67. The Nucleus of the Peripylea.—The nucleus of the Spumellaria or Peripylea shows in certain groups a very primitive arrangement, indeed the archaic structure from which the various forms of nuclei of other Radiolaria may be derived; but on the other hand, in other groups it exhibits very peculiar and remarkable differentiations. In the first place it may be noted that the monozootic or solitary Spumellaria usually possess a single serotinous nucleus, which only divides into numerous swarm-spores at a late period; whilst, on the contrary, the polyzootic colonial Spumellaria (or Polycyttaria) are uninuclear only in the young state (Pl. 3, fig. 12), and speedily present numerous small homogeneous nuclei, which have arisen by precocious division of a single nucleus; these are usually spherical and 0.008 to 0.012 mm. in diameter. The serotinous nucleus of the monozootic Spumellaria, in many divisions of this large legion, and especially in the simply constituted Sphæroidea, is a homogeneous sphere of nuclein, lying in the middle of the central capsule. In many other cases it assumes the form of a spherical vesicle ("Binnen-Bläschen"), whose fluid or semi-fluid contents are enclosed by a more or less firm membrane. This vesicle often contains a single central spherical nucleolus (Pl. 1, figs. 1l, 4l), but sometimes a variable number of small excentric nucleoli (Pl. 1, figs. 1a, 2a). The nuclear membrane is often somewhat thick, presenting a double contour, and in such cases may even exhibit a fine radial striation, the expression of minute pores (Pl. 1, fig. 2a). In the colossal nuclei (as much as 1 to 2 mm. in diameter) of certain large Thalassicollida the nucleolus presents a very remarkable form, becoming stellate by the protrusion of processes, which may again branch in a dendritic fashion (as in the common Thalassicolla nucleata), or it may develop into a very long cylindrical thread, which is disposed in serpentine coils, and in Thalassophysa pelagica passes into the different cæcal processes of the stellate nucleus. In many Sphæroidea, whose skeleton is composed of numerous concentric lattice spheres, the small central spherical nucleus lies at first within the innermost of these (the medullary shell); but afterwards it grows through the meshes of the lattice-work, and the radiating club-shaped processes thus formed (Pl. 11, fig. 5) unite with each other outside the medullary shell, and form an external nuclear sphere which completely encloses the latter. In the Polysphærida (with several concentric lattice-shells) and in the Spongosphærida (with spongy lattice-spheres), this process may be several times repeated, so that eventually the central spherical nucleus attains considerable dimensions, and encloses two or more concentric lattice-shells with their radial connecting rods. The nuclear membrane is in these cases usually penetrated by radial bars, which connect the outermost of the enclosed shells with the remaining cortical shells which surround the central capsule. The same remarkable arrangement is also very common among the Discoidea. The small spherical primary nucleus is in such instances immediately surrounded by the innermost earliest developed lattice-shell, around which the concentric rings are subsequently deposited; it then grows out through the meshes, and the processes fuse outside the ring to form a homogeneous lentiform nucleus (Pl. 43, fig. 15). The same process recurs in certain Prunoidea and Larcoidea, whilst in other Spumellaria of these groups (e.g., Pylonida) the lobate processes of the nucleus remain free.
Both the simple serotinous nucleus of the monozootic Spumellaria, and the numerous precocious nuclei of the Polycyttaria, were first described in my Monograph in 1862, the former as the "endocyst" ("Binnen-Bläschen"), the latter as "spherical transparent vesicles" ("Kugelige wasserhelle Bläschen"). I was in error, however, in regarding the latter as identical with the so-called "hyaline spherules" in the central capsule of many Monozoa, which rather belong to the category of intracapsular vacuoles (see § 72). The credit of recognising, by the aid of the modern methods of staining, the distinctness of these two structures, which may readily be mistaken for each other, and of demonstrating the true nature both of the serotinous and precocious nuclei, belongs to Richard Hertwig (1879, L. N. 33).
