Page:EB1911 - Volume 11.djvu/529

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represented by the genital duct. There is usually a penis in the male. The ctenidium is monopectinate and attached to the mantle along its whole length, except in Adeorbis and Valvata; in the latter alone it is bipectinate. There is a single well-developed, often pectinated osphradium. The eye is always a closed vesicle, and the internal cornea is extensive. In the radula there is a single central tooth or none.

EB1911 Gastropoda - Pyrula laevigata.jpg

Fig. 18.—Animal and shell of Pyrula laevigata. (From Owen.)

a, Siphon.

b, Head-tentacles.

C, Head, the letter placed near the right eye.

d, The foot, expanded as in crawling.

h, The mantle-skirt reflected over the sides of the shell.

The former classification into Holochlamyda, Pneumochlamyda and Siphonochlamyda has been abandoned, as it was founded on adaptive characters not always indicative of true affinities. The order is now divided into two sub-orders: the Taenioglossa, in which there are three teeth on each side of the median tooth of the radula, and the Stenoglossa, in which there is only one tooth on each side of the median tooth. In the latter a pallial siphon, a well-developed proboscis and an unpaired oesophageal gland are always present, in the former they are usually absent. The siphon is an incompletely tubular outgrowth of the mantle margin on the left side, contained in a corresponding outgrowth of the edge of the shell-mouth, and serving to conduct water to the respiratory cavity.

EB1911 Gastropoda - nature of so-called proboscides.jpg

Fig. 19.—Diagrams explanatory of the nature of so-called proboscides or “introverts.” (Lankester.)

A, Simple introvert completely introverted.

B, The same, partially everted by eversion of the sides, as in the Nemertine proboscis and Gastropod eye-tentacle = pleurecbolic.

C, The same, fully everted.

D, E, A similar simple introvert in course of eversion by the forward movement, not of its sides, but of its apex, as in the proboscidean Rhabdocoels = acrecbolic.

F, Acrecbolic ( = pleurembolic) introvert, formed by the snout of the proboscidiferous Gastropod. al, alimentary canal; d, the true mouth. The introvert is not a simple one with complete range both in eversion and introversion, but is arrested in introversion by the fibrous bands at c, and similarly in eversion by the fibrous bands at b.

G, The acrecbolic snout of a proboscidiferous Gastropod, arrested short of complete eversion by the fibrous band b.

H, The acrembolic ( = pleurecbolic) pharynx of a Chaetopod fully introverted. al, alimentary canal; at d, the jaws; at a, the mouth; therefore a to d is stomodaeum, whereas in the Gastropod (F) a to d is inverted body-surface.

I, Partial eversion of H.

K, Complete eversion of H.

The condition usually spoken of as a “proboscis” appears to be derived from the condition of a simple rostrum (having the mouth at its extremity) by the process of incomplete introversion of that simple rostrum. There is no reason in the actual significance of the word why the term “proboscis” should be applied to an alternately introversible and eversible tube connected with an animal’s body, and yet such is a very customary use of the term. The introversible tube may be completely closed, as in the “proboscis” of Nemertine worms, or it may have a passage in it leading into a non-eversible oesophagus, as in the present case, and in the case of the eversible pharynx of the predatory Chaetopod worms. The diagrams here introduced (fig. 19) are intended to show certain important distinctions which obtain amongst the various “introverts,” or intro- and e-versible tubes so frequently met with in animal bodies. Supposing the tube to be completely introverted and to commence its eversion, we then find that eversion may take place, either by a forward movement of the side of the tube near its attached base, as in the proboscis of the Nemertine worms, the pharynx of Chaetopods and the eye-tentacle of Gastropods, or by a forward movement of the inverted apex of the tube, as in the proboscis of the Rhabdocoel Planarians, and in that of Gastropods here under consideration. The former case we call “pleurecbolic” (fig. 19, A, B, C, H, I, K), the latter “acrecbolic” tubes or introverts (fig. 19, D, E, F, G). It is clear that, if we start from the condition of full eversion of the tube and watch the process of introversion, we shall find that the pleurecbolic variety is introverted by the apex of the tube sinking inwards; it may be called acrembolic, whilst conversely the acrecbolic tubes are pleurembolic. Further, it is obvious enough that the process either of introversion or of eversion of the tube may be arrested at any point, by the development of fibres connecting the wall of the introverted tube with the wall of the body, or with an axial structure such as the oesophagus; on the other hand, the range of movement of the tubular introvert may be unlimited or complete. The acrembolic proboscis or frontal introvert of the Nemertine worms has a complete range. So has the acrembolic pharynx of Chaetopods, if we consider the organ as terminating at that point where the jaws are placed and the oesophagus commences. So too the acrembolic eye-tentacle of the snail has a complete range of movement, and also the pleurembolic proboscis of the Rhabdocoel prostoma. The introverted rostrum of the Pectinibranch Gastropods presents in contrast to these a limited range of movement. The “introvert” in these Gastropods is not the pharynx as in the Chaetopod worms, but a prae-oral structure, its apical limit being formed by the true lips and jaws, whilst the apical limit of the Chaetopod’s introvert is formed by the jaws placed at the junction of pharynx and oesophagus, so that the Chaetopod’s introvert is part of the stomodaeum or fore-gut, whilst that of the Gastropod is external to the alimentary canal altogether, being in front of the mouth, not behind it, as is the Chaetopod’s. Further, the Gastropod’s introvert is pleurembolic (and therefore acrecbolic), and is limited both in eversion and in introversion; it cannot be completely everted owing to the muscular bands (fig. 19, G), nor can it be fully introverted owing to the bands (fig. 19, F) which tie the axial pharynx to the adjacent wall of the apical part of the introvert. As in all such intro- and e-versible organs, eversion of the Gastropod proboscis is effected by pressure communicated by the muscular body-wall to the liquid contents (blood) of the body-space, accompanied by the relaxation of the muscles which directly pull upon either the sides or the apex of the tubular organ. The inversion of the proboscis is effected directly by the contraction of these muscles. In various members of the Pectinibranchia the mouth-bearing cylinder is introversible (i.e. is a proboscis)—with rare exceptions these forms have a siphonate mantle-skirt. On the other hand, many which have a siphonate mantle-skirt are not provided with an introversible mouth-bearing