Chapter IX: On Concretions, Spicules, and Spicular Skeletons (2)

Furthermore, if these mechanical methods of conformation, and others like to these, be the true cause of the shapes which the spicules assume, it is plain that the production of these spicular shapes is not a specific function of sponges or of any particular sponge, but that we should expect {443} the same or very similar phenomena to occur in other organisms, wherever the conditions of inorganic secretion within closed cells was very much the same. As a matter of fact, in the group of Holothuroidea, where the formation of intracellular spicules is a characteristic feature of the group, all the principal types of conformation which we have just described can be closely paralleled. Indeed in many cases, the forms of the Holothurian spicules are identical and indistinguishable from those of the sponges[458]. But the Holothurian spicules are composed of calcium carbonate while those which we have just described in the case of sponges are usually, if not always, siliceous: this being just another proof of the fact that in such cases the form of the spicule is not due to its chemical nature or molecular structure, but to the external forces to which, during its growth, the spicule is submitted.

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So much for that comparatively limited class of sponge-spicules whose forms seem capable of explanation on the hypothesis that they are developed within, or under the restraint imposed by, the surface of a cell or vesicle. Such spicules are usually of small size, as well as of comparatively simple form; and they are greatly outstripped in number, in size, and in supposed importance as guides to zoological classification, by another class of spicules. This new class includes such as we have supposed to be capable of explanation on the assumption that they develop in association (of some sort or another) with the _lines of junction_ of contiguous cells. They include the triradiate spicules of the calcareous sponges, the quadriradiate or “tetractinellid” spicules which occur in the same group, but more characteristically in certain siliceous sponges known as the Tetractinellidae, and lastly perhaps (though these last are admittedly somewhat harder to understand) the six-rayed spicules of the Hexactinellids.

The spicules of the calcareous sponges are commonly triradiate, and the three radii are usually inclined to one another at equal, or nearly equal angles; in certain cases, two of the three rays are nearly in a straight line, and at right angles to the {444} third[459]. They are seldom in a plane, but are usually inclined to one another in a solid, trihedral angle, not easy of precise measurement under the microscope. The three rays are very often supplemented by a fourth, which is set tetrahedrally, making, that is to say, coequal angles with the other three. The calcareous spicule consists mainly of carbonate of lime, in the form of calcite, with (according to von Ebner) some admixture of soda and magnesia, of sulphates and of water. According to the same writer (but the fact, though it would seem easy to test, is still disputed) there is no organic matter in the spicule, either in the form of an axial filament or otherwise, and the appearance of stratification, often simulating the presence of an axial fibre, is due to “mixed crystallisation” of the various constituents. The spicule is a true crystal, and therefore its existence and its form are _primarily_ due to the molecular forces of crystallisation; moreover it is a single crystal and not a group of crystals, as is at once seen by its behaviour in polarised light. But its axes are not crystalline axes, and its form neither agrees with, nor in any way resembles, any one of the many polymorphic forms in which calcite is capable of crystallising. It is as though it were carved out of a solid crystal; it is, in fact, a crystal under restraint, a crystal growing, as it were, in an artificial mould; and this mould is constituted by the surrounding cells, or structural vesicles of the sponge.

We have already studied in an elementary way, but amply for our present purpose, the manner in which three or more cells, or bubbles, tend to meet together under the influence of surface-tension, and also the outwardly similar phenomena which may be brought about by a uniform distribution of mechanical pressure. We have seen that when we confine ourselves to a plane assemblage of such bodies, we find them meeting one another in threes; that in a section or plane projection of such an assemblage we see the partition-walls meeting one another at equal angles of 120°; that when the bodies are uniform in size, the partitions are straight lines, which combine to form regular hexagons; and that when {445} the bodies are unequal in size, the partitions are curved, and combine to form other and less regular polygons. It is plain, accordingly, that in any flattened or stratified assemblage of such cells, a solidified skeletal deposit which originates or accumulates either between the cells or within the thickness of their mutual partitions, will tend to take the form of triradiate bodies, whose rays (in a typical case) will be set at equal angles of 120° (Fig. 214, _F_). And this latter condition of equality will be open to modification in various ways. It will be modified by any inequality in the specific tensions of adjacent cells; as a special case, it will be apt to be greatly modified at the surface of the system, where a spicule happens to be formed in a plane perpendicular to the cell-layer, so that one of its three rays lies between two adjacent cells and the other two are associated with the surface of contact between the cells and the surrounding medium; in such a case (as in the cases considered in connection with the forms of the cells themselves {446} on p. 314), we shall tend to obtain a spicule with two equal angles and one unequal (Fig. 214, _A_, _C_). In the last case, the two outer, or superficial rays, will tend to be markedly curved. Again, the equiangular condition will be departed from, and more or less curvature will be imparted to the rays, wherever the cells of the system cease to be uniform in size, and when the hexagonal symmetry of the system is lost accordingly. Lastly, although we speak of the rays as meeting at certain definite angles, this statement applies to their _axes_, rather than to the rays themselves. For, if the triradiate spicule be developed in the _interspace_ between three juxtaposed cells, it is obvious that its sides will tend to be concave, for the interspace between our three contiguous equal circles is an equilateral, curvilinear triangle; and even if our spicule be deposited, not in the space between our three cells, but in the thickness of the intervening wall, then we may recollect (from p. 297) that the several partitions never actually meet at sharp angles, but the angle of contact is always bridged over by a small accumulation of material (varying in amount according to its fluidity) whose boundary takes the form of a circular arc, and which constitutes the “bourrelet” of Plateau.