68. The Nucleus of the Actipylea.—The nucleus of the Acantharia or Actipylea shows very peculiar relations in respect of structure and division, particularly special forms of lobular budding, which belong to the characteristic peculiarities of this singular legion, and are not found among other Radiolaria. The position of the nucleus is always excentric, even in the youngest Acantharia, for the centrogeneous formation of the skeleton, the constant development of the earliest radial portions of it in the middle of the central capsule, forces the nucleus from its normal central position. The majority of the Acantharia, like most Polycyttaria, are precocious, the primary nucleus early dividing into numerous small nuclei (see note A below). Nevertheless there are many exceptions to this rule in different families, e.g., Stauracantha, Xiphacantha, Phatnacantha, and Pristacantha among the Acanthometra, and Stauraspis, Echinaspis, Dodecaspis, and Phatnaspis among the Acanthophracta. In these instances the primary nucleus remains for a long time as a simple excentric ellipsoidal or irregularly round body, even in the fully developed stage, and only at a very late period (sometimes just before the formation of the spores) divides into many small nuclei. Since this serotinous division of the nucleus takes place in different genera of very various groups, it can only be decided by further investigations how widely it is spread among the Acantharia, and upon what circumstances it is dependent (see note B). The division of the nucleus appears to be precocious in the majority of this legion, and a number of small nuclei appear to be early formed by a peculiar process of budding; in most fully developed Acantharia these are disposed in one or two layers under the surface of the central capsule, but if their numbers increase to any considerable extent, the whole space between the skeletal rods becomes filled with small nuclei; sometimes these are homogeneous, sometimes vesicular, 0.002 to 0.012 mm. in diameter; usually they are spherical and have a small nucleolus (compare Pl. 129, figs. 6-11, and note C).
A. The numerous nuclei, which are to be found in the central capsule of most mature Acantharia, were first described in my Monograph (1862) as "spherical, transparent vesicles, provided with a small dark granule" (p. 374, Taf. xv. figs. 2, 5; Taf. xvi. figs. 2, 4; Taf. xxi. fig. 7, &c.). Their more minute constitution and peculiar origin were first accurately delineated by R. Hertwig (1879, loc. cit., pp. 11-24, Taf. i-iii.).
B. The fact that in a number of Acantharia the nucleus does not divide early as in the majority of the legion, but only at a later period, was first observed by R. Hertwig in a species of Acanthometra (Xiphacantha serrata), and a species of Acanthophracta (Phatnaspis mülleri = Haliommatidium mülleri) (loc. cit., pp. 11 and 27). This serotinous division of the nucleus seems, however, to be rather widely spread in both sublegions of the Acantharia; I have found, not only in the forms above mentioned, but also in several others belonging to different genera, a single large excentric nucleus, even in those individuals in which the skeleton was fully developed.
C. The peculiar mode of nuclear budding, by which these small nuclei arise, appears to proceed in the following manner (Pl. 129). The vesicular primary nucleus, which, in consequence of the centrogeneous development of the skeleton protrudes as it grows into irregular lobes (Pl. 129, fig. 9), assumes a peculiar concavo-convex form, sometimes that of a hood or dish, sometimes that of a kidney or sausage. The convex surface is apposed to the capsule-membrane, while the concave is turned towards the central star of the skeleton (fig. 6). There is now formed at the centre of the convex surface of the strong, doubly-contoured, nuclear membrane, a flask-shaped invagination with a narrow neck and expanded base; the membrane now becomes disposed in peculiar folds, which at the narrow aperture of invagination appear as folds, but on the expanded body of the flask take the form of concentric rings, laid closely side by side (Pl. 129, fig. 10). The convex bottom of the flask, which is directed towards the concave proximal side of the nucleus, becomes again invaginated by a central conical apophysis of the enlarged nucleolus, which is situated between them. Usually the nucleolus has already become flattened into a lentiform shape, and upon its distal face a conical apophysis has been developed, which is divisible into a darker proximal and clearer distal portion. The tip of the latter appears to be in direct connection with the nuclear membrane at the centre of the base of the flask-shaped invagination (figs. 6, 10). At this stage of development the nucleus of the Acantharia generally presents the characteristic form of a hood-shaped, concavo-convex vesicle, whose radial axis is also the axis of the flask-shaped distal invagination, and of the depressed conical nucleolus, which lies between the latter and the concave side of the nucleus. After this peculiar invagination has persisted for some time in connection with the enlarged nucleolus, both disappear, and then a remarkable growth of lobular processes takes place on the concave proximal side of the hood or kidney-shaped nucleus; from four to eight knobs of unequal size usually appear, and their thickened wall encloses a variable number of small of nucleoli; these are at first few but afterwards more numerous (fig. 7). Subsequently these knobs or lobes become completely separated by constriction from the original central mass of the nucleus, and appear as so many separate independent "sausage-shaped bodies" in the hollow central capsule (fig. 8). Each of the bodies now appears, and at first on its convex aspect, to form a large number of small nucleoli, which either separate by constriction from it or become free by its breaking up and lie in numbers in the central capsule. Finally the buds or lobes of the nucleus break up entirely into such nucleoli, which are evenly distributed in the central capsule, and become the nuclei of the swarm-spores (fig. 11). Compare R. Hertwig, L. N. 33, Taf. i.-iii. pp. 19-25.
