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On the Ciliary Mechanisms and Interrelationships of Lamellibranchs

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On the Ciliary Mechanisms and Interrelationships of Lamellibranchs On the Ciliary Mechanisms and Interrelationships of Lamellibranchs. PAET IV: Cuticular Fusion, with special reference to the Fourth Aperture in certain Lamelli- branchs. By Daphne Atkins, B.Sc. Marine Biological Laboratory, Plymouth. With 11 Text-figures. CONTENTS. PAGE INTRODUCTION 423 HISTOLOGICAL STRUCTURE OF THE JUNCTION .... 424 CUTICULAK FUSION IN RELATION TO THE FOURTH APERTURE IN CERTAIN LAMELLIBRANCHS ....... 436 Origin of the Fourth Aperture ....... 442 SUMMARY ........... 444 INTBODUCTION. A PECULIAR form of fusion, involving the cuticle only, has been found in a number of Lamellibranchs, but has been chiefly studied in the Solenidae. The positions so far discovered in which this type of fusion occurs are between the dorsal edges of the ascending lamellae of the outer demibranchs and the mantle or the visceral mass in Solen marginatus Montagu (=vagina), Ensis siliqua (L.), Ensis arcuatus (Jeffreys), Cultellus pellucidus (Pennant), Solecurtus scopula (Turton) (=candidus), Lutraria lutraria (L.) (=elliptica), and Tellina tenuis da Costa; between the dorsal edges of the ascending lamellae of the two inner demibranchs in Barnea parva (Pennant); and between the mantle lobes in the region between the pedal and fourth apertures in Ensis siliqua, Ensis arcuatus, and Cultellus pellucidus. 424 D. ATKINS HlSTOLOGICAL STRUCTURE OF THE JUNCTION. Material was fixed in Bouin-Duboscq's fluid with the following formula: saturated picric acid in 90 per cent, alcohol, 2 parts; saturated corrosive sublimate (water), 3 parts; 40 per cent, formalin, 1 part; glacial acetic acid, 2 parts. The stains used were Heidenhain's iron haematoxylin counterstained with acid m.f TEXT-FIG. 1 A. Transverse section showing the method of junction of the mantle lobes in the mid-ventral region between the pedal and fourth apertures. A. Ensis siliqua (adult). X980. fuchsin; Mallory's triple stain; and eosin and light (Gatenby, 1928, p. 432): the latter was preferred as it clearly differentiated the cuticle. The epithelium of the parts concerned in this cuticular type of junction has generally, in fixed material, a distinctive appear- ance, the cells being hourglass-shaped owing to the development of spaces between them (Text-figs. 1, 3, 4). It is uncertain, however, to what extent this may be due to shrinkage on fixation: THE CILIARY MECHANISMS OF LAMELLIBRANCHS 425 intercellular spaces were rare in the specimen of Cultellus pellucidus sectioned (Text-fig. 2A). Extremely fine muscle- fibrils penetrate the cells, giving to them an appearance rather similar to that of ciliated cells in which the ciliary rootlets are clearly visible (cf. Text-figs. 1 and 11): ciliated epithelia some- times also show intercellular spaces. B e. c. cu. TEXT-FIG. 1 B. B. Ensis arcuatus (young specimen, 4 cm. long), cu., cuticle: showing a dark median line in (A); ex., hourglass-shaped epithelial cell; m.f., muscle-fibres; sp., space between the cells. Eosin and light green. x980. The cells appear to increase in height with age, for in an Ensis siliqua, 1-8 cm. long, the cells of the epithelium concerned in the junction of the mantle lobes between the pedal and fourth aperture were 5 to 6^ high; in an Ensis arcua- tus, 4 cm. long, they were 7 to 10/u. high (see Text-fig. 1 B) ; and in an adult Ensis siliqua, about 16 cm. long, they were 18 to 30/x high (see Text-fig. 1 A). The combined layers of cuticle f:A J.EXT-ITO. Z. Cultellus pellucidus. Transverse sections of the junction of the mantle lobes in the mid-ventral region between the pedal and fourth apertures. A. About 216 p anterior to the fourth aperture. B. Showing breaking down of the cells; about 60 p. in front of the fourth aperture, cu., cuticle; e.c, epithelial cell; m.f., muscle- fibres ; o.s.m., outer surface of mantle; sp., space between cells. Eosin and light green, x 980. THE CILIARY MECHANISMS OF LAMELLIBRANCHS 427 also increase in thickness with age. This latter specimen had been in the laboratory four days, and the mantle had begun to rupture, so it is possible that the stretched appearance of the cells may be due to some extent to the fact that they had suffered tension owing to the gaping of the valves, and did not recover in the piece removed for fixing. However, in the junction between the dorsal edge of the ascending lamella of the outer demibraneh and the mantle (Text-figs. 3, 4) in Ensis arcua- tus (4 cm. long) the epithelial cells were rather higher than those of the junction between the mantle lobes in the same specimen, though in the former position they would suffer relatively less stress. It is noteworthy