On Growth and Form

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’s*. 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

* See Liquid Drops and Globules, 1914, p. 11. Robert Boyle used turpentine in much the same way. For other methods see Plateau, op. cit. p. 154. 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. 62 a); 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. 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 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 RF’ 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 2T/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.

The catenoid has the remarkable property that its curvature in one direction is precisely equal and opposite to its curvature in the other, this property holding good for all points of the surface. That is to say, R = — R’; and the expression becomes in other words, the surface, as in the case of the plane, has no curvature, and exercises no pressure. ‘There are no other surfaces, save these two, which share this remarkable property; and it follows, as a simple corollary, that we may expect at times to have the catenoid and the plane coexisting, as parts of one and the same boundary system; just as, in a cylindrical drop or cell, the cylinder is capped by portions of spheres, such that the cylindrical and spherical portions of the wall exert equal positive pressures.

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. 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

modified, and the cylindrical drop will assume the form of an unduloid (Fig. 64 a, B), with its dilated portion below or above, as the case may be; and our cylinder cases alike, the unduloid, like the A B original cylinder, will be capped by ae Oe 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. 658. This figure is a catenoid, which, as

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. 65 a).

It represents, in fact, that portion of the nodoid, which in Fig. 66 lies between such points as 0, Pp. While it is easy to draw the outline, or meridional - 0 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 dises 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 Nn. 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

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 prelimimary 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

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 (7') 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 (1/7 + 1/r’), in the case of the terminal spheres. Butin the cylinder 1/R’ = 0,and inthesphere 1/r = 1/r’. Therefore our relation of equality becomes 1/R = 2/r, or r = 2R; that is to say, the sphere in question has just twice the radius of the cylinder of which it forms a cap.

And if Ob, the radius of the sphere, be equal to twice the radius (Oa) of the cylinder, it follows that the angle aOb is an angle of ; 60°, and bOc is also an angle of 60°; that is to say, the are be is equal to ia. In other words, the spherical qi b — dise which (under the given conditions) caps our cylinder, is not a portion taken at haphazard, but is neither more nor less than that portion of a sphere which is subtended by a cone of 60°. Moreover, it is plain that the height of the spherical cap, de,

where R is the radius of our cylinder, or one-half the radius of our spherical cap: in other words the normal height of the spherical cap over the end of the cylindrical cell is just a very little more than oneeighth of the diameter of the cylinder, or of the radius of the sphere. And these are the proportions which we recognise, under normal circumstances, in such a case as the cylindrical cell of Spirogyra where its free end is capped by a portion of a sphere.

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

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 = 27R, or when the ratio of length to diameter is represented by z.

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 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 theoil-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 cohvex, 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; 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. 65 A).

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 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 Spirocheetes, 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 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 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 result*; 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