69. The Nucleus of the Monopylea.—The nucleus of the mature forms of the Nassellaria or Monopylea is generally simple or lobate, homogeneous or vesicular and excentric, and appears only to divide into numerous small nuclei just before the formation of the spores. Nevertheless I have sometimes, though not often, seen in representatives of very various families of the Monopylea, the central capsule filled with many small spherical homogeneous nuclei (Pl. 53, fig. 19). Hence all the families of this legion appear to be serotinous, their simple primitive nucleus persisting for a long period. It is commonly placed excentrically, and most usually in the apical or aboral portion of the central capsule, either between its apex and the podoconus, or quite excentrically on the dorsal aspect. The simple nucleus of the Nassellaria usually appears to be vesicular and to possess a somewhat firm membrane, clear contents, and a rather large, dark coloured nucleolus. In many Nassellaria the nucleus is spherical or ellipsoidal (Pl. 53, fig. 11); whilst in many Stephoidea and Spyroidea, where the central capsule is constricted by the sagittal ring and divided into two symmetrical lateral lobes, the nucleus partakes of the same mode of growth and appears in the middle of the capsule as a transversely placed ellipsoid or even as a short cylinder (Pl. 90, figs. 7, 9). The most remarkable modification in the form of the nucleus is to be found in the multi-articulate Cyrtoidea. Here it is usually enclosed in the cephalis and is spherical, ellipsoidal or spheroidal, often flattened almost into a disc. If now the central capsule increase greatly in size and put forth three or four clavate lobes which hang down through the pores of the cortinar septum into the thorax (or even into the succeeding joints), the nucleus usually undergoes similar modification, and three or four finger-like apophyses are developed from its base, which project into the corresponding lobes of the central capsule (Pl. 59, figs. 4, 12, 13).
The numerous small, spherical, homogeneous nuclei which are to be found in the central capsules of those Nassellaria, which are ripe and about to develop spores, were described in 1862 in my Monograph, as "numerous, small, transparent, spherical cells" in the case of various Cyrtoidea (Arachnocorys, Lithomelissa, Eucecryphalus, Eucyrtidium, &c.) (loc. cit., pp. 302, 305, 309, 321, &c.), and I find them of the same form and dimensions, but deeply stained with carmine in many preparations in the Challenger collection. R. Hertwig has delineated them very accurately in the case of Tridictyopus (1879, loc. cit., p. 84, Taf. vii. fig. 3). He was also the first to recognise the uninucleate condition of the Nassellaria, which is much more frequently observed than the serotinous multinucleate condition, and he described very clearly the peculiar lobed nuclei which arise in Cyrtoidea, owing to the protrusion of the nucleus through the cortinar septum (loc. cit., p. 85, Taf. viii. figs. 3-8).
