Page:EB1911 - Volume 24.djvu/553

SCYPHOMEDUSAE

523

After W. Schewiakoff, simplified from a coloured plate in Morphologisches Jahrbuch, xv., 1889, by permission of Wilhelm Engelmann.

Fig. 9.—Sensory cells from the retina of Charybdaea, highly magnified. c, Visual cone; n, nucleus; n.f, nerve fibril.

The ovum undergoes total cleavage, giving rise to a bastula which forms a gastrula (fig. 10, A) by invagination (see article Hydrozoa) This is a type of germ-layer formation never found in the Hydromedusae, though of universal occurrence in all groups of animals above the Coelentera. We may regard it as a form of unipolar immigration in which the immigrating cells pass into the interior in a connected epithelial layer, instead of going in singly and independently. The embryo is set free as a planula larva (fig. 10, B) in the gastrula stage, and the orifice of invagination or blastopore, which persists, is situated at the hinder pole. After a time the planula fixes itself by the anterior pole, with the blastopore uppermost. The larva after fixation changes into a polyp-like organism termed a scyphistoma or scyphopolyp (fig. 10, C, D). The body becomes in shape like a vase or urn attached by a narrow stalk, round which a chitinous membrane is secreted. From the edges of the vase the four primary tentacles grow out, each a slender filament with a solid endodermal axis. The tentacles border a broad, flattened peristome, from the centre of which arises the hypostome with the mouth at its extremity; the hypostome is at first low, but soon becomes a projecting, chimney-like tube. It has been sought to prove that the interior of the hypostome is lined by ectoderm, so as to form a stomodaeum or ectodermal oesophagus similar to that of the Anthozoa, but this has been disproved by the most recent investigations of Hein (4) and Friedemann (3), who have shown that the mouth at the extremity of the hypostome represents the persistent blastopore of the gastrula stage.

Fig. 10.—Four stages in the development of Chrysaora. From Balfour, after Claus.
A,	Diblastula stage.
B,	Stage after closure of blastopore.
C,	Fixed larva.
D,	Later stage with mouth, short tentacles, &c.
ep,	Ectoderm.
hy,	Endoderm.
pe,	Stomodaeum.
m,	Mouth.
bl,	Blastopore.

The internal gastric cavity of the scyphistoma is not a simple space as in the hydropolyp, but is subdivided by four ridges or taeniolae, arising one in each interradius (fig. 11, B). Each taeniola is similar in its anatomical relations to the similarly named structures in Haliclystus (fig. 1), and becomes perforated in the same way at its outer side by a “septal ostium,” forming as it were the rudiment of a ring-canal. Each taeniola bears a strongly developed longitudinal muscle-band, stated by Claus and Chun to be developed from the endoderm, like the retractor muscles of the anthopolyp, but by other investigators it is affirmed that each retractor muscle of the scyphistoma arises from the lining of a funnel-shaped ectodermal ingrowth (“Septaltrichter”) growing down from the peristome inside each taeniola, in a manner similar to the infundibular cavities of Lucernaria, which in their turn are homologous with the subgenital cavities of other Scyphomedusae. It is asserted, however, by Friedemann (3), a recent investigator of the subject, that the infundibular cavities appear late in the scyphistoma and have no relation either to the septal muscles or to the subgenital cavities of the adult. The muscle-bands are very contractile, rendering the scyphistoma one of the most difficult of all organisms to preserve in an expanded condition. By their contraction the muscles of the taeniolae drag the hypostome down and so produce the appearances which have been interpreted as a stomodaeal invagination.

As the scyphistoma grows the tentacles increase in number, four interradial and eight adradial being formed in addition to the four primary perradial tentacles (fig. 11, A, B, C). The animal may produce its like by lateral budding, or by budding from a basal stolon. The scyphistoma of Nausithoë forms a branching network which grows in the sponge Esperella and forms the colonial polypoid organism named by Schulze Spongicola fistularis, by Allman Stephanoscyphus mirabilis. Sooner or later, however, the scyphistoma produces free medusae by a process of transverse fission termed strobilization. In the simplest case one medusa, or at least one at a time, is produced in this way (monodisk strobilization); a circular furrow cuts off the upper, tentacle-bearing portion from the lower half of the scyphistoma (fig. 11, D, and fig. 12), and the upper part becomes detached and swims away, while the base regenerates a new crown. In most cases, however, many such furrows are formed (polydisk strobilization), so that the animal comes to resemble a pile of saucers one above the other (fig. 12). The uppermost saucers of the pile become detached successively and swim off. In this state the scyphistoma is termed a strobila.

Fig. 11.—Later development of Chrysaora and Aurelia. (After Claus.)
A,	Scyphistoma of Chrysaora, with four perradial tentacles and horny basal perisarc.
B,	Oral surface of later stage of scyphistoma of Aurelia, with commencement of four interradial tentacles. The quadrangular mouth is seen in the centre; the outline of the stomach wall, seen by transparency around it, is nipped in four places interradially to form the four gastric ridges.
C,	Oral surface of a sixteen-tentacled scyphistoma of Aurelia. The four gastric interradial ridges are seen through the mouth.
D,	First constriction of the Aurelia scyphistoma to form the pile of ephyrae or young medusae. The single ephyra carries the sixteen scyphistoma tentacles, which will atrophy and disappear. The four longitudinal gastric ridges are seen by transparency.
E,	Young ephyra just liberated, showing the eight bifurcate arms of the disk and the interradial single gastral filaments.
F,	Ephyra developing into a medusa by the growth of the adradial regions. The gastral filaments have increased to three in each of the four sets.
A,	Margin of the mouth.
Ad,	Adradial radius.
F,	Gastral filament.
In,	Interradial radius.
JG,	Adradial gastral canal.
JR = R³, Adradial lobe of the disk.
K,	Lappet of a per radial arm.
M,	Stomach wall.
Mst,	Muscle of the gastral ridge.
Mw,	Gastral ridge.
Ms,	Mesogloea.
O,	Tentaculocyst.
P,	Perradial radius.
R²,	Interradial radius.
R³,	Adradial radius.
SG,	Commencement of lateral vessel.

The medusae produced by strobilization of the scyphistoma are of a peculiar type termed Ephyrae (fig. 11, E, F). As preparations for their formation the margin of the peristome of the scyphistoma grows out into eight lobes, four perradial, four interradial. The sixteen tentacles of the scyphistoma disappear, and in the place of the four perradial and four interradial tentacles, the eight tentaculocysts of the adult are formed as outgrowths of the subumbral margin, independently of the tentacles of the scyphistoma (Friedemann). The septal ostia become widened and the gastral cavity flattened, whereby the taeniolae become comparatively shallow columns, similar to the septal nodes or cathammata of other forms.

The ephyra has a flat, disk-shaped body, with eight marginal lobes (four perradial, four interradial); a tentaculocyst is lodged in a deep notch at the apex of each lobe. Four groups of phacellae indicate the four interradii. The stomach has sixteen marginal pouches and the general anatomical structure recalls that of Pelagia. As the

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