilia. And here let me say, that a point or zone from which cilia arise would seem always to have a peculiar relation to the surrounding tensions. It usually forms a sharp salient, a prominent point or ridge, as in our little monads of Fig. 73; shewing that, in its formation, the surface tension had here locally diminished. But if such a ridge or fillet consolidate in the least degree, it becomes a source of strength, and a point d’appui for the adjacent film. We shall deal with this point again in the next chapter. {248}

Precisely the same series of unduloid forms may be traced in even greater variety among various other families or genera of the

Fig. 76. Various species of Salpingoeca.

Fig. 77. Various species of Tintinnus, Dinobryon and Codonella.
(After Saville Kent and others.)

Infusoria. Sometimes, as in Vorticella itself, the unduloid is seen merely in the contour of the soft semifluid body of the living animal. At other times, as in Salpingoeca, Tintinnus, and many

Fig. 78. Vaginicola.

other genera, we have a distinct membranous cup, separate from the animal, but originally secreted by, and moulded upon, its semifluid living surface. Here we have an excellent illustration of the contrast between the different ways in which such a structure may be regarded and interpreted. The teleological explanation is that it is developed for the sake of protection, as a domicile and shelter for the little organism within. The mechanical explanation of the physicist (seeking only after the “efficient,” and not the “final” cause), is that it is {249} present, and has its actual conformation, by reason of certain chemico-physical conditions: that it was inevitable, under the given

Fig. 79. Folliculina.

conditions, that certain constituent substances actually present in the protoplasm should be aggregated by molecular forces in its surface layer; that under this adsorptive process, the conditions continuing favourable, the particles should accumulate and concentrate till they formed (with the help of the surrounding medium) a pellicle or membrane, thicker or thinner as the case might be; that this surface pellicle or membrane was inevitably bound, by molecular forces, to become a surface of the least

Fig. 80. Trachelophyllum. (After Wreszniowski.)

possible area which the circumstances permitted; that in the present case, the symmetry and “freedom” of the system permitted, and ipso facto caused, this surface to be a surface of revolution; and that of the few surfaces of revolution which, as being also surfaces minimae areae, were available, the unduloid was manifestly the one permitted, and ipso facto caused, by the dimensions of the organisms and other circumstances of the case. And just as the thickness or thinness of the pellicle was obviously a subordinate matter, a mere matter of degree, so we also see that the actual outline of this or that particular unduloid is also a very subordinate matter, such as physico-chemical variants of a minute kind would suffice to bring about; for between the various unduloids which the various species of Vorticella represent, there is no more real difference than that difference of ratio or degree which exists between two circles of different diameter, or two lines of unequal length. {250}

In very many cases (of which Fig. 80 is an example), we have an unduloid form exhibited, not by a surrounding pellicle or shell, but by the soft, protoplasmic body of a ciliated organism. In such cases the form is mobile, and continually changes from one to another unduloid contour, according to the movements of the animal. We have here, apparently, to deal with an unstable equi­lib­rium, and also sometimes with the more complicated problem of “stream-lines,” as in the difficult problems suggested by the form of a fish. But this whole class of cases, and of problems, we can merely take note of in passing, for their treatment is too hard for us.

