Order xx. Phaeoconchia.—Shell of two valves, opening in the plane ("frontal") of the three openings of the capsule. (Families 83-85.)
We exclude Haeckel's Dictyochida, with a skeleton recalling that of the Stephoidea, but of the impure hollow substance of the Phaeodaria (p. [84]). They rank now as Silicoflagellates (p. [114]).
The Radiolarian is distinguished from all other Protozoa by the chitinous central capsule, so that its cytoplasm is separated into an outer layer, the extracapsular protoplasm (ectoplasm), and a central mass, the intracapsular, containing the nucleus.[[88]]
The extracapsular layer forms in its substance a gelatinous mass, of variable reaction, through which the plasma itself ramifies as a network of threads ("sarcodictyum"), uniting at the surface to constitute the foundation for the pseudopodia. This gelatinous matter constitutes the "calymma." It is largely vacuolated, the vacuoles ("alveoli"), of exceptional size, lying in the nodes of the plasmic network, and containing a liquid probably of lower specific gravity than seawater; and they are especially abundant towards the surface, where they touch and become polygonal. On mechanical irritation they disappear, to be formed anew after an interval, a fact that may explain the sinking from the surface in disturbed water. This layer may contain minute pigment granules, but the droplets of oil and of albuminous matter frequent in the central layer are rare here. The "yellow cells" of a symbiotic Flagellate or Alga, Zooxanthella, are embedded in the jelly of all except Phaeodaria, and the whole ectosarc has the average consistency of a firm jelly.
The pseudopodia are long and radiating, with a granular external layer, whose streaming movements are continuous with those of the inner network. In the Acantharia they contain a firm axial filament, like that of the Heliozoa, which is traceable to the central capsule; and occasionally a bundle of pseudopodia may coalesce to form a stout process like a flagellum ("sarcoflagellum"). Here, too, each spine, at its exit from the jelly, is surrounded by a little cone of contractile filaments, the myophrisks, whose action seems to be to pull up the jelly and increase the volume of the spherical body so as to diminish its density.
Fig. 25.—Skeletons of Radiolaria. A, Dorataspis; B, Theoconus. (After Haeckel.)
The intracapsular protoplasm is free from Zooxanthella except in the Acantharia. It is less abundantly vacuolated, and is finely granular. In the Porulosa it shows a radial arrangement, with pyramidal stretches of hyaline plasma separated by intervals rich in granules. Besides the alveoli with watery contents, others are present with albuminoid matter in solution. Oil-drops, often brilliantly coloured, occur either in the plasma or floating in either kind of vacuole; and they are often luminous at night. Added to these, the intracapsular plasm contains pigment-granules, most frequently red or orange, passing into yellow or brown, though violet, blue, and green also occur. The "phaeodium,"[[89]] however, that gives its name to the Phaeodaria, is an aggregate of dark grey, green, or brown granules which are probably formed in the endoplasm, but accumulate in the extracapsular plasm of the oral side of the central capsule. Inorganic concretions and crystals are also found in the contents of the central capsule, as well as aggregates of unknown composition, resembling starch-grains in structure.
In the Monopylaea, or Nassellaria (Figs. 25, B, 26, A), the endoplasm is differentiated above the perforated area of the central capsule into a cone of radiating filaments termed the "porocone," which may be channels for the communication between the exoplasm and the endoplasm, or perhaps serve, as Haeckel suggests, to raise, by their contraction, the perforated area: he compares them to the myophane striae of Infusoria. In the Phaeodaria (Fig. 26, B), a radiating laminated cone is seen in the outermost layer of the endoplasm above the principal opening ("astropyle"), and a fibrillar one around the two accessory ones ("parapyles"); and in some cases, continuous with these, the whole outer layer of the endoplasm shows a meridional striation.
The nucleus is contained in the endoplasm, and is always at first single, though it may divide again and again. The nuclear wall is a firm membrane, sometimes finely porous. If there are concentric shells it at first occupies the innermost, which it may actually come to enclose, protruding lobes which grow through the several perforations of the lattice-work, finally coalescing outside completely, so as to show no signs of the joins. In the Nassellaria a similar process usually results in the formation of a lobed nucleus, contained in an equally lobed central capsule. The chromatin of the nucleus may be concentrated into a central mass, or distributed into several "nucleoli," or it may assume the form of a twisted, gut-like filament, or, again, the nuclear plasm may be reticulated, with the chromatin deposited at the nodes of the network.