On the Development of the Plumes in Buds of Cephalodiscus. by W. O
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PLUMES OF CEPHALODISCUS. 221 On the Development of the Plumes in Buds of Cephalodiscus. By W. O. Ridcwood, D.Sc, Lecturer on Biology at St. Mary's Medical School, University of London. Witli 11 Text-flgures. PAGE Introduction ....... 221 Methods . .223 Development of the Plumes in Buds of Cephalodiscus hodgsoni ...... 224 Development of the Plumes in Buds of Cephalodiscus dodecalophus ...... 230 Development of the Plumes in Buds of Cephalodiscus nigrescens ...... 235 Development of the Plumes in Buds of Cepbalodiscus gilchristi . .242 General Remarks on the Collar and its Appendages . 245 Summary ....... 251 Explanation of the Abbreviations employed in Text-figures 2, 3,4, 6, 7, and 9 . .252 INTRODUCTION. During the progress of an investigation on the anatomy of three recently discovered species of Cephalodiscus (C. nigrescens, C. hodgsoni, and C. gilchristi), kindly entrusted to me for description by Prof. B. Ray Lankester, F.R.S., Director of the Natural History Museum, London, I noticed that in the development of the buds there is no torsion of the first and second plume-axes such as is de- scribed by Masterman as occurring in buds of C. dodeca- 222 W. G. MDEWOOD. lophus,1 and that the last plumes do not develop between the earlier plumes and the buccal shield, as recorded by the same writer, but on the side of those plumes remote from the buccal shield. His figures 64—68 are reproduced as text- fig. 1. I was unable to settle the point satisfactorily in time for the introduction of my results iuto the two papers describing the structure of the polypides and tubaria of the new species/ but since the completion of those papers I have made an exhaustive study of the growth, of the plumes in buds of TEXT-FIGUHE 1.—Diagrammatic transverse sections of plumes and buccal shield of Ceplialodiscus dodecaloplius at five stages of development. Copied from A. T. MasLermau's paper in the 'Trans. Roy. Soc. Edin.,' vol. xxxix, part 3, 1898, plate 4, figs. 64—68. The original numbering- has been retained. Oephalodiscus hodgsoni, 0. nigrescens, and C. gil- christi, aud having made from the abundant material at my disposal a well-graded series of buds of each species, I was enabled to confirm my earlier conclusions. I finally prepared a series of buds of the " Challenger " species, Cephalodiscus dodecalophus, with the result that I am satisfied that the buds of this species agree with 1 'Trans. Roy. Soc. Edin.,' vol. xxxix, part 3, 1898, p. 521. 3 '"Discovery" Expedition Reports,' vol. ii, 1007, and 'Marine Investi- gations in South Africa,' vol. iv, 190(5. PLUMES OF CEPHALODISOUS. 223 those of the other three species. The torsion described by Masterman does not occur, but the grooves of the first and second pairs of plumes, developed iu such a manner that they face ventrally towards the buccal shield, continue to face the shield. The last two pairs of plumes are not deve- loped between the first two pairs and the shield, but on the dorsal side of them, i.e. on the side remote from the shield. The gi-ooves of these last plumes (fifth and sixth pairs) are not directed towards the shield, but away from it. Although the four series of plumes of buds were prepared for the purpose oE deciding the points raised by Mastermau, they serve to show a number of other interesting features, such as (1) the difference of plume development in the buds of the four species; (2) the relation which the last-formed pair of plumes bear to the edge of the post-oral lamella; and (3) the conversion, iu the later stages of growth, of the line of the bases of the collar-outgrowths (plumes and post- oral lamella) from a circle to a double crescent, or two incomplete ellipses. I have selected from each of the four series a number of preparations to illustrate these points; they are reproduced in text-figures 2, 3, 4, 6, 7, aud 9, and are described under headings of the respective species. METHODS. In the case of the youngest stages the whole bud was mounted iu diluted glycerine, aud gold size was run round the edge of the cover-glass to keep it firmly iu position, and to prevent the glycerine from accumulating dust. Most of the buds were dissected, the shield being first removed by tearing through its stalk by the aid of fine mounted needles, and then the collar region, with plumes and post-oral lamella (e.g. text-fig. 2, J), was removed by carefully manipulating the needles between these parts and the " body" of the bud. The three parts, shield, collar region, aud " body " with its stalk, were then mounted on the same slide in dilute glyce- rine. 224 W. G. EIDEWOOD. The relation which the collar region, as dissected out, bears to the whole bud will be made clear by reference to text- fig. 5, A, a diagrammatic side view of a bud with three pairs of plumes, and text-fig. 10, a diagram of a longitudinal section of the adult polypide; in the latter figure the limit of the collar region is marked by two lines of heavy dashes. The results here recorded and illustrated are based upon 135 preparations of buds made and mounted as above ex- plained. In some cases (e.g. text-fig. 6, F) the three parts— " body," collar appendages, and shield—were drawn sepa- rately on tracing paper, and the perfect bud was recon- structed by a superposing of these transparent sheets. No staining fluids were used. The dissections were made under a Greenough binocular erecting microscope magni- fying 20 and 40 diameters; the drawings were made by the aid of a compound microscope with Zeiss apochromatic 8 mm. objective and No. 6 compensating ocular with eye-piece micrometer. The figures in text-figs. 2, 3, 4, 6, 7, and 9 are all reproduced to the same scale of enlargement, viz. 62 diameters. DEVELOPMENT OF THE PLUMES IN BUDS OF OEPHALODISCUS HODGSONI. In the earliest phase of development the bud is ovoid or pyriform (text-fig. 2, A) without signs of buccal shield or plumes. The shield and the first pair of plumes make their appearance simultaneously (B and c). The plumes are at first small mounds, later hemispherical (D), and still later longer than broad (K, F, etc.). The second pair of plumes appear on the external side of the first pair (D, 2), and in contact with them. The two plumes of the first pair are not in con- tact with one another at their bases. The third pair of plumes appear lateral to the second pair (E, 3), and in con- tact with them. The bud increases considerably in size before the fourth PLUMBS OF OEPHALODISCUS. 225 plumes appear; the buccal shield assumes its definitive shape by becoming flattened, and by the differentiation of the pos- b.s. ,-b.s. G. TEXT-FIGUHE 2.—Cephalodiscus hodgsoni. Figures illustrating the development of the plumes of the buds. C and L are side views, the others are dorsal views. J represents the collar and its appendages (plumes and post-oral lamella) dissected from a bud a little older than H. The Qrst pair of plumes are-just appearing in B and C, the second in D, the third in E, the fourth in H, and the fifth in K and L. The series is continued in text-fig. 3. All figures enlarged 62 diameters. Eor explanation of the lettering see end of paper. terior lobe. In G and H the dotted line marks the position of the free edge of the posterior lobe. The fourth plumes 226 "W. G. RIDEWOOD. appear when the bud has reached the stage of development represented in H. The structure which in G is marked po.l. might at first glance be taken for the developing fourth plume; it is the lateral part of the post-oral lamella. Figure J represents the collar region and its appendages (the plumes and post-oral lamella) of a bud a little older than that shown in figure H; the buccal shield has been dissected off from the one side, and the "body" and stalk from the other. The position of the mouth is indicated in the figure, but this region is always more or less damaged in making such a dissection as that described. The torn stalk of the buccal shield in this preparation—as also in the similar pre- parations shown in text-fig. 3, N, O, and u, and in other of the text-figures—occupies a position intermediate between the mouth aud the bases of the first pair of plumes. In figure J the fourth plumes are hemispherical projections at the bases of the third plumes, touching these on the one side, aud in continuity with the post-oral lamella on the other. Traces of pinnules are appearing on the first and second plume-axes, and the terminal end-bulbs of these same plumes are swelling out. In K the fourth plumes are now longer than broad, and the fifth pair are making their appearance, L is a side view of a bud of about the same stage as K, as may be determined by comparing the development of the fourth aud fifth plumes in the two buds. Pinnules are now appearing on the third plumes, and it will be noticed, in the case of the first, second, and third plumes, that the pinnules are not laterally placed on the plume-axes, but ventro-laterally, and the groove between the two rows of pinnules of each of the plumes in question opens ventrally, or ventro-laterally, towards the dorsal or stalked side of the shield.