Chapter V · THE FORMS OF CELLS (continued 2)

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).

To pass to a much more highly organised class of animals, we find the unduloid beautifully exemplified in the little flask-shaped shells of certain Pteropod mollusca, e.g. Cuvierina306. Here again the symmetry of the figure would at once lead us to suspect that the creature lived in a position of symmetry to the surrounding forces, as for instance if it floated in the ocean in an erect position, that is to say with its long axis coincident with the direction of gravity; and this we know to be actually the mode of life of the little Pteropod.

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}

In Mr Worthington’s experiment, there appears to be something of the nature of a viscous drag in the surface-pellicle; but whatever be the actual cause of variation of tension, it is not difficult to see that there must be in general a tendency towards longitudinal puckering or “fluting” in the case of a thin-walled cylindrical or other elongated body, rather than a tendency towards transverse puckering, or “pleating.” For let us suppose that some change takes place involving an increase of surface-tension in some small area of the curved wall, and leading therefore to an increase of pressure: that is to say let T become T + t, and P become P + p. Our new equation of equi­lib­rium, then, in place of P = T ⁄ r + T ⁄ r′ becomes

P + p = (T + t) ⁄ r + (T + t) ⁄ r′,

and by subtraction,

p = t ⁄ r + t ⁄ r′.

Now if

r < r′, t ⁄ r > t ⁄ r′.

Therefore, in order to produce the small increment of pressure p, it is easier to do so by increasing t ⁄ r than t ⁄ r′; that is to say, the easier way is to alter, or diminish r. And the same will hold good if the tension and pressure be diminished instead of increased.

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.

Two facts are especially noteworthy in connection with the form of the blood-corpuscle. In the first place, its form is only maintained, that is to say it is only in equi­lib­rium, in relation to certain properties of the medium in which it floats. If we add a little water to the blood, the corpuscle quickly loses its char­ac­ter­is­tic shape and becomes a spherical drop, that is to say a true surface of minimal area and of stable equi­lib­rium. If on the other hand we add a strong solution of salt, or a little glycerine, the corpuscle contracts, and its surface becomes puckered and uneven. In these phenomena it is so far obeying the laws of diffusion and of surface tension. {272}

In the second place, it can be exactly imitated artificially by means of other colloid substances. Many years ago Norris made the very interesting observation that in an emulsion of glue the drops assumed a biconcave form resembling that of the mammalian corpuscles317. The glue was impure, and doubtless contained lecithin; and it is possible (as Professor Waymouth Reid tells me) to make a similar emulsion with cerebrosides and cholesterin oleate, in which the same conformation of the drops or particles is beautifully shewn. Now such cholesterin bodies have an important place among those in which Lehmann and others have shewn and studied the formation of fluid crystals, that is to say of bodies in which the forces of cry­stal­li­sa­tion and the forces of surface tension are battling with one another318; and, for want of a better explanation, we may in the meanwhile suggest that some such cause is at the bottom of the conformation the explanation of which presents so many difficulties. But we must not, perhaps, pass from this subject without adding that the case is a difficult and complex one from the physiological point of view. For the surface of a blood-corpuscle consists of a “semi-permeable membrane,” through which certain substances pass freely and not others (for the most part anions and not cations), and it may be, accordingly, that we have in life a continual state of osmotic inequi­lib­rium, of negative osmotic tension within, to which comparatively simple cause the imperfect distension of the corpuscle may be also due319. The whole phenomenon would be comparatively easy to understand if we might postulate a stiffer peripheral region to the corpuscle, in the form for instance of a peripheral elastic ring. Such an annular thickening or stiffening, like the “collapse-rings” which an engineer inserts in a boiler, has been actually asserted to exist, but its presence is not authenticated.

But it is not at all improbable that we have still much to learn about the phenomena of osmosis itself, as manifested in the case of minute bodies such as a blood-corpuscle; and (as Professor Peddie suggests to me) it is by no means impossible that curvature {273} of the surface may itself modify the osmotic or perhaps the adsorptive action. If it should be found that osmotic action tended to stop, or to reverse, on change of curvature, it would follow that this phenomenon would give rise to internal currents; and the change of pressure consequent on these would tend to intensify the change of curvature when once started320.

Fig. 94. Sperm-cells of Decapod Crustacea (after Koltzoff). a, Inachus scorpio; b, Galathea squamifera; c, do. after maceration, to shew spiral fibrillae.

The sperm-cells of the Decapod crustacea exhibit various singular shapes. In the Crayfish they are flattened cells with stiff curved processes radiating outwards like a St Catherine’s wheel; in Inachus there are two such circles of stiff processes; in Galathea we have a still more complex form, with long and slightly twisted processes. In all these cases, just as in the case of the blood-corpuscle, the structure alters, and finally loses, its char­ac­ter­is­tic form when the nature or constitution (or as we may assume in particular—the density) of the surrounding medium is changed.

