coiled. When the spermatophore is expelled into the water the connective is extended and evaginated, and the sac containing the sperms bursts. In Nautilus the spermatophore when uncoiled is a little over 30 mm. in length. These spermatophores are somewhat similar to those formed in certain pulmonate Gastropods.
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Fig. 13.—Nervous system of Nautilus pompilius (from Genebaur, after Owen). t,t, Ganglion-like enlargements on nerves passing from the pedal ganglion to the inner series of tentacles.
t ′, Nerves to the tentacles of the outer or annular lobe.
b, Pedal ganglion-pair
a, Cerebral ganglion-pair.
c, Pleuro-visceral ganglionic band (fused pleural and visceral ganglion-pairs).
d, Genital ganglion placed on the course of the large visceral nerve, just before it gives off its branchial and its osphradial branches.
m, Nerves from the pleural ganglion to the mantle-skirt. |
The eggs are laid shortly after copulation. In Nautilus they are laid separately, each being about 4 cm. long and contained in two thick shells, the outer of which is partly open.
Nervous System.—Nautilus, like the other Cephalopoda, exhibits a great concentration of the typical Molluscan ganglia, as shown in fig. 13. The ganglia take on a band-like form, and are but little differentiated from their commissures and connectives—an archaic condition reminding us of Chiton. The special optic outgrowth of the cerebral ganglion, the optical ganglion (fig. 13, o), is characteristic. The cerebral ganglion-pair (a) lying above the oesophagus is connected with two suboesophageal ganglion-pairs, of band-like form. The anterior of these is the pedal b, b, and supplies the circumoral lobes and tentacles, and the funnel, a fact which proves the pedal origin of these organs. The hinder band is the visceral and pleural pair fused; from its pleural portion nerves pass to the mantle, from its visceral portion nerves to the branchiae and genital ganglion (fig. 13, d), and in immediate connexion with the latter is a nerve to the osphradium or olfactory papilla. A labial commissure arises by a double root from the cerebral ganglia and gives off a stomatogastric commissure, which passes under the pharynx immediately behind the radula and bears a buccal ganglion on either side.
Special Sense-Organs.—Nautilus possesses a pair of osphradial papillae (fig. 4, olf) corresponding in position and innervation to Spengel’s organ placed at the base of the ctenidia (branchiae) in all classes of Mollusca. This organ has not been detected in other Cephalopoda. Nautilus possesses other olfactory organs in the region of the head. Just below the eye is a small triangular process (not seen in our figures), having the structure of a shortened and highly-modified tentacle and sheath. By A. Valenciennes, who is followed by W. Keferstein, this is regarded as an olfactory organ. The large nerve which runs to this organ originates from the point of juncture of the pedal with the optic ganglion. The lamelliform organ upon the inner inferior tentacular lobe of Nautilus is possibly also olfactory in function. In Dibranchs behind the eye is a pit or open canal supplied by a nerve corresponding in origin to the olfactory nerve of Nautilus above mentioned. Possibly the sense of taste resides in certain processes within the mouth of Nautilus and other Cephalopoda.
The otocysts of Nautilus were discovered by J. D. Macdonald. Each lies at the side of the head, ventral to the eye, resting on the capito-pedal cartilage, and supported by the large auditory nerve which apparently arises from the pedal ganglion but originates in the cerebral. It has the form of a small sac, 1 to 2 mm. in diameter, and contains whetstone-shaped crystals, such as are known to form the otoliths of other Mollusca.
