Page:EB1911 - Volume 11.djvu/542

The adaptation of the Pulmonata to terrestrial life has entailed little modification of the internal organization. In one genus (Planorbis) the plasma of the blood is coloured red by haemoglobin, this being the only instance of the presence of this body in the blood of Glossophorous Mollusca, though it occurs in corpuscles in the blood of the bivalves Arca and Solen (Lankester).

Fig. 58.—Hermaphrodite Reproductive Apparatus of the Garden Snail (Helix hortensis).
z,	Ovo-testis.
ve,	Hermaphrodite duct.
Ed,	Albuminiparous gland.
u,	Uterine dilatation of the hermaphrodite duct.
d,	Digitate accessory glands on the female duct.
ps,	Calciferous gland or dart-sac on the female duct.
Rf,	Spermatheca or receptacle of the sperm in copulation, opening into the female duct.
vd,	Male duct (vas deferens).
p,	Penis.
fl,	Flagellum.

The generative apparatus of the snail (Helix) may serve as an example of the hermaphrodite apparatus common to the Pulmonata and Opisthobranchia (fig. 58). From the ovo-testis, which lies near the apex of the visceral coil, a common hermaphrodite duct ve proceeds, which receives the duct of the compact white albuminiparous gland, Ed, and then becomes much enlarged, the additional width being due to the development of glandular folds, which are regarded as forming a uterus u. Where these folds cease the common duct splits into two portions, a male and a female. The male duct vd becomes fleshy and muscular near its termination at the genital pore, forming the penis p. Attached to it is a diverticulum fl, in which the spermatozoa which have descended from the ovo-testis are stored and modelled into sperm ropes or spermatophores. The female portion of the duct is more complex. Soon after quitting the uterus it is joined by a long duct leading from a glandular sac, the spermatheca (Rf). In this duct and sac the spermatophores received in copulation from another snail are lodged. In Helix hortensis the spermatheca is simple. In other species of Helix a second duct (as large in Helix aspersa as the chief one) is given off from the spermathecal duct, and in the natural state is closely adherent to the wall of the uterus. This second duct has normally no spermathecal gland at its termination, which is simple and blunt. But in rare cases in Helix aspersa a second spermatheca is found at the end of this second duct. Tracing the widening female duct onwards we now come to the openings of the digitate accessory glands d, d, which probably assist in the formation of the egg-capsule. Close to them is the remarkable dart-sac ps, a thick-walled sac, in the lumen of which a crystalline four-fluted rod or dart consisting of carbonate of lime is found. It is supposed to act in some way as a stimulant in copulation, but possibly has to do with the calcareous covering of the egg-capsule. Other Pulmonata exhibit variations of secondary importance in the details of this hermaphrodite apparatus.

The nervous system of Helix is not favourable as an example on account of the fusion of the ganglia to form an almost uniform ring of nervous matter around the oesophagus. The pond-snail (Limnaeus) furnishes, on the other hand, a very beautiful case of distinct ganglia and connecting cords (fig. 59). The demonstration which it affords of the extreme shortening of the Euthyneurous visceral nerve-loop is most instructive and valuable for comparison with and explanation of the condition of the nervous centres in Cephalopoda, as also of some Opisthobranchia. The figure (fig. 59) is sufficiently described in the letterpress attached to it; the pair of buccal ganglia joined by the connectives to the cerebrals are, as in most of our figures, omitted. Here we need only further draw attention to the osphradium, discovered by Lacaze-Duthiers, and shown by Spengel to agree in its innervation with that organ in all other Gastropoda. On account of the shortness of the visceral loop and the proximity of the right visceral ganglion to the oesophageal nerve-ring, the nerve to the osphradium and olfactory ganglion is very long. The position of the osphradium corresponds more or less closely with that of the vanished right ctenidium, with which it is normally associated. In Helix and Limax the osphradium has not been described, and possibly its discovery might clear up the doubts which have been raised as to the nature of the mantle-chamber of those genera. In Planorbis, which is sinistral (as are a few other genera or exceptional varieties of various Anisopleurous Gastropods), instead of being dextral, the osphradium is on the left side, and receives its nerve from the left visceral ganglion, the whole series of unilateral organs being reversed. This is, as might be expected, what is found to be the case in all “reversed” Gastropods.

