of some days—possibly weeks—the ova of the second Aplysia
commence to descend the hermaphrodite duct; they become enclosed
in a viscid secretion at the point where the albuminiparous
gland opens into the duct intertwined with it; and on reaching the
point where the spermathecal duct debouches they are impregnated
by the spermatozoa which escape now
from the spermatheca and meet the ova.
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Fig. 42.—Follicles of the hermaphrodite gonads of Euthyneurous Gastropods. A, of Helix; B, of Eolis; a, ova; b, developing spermatozoa; c, common efferent duct. |
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Fig. 43.—Nervous system of Aplysia, as a type of the long-looped Euthyneurous condition. The untwisted visceral loop is lightly shaded. (After Spengel.) | |
| ce, | Cerebral ganglion. |
| pl, | Pleural ganglion. |
| pe, | Pedal ganglion. |
| ab. sp, | |
| Abdominal ganglion which represents also the supra-intestinal ganglion of Streptoneura and gives off the nerve to the osphradium (olfactory organ) o, and another to an unlettered so-called “genital” ganglion. The buccal nerves and ganglia are omitted. | |
The development of Aplysia from the egg presents many points of interest from the point of view of comparative embryology, but in relation to the morphology of the Opisthobranchia it is sufficient to point to the occurrence of a trochosphere and a veliger stage (fig. 36), and of a shell-gland or primitive shell-sac (fig. 36, shgr), which is succeeded by a nautiloid shell.
In the nervous system of Aplysia the great ganglion-pairs are well developed and distinct. The euthyneurous visceral loop is long, and presents only one ganglion (in Aplysia camelus, but two distinct ganglia joined to one another in Aplysia hybrida of the English coast), placed at its extreme limit, representing both the right and left visceral ganglia and the third or abdominal ganglion, which are so often separately present. The diagram (fig. 43) shows the nerve connecting this abdomino-visceral ganglion with the olfactory ganglion of Spengel. It is also seen to be connected with a more remote ganglion—the genital. Such special irregularities in the development of ganglia upon the visceral loop, and on one or more of the main nerves connected with it, are very frequent. Our figure of the nervous system of Aplysia does not give the small pair of buccal ganglia which are, as in all glossophorous Molluscs, present upon the nerves passing from the cerebral region to the odontophore.
For a comparison of various Opisthobranchs, Aplysia will be found to present a convenient starting-point. It is one of the more typical Opisthobranchs, that is to say, it belongs to the section Tectibranchia, but other members of the suborder, namely, Bulla and Actaeon (figs. 44 and 45), are less abnormal than Aplysia in regard to their shells and the form of the visceral hump. They have naked spirally twisted shells which may be concealed from view in the living animal by the expansion and reflection of the parapodia, but are not enclosed by the mantle, whilst Actaeon is remarkable for possessing an operculum like that of so many Streptoneura.
The great development of the parapodia seen in Aplysia is usual in Tectibranchiate Opisthobranchs. The whole surface of the body becomes greatly modified in those Nudibranchiate forms which have lost, not only the shell, but also the ctenidium. Many of these have peculiar processes developed on the dorsal surface (fig. 46, A, B), or retain purely negative characters (fig. 46, D). The chief modification of internal organization presented by these forms, as compared with Aplysia, is found in the condition of the alimentary canal. The liver is no longer a compact organ opening by a pair of ducts into the median digestive tract, but we find very numerous hepatic diverticula on a shortened axial tract (fig. 47). These diverticula extend usually one into each of the dorsal papillae or “cerata” when these are present. They are not merely digestive glands, but are sufficiently wide to act as receptacles of food, and in them the digestion of food proceeds just as in the axial portion of the canal. A precisely similar modification of the liver or great digestive gland is found in the scorpions, where the axial portion of the digestive canal is short and straight, and the lateral ducts sufficiently wide to admit food into the ramifications of the gland there to be digested; whilst in the spiders the gland is reduced to a series of simple caeca.
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Fig. 44.—Bulla vexillum (Chemnitz), as seen crawling. á, oral hood (compare with Tethys, fig. 46, B), possibly a continuation of the epipodia; b, b′, cephalic tentacles. (From Owen.) |
The typical character is retained by the heart, pericardium, and the communicating nephridium or renal organ in all Opisthobranchs. An interesting example of this is furnished by the fish-like transparent Phyllirhoë (fig. 37), in which it is possible most satisfactorily to study in the living animal, by means of the microscope, the course of the blood-stream, and also the reno-pericardial communication. In many of the Nudibranchiate Opisthobranchs the nervous system presents a concentration of the ganglia (fig. 48), contrasting greatly with what we have seen in Aplysia. Not only are the pleural ganglia fused to the cerebral, but also the visceral to these (see in further illustration the condition attained by the Pulmonate Limnaeus, fig. 59), and the visceral loop is astonishingly short and insignificant (fig. 48, e′). That the parts are rightly thus identified is probable from J. W. Spengel’s observation of the osphradium and its nerve-supply in these forms; the nerve to that organ, which is placed somewhat anteriorly—on the dorsal surface—being given off from the hinder part (visceral) of the right compound ganglion—the fellow to that marked A in fig. 48. The Eolid-like Nudibranchs, amongst other specialities of structure, possess (in some cases at any rate) apertures at the apices of the “cerata” or dorsal papillae, which lead from the exterior into the hepatic caeca. Some amongst them (Tergipes, Eolis) are also remarkable for possessing peculiarly modified cells placed in sacs (cnidosacs) at the apices of these same papillae, which resemble the “thread-cells” of the Coelentera. According to T. S. Wright and J. H. Grosvenor these nematocysts are derived from the hydroids on which the animals feed.
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Fig. 45.—Actaeon. h, shell; b, oral hood; d, foot; f, operculum. |
The development of many Opisthobranchia has been examined—e.g. Aplysia, Pleurobranchidium, Elysia, Polycera, Doris, Tergipes. All pass through trochosphere and veliger stages, and in all a nautiloid or boat-like shell is developed, preceded by a well-marked “shell-gland” (see fig. 36). The transition from the free-swimming veliger larva with its nautiloid shell (fig. 36) to the adult form has not been properly observed, and many interesting points as to the true nature of folds (whether parapodia or mantle or velum) have yet to be cleared up by a knowledge of such development in forms like Tethys, Doris, Phyllidia, &c. As in other Molluscan groups, we find even in closely-allied genera (for instance, in Aplysia and Pleurobranchidium, and other genera), the greatest differences as to the amount of food-material by which the egg-shell is encumbered. Some form their diblastula by emboly, others by epiboly; and in the later history of the further development of the enclosed cells (arch-enteron) very marked variations occur in closely-allied forms, due to the influence of a greater or less abundance of food-material mixed with the protoplasm of the egg.
Sub-order 1.—Tectibranchia. Opisthobranchs provided in the adult state with a shell and a mantle, except Runcina, Pleurobranchaea, Cymbuliidae, and some Aplysiomorpha. There is a ctenidium, except in some Thecosomata and Gymnosomata, and an osphradium.
Tribe 1.—Bullomorpha. The shell is usually well developed, except in Runcina and Cymbuliidae, and may be external or internal. No operculum, except in Actaeonidae and Limacinidae. The pallial cavity is always well developed, and contains the ctenidium, at least in part; ctenidium, except in Lophocercidae, of folded type. With