The eye of Nautilus is among the most interesting structures of that remarkable animal. No other animal which has the same bulk and general elaboration of organization has so simple an eye as that of Nautilus. When looked at from the surface no metallic lustre, no transparent coverings, are presented by it. It is simply a slightly projecting hemispherical box like a kettle-drum, half an inch in diameter, its surface looking like that of the surrounding integument, whilst in the middle of the drum-membrane is a minute hole (fig. 3, u). Sir R. Owen very naturally thought that some membrane had covered this hole in life, and had been ruptured in the specimen studied by him. It, however, appears from the researches of V. Hensen that the hole is a normal aperture leading into the globe of the eye, which is accordingly filled by sea-water during life. There is no dioptric apparatus in Nautilus, and in place of refracting lens and cornea we have actually here an arrangement for forming an image on the principle of “the pin-hole camera.” There is no other eye known in the whole animal kingdom which is so constructed. The wall of the eye-globe is tough, and the cavity is lined solely by the naked retina, which is bathed by sea-water on one surface and receives the fibres of the optic nerve on the other (see fig. 14, A). As in other Cephalopods (e.g. fig. 33, Ri, Re, p), the retina consists of two layers of cells, separated by a layer of dark pigment. The most interesting consideration connected with this eye of Nautilus is found when the further facts are noted—(1) that the elaborate lens-bearing eyes of Dibranchiata pass through a stage of development in which they have the same structure as the eye of Nautilus—namely, are open sacs (fig. 34); and (2) that amongst other Mollusca examples of cephalic eyes can be found which in the adult condition are, like the eye of Nautilus and the developing eye of Dibranchs, simple pits of the integument, the cells of which are surrounded by pigment and connected with the filaments of an optic nerve. Such is the structure of the eye of the limpet (Patella), and in such a simple eye we obtain the clearest demonstration of the fact that the retina of the Molluscan cephalic eye, like that of the Arthropod cephalic eye and unlike that of the vertebrate myelonic eye, is essentially a modified area of the general epiderm, and that the sensitiveness of its cells to the action of light and their relation to nerve-filaments is only a specialization and intensifying of a property common to the whole epiderm of the surface of the body. What, however, strikes us as especially remarkable is that the simple form of a pit, which in Patella serves to accumulate a secretion which acts as a refractive body, should in Nautilus be glorified and raised to the dignity of an efficient optical apparatus. In all other Mollusca, starting as we may suppose from the follicular or pit-like condition, the eye has proceeded to acquire the form of a closed sac, the cavity of the closed vesicle being then filled partially or completely by a refractive body (lens) secreted by its walls (fig. 14, B). This is the condition attained in most Gastropoda. It presents a striking contrast to the simple Arthropod eye, where, in consequence of the existence of a dense exterior cuticle, the eye does not form a vesicle, and the lens is always part of that cuticle.
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Fig. 14.—Diagrams of Sections of the Eyes of Mollusca. | |
A, Nautilus (and Patella).
B, Gastropod (Limax or Helix).
C, Dibranchiate Cephalopod (Oigopsid).
Pal, Eyelid (outermost fold).
Co, Cornea (second fold).
Ir, Iris (third fold).
Int 1,2,3,4, Different parts of
the integument. l, Deep portion of the lens. |
l1, Outer portion of the lens
Co.ep, Ciliary body.
R, Retina.
N.op, Optic nerve.
G.op, Optic ganglion.
x, Inner layer of the retina.
N.S., Nervous stratum of the retina. (From Balfour, after Grenacher.) |
The development of Nautilus is still entirely unknown. Dr Arthur Willey, during his sojourn in the East Indies, made special efforts to obtain fertilized eggs, both by offering rewards to the native fishermen and collectors and by keeping the living adults in captivity, but without success.
Phylogeny and Classification.—As Nautilus is the only living genus of the Tetrabranchiata, our knowledge of all the rest is based upon the study of their fossil shells. A vast number of species of shell similar in structure to that of Nautilus are known, chiefly from Primary and Secondary formations. These are divided into two sub-orders by differences in the form and structure of the initial chamber. In the Nautiloidea this chamber has the form of an obtuse cone, on the apex of which is a slit-like mark or cicatrix, elongated dorso-ventrally and placed opposite to the blind end of the siphuncle, which indents the front wall of the initial chamber but does not enter its cavity. In the Ammonoidea, on the other hand, the initial chamber is inflated, and is spheroidal, oval or pyriform in shape, with no cicatrix, and separated from the first air-chamber by a constriction. The siphuncle also commences with a dilatation which deeply indents the front wall of the initial chamber, called the protoconch, but does not penetrate into its cavity. Munier-Chalmas has shown that the cavity of the protoconch is traversed by a tubular organ, the “prosiphon,” which does not communicate with the true siphuncle, the place of which it is supposed to take in the early life of the animal. It is generally held, as suggested by