The shell of the Pulmonata, though always light and delicate, is in many cases a well-developed spiral “house” into which the creature can withdraw itself; and, although the foot possesses no operculum, yet in Helix the aperture of the shell is closed in the winter by a complete lid, the “hybernaculum” more or less calcareous in nature, which is secreted by the foot. In Clausilia a peculiar modification of this lid exists permanently in the adult, attached by an elastic stalk to the mouth of the shell, and known as the “clausilium.” In Limnaeus the permanent shell is preceded in the embryo by a well-marked shell-gland or primitive shell-sac (fig. 60), at one time supposed to be the developing anus, but shown by Lankester to be identical with the “shell-gland” discovered by him in other Mollusca (Pisidium, Pleurobranchidium, Neritina, &c.). As in other Gastropoda Anisopleura, this shell-sac may abnormally develop a plug of chitinous matter, but normally it flattens out and disappears, whilst the cap-like rudiment of the permanent shell is shed out from the dome-like surface of the visceral hump, in the centre of which the shell-sac existed for a brief period.

Fig. 59.—Nervous System of the Pond-Snail, Limnaeus stagnalis, as a type of the short-looped euthyneurous condition. The short visceral “loop” with its three ganglia is lightly-shaded.
ce,	Cerebral ganglion.
pe,	Pedal ganglion.
pl,	Pleural ganglion.
ab,	Abdominal ganglion.
sp,	Visceral ganglion of the left side; opposite to it is the visceral ganglion of the right side, which gives off the long nerve to the olfactory ganglion and osphradium o.
In Planorbis and in Auricula (Pulmonata, allied to Limnaeus) the olfactory organ is on the left side and receives its nerve from the left visceral ganglion. (After Spengel.)

In Clausilia, according to the observations of C. Gegenbaur, the primitive shell-sac does not flatten out and disappear, but takes the form of a flattened closed sac. Within this closed sac a plate of calcareous matter is developed, and after a time the upper wall of the sac disappears, and the calcareous plate continues to grow as the nucleus of the permanent shell. In the slug Testacella (fig. 56, C) the shell-plate never attains a large size, though naked. In other slugs, namely, Limax and Arion, the shell-sac remains permanently closed over the shell-plate, which in the latter genus consists of a granular mass of carbonate of lime. The permanence of the primitive shell-sac in these slugs is a point of considerable interest. It is clear enough that the sac is of a different origin from that of Aplysia (described in the section treating of Opisthobranchia), being primitive instead of secondary. It seems probable that it is identical with one of the open sacs in which each shell-plate of a Chiton is formed, and the series of plate-like imbrications which are placed behind the single shell-sac on the dorsum of the curious slug, Plectrophorus, suggest the possibility of the formation of a series of shell-sacs on the back of that animal similar to those which we find in Chiton. Whether the closed primitive shell-sac of the slugs (and with it the transient embryonic shell-gland of all other Mollusca) is precisely the same thing as the closed sac in which the calcareous pen or shell of the Cephalopod Sepia and its allies is formed, is a further question which we shall consider when dealing with the Cephalopoda. It is important here to note that Clausilia furnishes us with an exceptional instance of the continuity of the shell or secreted product of the primitive shell-sac with the adult shell. In most other Mollusca (Anisopleurous Gastropods, Pteropods and Conchifera) there is a want of such continuity; the primitive shell-sac contributes no factor to the permanent shell, or only a very minute knob-like particle (Neritina and Paludina). It flattens out and disappears before the work of forming the permanent shell commences. And just as there is a break at this stage, so (as observed by A. Krohn in Marsenia = Echinospira) there may be a break at a later stage, the nautiloid shell formed on the larva being cast, and a new shell of a different form being formed afresh on the surface of the visceral hump. It is, then, in this sense that we may speak of primary, secondary and tertiary shells in Mollusca recognizing the fact that they may be merely phases fused by continuity of growth so as to form but one shell, or that in other cases they may be presented to us as separate individual things, in virtue of the non-development of the later phases, or in virtue of sudden changes in the activity of the mantle-surface causing the shedding