In any sample of the triradiate spicules of Grantia, or in any series of careful drawings, such as those of Haeckel among others, we shall find that all these various configurations are precisely and completely illustrated.

The tetrahedral, or rather tetractinellid, spicule needs no explanation in detail (Fig. 214, _D_, _E_). For just as a triradiate spicule corresponds to the case of three cells in mutual contact, so does the four-rayed spicule to that of a solid aggregate of four cells: these latter tending to meet one another in a tetrahedral system, shewing four edges, at each of which four surfaces meet, the edges being inclined to one another at equal angles of about 109°. And even in the case of a single layer, or superficial layer, of cells, if the skeleton originate in connection with all the edges of mutual contact, we shall, in complete and typical cases, have a four-rayed spicule, of which one straight limb will correspond to the line of junction between the three cells, and the other three limbs (which will then be curved limbs) will correspond to the edges where two cells meet one another on the surface of the system. {447}

But if such a physical explanation of the forms of our spicules is to be accepted, we must seek at once for some physical agency by which we may explain the presence of the solid material just at the junctions or interfaces of the cells, and for the forces by which it is confined to, and moulded to the form of, these intercellular or interfacial contacts. It is to Dreyer that we chiefly owe the physical or mechanical theory of spicular conformation which I have just described,—a theory which ultimately rests on the form assumed, under surface-tension, by an aggregation of cells or vesicles. But this fundamental point being granted, we have still several possible alternatives by which to explain the details of the phenomenon.

Dreyer, if I understand him aright, was content to assume that the solid material, secreted or excreted by the organism, accumulated in the interstices between the cells, and was there subjected to mechanical pressure or constraint as the cells got more and more crowded together by their own growth and that of the system generally. As far as the general form of the spicules goes, such explanation is not inadequate, though under it we may have to renounce some of our assumptions as to what takes place at the outer surface of the system.

But in all (or most) cases where, but a few years ago, the concepts of secretion or excretion seemed precise enough, we are now-a-days inclined to turn to the phenomenon of adsorption as a further stage towards the elucidation of our facts. Here we have a case in point. In the tissues of our sponge, wherever two cells meet, there we have a definite _surface_ of contact, and there accordingly we have a manifestation of surface-energy; and the concentration of surface-energy will tend to be a maximum at the _lines_ or edges whereby the three, or four, such surfaces are conjoined. Of the micro-chemistry of the sponge-cells our ignorance is great; but (without venturing on any hypothesis involving the chemical details of the process) we may safely assert that there is an inherent probability that certain substances will tend to be concentrated and ultimately deposited just in these lines of intercellular contact and conjunction. In other words, adsorptive concentration, under osmotic pressure, at and in the surface-film which constitutes the mutual boundary between contiguous {448} cells, emerges as an alternative (and, as it seems to me, a highly preferable alternative) to Dreyer’s conception of an accumulation under mechanical pressure in the vacant spaces left between one cell and another.

But a purely chemical, or purely molecular adsorption, is not the only form of the hypothesis on which we may rely. For from the purely physical point of view, angles and edges of contact between adjacent cells will be _loci_ in the field of distribution of surface-energy, and any material particles whatsoever will tend to undergo a diminution of freedom on entering one of those boundary regions. In a very simple case, let us imagine a couple of soap bubbles in contact with one another. Over the surface of each bubble there glide in every direction, as usual, a multitude of tiny bubbles and droplets; but as soon as these find their way into the groove or re-entrant angle between the two bubbles, there their freedom of movement is so far restrained, and out of that groove they have little or no tendency to emerge. A cognate phenomenon is to be witnessed in microscopic sections of steel or other metals. Here, amid the “crystalline” structure of the metal (where in cooling its imperfectly homogeneous material has developed a cellular structure, shewing (in section) hexagonal or polygonal contours), we can easily observe, as Professor Peddie has shewn me, that the little particles of graphite and other foreign bodies common in the matrix, have tended to aggregate themselves in the walls and at the angles of the polygonal cells—this being a direct result of the diminished freedom which the particles undergo on entering one of these boundary regions[460].

It is by a combination of these two principles, chemical adsorption on the one hand, and physical quasi-adsorption or concentration of grosser particles on the other, that I conceive the substance of the sponge-spicule to be concentrated and aggregated at the cell boundaries; and the forms of the triradiate and tetractinellid spicules are in precise conformity with this hypothesis. A few general matters, and a few particular cases, remain to be considered.