70. The Nucleus of the Cannopylea.—The nucleus presents the same remarkable structures in all species of the Phæodaria or Cannopylea which have been examined, and closely resembles the germinal vesicle of an amphibian ovum, being a large spherical or spheroidal vesicle with numerous nucleoli. Its diameter usually amounts to half or two-thirds, sometimes even three-quarters, that of the central capsule. The vertical main axis of the latter is also that of the nucleus, which usually lies somewhat nearer to the aboral pole. The nucleus is generally rather more strongly compressed in the direction of the main axis than the capsule itself. The membrane of the vesicular nucleus is thin, but firm, and encloses a clear or finely granular mass of nuclein. The number and size of the contained nucleoli are variable even in one and the same species, and stand in inverse ratio to each other, an obvious result of the gradual process of division. Commonly from twenty to fifty roundish or spherical, strongly refracting nucleoli, are present; more rarely there are several hundred very small ones. Sometimes the nucleus is penetrated by fine trabeculæ, in whose meshes lie the nucleoli (Pl. 101, fig. 2). In certain nuclei, which contained a few large nucleoli, these were of irregular form, probably the result of amœboid movements (Pl. 101, fig. 1). In the formation of spores in the Cannopylea, the nucleus apparently becomes dissolved, and its numerous nucleoli develop directly into the nuclei or mother-nuclei, which produce the nuclei of the flagellate spores. Furthermore, many Phæodaria seem to multiply by simple cell-division, since very commonly (especially in the Phæocystina and Phæoconchia) two large nuclei (right and left), may be met with in one central capsule; sometimes also a single large nucleus, in which a sagittal constriction marks the commencing division of the capsule (Pl. 101, figs. 2, 36; Pl. 104, fig. 3; Pl. 124, fig. 6, &c.).
The large nucleus of the Phæodaria was first described in my Monograph in 1862, in the case of Aulacantha (p. 263), Aulosphæra (p. 359), and Cœlodendrum (p. 361), as a "large, spherical, thin-walled endocyst," from 0.1 to 0.2 mm. in diameter. More detailed descriptions, especially with respect to the behaviour of the nucleoli were given by R. Hertwig in 1879 (L. N. 33, p. 97).
71. The Endoplasm or Intracapsular Protoplasm.—In all Radiolaria the intracapsular protoplasm, which, for the sake of brevity, may be termed "endoplasm," constitutes originally, and especially in the earliest stages, the only important content of the central capsule, except the nucleus. In certain Spumellaria and Nassellaria, of simple structure and of small dimensions, this condition persists for a long period, and the endoplasm then appears as a homogeneous, colourless, turbid or finely granular, mucous, semi-solid mass, which cannot be distinguished from the ordinary undifferentiated protoplasm of young cells; no definite structure, and in particular, no fibrillar network, can be discovered in it even by the use of the customary reagents. In the great majority of the Radiolaria, however, this primitive homogeneous condition of the endoplasm is very transient, and it soon undergoes definite modifications, becoming differentiated into separate parts or producing new constituent contents. Such products of the internal protoplasm are in particular hyaline spheres (vacuoles and alveoles), oil-globules, pigment-bodies, crystals, &c. The most important of the differentiations which take place in the endoplasm is that into an internal, granular, medullary substance and an external, fibrillar, cortical substance; although the various legions behave somewhat differently in this respect (§§ 77-80).
