| Fig. 6.—Shell of Aspergillum vaginiferum. (From Owen.) | Fig. 7.—Shell of Aspergillum vaginiferum to show the original valves a, now embedded in a continuous calcification of tubular form. (From Owen.) |
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.
| Fig. 8.—Psammobia florida, right side, showing expanded foot e, and g incurrent and g′ excurrent siphons. (From Owen.) |
| 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 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.
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 intralamellar spongy growth becomes, the more do the original gill-filaments lose the character of blood-holding tubes, and tend to become dense elastic rods for the simple purpose of supporting the spongy growth. This is seen both in the section of Dreissensia gill (fig. 12) and in those of Anodonta (fig. 13, A, B, C). In the drawing of Dreissensia the individual filaments f, f, f are cut across in one lamella at the horizon of an inter-filamentar junction, in the other (lower in the figure) at a point where they are free. The chitinous substance ch is observed to be greatly thickened as compared with what it is in fig. 11, C, tending in fact to obliterate altogether the lumen of the filament. And in Anodonta (fig. 13, C) this obliteration is effected. In Anodonta, besides being thickened, the skeletal substance of the filament develops a specially dense, rod-like body on each side of each filament. Although the structure of the ctenidium is thus highly complicated in Anodonta, it is yet more so in some of the siphonate genera of Lamellibranchs. The filaments take on a secondary grouping, the surface of the lamella being thrown into a series of half-cylindrical ridges, each consisting of ten or twenty filaments; a filament of much greater strength and thickness than the others may be placed between each pair of groups. In Anodonta, as in many other Lamellibranchs, the ova and hatched embryos are carried for a time in the ctenidia or gill apparatus, and in this particular case the space between the two lamellae of the outer gill-plate is that which serves to receive the ova (fig. 13, A). The young are nourished by a substance formed by the cells which cover the spongy interlamellar outgrowths.
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| Fig. 10.—Structure of the Ctenidia of Nucula. (After Mitsukuri.)See also fig. 2. | |
A. Section across the axis of actenidium with a pair ofplates—flattened andshortened filaments—attached. i, j, k, g, Are placed on or near themembrane which attachesthe axis of the ctenidium tothe side of the body. a, b, Free extremities of the plates(filaments). d, Mid-line of the inferiorborder. e, Surface of the plate. t, Its upper border. h, Chitinous lining of the plate. r, Dilated blood-space. u, Fibrous tract. o, Upper blood-vessel of theaxis. n, Lower blood-vessel of theaxis. s, Chitinous framework of theaxis. cp, Canal in the same. A, B, Line along which the cross-sectionC of the plate istaken. B. Animal of a male Nuculaproxima, Say, as seen whenthe left valve of the shelland the left half of themantle-skirt are removed. | a, a, Anterior adductor muscle. p.a, Posterior adductor muscle. v.m, Visceral mass. f, Foot. g, Gill. l, Labial Tentacle. l.a, Filamentous appendage ofthe labial tentacle. lb, Hood-like appendage of thelabial tentacle. m, Membrane suspending thegill and attached to thebody along the line x, y, z,w. p, Posterior end of the gill(ctenidium). C. Section across one of the gill-plates(A, B, in A) comparablewith fig. 11 C. i.a, Outer border. d.a, Axial border. l.f, Latero-frontal epithelium. e, Epithelium of general surface. r, Dilated blood-space. h, Chitinous lining (compare A). |
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| Fig. 11.—Filaments of the Ctenidium of Mytilus edulis.(After R. H. Peck.) | |
A, Part of four filaments seenfrom the outer face in order toshow the ciliated junctions c.j. B, Diagram of the posterior faceof a single complete filament withdescending ramus and ascendingramus ending in a hook-like process;ep., ep., the ciliated junctions;il, j., interlamellar junction. | C, Transverse section of a filamenttaken so as to cut neithera ciliated junction nor an interlamellarjunction. f.e., Frontalepithelium; l.f.e′., l.f.e″., the tworows of latero-frontal epithelialcells with long cilia; ch, chitinoustubular lining of the filament;lac., blood lacuna traversed by afew processes of connective tissuecells; b.c., blood-corpuscle. |
Other points in the modification of the typical ctenidium must be noted in order to understand the ctenidium of Anodonta. The axis of each ctenidium, right and left, starts from a point well forward near the labial tentacles, but it is at first only a ridge, and does not project as a free cylindrical axis until the back part of the foot is reached. This is difficult to see in Anodonta, but if the mantle-skirt be entirely cleared away, and if the dependent lamellae which spring from the ctenidial axis be carefully cropped so as to leave the axis itself intact, we obtain the form shown in fig. 15, where g and h are respectively the left and the right ctenidial axes projecting freely beyond the body. In Arca this can be seen with far less trouble, for the filaments are more easily removed than are the consolidated lamellae formed by the filaments of Anodonta, and in Arca the free axes of the ctenidia are large and firm in texture (fig. 9, c, d).
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| Fig. 12.—Transverse Section of the Outer Gill-plate ofDreissensia polymorpha. (After R. H. Peck.) | |
f, Constituent gill-filaments. ff, Fibrous sub-epidermic tissue. ch, Chitonous substance of thefilaments. nch, Cells related to the chitonoussubstance. lac, Lacunar tissue. pig, Pigment-cells. | bc, Blood-corpuscles. fe, Frontal epithelium. lfe′, lfe″, Two rows of latero-frontalepithelial cells with longcilia. lrf, Fibrous, possibly muscular,substance of the inter-filamentarjunctions. |