In considering such series of forms as the various unduloids which we have just been regarding, we are brought sharply up (as in the case of our Bacteria or Micrococci) against the biological concept of organic species. In the intense clas­si­fi­ca­tory activity of the last hundred years, it has come about that every form which is apparently char­ac­ter­is­tic, that is to say which is capable of being described or portrayed, and capable of being recognised when met with again, has been recorded as a species,—for we need not concern ourselves with the occasional discussions, or individual opinions, as to whether such and such a form deserve “specific rank,” or be “only a variety.” And this secular labour is pursued in direct obedience to the precept of the Systema Naturae,—“ut sic in summa confusione rerum apparenti, summus conspiciatur Naturae ordo.” In like manner the physicist records, and is entitled to record, his many hundred “species” of snow-crystals299, or of crystals of calcium carbonate. But regarding these latter species, the physicist makes no assumptions: he records them simpliciter, as specific “forms”; he notes, as best he can, the circumstances (such as temperature or humidity) under which they occur, in the hope of elucidating the conditions determining their formation; but above all, he does not introduce {251} the element of time, and of succession, or discuss their origin and affiliation as an historical sequence of events. But in biology, the term species carries with it many large, though often vague assumptions. Though the doctrine or concept of the “permanence of species” is dead and gone, yet a certain definite value, or sort of quasi-permanency, is still connoted by the term. Thus if a tiny foraminiferal shell, a Lagena for instance, be found living to-day, and a shell in­dis­tin­guish­able from it to the eye be found fossil in the Chalk or some other remote geological formation, the assumption is deemed legitimate that that species has “survived,” and has handed down its minute specific character or characters, from generation to generation, unchanged for untold myriads of years300. Or if the ancient forms be like to, rather than identical with the recent, we still assume an unbroken descent, accompanied by the hereditary transmission of common characters and progressive variations. And if two identical forms be discovered at the ends of the earth, still (with occasional slight reservations on the score of possible “homoplasy”), we build hypotheses on this fact of identity, taking it for granted that the two appertain to a common stock, whose dispersal in space must somehow be accounted for, its route traced, its epoch determined, and its causes discussed or discovered. In short, the naturalist admits no exception to the rule that a “natural clas­si­fi­ca­tion” can only be a genealogical one, nor ever doubts that “The fact that we are able to classify organisms at all in accordance with the structural char­ac­teris­tics which they present, is due to the fact of their being related by descent301.” But this great generalisation is apt in my opinion, to carry us too far. It may be safe and sure and helpful and illuminating when we apply it to such complex entities,—such thousand-fold resultants of the combination and permutation of many variable characters,—as a horse, a lion or an eagle; but (to my mind) it has a very different look, and a far less firm foundation, when we attempt to extend it to minute organisms whose specific characters are few and simple, whose simplicity {252} becomes much more manifest when we regard it from the point of view of physical and math­e­mat­i­cal description and analysis, and whose form is referable, or (to say the least of it) is very largely referable, to the direct and immediate action of a particular physical force. When we come to deal with the minute skeletons of the Radiolaria we shall again find ourselves dealing with endless modifications of form, in which it becomes still more difficult to discern, or to apply, the guiding principle of affiliation or genealogy.

Fig. 81.

Among the more aberrant forms of Infusoria is a little species known as Trichodina pedicidus, a parasite on the Hydra, or fresh-water polype (Fig. 81.) This Trichodina has the form of a more or less flattened circular disc, with a ring of cilia around both its upper and lower margins. The salient ridge from which these cilia spring may be taken, as we have already said, to play the part of a strengthening “fillet.” The circular base of the animal is flattened, in contact with the flattened surface of the Hydra over which it creeps, and the opposite, upper surface may be flattened nearly to a plane, or may at other times appear slightly convex or slightly concave. The sides of the little organism are contracted, forming a symmetrical equatorial groove between the upper and lower discs; and, on account of the minute size of the animal and its constant movements, we cannot submit the curvature of this concavity to measurement, nor recognise by the eye its exact contour. But it is evident that the conditions are precisely similar to those described on p. 223, where we were considering the conditions of stability of the catenoid. And it is further evident that, when the upper disc is actually plane, the equatorial groove is strictly a catenoid surface of revolution; and when on the other hand it is depressed, then the equatorial groove will tend to assume the form of a nodoidal surface.

Another curious type is the flattened spiral of Dinenympha302 {253} which reminds us of the cylindrical spiral of a Spirillum among the bacteria. In Dinenympha we have a symmetrical figure, whose two opposite surfaces each constitute a surface of constant mean curvature; it is evidently a figure of equi­lib­rium under certain special conditions of restraint. The cylindrical coil of the Spirillum, on the other hand, is a surface of constant mean curvature, and therefore of equi­lib­rium, as truly, and in the same sense, as the cylinder itself.

Fig. 82. Dinenympha gracilis, Leidy.