It matters little or not at all, for the phenomenon in question, {449} what is the histological nature or “grade” of the vesicular structures on which it depends. In some cases (apart from sponges), they may be no more than the little alveoli of the intracellular protoplasmic network, and this would seem to be the case at least in one known case, that of the protozoan _Entosolenia aspera_, in which, within the vesicular protoplasm of the single cell, Möbius has described tiny spicules in the shape of little tetrahedra with concave sides. It is probably also the case in the small beginnings of the Echinoderm spicules, which are likewise intracellular, and are of similar shape. In the case of our sponges we have many varying conditions, which we need not attempt to examine in detail. In some cases there is evidence for believing that the spicule is formed at the boundaries of true cells or histological units. But in the case of the larger triradiate or tetractinellid spicules of the sponge-body, they far surpass in size the actual “cells”; we find them lying, regularly and symmetrically arranged, between the “pore-canals” or “ciliated chambers,” and it is in conformity with the shape and arrangement of these rounded or spheroidal structures that their shape is assumed.

Again, it is not necessarily at variance with our hypothesis to find that, in the adult sponge, the larger spicules may greatly outgrow the bounds not only of actual cells but also of the ciliated chambers, and may even appear to project freely from the surface of the sponge. For we have already seen that the spicule is capable of growing, without marked change of form, by further deposition, or crystallisation, of layer upon layer of calcareous molecules, even in an artificial solution; and we are entitled to believe that the same process may be carried on in the tissues of the sponge, without greatly altering the symmetry of the spicule, long after it has established its characteristic form of a system of slender trihedral or tetrahedral rays.

Neither is it of great importance to our hypothesis whether the rayed spicule necessarily arises as a single structure, or does so from separate minute centres of aggregation. Minchin has shewn that, in some cases at least, the latter is the case; the spicule begins, he tells us, as three tiny rods, separate from one another, each developed in the interspace between two sister-cells, which are themselves the results of the division of one of a {450} little trio of cells; and the little rods meet and fuse together while still very minute, when the whole spicule is only about 1/200 of a millimetre long. At this stage, it is interesting to learn that the spicule is non-crystalline; but the new accretions of calcareous matter are soon deposited in crystalline form.

This observation threw considerable difficulties in the way of former mechanical theories of the conformation of the spicule, and was quite at variance with Dreyer’s theory, according to which the spicule was bound to begin from a central nucleus coinciding with the meeting-place of the three contiguous cells, or rather the interspace between them. But the difficulty is removed when we import the concept of adsorption; for by this agency it is natural enough, or conceivable enough, that the process of deposition should go on at separate parts of a common system of surfaces; and if the cells tend to meet one another by their interfaces before these interfaces extend to the angles and so complete the polygonal cell, it is again conceivable and natural that the spicule should first arise in the form of separate and detached limbs or rays.

Among the tetractinellid sponges, whose spicules are composed of amorphous silica or opal, all or most of the above-described main types of spicule occur, and, as the name of the group implies, the four-rayed, tetrahedral spicules are especially represented. A somewhat frequent type of spicule is one in which one of the four rays is greatly developed, and the other three constitute small prongs diverging at equal angles from the main or axial ray. In all probability, as Dreyer suggests, we have here had to do with a group of four vesicles, of which three were large and co-equal, while a fourth and very much smaller one lay above and between the other three. In certain cases where we have likewise one large and three much smaller {451} rays, the latter are recurved, as in Fig. 215. This type, save for the constancy of the number of rays, and the limitation of the terminal ones to three, and save also for the more important difference that they occur only at one and not at both ends of the long axis, is similar to the type of spicule illustrated in Fig. 213, which we have explained as being probably developed within an oval cell, by whose walls its branches have been conformed to geodetic curves. But it is much more probable that we have here to do with a spicule developed in the midst of a group of three coequal and more or less elongated or cylindrical cells or vesicles, the long axial ray corresponding to their common line of contact, and the three short rays having each lain in the surface furrow between two out of the three adjacent cells.

Just as in the case of the little curved or S-shaped spicules, formed apparently within the bounds of a single cell, so also in the case of the larger tetractinellid and analogous types do we find among the Holothuroidea the same configurations reproduced as we have dealt with in the sponges. The holothurian spicules are a little less neatly formed, a little rougher, than the sponge-spicules; and certain forms occur among the former group which do not present themselves among the latter; but for the most part a community of type is obvious and striking (Fig. 216).

A curious and, physically speaking, strictly analogous formation to the tetrahedral spicules of the sponges is found in the {452} spores of a certain little group of parasitic protozoa, the Actinomyxidia. These spores are formed from clusters of six cells, of which three come to constitute the capsule of the spore; and this capsule, always triradiate in its symmetry, is in some species drawn out into long rays, of which one constitutes a straight central axis, while the others, coming off from it at equal angles, are recurved in wide circular arcs. The account given of the development of this structure by its discoverers[461] is somewhat obscure to me, but I think that, on physical grounds, there can be no doubt whatever that the quadriradiate capsule has been somehow modelled upon a group of three surrounding cells, its axis lying between the three, and its three radial arcs occupying the furrows between adjacent pairs.

