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Paleontological Contributions THE UNIVERSITY OF KANSAS PALEONTOLOGICAL CONTRIBUTIONS March 1, 1971 Paper 52 CONCEPT OF COMMON BUD AND RELATED PHENOMENA IN BRYOZOA MAXIM K. ELIAS University of Oklahoma, Norman, Oklahoma ABSTRACT The common bud of Stenolaemata (Cyclostomata of authors), as understood by BORG (1926), is analogous to the buds of higher plants in its terminal position on branches of the Bryozoa and in its vital activities. The common bud is truly common to all developing zooecia in a branch of a zoarium in a sense that its coelomic fluid remains undivided distally as long as a branch grows. As soon as a polypide bud originates, subtending to it, an oblique septum begins to rise proxi- mally by fission (BoRc's term) from the cylindrical wall of a branch; thus, a new zooecium with its own polypide is being initiated. All polypide buds originate at the edge of a transverse membrane that seals the coelomic fluid terminally. As the branch continues to grow the developing polypide falls behind the growing edge of the membrane, which BORG named growing zone. Some biologists (including HYMAN) erroneously consider growing zone synonymous with common bud— but the growing zone has the same relationship to the common bud as the meristem of plants has to their bud. The walls and septa of a zoaritnn which possesses a common bud combine their cal- careous layers into a single continuous structure, the colonial plexus (EtaAs' and CONDRA ' S term). The Bryozoa possessing common buds attain a higher level of colonial individualization (sensu BEKLEMISHEV) than those without it, and the greater the transverse expansion and complication of the common bud, the greater the colonial individualization—such as at- tained in more advanced Stenolaemata. Cheilostomata lack common buds, but in Membranipora and related crustose cheilo- stomes, whose colonies are built by contiguous rows of zooecia, each row develops by the means of its terminal linear bud (new term), with its own growing zone. The Ordovician "graptolite" Reticulograptus closely resembles the extant stenolaemate Stegoh ornera violacea (Sars) BORG INTRODUCTION The idea of the common bud, by means of (1926a, p. 254), SMITT believed "that in all Bryo- which the bryozoan colonies are developed, is zoa the developing margin of the zoarium is to be an imaginative concept conceived by F. A. SMITT regarded 'as a colonial bud, a common bud (sam- about a hundred years ago. As BORC briefly stated knopp), which separates [divides] itself by means 2 The University of Kansas Paleontological Contributions—Paper 52 of fission, and develops into separate zoids BORG also clarified biological understanding of [zooidsf " (Smurr, 1865, p. 6, translated by the "developing margin" in the Cyclostomata and BoRc). He erred, however, in his belief that all indicated it to be the growing zone of the com- Bryozoa developed in this manner; and it was mon bud. His most informative statement is NITSCHE (1871, sep. 31 ff.) who explained that quoted: it was not the case with Cheilostomata and Cteno- "At the edge of the terminal membrane the stomata. Accepting this correction, BORG worked ectodermal cells are higher, very closely packed out a revised concept of common bud, and estab- together without any intercellular spaces (pl. 3, lished on the evidence of its presence a "new fig. 12); their cytoplasm is densely granulated. order for the Cyclostomata, co-ordinate with two The mesodermic cells are also placed considerably old orders Gymnolaemata and Ctenostomata . ." closer together. The [ cellular] transition from to "be termed Stenolaemata" (BoRc, 1926a, p. this [marginal zone to the remaining part of 490). This taxonomic connotation ascribes to the the terminal membrane and to the lateral walls common bud foremost significance in a natural of the common bud [ below] is gradually effected. classification of Bryozoa. It is in this zone at the edge of the terminal mem- Whereas BORG described the stages of devel- brane that the growth of the common bud mainly opment of the modern Cyclostomata by the means takes place. Here new cells originate; here new of common bud, he noted that "certain differ- substance is stored in the cuticle, secreted from ences between the various cyclostomatous forms the ectodermal cells; here is also secreted calcare- present themselves even in the earliest develop- ous matter (chapter 4); and, finally, it is here, mental stages," although the common bud itself too, and only here, that the rudiments of the originates and develops in a similar manner from polypides [ polypide buds [ first become visible. the "semispherical primary disc characteristic for This zone in question may conveniently be termed all Cyclostomata" (1926a, p. 255). He also men- the growing zone of the common bud" (BoRc, tioned, in an unobtrusive way, his important 1926a, p. 321 [his underscoring). It is obvious discovery of an optical orientation of the minute that in this statement BORG described the structure particles comprising the calcified layer in the and activities of only a part—even if most essen- Crisiidae, which