72. Intracapsular Hyaline Spheres.—The central capsule of very many Radiolaria contains in its endoplasm numerous spherical bodies of varying size, which consist of watery or albuminous fluid, and have previously been regarded as nuclei, or described as products of the internal protoplasm, under various names, such as "spherical transparent vesicles" (see note A, below), "albumen spheres" (see B), "gelatinous spheres" (see C), "alveolar cells" (see D), &c. Some of these spheres are perfectly transparent, structureless and of varying refractive power, producing the impression of drops of fluid; others contain various formed constituents, such as oil-globules, fat-granules, pigment-granules, concretions, crystals, &c. From a morphological point of view they may all be divided into two categories, membraneless vacuoles and vesicular alveoles. The vacuoles are simple spherical drops of fluid or of gelatinous material, devoid of a special envelope, but immediately surrounded by the endoplasm. The alveoles, on the other hand, are true vesicles with a thin spherical envelope, enclosing a drop of fluid or jelly. This envelope is commonly very thin, homogeneous, and often scarcely discernible, so that in practice a sharp line of demarcation cannot be drawn between alveoles and vacuoles; the former are usually somewhat larger than the latter. The fact is, nevertheless, certain that the hyaline spheres, which may be isolated on rupturing the central capsule of many Radiolaria, in certain cases, particularly in large species, possess a clear, anatomically demonstrable membrane, whilst in others no such appearance is presented. It may be assumed that the vesicular alveoles are developed from the drop-like vacuoles by increase in size, and by the precipitation of a delicate envelope from the endoplasm. The character common to all these hyaline spheres, whether vacuoles or alveoles, is found in their aqueous, not adipose, constitution, and in their clear transparent appearance, which allows of no structure (the above-mentioned contained bodies excepted) being recognised. Their refractive power and consistency vary somewhat, and probably their chemical constitution still more. Sometimes they are strongly refractive and shining, and sometimes feebly refractive and pale; their consistency shows all intermediate stages between a thin fluid, which readily disappears in water, and a firm, insoluble jelly. As regards their chemical composition (which is probably very variable), the hyaline spheres may be best divided into two groups, the organic and inorganic. The inorganic hyaline spheres are simple drops of saline solution without any carbonaceous constituent; the organic, on the other hand, contain a small quantity of organic matter dissolved in the watery fluid, and may be either albuminous or gelatinous spheres. The formed contents which are commonly present are of very various natures, usually small fat-granules, more rarely larger fat-granules or pigment-granules, sometimes concretions or crystals. In many groups, especially among the large Phæodaria and Collodaria, the numerous hyaline spheres are remarkable for their equal size and even distribution throughout the endoplasm (Pl. 1, figs. 1, 4; Pl. 104, fig. 2, &c.). In some genera belonging to the Thalassicollida the alveoles are of enormous size (Pl. 1, figs. 2, 3); they then become flattened by mutual pressure into polyhedra and distend the central capsule to unusual dimensions (in Physematium and Thalassolampe 8 to 12 mm.).
A. The "spherical hyaline vesicles," which I described in my Monograph (1862, p. 71) as among the most important and constant contents of the central capsule, are partly vacuoles, partly homogeneous nuclei. Most recent investigators, Bütschli in particular (1882, L. N. 41), have pointed out and rightly criticised this confusion. The criticism might, however, have been more justly expressed by stating that, in the preparation of my Monograph (1859-1862), I did not make use of modern methods of demonstrating the nucleus by staining fluids, which were quite unknown at the time, and only discovered a decade later. In fact, without the aid of such reagents, it is quite impossible to distinguish between the various "spherical transparent vesicles," of which those found in the central capsule of the Phæodaria and many monozootic Collodaria are simple vacuoles lying in the endoplasm, whilst, on the other hand, those of the Polycyttaria and many other Radiolaria are true homogeneous nuclei. For not only are the general appearance of the small clear spheres, their refractive power, and regular distribution in the endoplasm quite similar, but they are also of much the same size, for the diameter ranges from 0.005 to 0.015 mm., being generally between 0.008 and 0.012 mm. In addition to this there is generally in each hyaline sphere a dark brightly shining granule, which, in the case of the vacuole, is simply a fat-granule, whilst in the case of the nucleus, it is a true nucleolus. The small hyaline spheres in the young uninucleate capsules of the Polycyttaria are simple vacuoles (Pl. 3, fig. 12), whilst in the ripe multinucleate capsules they are true nuclei (Pl. 3, figs. 3, 8, 9), and it is quite impossible to discriminate between these two conditions without the use of reagents. This has been expressly recognised by R. Hertwig, who has the merit of having been the first to clearly distinguish, by the aid of staining fluids, between these two different constituents (1879, L. N. 33, p. 108).
B. The "albumen spheres," which were first observed by A. Schneider in 1858 in the common cosmopolitan Thalassicolla nucleata (L. N. 13, p. 40), and which appear to occur in only a few other Thalassicollida, are distinguished from the ordinary hyaline spheres of about the same size by their higher refractive power and by certain albuminoid reactions, especially the coagulation of a membranous envelope under the influence of certain reagents (see my Monograph, p. 250, and Hertwig, L. N. 26, 1876, p. 46). They often enclose various formed contents, and require further investigation.