A very curious conformation is that of the vibratile “collar,” found in Codosiga and the other “Choanoflagellates,” and which we also meet with in the “collar-cells” which line the interior cavities of a sponge. Such collar-cells are always very minute, and the collar is constituted of a very delicate film, which shews an undulatory or rippling motion. It is a surface of revolution, and as it maintains itself in equi­lib­rium (though a somewhat unstable and fluctuating one), it must be, under the restricted circumstances of its case, a surface of minimal area. But it is not so easy to see what these special circumstances are; and it is obvious that the collar, if left to itself, must at once {254} contract downwards towards its base, and become confluent with

Fig. 83.

the general surface of the cell; for it has no longitudinal supports and no strengthening ring at its periphery. But in all these collar-cells, there stands within the annulus of the collar a large and powerful cilium or flagellum, in constant movement; and by the action of this flagellum, and doubtless in part also by the intrinsic vibrations of the collar itself, there is set up a constant steady current in the surrounding water, whose direction would seem to be such that it passes up the outside of the collar, down its inner side, and out in the middle in the direction of the flagellum; and there is a distinct eddy, in which foreign particles tend to be caught, around the peripheral margin of the collar. When the cell dies, that is to say when motion ceases, the collar immediately shrivels away and disappears. It is notable, by the way, that the edge of this little mobile cup is always smooth, never notched or lobed as in the cases we have discussed on p. 236: this latter condition being the outcome of a definite instability, marking the close of a period of equi­lib­rium; while in the vibratile collar of Codosiga the equi­lib­rium, such as it is, is being constantly renewed and perpetuated like that of a juggler’s pole, by the motions of the system. I take it that, somehow, its existence (in a state of partial equi­lib­rium) is due to the current motions, and to the traction exerted upon it through the friction of the stream which is constantly passing by. I think, in short, that it is formed very much in the same way as the cup-like ring of streaming ribbons, which we see fluttering and vibrating in the air-current of a ventilating fan.

It is likely enough, however, that a different and much better explanation may yet be found; and if we turn once more to Mr Worthington’s Study of Splashes, we may find a curious suggestion of analogy in the beautiful craters encircling a central jet (as the collar of Codosiga encircles the flagellum), which we see produced in the later stages of the splash of a pebble303. {255}

Among the Foraminifera we have an immense variety of forms, which, in the light of surface tension and of the principle of minimal area, are capable of explanation and of reduction to a small number of char­ac­ter­is­tic types. Many of the Foraminifera are composite structures, formed by the successive imposition of cell upon cell, and these we shall deal with later on; let us glance here at the simpler conformations exhibited by the single chambered or “monothalamic” genera, and perhaps one or two of the simplest composites.

We begin with forms, like Astrorhiza (Fig. 219, p. 464), which are in a high degree irregular, and end with others which manifest a perfect and math­e­mat­i­cal regularity. The broad difference between these two types is that the former are characterised, like Amoeba, by a variable surface tension, and consequently by unstable equi­lib­rium; but the strong contrast between these and the regular forms is bridged over by various transition-stages, or differences of degree. Indeed, as in all other Rhizopods, the very fact of the emission of pseudopodia, which reach their highest development in this group of animals, is a sign of unstable surface-equi­lib­rium; and we must therefore consider that those forms which indicate symmetry and equi­lib­rium in their shells have secreted these during periods when rest and uniformity of surface conditions alternated with the phases of pseudopodial activity. The irregular forms are in almost all cases arenaceous, that is to say they have no solid shells formed by steady adsorptive secretion, but only a looser covering of sand grains with which the protoplasmic body has come in contact and cohered. Sometimes, as in Ramulina, we have a calcareous shell combined with irregularity of form; but here we can easily see a partial and as it were a broken regularity, the regular forms of sphere and cylinder being repeated in various parts of the ramified mass. When we look more closely at the arenaceous forms, we find that the same thing is true of them; they represent, either in whole or part, approximations to the form of surfaces of equi­lib­rium, spheres, cylinders and so forth. In Aschemonella we have a precise replica of the calcareous Ramulina; and in Astrorhiza itself, in the forms distinguished by naturalists as A. crassatina, what is described as the “subsegmented interior304” {256} seems to shew the natural, physical tendency of the long semifluid cylinder of protoplasm to contract, at its limit of stability, into unduloid constrictions, as a step towards the breaking up into separate spheres: the completion of which process is restrained or prevented by the rigidity and friction of the arenaceous covering.

Fig. 84. Various species of Lagena. (After Brady.)