Here again, as in the blood-corpuscle, we have to do with a very important force, which we had not hitherto considered in this connection,—the force of osmosis, manifested under conditions similar to those of Pfeffer’s classical experiments on the plant-cell. The surface of the cell acts as a “semi-permeable membrane,” {274} permitting the passage of certain dissolved substances (or their “ions”) and including or excluding others; and thus rendering manifest and measurable the existence of a definite “osmotic pressure.” In the case of the sperm-cells of Inachus, certain quantitative experiments have been performed321. The sperm-cell exhibits its char­ac­ter­is­tic conformation while lying in the serous fluid of the animal’s body, in ordinary sea-water, or in a 5 per cent. solution of potassium nitrate; these three fluids being all “isotonic” with one another. As we alter the concentration of potassium nitrate, the cell assumes certain definite forms cor­re­spon­ding to definite concentrations of the salt; and, as a further and final proof that the phenomenon is entirely physical, it is found that other salts produce an identical effect when their concentration is proportionate to their molecular weight, and whatever identical effect is produced by various salts in their respective concentrations, a similarly identical effect is produced when these concentrations are doubled or otherwise proportionately changed322.

Fig. 95. Sperm-cells of Inachus, as they appear in saline solutions of varying density. (After Koltzoff.)

Thus the following table shews the percentage concentrations of certain salts necessary to bring the cell into the forms a and c of Fig. 95; in each case the quantities are proportional to the molecular weights, and in each case twice the quantity is necessary to produce the effect of Fig. 95c compared with that which gives rise to the all but spherical form of Fig. 95a. {275}

% concentration of salts in which the sperm-cell of Inachus assumes the form of

fig. a fig. c

Sodium chloride 0·6 1·2

Sodium nitrate 0·85 1·7

Potassium nitrate 1·0 2·0

Acetic acid 2·2 4·5

Cane sugar 5·0 10·0

If we look then, upon the spherical form of the cell as its true condition of symmetry and of equi­lib­rium, we see that what we call its normal appearance is just one of many intermediate phases of shrinkage, brought about by the abstraction of fluid from its interior as the result of an osmotic pressure greater outside than inside the cell, and where the shrinkage of volume is not kept pace with by a contraction of the surface-area. In the case of the blood-corpuscle, the shrinkage is of no great amount, and the resulting deformation is symmetrical; such structural inequality as may be necessary to account for it need be but small. But in the case of the sperm-cells, we must have, and we actually do find, a somewhat complicated arrangement of more or less rigid or elastic structures in the wall of the cell, which like the wire framework in Plateau’s experiments, restrain and modify the forces acting on the drop. In one form of Plateau’s experiments, instead of

Fig. 96. Sperm-cell of Dromia. (After Koltzoff.)

supporting his drop on rings or frames of wire, he laid upon its surface one or more elastic coils; and then, on withdrawing oil from the centre of his globule, he saw its uniform shrinkage counteracted by the spiral springs, with the result that the centre of each elastic coil seemed to shoot out into a prominence. Just such spiral coils are figured (after Koltzoff) in Fig. 96; and they may be regarded as precisely akin to those local thickenings, spiral and other, to which we have already ascribed the cylindrical form of the Spirogyra cell. In all probability we must in like manner attribute the peculiar spiral and other forms, for instance of many Infusoria, to the {276} presence, among the multitudinous other differentiations of their protoplasmic substance, of such more or less elastic fibrillae, which play as it were the part of a microscopic skeleton323.

But these cases which we have just dealt with, lead us to another consideration. In a semi-permeable membrane, through which water passes freely in and out, the conditions of a liquid surface are greatly modified; and, in the ideal or ultimate case, there is neither surface nor surface tension at all. And this would lead us somewhat to reconsider our position, and to enquire whether the true surface tension of a liquid film is actually responsible for all that we have ascribed to it, or whether certain of the phenomena which we have assigned to that cause may not in part be due to the contractility of definite and elastic membranes. But to in­ves­ti­gate this question, in particular cases, is rather for the physiologist: and the morphologist may go on his way, paying little heed to what is no doubt a difficulty. In surface tension we have the production of a film with the properties of an elastic membrane, and with the special peculiarity that contraction continues with the same energy however far the process may have already gone; while the ordinary elastic membrane contracts to a certain extent, and contracts no more. But within wide limits the essential phenomena are the same in both cases. Our fundamental equations apply to both cases alike. And accordingly, so long as our purpose is morphological, so long as what we seek to explain is regularity and definiteness of form, it matters little if we should happen, here or there, to confuse surface tension with elasticity, the contractile forces manifested at a liquid surface with those which come into play at the complex internal surfaces of an elastic solid.