The typically six-rayed siliceous spicules of the hexactinellid sponges, while they are perhaps the most regular and beautifully formed spicules to be found within the entire group, have been found very difficult to explain, and Dreyer has confessed his complete inability to account for their conformation. But, though it is doubtless only throwing the difficulty a little further back, we may so far account for them by considering that the cells or vesicles by which they are conformed are not arranged in {453} what is known as “closest packing,” but in linear series; so that in their arrangement, and by their mutual compression, we tend to get a pattern, not of hexagons, but of squares: or, looking to the solid, not of dodecahedra but of cubes or parallelopipeda. This indeed appears to be the case, not with the individual cells (in the histological sense), but with the larger units or vesicles which make up the body of the hexactinellid. And this being so, the spicules formed between the linear, or cubical series of vesicles, will have the same tendency towards a “hexactinellid” shape, corresponding to the angles and adjacent edges of a system of cubes, as in our former case they had to a triradiate or a tetractinellid form, when developed in connection with the angles and edges of a system of hexagons, or a system of dodecahedra.

Histologically, the case is illustrated by a well-known phenomenon in embryology. In the segmenting ovum, there is a tendency for the cells to be budded off in linear series; and so they often remain, in rows side by side, at least for a considerable time and during the course of several consecutive cell divisions. Such an arrangement constitutes what the embryologists call the “radial type” of segmentation[462]. But in what is described as the “spiral type” of segmentation, it is stated that, as soon as the first horizontal furrow has divided the cells into an upper and a lower layer, those of “the upper layer are shifted in respect to the lower layer, by means of a rotation about the vertical axis[463].” It is, of course, evident that the whole process is merely that which is familiar to physicists as “close packing.” It is a very simple case of what Lord Kelvin used to call “a problem in tactics.” It is a mere question of the rigidity of the system, of the freedom of movement on the part of its constituent cells, whether or at what stage this tendency to slip into the closest propinquity, or position of minimum potential, will be found to manifest itself.

However the hexactinellid spicules be arranged (and this is {454} not at all easy to determine) in relation to the tissues and chambers of the sponge, it is at least clear that, whether they be separate or be fused together (as often happens) in a composite skeleton, they effect a symmetrical partitioning of space according to the cubical system, in contrast to that closer packing which is represented and effected by the tetrahedral system[464].

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This question of the origin and causation of the forms of sponge-spicules, with which we have now briefly dealt, is all the more important and all the more interesting because it has been discussed time and again, from points of view which are characteristic of very different schools of thought in biology. Haeckel found in the form of the sponge-spicule a typical illustration of his theory of “bio-crystallisation”; he considered that these “biocrystals” represented “something midway—_ein Mittelding_—between an inorganic crystal and an organic secretion”; that there was a “compromise between the crystallising efforts of the calcium carbonate and the formative activity of the fused cells of the syncytium”; and that the semi-crystalline secretions of calcium carbonate “were utilised by natural selection as ‘spicules’ for building up a skeleton, and afterwards, by the interaction of adaptation and heredity, became modified in form and differentiated in a vast variety of ways in the struggle for existence[465].” What Haeckel precisely signified by these words is not clear to me.

F. E. Schultze, perceiving that identical forms of spicule were developed whether the material were crystalline or non-crystalline, abandoned all theories based upon crystallisation; he simply saw in the form and arrangement of the spicules something which was “best fitted” for its purpose, that is to say for the support and strengthening of the porous walls of the sponge, and found clear evidence of “utility” in the specific structure of these skeletal elements. {455}

Sollas and Dreyer, as we have seen, introduced in various ways the conception of physical causation,—as indeed Haeckel himself had done in regard to one particular, when he supposed the _position_ of the spicules to be due to the constant passage of the water-currents. Though even here, by the way, if I understand Haeckel aright, he was thinking not merely of a direct or immediate physical causation, but of one manifesting itself through the agency of natural selection[466]. Sollas laid stress upon the “path of least resistance” as determining the direction of growth; while Dreyer dealt in greater detail with the various tensions and pressures to which the growing spicule was exposed, amid the alveolar or vesicular structure which was represented alike by the chambers of the sponge, by the reticulum of constituent cells, or by the minute structure of the intracellular protoplasm. But neither of these writers, so far as I can discover, was inclined to doubt for a moment the received canon of biology, which sees in such structures as these the characteristics of true organic species, and the indications of an hereditary affinity by which blood-relationship and the succession of evolutionary descent throughout geologic time can be ultimately deduced.

Lastly, Minchin, in a well-known paper[467], took sides with Schultze, and gave reasons for dissenting from such mechanical theories as those of Sollas and of Dreyer. For example, after pointing out that all protoplasm contains a number of “granules” or microsomes, contained in the alveolar framework and lodged at the nodes of the reticulum, he argued that these also ought to acquire a form such as the spicules possess, if it were the case that these latter owed their form to their very similar or identical position. “If vesicular tension cannot in any other instance cause the granules at the nodes to assume a tetraxon form, why should it do so for the sclerites?” In all probability the answer to this question is not far to seek. If the force which the “mechanical” hypothesis has in view were simply that of mechanical _pressure_, {456} as between solid bodies, then indeed we should expect that any substances whatsoever, lying between the impinging spheres, would tend (unless they were infinitely hard) to assume the quadriradiate or “tetraxon” form; but this conclusion does not follow at all, in so far as it is to _surface-energy_ that we ascribe the phenomenon. Here the specific nature of the substances involved makes all the difference. We cannot argue from one substance to another; adsorptive attraction shews its effect on one and not on another; and we have not the least reason to be surprised if we find that the little granules of protoplasmic material, which as they lie bathed in the more fluid protoplasm have (presumably, and as their shape indicates) a strong surface-tension of their own, behave towards the adjacent vesicles in a very different fashion to the incipient aggregations of calcareous or siliceous matter in a colloid medium. “The ontogeny of the spicules,” says Professor Minchin, “points clearly to their regular form being a _phylogenetic adaptation, which has become fixed and handed on by heredity, appearing in the ontogeny as a prophetic adaptation_.” And again, “The forms of the spicules are the result of adaptation to the requirements of the sponge as a whole, produced by _the action of natural selection upon variation in every direction_.” It would scarcely be possible to illustrate more briefly and more cogently than by these few words (or the similar words of Haeckel quoted on p. 454), the fundamental difference between the Darwinian conception of the causation and determination of Form, and that which is characteristic of the physical sciences.