C. The gelatinous spheres of various sizes, found in the endoplasm of the Radiolaria, agree in their reactions (especially in staining by certain reagents) with the common extracapsular jelly of the calymma, and are hence distinguishable both from the true (coagulable) "albumen sphere," and from the ordinary watery vacuoles.
D. The alveoles, which are only accurately known in the case of certain large monozootic Collodaria, but which also seem to occur in the central capsule of other remarkably large Radiolaria, were described in my Monograph in the case of Thalassolampe margarodes and Physematium mülleri, under the name "intracapsular alveolar cells" (1862, pp. 77, 254, 257). They are not, however, true nucleated cells, and the body described as a nucleus is not such in reality. Nevertheless these large hyaline spheres do possess a special envelope, as I have recently convinced myself by the examination of ruptured central capsules of Thalassolampe maxima, Thalassopila cladococcus, and Physematium atlanticum (Pl. 1, figs. 2, 3). The central capsule of these Collodaria becomes distended to most unusual dimensions (2 to 12 mm. in diameter) by the great development of these large hyaline vesicles, each of which measure from 0.1 to 0.5 mm. in diameter.
73. The Intracapsular Fat-Globules.—Fat is present in the central capsule of all Radiolaria in larger or smaller quantities, and generally appears in the form of very numerous, small, spherical granules, which are either distributed evenly in the endoplasm (as an emulsion) or enclosed in the vacuoles; the latter, in particular, is the case in most Phæodaria, perhaps generally. In this group each vacuole contains as a rule a single dark, shining fat-granule, and sometimes also an irregular bunch composed of from two to five or more granules. In addition to these small fat-granules (granula adiposa) which are always present, the central capsule of many Radiolaria contains also larger fat-globules (globuli adiposi). These appear to be generally wanting in the Phæodaria, and are on the whole rare in the Acantharia; whilst, on the contrary, they are very common in the Nassellaria and Spumellaria. The Polycyttaria or social Radiolaria are as a rule distinguished by the possession of a single large central oil-globule, which lies in the centre of the central capsule, and is on an average about one-third of it in diameter (Pl. 3, figs. 4, 5). This is absent, however, in those young capsules of the Polycyttaria in which the primary nucleus is centrally situated (Pl. 3, fig. 12). Those species of Polycyttaria whose central capsule reaches a considerable size, often enclose numerous oil-globules, and in Collophidium (species of Collozoum with an elongated cylindrical capsule, Pl. 3, figs. 1, 3) the axis of each capsule is occupied by a row of numerous oil-globules. In the monozootic Spumellaria, in which the nucleus is always centrally situated, the large oil-globules are, of course, excentric, being in apposition to the inner surface of the capsule-membrane (Pl. 1, fig. 3; Pl. 2, figs. 2, 5). In the Discoidea the oil-globules, which are often present in large numbers, form elegant concentric rings around the central nucleus, and in those species with segmented arms, there are one or more transverse rows in each segment (Pl. 43, fig. 15). In the Nassellaria the number and distribution of the oil-globules are dependent upon the form of the central capsule. When this is simple, without lobes, and ovoid or conical, they generally lie in its aboral half above the podoconus (Pl. 51, figs. 5, 13; Pl. 97, fig. 1). When, on the contrary, the basal portion of the capsule sends out three or four dependent processes (as in the majority of the Cyrtoidea), a large globule may generally be seen in the swollen distal part of each conical or ovoid lobe (Pl. 53, fig. 19; Pl. 60, figs. 4-7). In many Stephoidea and Spyroidea, whose central capsule is separated into two lateral portions by the constriction corresponding to the sagittal ring, each of these contains either a single large globule or a group of small ones (Pl. 90, figs. 7, 10). These oil-globules are usually colourless and highly refractive; rarely they are yellow or brown, sometimes rose-coloured, or an intense blood-red (e.g., in Thalassophysa sanguinolenta) or even orange (in Physematium mülleri). In many Spumellaria, and particularly in the Polycyttaria, an albuminous substratum may be recognised in them, which is sometimes disposed in layers, and after extraction of the fat presents the appearance of a laminated sphere. The physiological significance of the oil-globules is twofold; in the first place they tend to diminish the specific gravity of the organism; in the second they may be utilised as a reserve store of nutriment. In the latter respect they are of special importance in the process of spore-formation, each flagellate spore usually containing a fat-granule.