Passing to the typical, calcareous-shelled Foraminifera, we have the most symmetrical of all possible types in the perfect sphere of Orbulina; this is a pelagic organism, whose floating habitat places it in a position of perfect symmetry towards all external forces. Save for one or two other forms which are also spherical, or ap­prox­i­mate­ly so, like Thurammina, the rest of the monothalamic calcareous Foraminifera are all comprised by naturalists within the genus Lagena. This large and varied genus consists of “flask-shaped” shells, whose surface is simply that of an unduloid, or more frequently, like that of a flask itself, an unduloid combined with a portion of a sphere. We do not know the circumstances {257} under which the shell of Lagena is formed, nor the nature of the force by which, during its formation, the surface is stretched out into the unduloid form; but we may be pretty sure that it is suspended vertically in the sea, that is to say in a position of symmetry as regards its vertical axis, about which the unduloid surface of revolution is symmetrically formed. At the same time we have other types of the same shell in which the form is more or less flattened; and these are doubtless the cases in which such symmetry of position was not present, or was replaced by a broader, lateral contact with the surface pellicle305.

Fig. 85. (After Darling.)

While Orbulina is a simple spherical drop, Lagena suggests to our minds a “hanging drop,” drawn out to a long and slender neck by its own weight, aided by the viscosity of the material. Indeed the various hanging drops, such as Mr C. R. Darling shews us, are the most beautiful and perfect unduloids, with spherical ends, that it is possible to conceive. A suitable liquid, a little denser than water and incapable of mixing with it (such as ethyl benzoate), is poured on a surface of water. It spreads {258} over the surface and gradually forms a hanging drop, ap­prox­i­mate­ly hemispherical; but as more liquid is added the drop sinks or rather grows downwards, still adhering to the surface film; and the balance of forces between gravity and surface tension results in the unduloid contour, as the increasing weight of the drop tends to stretch it out and finally break it in two. At the moment of rupture, by the way, a tiny droplet is formed in the attenuated neck, such as we described in the normal division of a cylindrical thread (p. 233).

Many species of Lagena are complicated and beautified by a pattern, and some by the superaddition to the shell of plane extensions or “wings.” These latter give a secondary, bilateral symmetry to the little shell, and are strongly suggestive of a phase or period of growth in which it lay horizontally on the surface, instead of hanging vertically from the surface-film: in which, that is to say, it was a floating and not a hanging drop. The pattern is of two kinds. Sometimes it consists of a sort of fine reticulation, with rounded or more or less hexagonal interspaces: in other cases it is produced by a symmetrical series of ridges or folds, usually longitudinal, on the body of the flask-shaped cell, but occasionally transversely arranged upon the narrow neck. The reticulated and folded patterns we may consider separately. The netted pattern is very similar to the wrinkled surface of a dried pea, or to the more regular wrinkled patterns upon many other seeds and even pollen-grains. If a spherical body after developing a “skin” begin to shrink a little, and if the skin have so far lost its elasticity as to be unable to keep pace with the shrinkage of the inner mass, it will tend to fold or wrinkle; and if the shrinkage be uniform, and the elasticity and flexibility of the skin be also uniform, then the amount of {259} folding will be uniformly distributed over the surface. Little concave depressions will appear, regularly interspaced, and separated by convex folds. The little concavities being of equal size (unless the system be otherwise perturbed) each one will tend to be surrounded by six others; and when the process has reached its limit, the intermediate boundary-walls, or raised folds, will be found converted into a regular pattern of hexagons.

But the analogy of the mechanical wrinkling of the coat of a seed is but a rough and distant one; for we are evidently dealing with molecular rather than with mechanical forces. In one of Darling’s experiments, a little heavy tar-oil is dropped onto a saucer of water, over which it spreads in a thin film showing beautiful interference colours after the fashion of those of a soap-bubble. Presently tiny holes appear in the film, which gradually increase in size till they form a cellular pattern or honeycomb, the oil gathering together in the meshes or walls of the cellular net. Some action of this sort is in all probability at work in a surface-film of protoplasm covering the shell. As a physical phenomenon the actions involved are by no means fully understood, but surface-tension, diffusion and cohesion doubtless play their respective parts therein307. The very perfect cellular patterns obtained by Leduc (to which we shall have occasion to refer in a subsequent chapter) are diffusion patterns on a larger scale, but not essentially different.

Fig. 86.

