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CEPHALOPODA

of these parts in the Dibranchiata. The ganglia are more distinctly swollen than in Nautilus. In Octopus an infra-buccal ganglion-pair are present, corresponding to the buccal ganglion-pair of Gastropoda. In Decapoda a supra-buccal ganglion-pair connected with these are also developed. Instead of the numerous radiating pallial nerves of Nautilus, we have in the Dibranchiata on each side (right and left) a large pleural nerve passing from the pleural portion of the pleuro-visceral ganglion to the mantle, where it enlarges to form the stellate ganglion. From each stellate ganglion nerves radiate to supply the powerful muscles of the mantle-skirt. The two stellate ganglia are connected, except in Sepiola, by a transverse supra-oesophageal commissure, which represents the pallial cords united by a commissure above the intestine in Amphineura. The nerves from the visceral portion of the pleuro-visceral ganglion have the same course as in Nautilus, but no osphradial papilla is present. An enteric nervous system is richly developed in the Dibranchiata, connected with the somatic nervous centres through the buccal ganglia, as in the Arthropoda through the stomato-gastric ganglia, and anastomozing with deep branches of the visceral nerves of the viscero-pleural ganglion-pair. It has been especially described by A. Hancock in Ommatostrephes. Upon the stomach it forms a single large and readily detected gastric ganglion.

EB1911 Cephalopoda Fig. 33.—Horizontal section of the eye of Sepia (Myopsid).jpg
Fig. 33.—Horizontal section of the eye of Sepia (Myopsid). (From Gegenbaur, after Hensen.)

KK, Cephalic cartilages (see fig. 8).

C, Cornea (closed).

L, Lens.

ci, Ciliary body.

Ri, Internal layer of the retina.

Re, External layer of the retina.

p, Pigment between these.

o, Optic nerve.

go, Optic ganglion.

k and k′, Capsular cartilage.

ik, Cartilage of the iris.

w, White body.

ae, Argentine integument.

EB1911 Cephalopoda Fig. 34.—Diagrams of sections showing the early stage of development of the eye of Loligo.jpg
Fig. 34.—Diagrams of sections showing the early stage of development of the eye of Loligo when it is, like the permanent eye of Nautilus and of Patella, an open sac. (From Lankester.)

A, First appearance of the eye as a ring-like upgrowth.

B, Ingrowth of the ring-like wall so as to form a sac, the primitive optic vesicle of Loligo.

In the Dibranchiate division of the Cephalopoda the greatest elaboration of the dioptric apparatus of the eye is attained, so that we have in this class the extremes of the two lines of development of the Molluscan eye, those two lines being the punctigerous and the lentigerous. The structure of the Dibranchiate’s eye is shown in section in fig. 14, C, and in fig. 33, and its development in figs. 34 and 37. The open sac which forms the retina of the young Dibranchiate closes up, and constitutes the posterior chamber of the eye, or primitive optic vesicle (fig. 37, A, poc). The lens forms as a structureless growth, secreted by both the internal and external surfaces of the front wall of the optic vesicle (fig. 37, B, l). The integument around the primitive optic vesicle which has sunk below the surface now rises up and forms firstly nearest the axis of the eye the iridian folds (if in B, fig. 37; ik in fig. 33; Ir in fig. 14), and then secondly an outer circular fold grows up like a wall and completely closes over the iridian folds and the axis of the primitive vesicle (fig. 33, C). This covering is transparent, and is the cornea. In the oceanic Decapoda the cornea does not completely close, but leaves a central aperture traversed by the optic axis. These forms are termed Oigopsidae by C. d’Orbigny, whilst the Decapoda with closed cornea are termed Myopsidae. In the Octopoda the cornea is closed, and there is yet another fold thrown over the eye. The skin surrounding the cornea presents a free circular margin, and can be drawn over the surface of the cornea by a sphincter muscle. It thus acts as an adjustable diaphragm, exactly similar in movement to the iris of Vertebrates. Sepia and allied Decapods have a horizontal lower eyelid, that is to say, only one-half of the sphincter-like fold of integument is movable. The statocysts are situated ventrally between the pedal and visceral ganglia, and are entirely enclosed in the cranial cartilage. The cavity of each is continued into a small blind process which is the remnant of the embryonic connexion of the vesicle with the external surface. The sensory epithelium is at the anterior end of the vesicle forming a macula acustica, and in the cavity is a single otolith, partly calcareous and partly organic except in Eledone, in which it is entirely organic. The nerve arises from the cerebral ganglion on each side and passes through the pedal ganglion.