74. The Intracapsular Pigment-Bodies.—In the majority of Radiolaria when observed alive, the central capsule is coloured, only in the minority is it colourless. The colour is never diffuse, but always due to the formation of definite pigment granules or vesicles, which are sometimes distributed evenly throughout the endoplasm, sometimes aggregated in the central or peripheral regions. Their form may be either spherical, irregularly rounded, or polyhedral. They vary much in dimensions, but in most cases are immeasurably small, and appear under a high magnifying power as fine dust; occasionally, however, their diameter may amount to from 0.001 to 0.005 or more. The chemical constitution of the intracapsular pigment is unknown in most Radiolaria, and is probably very various. In many instances the pigment-granules consist of fat, in others not. The commonest colours are yellow, red, and brown; violet and blue are rare, and green still rarer. Sometimes a definite tone of colour prevails throughout a whole group, and may then be attributed to inheritance, e.g., red is found in most Sphæroidea, and blue in the Polycyttaria (see note A). One colour is almost always constant in the members of the same species. True pigment-cells, belonging to the Radiolarian organism, do not occur within the central capsule. The peculiar yellow cells which are found in the central capsule of many Acantharia are symbiotic xanthellæ (see § 76).
A. The number of Radiolaria whose pigment has been examined in the living state, is too small to allow of any general conclusions being drawn. Regarding the different colours known, see my Monograph, L. N. 16, p. 76.
75. The Intracapsular Crystals.—The crystals found in the central capsule of many Radiolaria may be divided into two groups, of very different significance; small crystals, which are very widely distributed, and large crystals, which occur in only a few genera. The small crystals may also be termed "spore-crystals," since each swarm-spore often contains such a crystal. They are rod-like or spindle-shaped, and consist of an organic substance which probably serves as a reserve of nutriment for the developing spores. Such spore-crystals have been observed in numerous Spumellaria and Acantharia belonging to various families, and are probably present throughout the two legions which make up the Porulosa. On the other hand, they have not been noticed in the Osculosa (Nassellaria and Phæodaria), the few swarm-spores belonging to these groups which have been observed not exhibiting any crystals. The large crystals, which occur in small numbers in the endoplasm, have hitherto only been observed in a few species of Spumellaria, belonging to the Polycyttaria. They were first noticed in the common Collosphæra huxleyi, and regarded as cœlestin. They are also found in the central capsule of many other Collosphærida, e.g., Buccinosphæra (Pl. 5, figs. 11, 12). Crystal-masses, crystal-sheaves, or spherical masses of radiating acicular crystals are enclosed in the vacuoles or "albumen globules" of Thalassicola nucleata and other Thalassicollida, as well as in the central capsule of Cœlographis and some other Phæodaria (Pl. 127, figs. 4-7). All these large crystals are probably to be regarded as excretory products.
75A. The Intracapsular Concrements.—Concretions, either mineral or organic, of varying form and constitution, are to be found in the endoplasm of Radiolaria belonging to very different families. They are most abundant and multiform in Thalassicolla nucleata, being usually circular or elliptical discs, which are concentrically laminated and highly refractive, resembling starch-grains. Among them twin forms may frequently be observed, as though the concrements were in process of division (see note A). Similar amyloid concretions are to be seen in the central capsule of different Spumellaria and Nassellaria, e.g., in Cephalospyris triangulata (Pl. 96, fig. 28). Violin-shaped, highly refractive concrements have been observed in the central capsule of numerous Spumellaria, Nassellaria, and Acantharia, e.g., Thalassosphæra, Spongosphæra, Plegmosphæra, Cyrtocalpis, Peripyramis, Botryocella, &c. (see note B). The chemical constitution of these concrements is insufficiently known.
A. The amyloid concretions of Thalassicolla nucleata have been described in detail in my Monograph (pp. 80, 250, Taf. iii. figs. 2, 3), and by R. Hertwig in the Histologie der Radiolarien (1876, p. 47, Taf. iii. figs. 9-13).