The folded or pleated pattern is doubtless to be explained, in a general way, by the shrinkage of a surface-film under certain {260} conditions of viscous or frictional restraint. A case which (as it seems to me) is closely analogous to that of our foraminiferal shells is described by Quincke308, who let a film of albumin or of resin set and harden upon a surface of quicksilver, and found that the little solid pellicle had been thrown into a pattern of symmetrical folds. If the surface thus thrown into folds be that of a cylinder, or any other figure with one principal axis of symmetry, such as an ellipsoid or unduloid, the direction of the folds will tend to be related to the axis of symmetry, and we might expect accordingly to find regular longitudinal, or regular transverse wrinkling. Now as a matter of fact we almost invariably find in the Lagena the former condition: that is to say, in our ellipsoid or unduloid cell, the puckering takes the form of the vertical fluting on a column, rather than that of the transverse pleating of an accordion. And further, there is often a tendency for such longitudinal flutings to be more or less localised at the end of the ellipsoid, or in the region where the unduloid merges into its spherical base. In this latter region we often meet with a regular series of short longitudinal folds, as we do in the forms of Lagena denominated L. semistriata. All these various forms of surface can be imitated, or rather can be precisely reproduced, by the art of the glass-blower309.

Furthermore, they remind one, in a striking way, of the regular ribs or flutings in the film or sheath which splashes up to envelop a smooth ball which has been dropped into a liquid, as Mr Worthington has so beautifully shewn310. {261}

This is as much as to say that, when corrugation or “rippling” of the walls takes place owing to small changes of surface-tension, and consequently of pressure, such corrugation is more likely to take place in the plane of r,—that is to say, in the plane of greatest curvature. And it follows that in such a figure as an ellipsoid, wrinkling will be most likely to take place not only in a longitudinal direction but near the extremities of the figure, that is to say again in the region of greatest curvature.

Fig. 87. Nodosaria scalaris, Batsch.

Fig. 88. Gonangia of Campanularians. (a) C. gracilis; (b) C. grandis. (After Allman.)

The longitudinal wrinkling of the flask-shaped bodies of our Lagenae, and of the more or less cylindrical cells of many other Foraminifera (Fig. 87), is in complete accord with the above theoretical con­si­de­ra­tions; but nevertheless, we soon find that our result is not a general one, but is defined by certain limiting conditions, and is accordingly subject to what are, at first sight, important exceptions. For instance, when we turn to the narrow neck of the Lagena we see at once that our theory no longer holds; for {262} the wrinkling which was invariably longitudinal in the body of the cell is as invariably transverse in the narrow neck. The reason for the difference is not far to seek. The conditions in the neck are very different from those in the expanded portion of the cell: the main difference being that the thickness of the wall is no longer insignificant, but is of considerable magnitude as compared with the diameter, or circumference, of the neck. We must accordingly take it into account in considering the bending moments at any point in this region of the shell-wall. And it is at once obvious that, in any portion of the narrow neck, flexure of a wall in a transverse direction will be very difficult, while flexure in a longitudinal direction will be comparatively easy; just as, in the case of a long narrow strip of iron, we may easily bend it into folds running transversely to its long axis, but not the other way. The manner in which our little Lagena-shell tends to fold or wrinkle, longitudinally in its wider part, and transversely or annularly in its narrow neck, is thus completely and easily explained.

An identical phenomenon is apt to occur in the little flask-shaped gonangia, or reproductive capsules, of some of the hydroid zoophytes. In the annexed drawings of these gonangia in two species of Campanularia, we see that in one case the little vesicle {263} has the flask-shaped or unduloid configuration of a Lagena; and here the walls of the flask are longitudinally fluted, just after the manner we have witnessed in the latter genus. But in the other Campanularian the vesicles are long, narrow and tubular, and here a transverse folding or pleating takes the place of the longitudinally fluted pattern. And the very form of the folds or pleats is enough to suggest that we are not dealing here with a simple phenomenon of surface-tension, but with a condition in which surface-tension and stiffness are both present, and play their parts in the resultant form.

Fig. 89. Various Foraminifera (after Brady), a, Nodosaria simplex; b, N. pygmaea; c, N. costulata; e, N. hispida; f, N. elata; d, Rheophax (Lituola) distans; g, Sagrina virgata.

Passing from the solitary flask-shaped cell of Lagena, we have, in another series of forms, a constricted cylinder, or succession of unduloids; such as are represented in Fig. 89, illustrating certain species of Nodosaria, Rheophax and Sagrina. In some of these cases, and certainly in that of the arenaceous genus Rheophax, we have to do with the ordinary phenomenon of a segmenting or partially segmenting cylinder. But in others, the structure is not developed out of a continuous protoplasmic cylinder, but as we can see by examining the interior of the shell, it has been formed in successive stages, beginning with a simple unduloid “Lagena,” about whose neck, after its solidification, another drop of protoplasm accumulated, and in turn assumed the unduloid, or lagenoid, form. The chains of interconnected bubbles which {264} Morey and Draper made many years ago of melted resin are a very similar if not identical phenomenon311.