that the mantle is pro- duced into a ridge of varying height along the line of attachment of the demibranchs (Text-fig. 3): a similar ridge is found in Solen marginatus. In an adult Solen marginatus the cells of the epithelium concerned in the union between the outer demibranchs and the mantle were high and the fibrils running through them especially clear: this was also found in the same position in Solecurtus s c o p u 1 a. The epithelial cells of the junction between the two inner demibranchs in Barnea parva were tall and slender and the cuticle clearly visible. In Tellina tenuis though the junction between the upturned outer demibranchs and the visceral mass is of the cuticular type, at least in the posterior region, yet the cells are small, have not the distinctive hourglass- shape, and the cuticle between the two parts is extremely thin and difficult to distinguish. The chief interest of this type of junction lies in the cuticle of the two opposed surfaces which appear to have undergone fusion, the sinuosity of the cuticle of one side being faithfully followed by that of the opposite side (Text-figs. 1, 2, 4). A small cavity between the two layers in the material of Ensis siliqua sectioned was probably pathological, for it contained some amorphous material. In some instances a darker median line was visible in the cuticle between the two opposed epithelial layers, when stained with eosin and light green. This was observed in the junction of the mantle lobes between the pedal and fourth apertures in an adult Ensis siliqua (Text-fig. 1 A) 428 D. ATKINS TEXT-FIG. 3. Ensis arcuatus (4cm.long). Section showing the attachment of the dorsal edge of the ascending lamella of the outer demibranch to the ridge of the mantle, asc.l., ascending lamella; m., mantle; m./., muscle-fibres; m.r., mantle ridge. Eosin and light green. X 450. and in the union of the outer demibranchs and the mantle in an Ensis arcuatus 4 cm. long (Text-fig. 4): curiously enough it was not visible in the junction between the mantle lobes in this small specimen, the appearance being as indicated THE CILIABY MECHANISMS OF LAMELLIBRANCHS 429 in Text-fig. 1 B, the median region of the cuticle staining less heavily than that on each side. A tendency for the cuticle to be faintly striated, with striae at right angles to the fused sur- face, and to separate in blocks corresponding to the cells is also evident. From the examination of sections, and from observations on living animals which will be given later, it is evident that the — m.f TEXT-FIG. 4. Ensis areuatus. Enlarged sketch of the junction of the dorsal edge of the ascending lamella of the outer demibranch and the mantle shown in Text-fig. 3. asc.L, ascending lamella; cu., cuticle, showing a dark median line; e.c., epithelial cell; m./., muscle-fibres; m.r., mantle ridge; sp., space between cells. Eosin and light green. X980. cuticular type of junction does not separate normally; when separation does occur it is due to accident or to tissue deteriora- tion on the approach of death of the animal. Rupture seems to be usually due to breaking down of the cells (Text-fig. 2 B) and not to separation of the two cuticular layers. Rupture of the junction between the outer demibranehs and the mantle, or the visceral mass, and between the two inner demibranehs has not been observed, only of that between the mantle lobes in Ensis and Cultellus on unhealthy gaping of the valves, when the mantle in the ventral region would be liable to suffer considerable strain. In Ensis siliqua the ascending lamella 430 D. ATKINS of the outer demibranch could not be separated from the mantle by pulling the demibranch. The difference in the form of the junction between the mantle lobes, and in the arrangement of the pallial muscles, in C u 11 e 1 - B TEXT-FIG. 0. Cultellus pellucid u a. Transverse sections of the ventral region of the mantle to show the general appearance: A. About 200/x anterior to the fourth aperture. B. About 150 /J. posterior to the aperture (ciliated grooves are not present in sections close to the aperture), m.l., longitudinal muscles; m.r., radial muscles; m.t., transverse muscles; n.p., pallial nerve; p., periostracum. Eosin and light green, x 70. lus pellucidus in the mid-ventral line in front of, and behind, the fourth aperture is evident from Text-fig. 5; A being a section about 200/LI anterior and B one about 150^. posterior to it. Anteriorly the junction is cuticular, the epithelium of the two opposed surfaces being clearly visible, and the muscles of the one lobe not penetrating into the other; while posteriorly THE CILIAEY MECHANISMS OF LAMELLIBRANCHS 481 fusion is complete, no trace of fused epithelial surfaces being discernible, and muscles crossing the ventral region from one side to the other.
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