A. The extracapsular vacuoles in the calymma were first observed in 1851 by Huxley, in Thalassicolla and Sphærozoum, and compared with Dujardin's sarcode vacuoles (L. N. [5]). Afterwards J. Müller noticed that generally these "large clear vesicles are covered by a fine membrane," and hence he called them "alveoles" (L. N. [12], pp. 3, 7, &c.). In my Monograph I have described them more in detail as "extracapsular alveoles" (1862, p. 88, Tafs. i.-iii. xxxii.-xxxv.). Ever since then the point has been debated whether these clear spaces are simple vacuoles in the sense of Huxley or vesicular alveoles as stated by J. Müller. This contention is unnecessary, for both varieties are present, and often no sharp line can be drawn between them. R. Hertwig has recently come to the conclusion that they are as a rule "membraneless vacuoles," but that they "sometimes become surrounded by a special envelope" (L. N. [33], p. 31). He even succeeded "in extracting from a Collosphæra the large vesicle which lies in the centre of many colonies and removing its covering of central capsules and jelly."
B. The mechanical importance of the alveolar structure, which certainly increases the elasticity and mechanical resistance of the voluminous calymma, has not yet been sufficiently realised; in the case of those Radiolaria which have no skeleton, or at all events no lattice-shell, it may take the place of this as a protective envelope. Furthermore, by taking in and giving out water it may discharge a hydrostatic function, causing the organism to rise or sink in the water.
C. The large central alveole found in the colonies of many Polycyttaria (especially Collosphærida) and first described in my Monograph (Taf. xxxiv. fig. 1), has since then been observed by Hertwig, Bütschli, and other investigators, and recognised as the "central support of the whole colony, surrounded by a delicate membrane" (compare L. N. [33], p. 31, and L. N. [41], p. 436). In a colony of Trypanosphæra transformata (Pl. [5], fig. 1), which I observed living while in Ceylon in 1881, the membrane of the large central alveole was surrounded by a firm network of sarcoplegma, and could be mechanically isolated from the central jelly-sphere which it enclosed.
D. The pericapsular alveoles, figured in Pl. [4], figs. 2, 3, from a Sphærozoum, and in Pl. [6], fig. 2, from a Siphonosphæra, were very well preserved in some preparations in the Challenger collection; perhaps their development coincides with the formation of spores, and may be regarded as an encystation.
87. The Extracapsular Fat-Globules.—Fat is probably as widely distributed in the exoplasm as in the endoplasm of the Radiolaria; a considerable proportion of the small, dark, highly refractive granules appear to consist of fat; most likely they are for the most part direct products of metastasis. These widely-spread granules, which are sometimes coloured, and which by their passive motion produce the phenomenon of granular circulation in the exoplasm, are not the only fatty structures in the extracapsulum; larger globules sometimes occur. In certain large Collodaria (e.g., Thalassicolla melacapsa, Pl. [1], fig. 5; Thalassophysa sanguinolenta, &c.) radial series of oil-globules are found in the calymma, especially in its proximal portion; in others the central capsule is surrounded by a layer of oil-globules (situated in the sarcomatrix). In the Phæodaria a part of the phæodium appears to consist of fat-globules.
88. The Extracapsular Pigment.—The formation of colouring matters in the extracapsulum is on the whole rare in the Radiolaria, apart from the "yellow cells" (see § [91]) and from the peculiar phæodium of the Phæodaria, which will be separately treated of in the next paragraph. Considerable masses of extracapsular pigment, usually black or blue, rarely brown or red, are found only in a few Radiolaria belonging to the first three legions; most often in the Spumellaria. Some large Collodaria, e.g., the common Thalassicolla nucleata and a few other species of this genus (Pl. [1], fig. 4), are characterised by a rich deposit of black or blue pigment in the sarcomatrix and in the proximal portion of the calymma. Brown pigment is deposited in the calymma of many Sphæroidea and Discoidea, as well as of some Nassellaria (Cystidium, Tridictyopus, &c.). In a part of the Acantharia red pigment granules are thickly strewn in the sarcoplegma and pass along the free pseudopodia, as for example in Actinelius purpureus and Acanthostaurus purpurascens. The composition and significance of these extracapsular pigments are not completely known.
On the extracapsular pigment of Thalassicolla nucleata, compare my Monograph, pp. 87, 251. On the red extracapsular pigment-granules of the Acantharia, see L. N. [19], pp. 345, 364, &c.
89. The Phæodium of the Phæodaria.—The Phæodaria, which are distinguished from the other three legions of Radiolaria by the double membrane of the central capsule, and the peculiar structure of the main-opening (astropyle), differ also in other points, the most important of which is the constant presence of a voluminous mass of extracapsular pigment. This possesses a peculiar constitution and special significance, and is not to be confounded with the extracapsular pigment-granules of other Radiolaria (e.g., Thalassicolla), and hence it has been distinguished by the name "Phæodium," and the individual granules which compose it as "Phæodella" (see note A). The phæodium is always excentric in position relatively to the central capsule, of which it surrounds the oral half in the form of a voluminous concavo-convex cap, hiding the astropyle at its basal pole so completely that the latter is rarely visible until the phæodium has been removed (Pls. [99]-[104]; Pl. [115], fig. 8; Pl. [123], &c.). The central capsule is generally almost completely embedded in the phæodium, so that only its aboral pole (with the two parapylæ in the Tripylea) projects. In the Phæogromia, in which the lattice-shell possesses a special opening and the central capsule lies excentrically in the aboral position of its interior, the phæodium occupies the oral aspect, between the capsule and the aperture (Pls. [99], [100], [118]-[120], &c.). In the peculiar family Cœlographida (Pls. [126]-[128]) a special receptacle (galea with its rhinocanna) for the phæodium is developed outside the bivalve shell, within which the central capsule lies. The proboscis, which in all Phæodaria arises from the centre of the astropyle, lies in the vertical axis of the phæodium and is entirely surrounded by it. The volume of the phæodium in the majority of the Phæodaria may be said to be about as great as that of the central capsule, although in some species it is considerably larger. Its colour is always dark, usually between green and brown, commonly olive-green or blackish-brown, rarely reddish-brown or black. The phæodellæ or pigment-granules which make up the greater part of the phæodium (see note B) are irregular in form and unequal in size and show no definite structure; usually they are spherical or ellipsoidal, and exhibit fine parallel striæ which run transversely or obliquely (Pl. [101], fig. 3, 6, 10; Pl. [103], fig. 1, &c.). Between the larger granules is usually found a thick dust-like mass of innumerable very small grains. The physiological significance of this peculiar phæodium is still unknown, but is probably considerable, judging from its large size and especially from its constant topographical relation to the astropyle; the latter consideration would lead to the supposition that it plays an important part in the nutrition and metastasis of the Phæodaria (see note C).
A. The phæodium of Aulacantha, Thalassoplancta, and Cœlodendrum was first described in 1862, in my Monograph, as an excentric extracapsular mass of pigment of blackish-brown or olive-green colour (pp. 87, 262, 264, 361, Taf. ii. iii. xxxii.). Since then John Murray, who investigated many living Phæodaria during the Challenger expedition, has shown its general distribution in this legion (Proc. Roy. Soc. Lond., vol. xxiv. p. 536, 1876). From the constancy of its presence I gave the legion the name Phæodaria in 1879 (L. N. [34]).
B. With regard to the special composition of the phæodium and the constitution of the phæodellæ, see the general description of the Phæodaria, pp. [1533]-[1537].