There now remain for our consideration, among the Protozoa, the great oceanic group of the Radiolaria, and the little group of their freshwater allies, the Heliozoa. In nearly all these forms we have this specific chemical difference from the Foraminifera, that when they secrete, as they generally do secrete, a hard skeleton, it is composed of silica instead of lime. These organisms and the various beautiful and highly complicated skeletal fabrics which they develop give us many interesting illustrations of physical phenomena, among which the manifestations of surface-tension are very prominent. But the chief phenomena connected with their skeletons we shall deal with in another place, under the head of spicular concretions.

In a simple and typical Heliozoan, such as the Sun-animalcule, Actinophrys sol, we have a “drop” of protoplasm, contracted by its surface tension into a spherical form. Within the heterogeneous protoplasmic mass are more fluid portions, and at the surface which separates these from the surrounding protoplasm a similar surface tension causes them also to assume the form of spherical “vacuoles,” which in reality are little clear drops within the big one; unless indeed they become numerous and closely packed, in which case, instead of isolated spheres or droplets they will constitute a “froth,” their mutual pressures and tensions giving rise to regular con­fi­gur­a­tions such as we shall study in the next chapter. One or more of such clear spaces may be what is called a “contractile vacuole”: that is to say, a droplet whose surface tension is in unstable equi­lib­rium and is apt to vanish altogether, so that the definite outline of the vacuole suddenly disappears312. Again, within the protoplasm are one or more nuclei, whose own surface tension (at the surface between the nucleus and the surrounding protoplasm), has drawn them in turn into the shape {265} of spheres. Outwards through the protoplasm, and stretching far beyond the spherical surface of the cell, there run stiff linear threads of modified or differentiated protoplasm, replaced or reinforced in some cases by delicate siliceous needles. In either case we know little or nothing about the forces which lead to their production, and we do not hide our ignorance when we ascribe their development to a “radial polarisation” of the cell. In the case of the protoplasmic filament, we may (if we seek for a hypothesis), suppose that it is somehow comparable to a viscid stream, or “liquid vein,” thrust or squirted out from the body of the cell. But when it is once formed, this long and comparatively rigid filament is separated by a distinct surface from the neighbouring protoplasm, that is to say from the more fluid surface-protoplasm of the cell; and the latter begins to creep up the filament, just as water would creep up the interior of a glass tube, or the sides of a glass rod immersed in the liquid. It is the simple case of a balance between three separate tensions: (1) that between the filament and the adjacent protoplasm, (2) that between the filament and the adjacent water, and (3) that between the water and the protoplasm. Calling these tensions respectively T_fp, T_fw, and T_wp, equi­lib­rium will be attained when the angle of contact between the fluid protoplasm and the filament is such that cos α = (T_fw − T_wp) ⁄ T_fp. It is evident in this case that the angle is a very small one. The precise form of the curve is somewhat different from that which, under ordinary circumstances, is assumed by a liquid which creeps up a solid surface, as water in contact with air creeps up a surface of glass; the difference being due to the fact that here, owing to the density of the protoplasm being practically identical with that of the surrounding medium, the whole system is practically immune from gravity. Under normal circumstances the curve is part of the “elastic curve” by which that surface of revolution is generated which we have called, after Plateau, the nodoid; but in the present case it is apparently a catenary. Whatever curve it be, it obviously forms a surface of revolution around the filament.

Since the attraction exercised by this surface tension is symmetrical around the filament, the latter will be pulled equally {266} in all directions; in other words it will tend to be set normally to the surface of the sphere, that is to say radiating directly outwards from the centre. If the distance between two adjacent filaments be considerable, the curve will simply meet the filament at the angle α already referred to; but if they be sufficiently near together, we shall have a continuous catenary curve forming a hanging loop between one filament and the other. And when this is so, and the radial filaments are more or less symmetrically interspaced, we may have a beautiful system of honeycomb-like depressions over the surface of the organism, each cell of the honeycomb having a strictly defined geometric configuration.

Fig. 90. A, Trypanosoma tineae (after Minchin); B, Spirochaeta anodontae (after Fantham).