B. The violin-shaped concretions of Thalassosphæra bifurca have been figured in my Monograph (pp. 80, 261, Taf. xii. fig. 1).
76. The Intracapsular Xanthellæ.—The xanthellæ, zooxanthellæ, or symbiotic "yellow cells" are found within the central capsule only in the Acantharia, whilst in other Radiolaria they only occur in the extracapsulum. They are most frequent in the Acanthometra, rarer in the Acanthophracta, but even in the former they are often wanting. Their number is very variable, but usually small, from ten to thirty in one capsule. They lie for the most part immediately below the capsule membrane, in the cortical layer of the endoplasm. The form of the yellow cells is either spherical or ellipsoidal, often also spheroidal or even lentiform. The diameter varies from 0.01 to 0.03 mm. They possess a distinct membrane and an excentric nucleus, and contain numerous yellow pigment-granules in the endoplasm. This yellow pigment dissolves in mineral acids to form a green fluid, and in other respects also behaves somewhat differently from the yellow pigment in the extracapsular yellow cells of the Spumellaria and Nassellaria. In both cases, however, the xanthellæ are not integral portions of the organism, but unicellular algae, living as parasites or symbiontes in the body.
A. The yellow cells in the central capsule of the Acantharia were first observed by Joh. Müller (L. N. 12, pp. 14, 47). In my Monograph I described them at greater length, and indicated their differences from the extracapsular yellow cells of other Radiolaria (L. N. 16, pp. 77, 86). Since then, R. Hertwig has demonstrated their cellular nature (L. N. 33, pp. 12, 113), and still more recently Brandt has given further accurate information regarding their occurrence, constitution, and physiological significance (L. N. 39, ii. Art., p. 235, figs. 62-73).
77. The Endoplasm of the Peripylea.—The intracapsular protoplasm of the Spumellaria or Peripylea is usually distinguished by a more or less complete radial arrangement, which does not occur in the same form in other Radiolaria; it may be regarded as characteristic of this legion, for it probably occurs in all the species at some period of life or other, and stands in a direct causal relationship with the typical structure of the capsule-membrane in all the "Peripylea" (see note A). For as this is commonly perforated by very numerous pores distributed at equal intervals over the whole surface of the capsule, and since a communication between the intra- and extracapsular sarcode takes place through these, the radiate structure of the endoplasm may be readily explained as due to the influence of radial currents which take place continuously or intermittently in the endoplasm. This radiate structure is most obvious when the endoplasm contains no secondary products or only an insignificant amount of these, and thus appears colourless and almost homogeneous, or only finely granular. Under these circumstances, an optical section of the central capsule usually reveals a distinct radial striation; numerous narrow, straight, dark streaks alternating regularly with still narrower clear ones; the latter consist of homogeneous, the former of more or less granular protoplasm (Pl. 20, fig. 1a). Often there may be distinguished in each darker streak a single straight row of strongly refracting (fat?) granules, sometimes several such rows. Occasionally the whole endoplasm becomes divided up into a number of large "radial wedges," club-shaped, conical or pyramidal masses of granular protoplasm, separated by clear divisions of hyaline plasma (e.g., in Actissa radiata, p. 14, where in the optical section of the central capsule, between the membrane and the nucleus, twenty-five dark radial wedges of equal size were separated by thick clear partitions of hyaline protoplasm). In the majority of the Spumellaria this radial striation is partially or entirely concealed by the formation of pigment or of other products. Very often it is only visible in the cortical layer, which lies immediately below the capsule-membrane (Pl. 1, figs. 1, 3). The remarkable "centripetal cones" which characterise the Thalassicollid genus Physematium, and were formerly described as "centripetal cell-groups," are probably a special development of these cortical radial wedges; they are conical cortical bodies, regularly distributed on the inner surface of the membrane of the central capsule, and disposed with the apex turned towards the centre (see note B). More rarely than in the cortical layer, a similar radial structure is to be found in the innermost medullary layer immediately surrounding the nucleus. Here the endoplasm sometimes breaks up into fine radial threads, which are anatomically separable and hang down from the free nucleus as thin processes (see note C). In some cases it is also possible to isolate radial rods from the cortical layer of teased out central capsules.