may increase in size by the fusion (“plastogamic”) of small individuals. If a large specimen be cut up or fragment itself under irritation, the small ones so produced soon approach one another and fuse completely.
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Fig. 2.—Heliozoa. 1. Actinosphaerium eichhornii, Ehr.; a, nuclei; b, deeper protoplasm with smaller vacuoles and numerous nuclei; c, contractile vacuoles; d, peripheral protoplasm with larger vacuoles. 2. A portion of the same specimen more highly magnified and seen in optical section. a, Nuclei; b, deeper protoplasm (so-called endosarc); d, peripheral protoplasm (so-called ectosarc); e, pseudopodia showing the granular protoplasm streaming over the stiff axial filament: f, food-particle in a good-vacuole. 3, 4. Nuclei of Actinosphaerium in the resting condition. 5-13. Successive stages in the division of a nucleus of Actinosphaerium, showing fibrillation, and in 7 and 8 formation of an equatorial plate of chromatin substance (after Hertwig). 14. Cyst-phase of Actinosphaerium eichhornii, showing the protoplasm divided into twelve chlamydospores, each of which has a siliceous coat; a, nucleus of the spore; g, gelatinous wall of the cyst; h, siliceous coat of the spore. |
Reproduction.—Binary fission has been repeatedly observed; in some cases one or both of the daughter cells may swim for a time as a biflagellate zoospore (fig. 1 (6, 7)). The process may take place when the cell is naked or after preliminary encystment. Budding has been well studied in Acanthocystis; the cell nucleus divides repeatedly and most of the daughter nuclei pass to the periphery, aggregate part of the cytoplasm, and with it are constricted off as independent cells; one nucleus remains central and the process may be repeated. The detached bud may assume the typical character after a short amoeboid (lobose) stage, sometimes preceded by rest, or it may develop 2 flagella and swim off (fig. 1 (6)).
Brood formation is only known here in relation to a syngamic process; this is a sharp contrast to Proteomyxa (q.v.) where brood formation is the commonest mode of reproduction, and plasmodium-formation, rare indeed, is the nearest approach to syngamy observed. Indeed, if we knew the life-history of all the species this difference in the life cycle would be a convenient critical character.
Equal conjugation was demonstrated fully by F. Schaudinn in Actinophrys; two individuals approach and enter into close contact, and are surrounded by a common cyst wall. The nucleus of either male divides; and one nucleus passes to the surface at either side, and is budded off with a small portion of the cytoplasm as an abortive cell; the two remaining nuclei which are “first cousins” in cellular relationship now fuse, as is the case with the cytoplasts. The resulting coupled cell or zygote divides into two, which again encyst.
Actinosphaerium (fig. 2) shows a still more remarkable process, fully studied by R. Hertwig. The large multinucleate animal withdraws its pseudopods, its vacuoles disappear, it encysts and its nuclei diminish in number to about 120th partly by fusion, 2 and 2, probably by digestion of the majority. Within the primary cyst the body is now resolved into nuclear cells, which again surround themselves with secondary cysts. The cell in each secondary cyst divides (by karyokinesis), and these sister cells, or rather their offspring, pair in much the same way as the individual cells of Actinophrys—the chief difference is that after the first division and budding off of a rudimentary cell, a second division of the same character takes place, with the formation of a second rudimentary cell, which is the niece of the first, absolutely in the same way as the 1st and 2nd polar bodies are formed in the maturation of the ovum in Metazoa. The actual pairing cells are thus second cousins, great-granddaughters of the original cell of the secondary cysts. Complete fusion now takes place to form the coupled cell, which is now contracted and forms a gelatinous wall within the siliceous secondary cyst wall (fig. 2 (14)), During a resting stage nuclear divisions occur and finally a brood of young 1-nuclear Actinosphaerium leave the cyst.
Classification.
Aphrothoraca. Body naked. Actinophrys Ehrb. (fig. 1 (1)) (nucleate), Actinosphaerium Stein plurinucleate (fig. 2 (1)), Camptonema (plurinucleate) Schaud., Dimorpha Gruber (sometimes 2 flagellate).
I. Chlamydophora. Investment gelatinous. Astrodiscus.
II. Chalarothoraca. Body protected by an investment of spicules or fibre scattered or approximated, never fused into a continuous skeleton.
- § 1. Spicules netted or free in the protoplasm. Heterophrys Arch. (fig. 1 (3)), Raphidiophrys Arch. (fig. 1 (4)), Pinacodocystis, Hertw. and Less.
- § 2. Spicules approximated radially. Pinaciophora Greeff, Pompholyxophrys Arch., Lithocolla F. E. Schultze, Elaeorhanis Greeff (in the two foregoing genera the spicules represented by sand granules), Acanthocystis Carter (fig. 1 (5)), Pinacocystis (?) Hertw. and Less, Myriophrys Penard. (Astrodisculus).
III. Desmothoraca. § 1 attached by a stalk. Clathrulina Cienk. (fig. 1 (2, 7)), Hedriocystis, Hertw. and Less.
- § 2. Free Elaster, Grimin, Choanocystis.
Literature.—The most important English original papers on this group are those by W. Archer, “On some Freshwater Rhizopoda, new, or little known,” Quarterly Journal of Microscopic Science, N.S. ix.-xi. (1869–1871), and “Résumé of Recent Contributions to the Knowledge of Freshwater Rhizopods,” ibid. xvi., xvii. (1876–1877). See also R. Hertwig and Lesser, “Über Rhizopoda und denselben nahestehenden Organismen,” in Archiv für mikroscopische Anatomie, x. (1874), p. 35; R. Schaudinn, “Heliozoa” in Tierreich (1896); E. Penard, Les Héliozoaires d’eau douce (1904); the two last named contain full bibliographies. (M. Ha.)
HELIUM (from Gr. ἥλιος, the sun), a gaseous chemical element, the modern discovery of which followed closely on that of argon (q.v.). The Investigations of Lord Rayleigh and Sir William Ramsay had shown that indifference to chemical reagents did not sufficiently characterize an unknown gas as nitrogen, and it became necessary to reinvestigate other cases of the occurrence of “nitrogen” in nature. H. Miers drew Ramsay’s attention to the work of W. F. Hillebrand, who had noticed, in examining the mineral uraninite, that an inert gas was evolved when the mineral was decomposed with acid. Ramsay, repeating these experiments, found that the inert gas emitted refused