In the simpler Radiolaria, the spherical form of the entire organism is equally well-marked; and here, as also in the more complicated Heliozoa (such as Actinosphaerium), the organism is differentiated into several distinct layers, each boundary surface tending to be spherical, and so constituting sphere within sphere. One of these layers at least is close packed with vacuoles, forming an “alveolar meshwork,” with the con­fi­gur­a­tions of which we shall attempt in another chapter to correlate the char­ac­ter­is­tic structure of certain complex types of skeleton.

An exceptional form of cell, but a beautiful manifestation of surface-tension (or so I take it to be), occurs in Trypanosomes, those tiny parasites of the blood that are associated with sleeping-sickness and many other grave or dire maladies. These tiny organisms consist of elongated solitary cells down one side of which runs a very delicate frill, or “undulating membrane,” the free edge of which is seen to be slightly thickened, and the whole of {267} which undergoes rhythmical and beautiful wavy movements. When certain Trypanosomes are artificially cultivated (for instance T. rotatorium, from the blood of the frog), phases of growth are witnessed in which the organism has no undulating membrane, but possesses a long cilium or “flagellum,” springing from near the front end, and exceeding the whole body in length313. Again, in T. lewisii, when it reproduces by “multiple fission,” the products of this division are likewise devoid of an undulating membrane, but are provided with a long free flagellum314. It is a plausible assumption to suppose that, as the flagellum waves about, it comes to lie near and parallel to the body of the cell, and that the frill or undulating membrane is formed by the clear, fluid protoplasm of the surface layer springing up in a film to run up and along the flagellum, just as a soap-film would be formed in similar circumstances.

Fig. 91. A, Trichomonas muris, Hartmann; B, Trichomastix serpentis, Dobell; C, Trichomonas angusta, Alexeieff. (After Kofoid.)

This mode of formation of the undulating membrane or frill appears to be confirmed by the appearances shewn in Fig. 91. {268} Here we have three little organisms closely allied to the ordinary Trypanosomes, of which one, Trichomastix (B), possesses four flagella, and the other two, Trichomonas, apparently three only: the two latter possess the frill, which is lacking in the first315. But it is impossible to doubt that when the frill is present (as in A and C), its outer edge is constituted by the apparently missing flagellum (a), which has become attached to the body of the creature at the point c, near its posterior end; and all along its course, the superficial protoplasm has been drawn out into a film, between the flagellum (a) and the adjacent surface or edge of the body (b).

Fig. 92. Her­pe­to­mo­nas as­sum­ing the un­du­la­tory mem­brane of a Try­pa­no­some. (After D. L. Mac­kin­non.)

Moreover, this mode of formation has been ac­tual­ly wit­nessed and de­scribed, though in a some­what ex­cep­tional case. The little fla­gel­late monad Her­pe­to­mo­nas is nor­mal­ly des­ti­tute of an un­du­la­ting membrane, but possesses a single long terminal flagellum. According to Dr D. L. Mackinnon, the cyto­plasm in a certain stage of growth becomes somewhat “sticky,” a phrase which we may in all probability interpret to mean that its surface tension is being reduced. For this stickiness is shewn in two ways. In the first place, the long body, in the course of its various bending movements, is apt to adhere head to tail (so to speak), giving a rounded or sometimes annular form to the organism, such as has also been described in certain species or stages of Trypanosomes. But again, the long flagellum, if it get bent backwards upon the body, tends to adhere to its surface. “Where the flagellum was pretty long and active, its efforts to continue movement under these abnormal conditions resulted in the gradual lifting up from the cytoplasm of the body of a sort of pseudo-undulating membrane (Fig. 92). The movements of this structure were so exactly those of a true undulating membrane that it was {269} difficult to believe one was not dealing with a small, blunt trypanosome316.” This in short is a precise description of the mode of development which, from theoretical con­si­de­ra­tions alone, we should conceive to be the natural if not the only possible way in which the undulating membrane could come into existence.

There is a genus closely allied to Trypanosoma, viz. Trypanoplasma, which possesses one free flagellum, together with an undulating membrane; and it resembles the neighbouring genus Bodo, save that the latter has two flagella and no undulating membrane. In like manner, Trypanosoma so closely resembles Herpetomonas that, when individuals ascribed to the former genus exhibit a free flagellum only, they are said to be in the “Herpetomonas stage.” In short all through the order, we have pairs of genera, which are presumed to be separate and distinct, viz. Trypanosoma-Herpetomonas, Trypanoplasma-Bodo, Trichomastix-Trichomonas, in which one differs from the other mainly if not solely in the fact that a free flagellum in the one is replaced by an undulating membrane in the other. We can scarcely doubt that the two structures are essentially one and the same.