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If I have dealt comparatively briefly with the inorganic skeleton of sponges, in spite of the obvious importance of this part of our subject from the physical or mechanical point of view, it has been owing to several reasons. In the first place, though the general trend of the phenomena is clear, it must be at once admitted that many points are obscure, and could only be discussed at the cost of a long argument. In the second place, the physical theory is (as I have shewn) in manifest conflict with the accounts given by various embryologists of the development of the spicules, and of the current biological theories which their descriptions embody; it is beyond our scope to deal with such descriptions {457} in detail. Lastly, we find ourselves able to illustrate the same physical principles with greater clearness and greater certitude in another group of animals, namely the Radiolaria. In our description of the skeletons occurring within this group we shall by no means abandon the preliminary classification of microscopic skeletons which we have laid down; but we shall have occasion to blend with it the consideration of certain other more or less correlated phenomena.

The group of microscopic organisms known as the Radiolaria is extraordinarily rich in diverse forms, or “species.” I do not know how many of such species have been described and defined by naturalists, but some thirty years ago the number was said to be over four thousand, arranged in more than seven hundred genera[468]. Of late years there has been a tendency to reduce the number, it being found that some of the earlier species and even genera are but growth-stages of one and the same form, sometimes mere fragments or “fission-products” common to several species, or sometimes forms so similar and so interconnected by intermediate forms that the naturalist denominates them not “species” but “varieties.” It has to be admitted, in short, that the conception of species among the Radiolaria has not hitherto been, and is not yet, on the same footing as that among most other groups of animals. But apart from the extraordinary multiplicity of forms among the Radiolaria, there are certain other features in this multiplicity which arrest our attention. For instance, the distribution of species in space is curious and vague; many species are found all over the world, or at least every here and there, with no evidence of specific limitations of geographical habitat; others occur in the neighbourhood of the two poles; some are confined to warm and others to cold currents of the ocean. In time also their distribution is not less vague: so much so that it has been asserted of them that “from the Cambrian age downwards, the families and even genera appear identical with those now living.” Lastly, except perhaps in the case of a few large “colonial forms,” we seldom if ever find, as is usual {458} in most animals, a local predominance of one particular species. On the contrary, in a little pinch of deep-sea mud or of some fossil “Radiolarian earth,” we shall probably find scores, and it may be even hundreds, of different forms. Moreover, the radiolarian skeletons are of quite extraordinary delicacy and complexity, in spite of their minuteness and the comparative simplicity of the “unicellular” organisms within which they grow; and these complex conformations have a wonderful and unusual appearance of geometric regularity. All these _general_ considerations seem such as to prepare us for the special need of some physical hypothesis of causation. The little skeletal fabrics remind us of such objects as snow-crystals (themselves almost endless in their diversity), rather than of a collection of distinct animals, constructed in apparent accordance with functional needs, and distributed in accordance with their fitness for particular situations. Nevertheless great efforts have been made of recent years to attach “a biological meaning” to these elaborate structures; and “to justify the hope that in time the utilitarian character [of the skeleton] will be more completely recognised[469].”

In the majority of cases, the skeleton of the Radiolaria is composed, like that of so many sponges, of silica; in one large family, the Acantharia (and perhaps in some others), it is composed, in great part at least, of a very unusual constituent, namely strontium sulphate[470]. There is no fundamental or important morphological character in which the shells formed of these two constituents differ from one another; and in no case can the chemical properties of these inorganic materials be said to influence the form of the complex skeleton or shell, save only in this general way that, by their rigidity and toughness, they may give rise to a fabric far more delicate and slender than we find developed among calcareous organisms.

A slight exception to this rule is found in the presence of true crystals, which occur within the central capsules of certain {459} Radiolaria, for instance the genus Collosphaera[471]. Johannes Müller (whose knowledge and insight never fail to astonish us) remarked that these were identical in form with crystals of celestine, a sulphate of strontium and barium; and Bütschli’s discovery of sulphates of strontium and of barium in kindred forms render it all but certain that they are actually true crystals of celestine[472].