There is no branchial osphradium in the Dibranchiata corresponding to that of Nautilus, but the olfactory organ or rhinophore near the eye is present. In Sepia and the majority of the Dibranchiata it is a simple pit, in some of the Oigopsida it is a projection which may be stalked.

Reproduction and Development.—The modification of one or a pair of the arms in the male for purposes of copulation has already been described. In many genera the sexes differ from one another in other characters also. As a rule the males are more slender or smaller than the females. The maximum degree of sexual dimorphism occurs in Argonauta among the Octopods; in this genus the female may be fifteen times as large as the male, and the peculiar modification of the dorsal arms for the secretion of the shell occurs in the female only, no shell being formed in the male. In most cases the females are much more numerous than the males, but the opposite relation appears to exist in those Octopoda in which the hectocotylus is autotomous, for as many as four hectocotyli have been found in the pallial cavity of a single female. When the hectocotylus is not detached it is usually inserted into the pallial cavity of the female so as to deposit the spermatophores in or near the aperture of the oviduct, but in Sepia and Loligo they are merely deposited on the ventral lobes of the buccal membrane.

The eggs are laid shortly after copulation. In the Octopoda and in Sepia, Sepiola and Rossia, each egg has a separate envelope continued into a long stalk by which it is attached with several others in a cluster. In Argonauta the eggs are carried by the female in the cavity of the shell. In Loligo the eggs are very numerous, and are enclosed in cylindrical transparent gelatinous strings united at one end into a cluster.

The Cephalopoda appear to be the only Invertebrates in which the egg is mesoblastic and telolecithal like that of Vertebrata. This is the result of the large quantity of the yolk, and the position the latter assumes in relation to the blastoderm. In all other Mollusca the segmentation is complete though in some cases very unequal. In the egg of Loligo, which has been chiefly studied (fig. 35), the protoplasmic pole is at the narrower end of the egg, and segmentation is restricted to this end, forming a layer of ectoderm cells. From one part of the periphery of the ectoderm proliferation of cells takes place and gives rise to a layer of scattered nuclei over the whole surface of the yolk. The region of proliferation marks the anal side of the ectoderm, and the layer of nuclei forms the perivitelline membrane. This process must be regarded as equivalent to the first stage of invagination, the yolk being surrounded by hypoblast cells or their nuclei. Later on the same anal edge of the ectoderm forms another cellular layer, the endoderm proper, which forms a continuous sheet below the ectoderm.

The mesoderm also originates at the anal side of the ectoderm and extends in two bands right and left between ectoderm and endoderm. After the mesoderm is thus established, a little vesicle lying upon and open to the yolk is formed from the endoderm, and this vesicle ultimately gives rise to the stomach, the two lobes of the liver and the intestine. The buccal mass and oesophagus arise from a stomodaeal invagination, and the anus is formed later from a short proctodaeal invagination.

The external changes of form are as follows:—The mantle is the middle of the embryonic area, and in its centre is the shell-gland, which, however, behaves in a different way from that seen in other Molluscs. Its borders grow inwards and approach each other to form the shell-sac. E. Ray Lankester showed that in Argonauta and other Octopods the shell-sac disappears before it is closed up, but in other forms except Spirula it closes completely and the shell develops within it. The lateral and posterior borders of the embryo form the foot, and these borders grow out into ten or eight lobes which become the arms, and which at first, as seen in fig. 35 (8), are entirely posterior to the mouth. Development actually shows the anterior arms

gradually growing round the mouth and uniting in front of it.