The undulating membrane of a Trypanosome, then, according to our interpretation of it, is a liquid film and must obey the law of constant mean curvature. It is under curious limitations of freedom: for by one border it is attached to the comparatively motionless body, while its free border is constituted by a flagellum which retains its activity and is being constantly thrown, like the lash of a whip, into wavy curves. It follows that the membrane, for every alteration of its longitudinal curvature, must at the same instant become curved in a direction perpendicular thereto; it bends, not as a tape bends, but with the accompaniment of beautiful but tiny waves of double curvature, all tending towards the establishment of an “equipotential surface”; and its char­ac­ter­is­tic undulations are not originated by an active mobility of the membrane but are due to the molecular tensions which produce the very same result in a soap-film under similar circumstances.

In certain Spirochaetes, S. anodontae (Fig. 90) and S. balbiani {270} (which we find in oysters), a very similar undulating membrane exists, but it is coiled in a regular spiral round the body of the cell. It forms a “screw-surface,” or helicoid, and, though we might think that nothing could well be more curved, yet its math­e­mat­i­cal properties are such that it constitutes a “ruled surface” whose “mean curvature” is everywhere nil; and this property (as we have seen) it shares with the plane, and with the plane alone. Precisely such a surface, and of exquisite beauty, may be produced by bending a wire upon itself so that part forms an axial rod and part a spiral wrapping round the axis, and then dipping the whole into a soapy solution.

These undulating and helicoid surfaces are exactly reproduced among certain forms of spermatozoa. The tail of a spermatozoon consists normally of an axis surrounded by clearer and more fluid protoplasm, and the axis sometimes splits up into two or more slender filaments. To surface tension operating between these and the surface of the fluid protoplasm (just as in the case of the flagellum of the Trypanosome), I ascribe the formation of the undulating membrane which we find, for instance, in the spermatozoa of the newt or salamander; and of the helicoid membrane, wrapped in a far closer and more beautiful spiral than that which we saw in Spirochaeta, which is char­ac­ter­is­tic of the spermatozoa of many birds.

Before we pass from the subject of the conformation of the solitary cell we must take some account of certain other exceptional forms, less easy of explanation, and still less perfectly understood. Such is the case, for instance, with the red blood-corpuscles of man and other vertebrates; and among the sperm-cells of the decapod crustacea we find forms still more aberrant and not less perplexing. These are among the comparatively few cells or cell-like structures whose form seems to be incapable of explanation by theories of surface-tension.

In all the mammalia (save a very few) the red blood-corpuscles are flattened circular discs, dimpled in upon their two opposite sides. This configuration closely resembles that of an india-rubber ball when we pinch it tightly between finger and thumb; and we may also compare it with that experiment of Plateau’s {271} (described on p. 223), where a flat cylindrical oil-drop, of certain relative dimensions, can, by sucking away a little of the contained oil, be made to assume the form of a biconcave disc, whose periphery is part of a nodoidal surface. From the relation of the nodoid to the “elastic curve,” we perceive that these two examples are closely akin one to the other.

Fig. 93.

The form of the corpuscle is symmetrical, and its surface is a surface of revolution; but it is obviously not a surface of constant mean curvature, nor of constant pressure. For we see at once that, in the sectional diagram (Fig. 93), the pressure inwards due to surface tension is positive at A, and negative at C; at B there is no curvature in the plane of the paper, while perpendicular to it the curvature is negative, and the pressure therefore is also negative. Accordingly, from the point of view of surface tension alone, the blood-corpuscle is not a surface of equi­lib­rium; or in other words, it is not a fluid drop suspended in another liquid. It is obvious therefore that some other force or forces must be at work, and the simple effect of mechanical pressure is here excluded, because the blood-corpuscle exhibits its char­ac­ter­is­tic shape while floating freely in the blood. In the lower vertebrates the blood-corpuscles have the form of a flattened oval disc, with rather sharp edges and ellipsoidal surfaces, and this again is manifestly not a surface of equi­lib­rium.