In its typical form, the Radiolarian body consists of a spherical mass of protoplasm, around which, and separated from it by some sort of porous “capsule,” lies a frothy mass, composed of protoplasm honeycombed into a multitude of alveoli or vacuoles, filled with a fluid which can scarcely differ much from sea-water[473]. According to their surface-tension conditions, these vacuoles may appear more or less isolated and spherical, or joining together in a “froth” of polygonal cells; and in the latter, which is the commoner condition, the cells tend to be of equal size, and the resulting polygonal meshwork beautifully regular. In many cases, a large number of such simple individual organisms are associated together, forming a floating colony, and it is highly probable that many other forms, with whose scattered skeletons we are alone acquainted, had in life formed part likewise of a colonial organism.

In contradistinction to the sponges, in which the skeleton always begins as a loose mass of isolated spicules, which only in a few exceptional cases (such as Euplectella and Farrea) fuse into a continuous network, the characteristic feature of the Radiolarians lies in the possession of a continuous skeleton, in the form of a netted mesh or perforated lacework, sometimes however replaced by and often associated with minute independent spicules. Before we proceed to treat of the more complex skeletons, we may begin, then, by dealing with these comparatively simple cases where either the entire skeleton or a considerable part of it is represented, not by a continuous fabric, but by a quantity of loose, separate spicules, or aciculae, which seem, like the spicules of Alcyonium, {460} to be developed as free and isolated formations or deposits, precipitated in the colloid matrix, with no relation of form to the cellular or vesicular boundaries. These simple acicular spicules occupy a definite position in the organism. Sometimes, as for instance among the fresh-water Heliozoa (e.g. Raphidiophrys), they lie on the outer surface of the organism, and not infrequently (when the spicules are few in number) they tend to collect round the bases of the pseudopodia, or around the large radiating spicules, or axial rays, in the cases where these latter are present. When the spicules are thus localised around some prominent centre, they tend to take up a position of symmetry in regard to it; instead of forming a tangled or felted layer, they come to lie side by side, in a radiating cluster round the focus. In other cases (as for instance in the well-known Radiolarian _Aulacantha scolymantha_) the felted layer of aciculae lies at some depth below the surface, forming a sphere concentric with the entire spherical organism. In either case, whether the layer of spicules be deep or be superficial, it tends to mark a “surface of discontinuity,” a meeting place between two distinct layers of protoplasm or between the protoplasm and the water around; and it is obvious that, in either case, there are manifestations of surface-energy at the boundary, which cause the spicules to be retained there, and to take up their position in its plane. The case is somewhat, though not directly, analogous to that of a cirrus cloud, which marks the place of a surface of discontinuity in a stratified atmosphere.

We have, then, to enquire what are the conditions which shall, apart from gravity, confine an extraneous body to a surface-film; and we may do this very simply, by considering the surface-energy of the entire system. In Fig. 218 we have two fluids in contact with one another (let us call them water and protoplasm), and a body (_b_) which may be immersed in either, or may be restricted to the boundary {461} between. We have here three possible “interfacial contacts” each with its own specific surface-energy, per unit of surface area: namely, that between our particle and the water (let us call it α), that between the particle and the protoplasm (β), and that between water and protoplasm (γ). When the body lies in the boundary of the two fluids, let us say half in one and half in the other, the surface-energies concerned are equivalent to (_S_/2)α + (_S_/2)β; but we must also remember that, by the presence of the particle, a small portion (equal to its sectional area _s_) of the original contact-surface between water and protoplasm has been obliterated, and with it a proportionate quantity of energy, equivalent to _s_γ, has been set free. When, on the other hand, the body lies entirely within one or other fluid, the surface-energies of the system (so far as we are concerned) are equivalent to _S_α + _s_γ, or _S_β + _s_γ, as the case may be. According as α be less or greater than β, the particle will have a tendency to remain immersed in the water or in the protoplasm; but if (_S_/2)(α + β) − _s_γ be less than either _S_α or _S_β, then the condition of minimal potential will be found when the particle lies, as we have said, in the boundary zone, half in one fluid and half in the other; and, if we were to attempt a more general solution of the problem, we should evidently have to deal with possible conditions of equilibrium under which the necessary balance of energies would be attained by the particle rising or sinking in the boundary zone, so as to adjust the relative magnitudes of the surface-areas concerned. It is obvious that this principle may, in certain cases, help us to explain the position even of a _radial_ spicule, which is just a case where the surface of the solid spicule is distributed between the fluids with a minimal disturbance, or minimal replacement, of the original surface of contact between the one fluid and the other.

In like manner we may provide for the case (a common and an important one) where the protoplasm “creeps up” the spicule, covering it with a delicate film. In Acanthocystis we have yet another special case, where the radial spicules plunge only a certain distance into the protoplasm of the cell, being arrested at a boundary-surface between an inner and an outer layer of cytoplasm; here we have only to assume that there is a tension {462} at this surface, between the two layers of protoplasm, sufficient to balance the tensions which act directly on the spicule[474].

