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Encyclopedia Britannica



HELIOZOA

This article appears in Volume V13, Page 233 of the Encyclopedia Britannica.

Encyclopedia Britannica - Main :: HEG-HIG
HELIOZOA , in
zoology
 , a group of the Sarcodina (q.v.) so named by E. Haeckel, 1866. They are characterized by the radiate pseudopods, finely tapering at the apex, springing abruptly from the superficial protoplasm, containing a denser, rather permanent axial rod (figs. I (I), 2 (2); protoplasm without a clear ectoplasm or pellicle, often frothy with large vacuoles, like the alveoli of Radiolaria; nucleus I or numerous; skeleton absent, gelatinous or of separate siliceous fibres, plates or spicules, rarely complete and latticed; reproduction by simple fission or by brood-formation, often syngamous; form usually nearly spherical, rarely changing slowly. This group was formerly included with the Rhizopoda; but was separated from it by Haeckel on account of the character of its pseudopods, and its general adaptation to a semipelagic existence correlated with the frothy cytoplasm (fig. I (I)). Actinophrys sol and Actinosphaerium eichhornii (fig. 2), known as sun animalcules to the older microscopists, float freely in stagnant or slow-flowing waters, and Alyriophrys is able by an investment of long fiagelliform cilia to swim freely. The majority, however, lurk among confervae or the light debris of the bottom
ooze
 ; and come under the head of " sapropelic " rather than pelagic organisms. The
body
  is usually of constant spherical form in relation to the floating habit. Nuclearia, however, shows amoeboid changes of general outline. The pseudopods are retractile, the axial filament being absorbed as the filament grows shorter and thicker and disappearing when the pseudopod merges into the ectoplasm, to he reformed at the same time with the pseudopod. There is often a distinction, clear, but never
sharp
 , between the richly vacuolate, almost frothy ectoplasm and the denser endoplasm. One or more contractile vacuoles may protrude from the ectoplasm. The endoplasm contains the nucleus or nuclei. The nucleus when single may be central or excentric: in the latter case, the endoplasm contains a clear central sphere (" centrosome ") on which abut the axial filaments of the pseudo-pods. The ectoplasm contains, in some species, constantly (Raphidiophrys viridis) or occasionally (A ctinosphaerium), green cells belonging to the genera Zoochlorella and Sphaerocystis, both probablythe latter certainlyvegetative stages of a Chlamydomonad (FLAGELLATA, q.v.) and of symbiotic significance.
The Heliozoa can move by rolling over on their extended pseudo-pods; Acanthocystis ludibunda traversing a path of as much as twenty times its
diameter
  in a minute, according to Penard. several species (e.g. Raphidiophrys elegans) remain associated by the union of their pseudopods, whether into social aggregates (due to approximation) or " colonies " due to lack of separation1 ....a 8
after fission, is not accurately known. The multinuclear species Actinosphaerium eichhornii (fig. 2), normally apocytial (i.e. the nuclei divide repeatedly without division of the cytoplasm),
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.
Reproduction.Binary fission has been repeatedly observed; in some cases one or both of the daughter cells may swim for a timeas a biflagellate zoospore (fig. i (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. T (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 th 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 Actinophrysthe
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 (I)) (nucleate), Actinosphaerium Stein plurinucleate (fig. 2 (I)), Camptonema (plurinucleate) Schaud., Dimorpha Gruber (some-times 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. i (5)), Pinacocystis (?) Hertw. and Less, Myriophrys Penard. (Astrodisculus).
2. Free Elaster, Grimin, Choanocystis.


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