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Volume 75, 1945-46
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The Vestigial Pluteus.

The embryo is gradually growing larger, there being a notable increase in the amount of the mesenchyme, especially at the aboral (posterior) pole. This tissue comes to surround completely both the oesophagus and hydrocoel as well as the archenteron, filling up the space between these organs and the ectoderm. At the same time the mesenchyme cells become more compacted together, so as to eliminate the intercellular spaces originally present. Fig. 11 is an oblique

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Fig. 11.—Oblique section of later larva, showing five-lobed hydrocoe.
Arch., archenteron; Hydr., hydrocoel; Rud.Amb.C., rudiment of ambulacral canal; Oes., oesophageal sac; Mesch., mesenchyme; Ect., ectoderm.

section through the larva, so directed as to pass through all lobes of the hydrocoel—these having now reached the maximum number of five; for no further differentiation of the organ occurs till the pluteus stage has been superseded. Each lobe is a rudiment of a future radial ambulacral canal, and at present they are placed in linear series in an antero-posteral direction on the left side of the oesophagus. The cells of the archenteron—or stomach as it may now be termed—have multiplied to form a wall two cells deep. In Fig. 11 the section, being oblique so as to show all lobes of the hydrocoel, does not pass through the centre of the stomach, and so only a small part of it is seen in the section. The wall tissue of the oesophagus

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has also increased in depth, reaching two or three cells thickness, while the central lumen has assumed a three-lobed shape. The embryo is now becoming more transparent, so that it is possible to utilise whole mounts if well cleared. A slightly older stage, when the larva has reached its fullest development as a pluteus, is drawn

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Fig. 12.—Fully-developed larva, with embryonic attachment.
Post.P., posterior pole; Rt., Rt., Skel.Pl., left, and right skeletal plate of larva; Stom., stomach; Hydr., hydrocoel; Ant.P., anterior pole; Oes., oesophageal sac; Mesench., mesenchyme; Skel.Rud.Ad., skeletal rudiment of the adult; Emb.Al., embryonic attachment.

in Fig. 12, from a cleared whole object. The pluteus has a somewhat pyriform shape, rather pointed at the anterior (oral) pole, and rounded in the posterior part to which is joined the embryonic attachment. The oesophagus in the specimen figured is somewhat elongate in form, but this character is variable, some specimens having the organ rounded. The wall of the oesophagus is thickened and its lumen consequently very narrow. To the left is seen the five-lobed hydrocoel with the rudiments of the ambulacral canals visible within the five lobes. The large, spherical stomach occupies most of the aboral part of the larval body. The ectoderm is in the form of a thin sheet of investing cells of regular arrangement. On either side of the stomach extend forwards the two vestigial skeletal rods of the pluteus, whilst triradiate rudiments of some of the plates of the future adult are to be seen scattered in the more superficial regions of the mesenchyme just below the ectoderm.

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The relationship between this peculiar reduced and vestigial larval stage and a typical pelagic Ophiopluteus is illustrated in diagrammatic form in Fig. 13, where it can readily be seen that the pluteus found in Amphipholis squamata is in reality an armless version of the strongly bilaterally symmetrical larva of those types of ophiuroids which have an indirect, pelagic development. As is shown elsewhere (Fell, 1945), this vestigial larva of Amphipholis can be regarded as but one of a series of forms which illustrate various stages in the gradual reduction and disappearance of the larva in the ontogeny of ophiuroids. It has been one of the purposes of this study

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Fig. 13.—Diagram illustrating the relationship of the vestigial pluteus of Amphipholis squamata (right) to the fully-developed ophiopluteus of ophiuroids with indirect development (left).

to obtain information on such a reduced larva as this in order to achieve some further degree of understanding of the manner in which the larva has been so completely lost by Kirk's ophiuroid. While the general relationships of the reduced pluteus of Amphipholis squamata are more fully discussed elsewhere, it will be useful to note here the more salient points in which it differs from the pelagic Ophiopluteus. These are:—

  • (1) All four pairs of larval arms have disappeared, leaving only traces of their skeletal rods in the two calcareous plates on either side of the stomach.

  • (2) The pluteus here is not free-living, being formed within the bursa of the parent, and attached to it by an outgrowth of cells from the wall of the bursa. This organ is termed here the “embryonic attachment.”

  • (3) The tissues are more heavily charged with yolk material. Nutrition is obtained partly from this substance, and partly from secretions of the bursal wall (see below), and thus a pelagic food-gathering stage is rendered physiologically unnecessary.

  • (4) As a condition correlated to the previous one, the alimentary system is not yet functional. The oesophageal sac is still closed off from the exterior, and there is no anus developed.

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  • (5) The only part of the original coelomic sacs to survive is the well-developed hydrocoel. This is interesting in view of the conditions found in Kirk's ophiuroid, where the hydrocoel is the part of the definitive coelom which is first brought into being. Here also the hydrocoel is the first part of the definitive coelom to form, there being no general coelom present.

  • (6) The mesenchyme is very greatly developed, a condition also paralleled in Kirk's ophiuroid. In the present instance, as in Kirk's ophiuroid, the mesenchyme is destined to play a most important part in the subsequent differentiation of the coelom of the adult.

Before passing on to describe the metamorphosis, it is convenient at this point to consider in greater detail the larval skeleton, the embryonic attachment, and the mode of nutrition.