Let us now examine the organs which lie beneath the mantle-skirt
of Anodonta, and are bathed by the current of water which circulates
through it. This can be done by lifting up and throwing back the left
half of the mantle-skirt as is represented in fig. 1 (3). We thus expose
the plough-like foot (f), the two left labial tentacles, and the two left
gill-plates or left ctenidium. In fig. 1 (5), one of the labial tentacles n
is also thrown back to show the mouth w, and the two left gill-plates
are reflected to show the gill-plates of the right side (rr, rq) projecting
behind the foot, the inner or median plate of each side being
united by concrescence to its fellow of the opposite side along a
continuous line (aa). The left inner gill-plate is also snipped to show
the subjacent orifices of the left renal organ
x, and of the genital gland (testis or ovary) y.
The foot thus exposed in Anodonta is a simple
muscular tongue-like organ. It can be protruded
between the flaps of the mantle (fig. 1
[1] [2]) so as to issue from the shell, and by
its action the Anodonta can slowly crawl or
burrow in soft mud or sand. Other Lamellibranchs
may have a larger foot relatively than
has Anodonta. In Arca it has a sole-like
surface. In Arca too and many others it
carries a byssus-forming gland and a byssus-cementing
gland. In the cockles, in Cardium
and in Trigonia, it is capable of a sudden
stroke, which causes the animal to jump when
out of the water, in the latter genus to a
height of four feet. In Mytilus the foot is reduced to little more
than a tubercle carrying the apertures of these glands. In the
oyster it is absent altogether.
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The labial tentacles or palps of Anodonta (n, o in fig. 1 [3], [5]) are highly vascular flat processes richly supplied with nerves. The left anterior tentacle (seen in the figure) is joined at its base in front of the mouth (w) to the right anterior tentacle, and similarly the left (o) and right posterior tentacles are joined behind the mouth. Those of Arca (i, k in fig. 9) show this relation to the mouth (a). These organs are characteristic of all Lamellibranchs; they do not vary except in size, being sometimes drawn out to streamer-like dimensions. Their appearance and position suggest that they are in some way related morphologically to the gill-plates, the anterior labial tentacle being a continuation of the outer gill-plate, and the posterior a continuation of the inner gill-plate. There is no embryological evidence to support this suggested connexion, and, as will appear immediately, the history of the gill-plates in various forms of Lamellibranchs does not directly favour it. The palps are really derived from part of the velar area of the larva.
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Fig. 8.—Psammobia florida, right side, showing expanded foot e, and g incurrent and g′ excurrent siphons. (From Owen.) |
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Fig. 9.—View from the ventral (pedal) aspect of the animal of Arca noae, the mantle-flap and gill-filaments having been cut away. (Lankester.) | |
| a, | Mouth. |
| b, | Anus. |
| c, | Free spirally turned extremity of the gill-axis or ctenidial axis of the right side. |
| d, | Do. of the left side. |
| e, f, | Anterior portions of these axes fused by concrescence to the wall of the body. |
| g, | Anterior adductor muscle. |
| h, | Posterior adductor. |
| i, | Anterior labial tentacle. |
| k, | Posterior labial tentacle. |
| l, | Base line of the foot. |
| m, | Sole of the foot. |
| n, | Callosity. |
The gill-plates have a structure very different from that of the labial tentacles, and one which in Anodonta is singularly complicated as compared with the condition presented by these organs in some other Lamellibranchs, and with what must have been their original condition in the ancestors of the whole series of living Lamellibranchia. The phenomenon of “concrescence” which we have already had to note as showing itself so importantly in regard to the free edges of the mantle-skirt and the formation of the siphons, is what, above all things, has complicated the structure of the Lamellibranch ctenidium. Our present knowledge of the interesting series of modifications through which the Lamellibranch gill-plates have developed to their most complicated form is due to R. H. Peck, K. Mitsukuri and W. G. Ridewood. The Molluscan ctenidium is typically a plume-like structure, consisting of a vascular axis, on each side of which is set a row of numerous lamelliform or filamentous processes. These processes are hollow, and receive the venous blood from, and return it again aerated into, the hollow axis, in which an afferent and an efferent blood-vessel may be differentiated. In the genus Nucula (fig. 10) we have an example of a Lamellibranch retaining this plume-like form of gill. In the Arcacea (e.g. Arca and Pectunculus) the lateral processes which are set on the axis of the ctenidium are not lamellae, but are slightly flattened, very long tubes or hollow filaments. These filaments are so fine and are set so closely together that they appear to form a continuous membrane until examined with a lens. The microscope shows that the neighbouring filaments are held together by patches of cilia, called “ciliated junctions,” which interlock with one another just as two brushes may be made to do. In fig. 11, A a portion of four filaments of a ctenidium of the sea-mussel (Mytilus) is represented, having precisely the same structure as those of Arca. The filaments of the gill (ctenidium) of Mytilus and Arca thus form two closely set rows which depend from the axis of the gill like two parallel plates. Further, their structure is profoundly modified by the curious condition of the free ends of the depending filaments. These are actually reflected at a sharp angle—doubled on themselves in fact—and thus form an additional row of filaments (see fig. 11 B). Consequently, each primitive filament has a descending and an ascending ramus, and instead of each row forming a simple plate, the plate is double, consisting of a descending and an ascending lamella. As the axis of the ctenidium lies by the side of the body, and is very frequently connate with the body, as so often happens in Gastropods also, we find it convenient to speak of the two plate-like structures formed on each ctenidial axis as the outer and the inner gill-plate; each of these is composed of two lamellae, an outer (the reflected) and an adaxial in the case of the outer gill-plate, and an adaxial and an inner (the reflected) in the case of the inner gill-plate. This is the condition seen in Arca and Mytilus, the so-called plates dividing upon the slightest touch into their constituent filaments, which are but loosely conjoined by their “ciliated junctions.” Complications follow upon this in other forms. Even in Mytilus and Arca a connexion is here and there formed between the ascending and descending rami of a filament by hollow extensible outgrowths called “interlamellar junctions” (il. j in B, fig. 11). Nevertheless the filament is a complete tube formed of chitinous substance and clothed externally by ciliated epithelium, internally by endothelium and lacunar tissue—a form of connective tissue—as shown in fig. 11, C. Now let us suppose as happens in the genus Dreissensia—a genus not far removed from Mytilus—that the ciliated inter-filamentar junctions (fig. 12) give place to solid permanent inter-filamentar junctions, so that the filaments are converted, as it were, into a trellis-work. Then let us suppose that the interlamellar junctions already noted in Mytilus become very numerous, large and irregular; by them the two trellis-works of filaments would be united so as to leave only a sponge-like set of spaces between them. Within the trabeculae of the sponge-work blood circulates, and between the trabeculae the water passes, having entered by the apertures left in the trellis-work formed by the united gill-filaments (fig. 14). The larger the