In various Acanthometridae, besides such typical characters as the radial symmetry, the concentric layers of protoplasm, and the capillary surfaces in which the outer, vacuolated protoplasm is festooned upon the projecting radii, we have another curious feature. On the surface of the protoplasm where it creeps up the sides of the long radial spicules, we find a number of elongated bodies, forming in each case one or several little groups, and lying neatly arranged in parallel bundles. A Russian naturalist, Schewiakoff, whose views have been accepted in the text-books, tells us that these are muscular structures, serving to raise or lower the conical masses of protoplasm about the radial spicules, which latter serve as so many “tent-poles” or masts, on which the protoplasmic membranes are hoisted up; and the little elongated bodies are dignified with various names, such as “myonemes” or “myophriscs,” in allusion to their supposed muscular nature[475]. This explanation is by no means convincing. To begin with, we have precisely similar festoons of protoplasm in a multitude of other cases where the “myonemes” are lacking; from their minute size (·006–·012 mm.) and the amount of contraction they are said to be capable of, the myonemes can hardly be very efficient instruments of traction; and further, for them to act (as is alleged) for a specific purpose, namely the “hydrostatic regulation” of the organism giving it power to sink or to swim, would seem to imply a mechanism of action and of coordination which is difficult to conceive in these minute and simple organisms. The fact is (as it seems to me), that the whole method of explanation is unnecessary. Just as the supposed “hauling up” of the protoplasmic festoons is at once explained by capillary phenomena, so also, in all probability, is the position and arrangement of the little elongated bodies. Whatever the actual nature of these bodies may be, whether they are truly portions of differentiated protoplasm, or whether they are foreign bodies or spicular structures (as bodies occupying a similar position in other cases undoubtedly are), we can explain their situation on the surface {463} of the protoplasm, and their arrangement around the radial spicules, all on the principles of surface-tension[476].

This last case is not of the simplest; and I do not forget that my explanation of it, which is wholly theoretical, implies a doubt of Schewiakoff’s statements, which are founded on direct personal observation. This I am none too willing to do; but whether it be justly done in this case or not, I hold that it is in principle justifiable to look with great suspicion upon a number of kindred statements where it is obvious that the observer has left out of account the purely physical aspect of the phenomenon, and all the opportunities of simple explanation which the consideration of that aspect might afford.

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Whether it be wholly applicable to this particular and complex case or no, our general theorem of the localisation and arrestment of solid particles in a surface-film is of very great biological importance; for on it depends the power displayed by many little naked protoplasmic organisms of covering themselves with an “agglutinated” shell. Sometimes, as in _Difflugia_, _Astrorhiza_ (Fig. 219) and others, this covering consists of sand-grains picked up from the surrounding medium, and sometimes, on the other hand, as in _Quadrula_, it consists of solid particles which are said to arise, as inorganic deposits or concretions, within the protoplasm itself, and which find their way outwards to a position of equilibrium in the surface-layer; and in both cases, the mutual capillary attractions between the particles, confined to the boundary-layer but enjoying a certain measure of freedom therein, tends to the orderly arrangement of the particles one with another, and even to the appearance of a regular “pattern” as the result of this arrangement.

The “picking up” by the protoplasmic organism of a solid particle with which “to build its house” (for it is hard to avoid this customary use of anthropomorphic figures of speech, misleading though they be), is a physical phenomenon kindred to that by which an Amoeba “swallows” a particle of food. This latter process has been reproduced or imitated in various pretty experimental {465} ways. For instance, Rhumbler has shewn that if a thread of glass be covered with shellac and brought near a drop of chloroform suspended in water, the drop takes in the spicule, robs it of its shellac covering, and then passes it out again[477]. It is all a question of relative surface-energies, leading to different degrees of “adhesion” between the chloroform and the glass or its covering. Thus it is that the Amoeba takes in the diatom, dissolves off its proteid covering, and casts out the shell.

Furthermore, as the whole phenomenon depends on a distribution of surface-energy, the amount of which is specific to certain particular substances in contact with one another, we have no difficulty in understanding the _selective action_, which is very often a conspicuous feature in the phenomenon[478]. Just as some caddis-worms make their houses of twigs, and others of shells and again others of stones, so some Rhizopods construct their agglutinated “test” out of stray sponge-spicules, or frustules of diatoms, or again of tiny mud particles or of larger grains of sand. In all these cases, we have apparently to deal with differences in specific {466} surface-energies, and also doubtless with differences in the total available amount of surface-energy in relation to gravity or other extraneous forces. In my early student days, Wyville Thomson used to tell us that certain deep-sea “Difflugias,” after constructing a shell out of particles of the black volcanic sand common in parts of the North Atlantic, finished it off with “a clean white collar” of little grains of quartz. Even this phenomenon may be accounted for on surface-tension principles, if we assume that the surface-energy ratios have tended to change, either with the growth of the protoplasm or by reason of external variation of temperature or the like; and we are by no means obliged to attribute the phenomenon to a manifestation of volition, or taste, or aesthetic skill, on the part of the microscopic organism. Nor, when certain Radiolaria tend more than others to attract into their own substance diatoms and such-like foreign bodies, is it scientifically correct to speak, as some text-books do, of species “in which diatom selection has become _a regular habit_.” To do so is an exaggerated misuse of anthropomorphic phraseology.