he examined in polarized light. tial—of the common bud: its growing zone. He found that these particles "are located in a However, because he did not attempt a formal certain direction; namely parallel to the longi- definition of the common bud and limited himself tudinal [along the branches] axis of the [ com- to specific descriptions of its occurrence in various mon} bud"; and, he noted, "the same is true also cyclostomes, he left his concept exposed to mis- of the calcified walls of the zoids [zooids]." No understandings and misinterpretations. observation of similar nature has ever been made elsewhere in the description of other groups of ACKNOWLEDGMENTS In view of the taxonomic the Cyclostomata in this or other books by BORG, connotation of the but greater biological and taxonomical significance concept of common bud, implied in its revision to the optical orientation of the particles in the by BORG (1926a), the present review of the con- calcified layer in Crisiidae was added by the dis- cept became an essential part in biometric research covery of a similar optical orientation of the calcite on fenestrate bryozoans, where an evidence of the in the unit-crystal primary skeleton in fenestrate presence of a common bud was reported by ELIAS bryozoans, hence suggestive of the presence in & CONDRA (1957). The whole research has been them of the common bud.' supported by several grants from the National Science Foundation, the last one in 1967-68, no. I resolutely disagree with the speculative concept of growth GA-817. in fenestellid branches by means of a "conveyor belt" principle, which was adapted for this purpose by TAVENER-SMITH (1969) in Parts of the manuscript have been read by Dr. preference of the natural (for them) growth by means of common HELEN DUNCAN, bud. It seems rather ridiculous to imagine that the same mech- U.S. Geological Survey, and the anism which was imaginatively conceived to explain the growth whole, in its final state, by of certain brachiopod shells by ALWYN WILLIAMS (1966, 1968), Dr. RAYMOND C. would need to be proposed for explanation of the growth of a MOORE, University of Kansas. I am indebted to colonial organism. Even HYMAN (1959), who with the help of her academic redefinition of the well recognized concept of Lophophore, ROGER B. WILLIAMS, of the Paleontological Insti- united into her "Lophophorate Coelomates" (1959, p. 228 ff.) the phyla Phoronida, Ectoprocta, and Brachiopoda, was avoiding any tute, for skillful preparation of new illustrations comparison of growth of the bryozoan colonies to that of brachio- pod shells. after BORG. Elias—Common Bud and Related Phenomena in Bryozoa 3 COMPONENTS OF COMMON BUD The following components are present in the and drag along their sides, as it were, the ecto- common bud of all stenolaemate bryozoan col- derm and mesoderm, which cover the calcareous onies: wall (or septum), from which they originate. ) Terminal membrane which stretches across Thus each septum consists of a central calcareous and covers the bud cavity at the distal margin layer, which is flanked on either side by ectoder- (or margins) in a developing colony. The ter- mal and mesodermal layers. This manner of minal membrane consists (from outside inward) origin "by fission" makes all septa inherently of a very thin cuticle, a one-cell thick layer of possessive of a single primary calcareous layer, ectoderm, and a one-cell thick layer of diffusely directly continuing from the calcareous layer of spaced cells of mesoderm. the zoarial wall, from which they originate. It 2) Coelomic cavity, which is filled with co- is this uniform, one-layered, calcareous structure elomic fluid, sealed by a terminal membrane of all walls and septa which is highly character- above, and contained laterally and proximally by istic of stenolaemates. And because it is a result an outer wall and oblique septa or both. of colonial development through mechanism of bud, the definition of the latter must take 3) Colonial wall and partitions (septa), all of common which (from outside inward) consist of a very thin this into account. cuticle comprising a calcareous layer (precipitated 6) Otherwise oriented calcareous septa, which by ectodermal epithelium), ectoderm, and meso- originate from the oblique septa, such as "median derm (Fig. 1). vertical" (BoRG, 1926a, p. 270) septa rising along the two opposite oblique 4) Growing zone, a ringlike area along the the linear coalescence of septa in Acamptostega (Stomatopora, Tubuli- contact of the terminal membrane and the colonial pora), as explained below. wall, where new cells of ectoderm and mesoderm originate and polypi& buds make their "first appearance." SPECIAL TERMS PERTAINING TO 5) Oblique calcareous septa, which arise in STENOLAEMATA (CYCLOSTO- part of the coelomic cavity by send- the proximal MATA) AND FENESTRATA ing out from the colonial wall, as if by fission, oblique ingrowths (internal outgrowths) which Proper understanding of special terms which ascend subparallel to the wall toward the terminal indicate the peculiar processes in growth by means membrane.
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