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On the Mantle Cavity and Its Contained Organs in the Loricata (Placophora)

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On the Mantle Cavity and Its Contained Organs in the Loricata (Placophora) On the Mantle Cavity and its Contained Organs in the Loricata (Placophora). By C. M. Yonge, DJSe., University of Bristol. With 6 Text-figures. CONTENTS. PAGE 1. INTRODUCTION 367 2. THE MANTLE CAVITY 368 (a) Lepidochitona oinereus, p. 370; (6) Tonieella marmorea and AcanthocMtona crinitus, p. 373; (c) Lepidopleurus asellus, p. 374. 3. THE GILLS 376 4. Mucous GLANDS 382 5. OSPHRADIA AND OTHER SENSE OBGANS 383 6. DISCUSSION 386 7. SUMMARY 388 8. BEFERENCES 389 1. INTRODUCTION. The investigations described in this paper represent a con- tinuation of previous work on the mantle cavity and its con- tained organs in the Gastropoda (Yonge, 1937 c, 1938) and in the Scaphopoda (Yonge, 19376). They are intended to form part of a comprehensive survey, from the functional aspect, of these organs throughout the Mollusca. The structure and the anatomical relations of the organs contained in the mantle cavity of the Loricata are well known, as a result, in the main, of the morphological investigations of Haller (1882, 1884), Pelseneer (1898, 1899), and Plate (1898-1901). But only Arey and Crozier (1919) have made observations on the water cur- rents in the mantle cavity, and they failed to point out how these were brought about, giving no account either of the ciliation of the gills or of the manner in which the mantle cavity is divided into inhalant and exhalant chambers. 368 0. M. YONGB The greater part of this work has been carried out on two common British species, Lepidochitona cinereus (Lepi- dochitonidae) and Lepidopleurus asellus, which is one of the few representatives of the primitive order Lepidopleurida. Comparative data only has been obtained from Tonicella marmorea (Lepidochitonidae) and Acanthochitona crinitus (Oryptoplacidae).1 All of these species live between tide-marks with the exception of Lepidopleurus asellus, which, in the. English Channel, occurs at depths ranging from 15 to 43 fathoms. Comparative observations on the gills of the Gastropod, Haliotis tuberculata, were carried out at Naples during the course of other work. Acknowledgements are due to Dr. S. Kemp, F.E.S., and members of the staff of the Plymouth Laboratory, for assistance during a period spent there in connexion with this work, to Professor E. Dohm, Director of the Stazione Zoologica, Naples, to the Colston Eesearch Society of the University of Bristol for financial assistance, also to Mr. H. P. Steedman, Laboratory Steward in the Department of Zoology, University of Bristol, for cutting sections. 2. THE MANTLE CAVITY. The mantle cavity in the Loricata consists of lateral pallial grooves in communication posteriorly, and bounded internally by the sides of the foot and externally by the inner margin of the girdle. The mouth opens anteriorly and the anus posteriorly (Text-figs. 1 and 3, A.), both in the middle line. The sole of the foot frequently extends under the anus in life. Eeproductive and excretory pores open into the pallial groove on either side, near the posterior end but always in the region occupied by gills. The former openings are always the more anterior. The gills vary widely in number throughout the Loricata and are actually not constant for any one species. Thus, of the species here examined, Tonicella marmorea has between 19 and 26, Lepidochitona cinereus between 16 and 19, Acanthochitona crinitus about 15, and Lepido- pleurus asellus between 11 and 13. The gill series may be 1 The names used are those adopted by Wlnckworth (1932). MANTLE CAVITY IN LORICATA 869 holobranch or merobranch according as to whether or not they extend the full length of the pallial grooves. None of the species examined possesses gill series of the former type, although Tonicella marmorea and Lepidochitona cinereus (Text-fig. 1) approach this condition. The first-formed gill occurs in the region between plates seven and eight. According to Pelseneer (1898, 1899) the excretory pore invariably opens immediately anterior to this gill, which is morphologically, and also functionally as the present work reveals, of great significance. It will be termed throughout the post-renal gill. Pelseneer also states that it is invariably the largest gill, but, as Plate (1901) had pointed out, this is not always the case, although, if not actually the largest, it is always one of a group of especially large gills. This post-renal gill may be the last of the series, as in Lepidochitona cinereus (Text-fig. 1, GP.), in which case the condition is known as abanal. But, during development, gills may be added posteriorly as well as anteriorly to this gill. The former are termed adanal gills. There may or may not be a space between the last adanal gill and the anus. Of the species examined only Lepidopleurus asellus (Text-fig. 3) possesses adanal gills, and these extend up to the anus. Finally, the thin in- wardly projecting ridge which bounds the inner surface of the girdle is dilated on each side in the region opposite to the posterior margin of the foot. The pair of inwardly projecting mantle folds (Text-figs. 1-3, GP.) SO formed have been described by both Pelseneer and Plate.1 This investigation has revealed their function. Observations on the nature of the currents in the mantle cavity were carried out by placing animals on glass slides. When the animals had attached themselves the slides were inverted on. two pillars of plasticine in a shallow glass dish, the water in which just covered the slides. The dish was then placed on the 1 Plate (1901) states that the folds are absent in some species, but he includes amongst these Lepidopleurus asellus and Tonieella marmorea where it certainly occurs. It is probably of universal occur- rence, Plate being misled by contraction in the fixed material which he exclusively studied. NO. 323 B b 870 0. M. YONGE stage of a binocular microscope, carmine added to the water, and the nature of the water currents observed with the animal ventral side uppermost. The general observations of Arey and Crozier (1919) on Chiton tuberculatuj (a Bermudan species attaining a length of 9 cm.) were confirmed. The gills create a current of water which runs backward along the pallial grooves and out in the mid-line posteriorly (Text-figs. 1 and 8). The regions of intake vary, being created by local liftings of the girdle. When the animals are completely submerged these are usually anterior (Text-figs. 1 and 8, i.) but may be lateral, and there may be more than one opening on either side. When the anterior ©nd of an animal is out of water inhalant openings are created by a lifting of the girdle in the region still submerged. In this way a shorter, but still efficient, respiratory current is produced. The exhalant opening (Text-figs. 1-8, E.) is also created by a local raising of the girdle, in this case always at the posterior end. The full discussion of the mechanisms con- cerned in the maintenance of the respiratory currents demands separate consideration of the various species. (a) L©pidochitona cinereus.—Although this species belongs to the order Chitonida, and so is less primitive than Lepidopleurus asellus (order Lepidopleurida), the de- scription of conditions in both species will be easier if Lepi- dochitona cinereus is considered first. In the animal shown in Text-fig. 1 there are seventeen gills on each side extending along some four-fifths of the pallial grooves. They are attached to the roof of the grooves. All but the last and largest of these, the post-renal gill (GP.), bend inwards towards the sides of the foot, but the post-renal gill extends backwards with its (morphologically) posterior surface applied to the side of the girdle fold (GF.) and its anterior surface to the side of the foot. It thus blocks the pallial groove in this region. The five gills immediately anterior to it also bend backwards to a greater or less extent, but they lie against the sides of the foot only. Throughout the whole series the sides of adjacent gills are alwayi closely applied. Lateral cilia on the gills (described later) create a current of water which passes from their outer to their inner surfaces. IC o GF TBXT-HG. 1. Lepidochitona cinereus, ventral aspect, drawn from life: gills and boundaries of shell plates shown in left pallial groove, osphradium and division between inhalant and exhalant chambers (denoted by broken line) shown in right pallial groove, x 10. A., anus; E., exhalant current; EC, exhalant chamber; p., foot; o., girdle; GI., most anterior gill; GF., girdle fold; GP., post-renal gill; I., inhalant currents; M., mouth; o., osphradium. Arrows indicate direction of currents, broken arrows those in exhalant ohamber. 372 C. M. YONGB Owing to the arrangement of the post-renal gill across the pallial groove there is a complete separation of the mantle cavity into inhalant and exhalant chambers (Text-fig. 1, ic, BO.). The former consists of the pallial groove anterior to the first gill and the region outside the gills behind this. The exhalant chamber consists anteriorly of the region between the gills and the sides of the foot (and so bounded ventrally by the lateral extensions of the sole) and posteriorly of the entire pallial groove behind the post-renal gills. The boundary between these chambers is indicated by the broken line in the right pallial groove in Text-fig. 1. A powerful backwardly directed current runs along the in- halant chamber, water being drawn through the gills laterally into the exhalant chamber and also posteriorly between the filaments of the post-renal gills. This inhalant current is so powerful that suspended particles do not tend to settle on to the roof of the pallial groove when the animal is inverted for inspection.
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