The formation of an “agglutinated” shell is thus seen to be a purely physical phenomenon, and indeed a special case of a more general physical phenomenon which has many other important consequences in biology. For the shell to assume the solid and permanent character which it acquires, for instance, in Difflugia, we have only to make the additional assumption that some small quantities of a cementing substance are secreted by the animal, and that this substance flows or creeps by capillary attraction between all the interstices of the little quartz grains, and ends by binding them all firmly together. Rhumbler[479] has shewn us how these agglutinated tests, of spicules or of sand-grains, can be precisely imitated, and how they are formed with greater or less ease, and greater or less rapidity, according to the nature of the materials employed, that is to say, according to the specific surface-tensions which are involved. For instance if we mix up a little powdered glass with chloroform, and set a drop of the mixture in water, the glass particles gather neatly round the surface of the drop so quickly that the eye cannot follow the {467} operation. If we perform the same experiment with oil and fine sand, dropped into 70 per cent. alcohol, a still more beautiful artificial Rhizopod shell is formed, but it takes some three hours to do.

It is curious that, just at the very time when Rhumbler was thus demonstrating the purely physical nature of the Difflugian shell, Verworn was studying the same and kindred organisms from the older standpoint of an incipient psychology[480]. But, as Rhumbler himself admits, Verworn was very careful not to overestimate the apparent signs of volition, or selective choice, in the little organism’s use of the material of its dwelling.

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This long parenthesis has led us away, for the time being, from the subject of the Radiolarian skeleton, and to that subject we must now return. Leaving aside, then, the loose and scattered spicules, which we have sufficiently discussed, the more perfect Radiolarian skeletons consist of a continuous and regular structure; and the siliceous (or other inorganic) material of which this framework is composed tends to be deposited in one or other of two ways or in both combined: (1) in the form of long spicular axes, usually conjoined at, or emanating from, the centre of the protoplasmic body, and forming a symmetric radial system; (2) in the form of a crust, developed in various ways, either on the outer surface of the organism or in relation to the various internal surfaces which separate its concentric layers or its component vesicles. Not unfrequently, this superficial skeleton comes to constitute a spherical shell, or a system of concentric or otherwise associated spheres.

We have already learned that a great part of the body of the Radiolarian, and especially that outer portion to which Haeckel has given the name of the “calymma,” is built up of a great mass of “vesicles,” forming a sort of stiff froth, and equivalent in the physical sense (though not necessarily in the biological sense) to “cells,” inasmuch as the little vesicles have their own well-defined boundaries, and their own surface phenomena. In short, all that we have said of cell-surfaces, and cell conformations, in our discussion of cells and of tissues, will apply in like manner, and under appropriate conditions, to these. In certain cases, even in {468} so common and simple a one as the vacuolated substance of an Actinosphaerium, we may see a very close resemblance, or formal analogy, to an ordinary cellular or “parenchymatous” tissue, in the close-packed arrangement and consequent configuration of these vesicles, and even at times in a slight membranous hardening of their walls. Leidy has figured[481] some curious little bodies, like small masses of consolidated froth, which seem to be nothing else than the dead and empty husks, or filmy skeletons, of Actinosphaerium. And Carnoy[482] has demonstrated in certain cell-nuclei an all but precisely similar framework, of extreme delicacy and minuteness, as the result of partial solidification of interstitial matter in a close-packed system of alveoli (Fig. 220).

Let us now suppose that, in our Radiolarian, the outer surface of the animal is covered by a layer of froth-like vesicles, uniform or nearly so in size. We know that their tensions will tend to conform them into a “honeycomb,” or regular meshwork of hexagons, and that the free end of each hexagonal prism will be a little spherical cap. Suppose now that it be at the outer surface of the protoplasm (that namely which is in contact with the surrounding sea-water), that the siliceous particles have a tendency to be secreted or adsorbed; it will at once follow that they will show a tendency to aggregate in the grooves which separate the vesicles, and the result will be the development of a most delicate sphere composed of tiny rods arranged in a regular hexagonal network (e.g. _Aulonia_). Such a conformation is {469} extremely common, and among its many variants may be found cases in which (e.g. _Actinomma_), the vesicles have been less regular in size, and some in which the hexagonal meshwork has been developed not only on one outer surface, but at successive {470} surfaces, producing a system of concentric spheres. If the siliceous material be not limited to the linear junctions of the cells, but spread over a portion of the outer spherical surfaces or caps, then we shall have the condition represented in Fig. 223 (_Ethmosphaera_), where the shell appears perforated by circular instead of hexagonal apertures, and the circular pores are set on slight spheroidal eminences; and, interconnected with such types as this, we have others in which the accumulating pellicles of skeletal matter have extended from the edges into the substance of the boundary walls and have so produced a system of films, normal to the surface of the sphere, constituting a very perfect honeycomb, as in _Cenosphaera favosa_ and _vesparia_[483].

In one or two very simple forms, such as the fresh-water _Clathrulina_, just such a spherical perforated shell is produced out of some organic, acanthin-like substance; and in some examples of _Clathrulina_ the chitinous lattice-work of the shell is just as {471} regular and delicate, with the meshes just as beautifully hexagonal, as in the siliceous shells of the oceanic Radiolaria. This is only another proof (if proof be needed) that the peculiar conformation of these little skeletons is not due to the material of which they are composed, but to the moulding of that material upon an underlying vesicular structure.