the apical plate (1). This usually carries a tuft of long cilia or sensory
hairs, and sometimes rudimentary visual organs. Mesoblastic
bands are present, proceeding a short distance forwards from the
anus on each side of the middle ventral line (6), and at the anterior
end of each of these structures is a tube (5) which more or less
branches internally and opens on the ventral surface. The branches
of this tube end internally in peculiar cells containing a
flame-shaped
flagellum and
floating in the so-called
body cavity, into
which, however, they
do not open. These
are the primitive kidneys.
The body
cavity, which is a
space between the
ectoderm and alimentary
canal, is not
lined by mesoderm
and is traversed by a
few muscular fibres.
Such a larva is found,
almost as described,
in many Chaetopods
(fig. 1), in Echiurus (fig.
2), in many Gastropods
(fig. 3), and
Lamellibranchiates
(fig. 4). This typical
structure of the larva
is often departed from,
and the molluscan trochosphere
can be distinguished
from the
annelidan by the possession
of a rudiment
at least of the shell-gland
and foot (figs. 3
and 4); but in all
cases in which the
young leaves the egg
at an early stage of
development it has
a form which can
be referred without
much difficulty to the
trochosphere type just described. A larva similar to the trochosphere
in some features, particularly in possessing a preoral
ring of cilia and an apical plate, is found in the Polyzoa, and
in adult Rotifera, which latter, in their ciliary ring and excretory
organs, present some
resemblance to the trochosphere,
and are sometimes described
as permanent adult
trochospheres. But in these
phases the resemblance to the
typical forms is not nearly so close as it is in the case of the larva
of Annelida and Mollusca.
|
| After V. Drasche in Beiträge zur Entwickelung der Polychaeten, Entwickelung von Pomatoceros. |
| Fig. 1.—Trochosphere Larva of the Chaetopod Pomatoceros trigueter, L. (Osmic acid preparation.) |
| 1. The apical plate. |
| 2. Long cilia of preoral band (velum). |
| 3. Long cilia of postoral band. |
| 4. Mouth. |
| 5. Excretory organ. |
| 6. Mesoblastic band. |
| 7. Anus. |
|
| |
| After Hatschek, “Echiurus” in Claus’s Arbeiten aus dem zoolog. Institut der Wien. | After Patten, “Patella” in Claus’s Arbeiten aus dem zoolog. Institut der Wien. | |
| Fig. 2.—Young Trochosphere Larva of the Gephyrean Echiurus, seen in optical section. | Fig. 3.—Larva of the Gastropod Patella, seen in longitudinal vertical section. | |
| 1. Apical plate. | 1. Apical plate. | |
| 2. Muscle-bands. | 2. Cilia of preoral circlet (velum). | |
| 3. Preoral band of cilia (velum). | 3. Mouth. | |
| 4. Mouth. | 4. Foot. | |
| 5. Mesoblastic band. | 5. Anal tuft of cilia. | |
| 6. Anus. | 6. Shell-gland covered by shell. |
In the Echinodermata there are two distinct larval forms which cannot be brought into relation with one another. The one of these is found in the Asteroids, Ophiuroids, Echinoids and Holothuroids; the other in the Crinoids.
 |
| After Hatschek on “Teredo” in Claus’ Arbeiten aus dem zoolog. Institut der Wien. |
| Fig. 4.—A, Embryo, and B, Young Trochosphere Larva of the Lamellibranch Teredo. |
|
In A the shell-gland (1) and the mouth (2) and the rudiment of the enteron (3) are shown; (4) primitive mesoderm cells. In B the shell-gland has flattened out and the shell is formed. The cilia of the preoral and postoral bands are not clearly differentiated at this stage. |
The first is, in its most primitive form, a small transparent creature, with a mouth and anus and a postoral longitudinal ciliated band (fig. 5, A). In Asteroids the band of cilia becomes divided in such a way as to give rise to two bands, the one preoral, encircling the preoral lobe, and the other remaining postoral (fig. 5, B). In the other groups the band remains single and longitudinal. In all cases the edges of the body carrying the ciliary bands become sinuous (fig. 6) and sometimes prolonged into arms (figs. 7-9), and each of the four groups has its own type of larva. In Asteroids, in which the band divides, the larva is known as the bipinnaria (fig. 7); in Holothurians it is called the auricularia (fig. 6); in Echinoids and Ophiuroids, in which the arms are well marked, it is known as the pluteus, the echinopluteus (fig. 9) and ophiopluteus (fig. 8) respectively.
All these forms were obviously distinct but as obviously modifications of a common type and related to one another. They present certain remarkable structural features which differentiate them from other larval types except the tornaria larvae of the Enteropneusta. They possess an alimentary canal with a mouth and anus as does the trochosphere, but they differ altogether from that larva in having a diverticulum of the alimentary canal which gives rise to the coelom and to a considerable part of the mesoblast. Further, they are without an apical plate with its tuft of sensory hairs.
In Crinoids the type is different (fig. 10), and might belong to a different phylum. The body is opaque, and encircled by five ciliary bands, and is without either mouth, anus or arms, and there is a tuft of cilia on the preoral lobe. A resemblance to the other Echinoderm larvae is found in the fact that coelomic diverticula of the enteron are present.
|
|  |
| From Balfour’s Comparative Embryology, by permission of Macmillan & Co., Ltd. |
After J. Müller. | |
Fig. 5.—Diagrams of side views of two young Echinoderm Larvae, showing the course of the ciliary bands. A, auricularia larva of a Holothurian; B, bipinnaria larva of an Asteroid; a, anus; l.c, in A primitive longitudinal ciliary band, in B postoral longitudinal ciliary band; m, mouth; pr.c, preoral ciliary band; st, stomach. |
Fig. 6.—Auricularia stelligera, ventral view, somewhat diagrammatic. The larva of a Holothurian.
1. Frontal area. |
The larvae of two other groups present certain resemblances to the typical Echinoderm larvae. The one of these is the tornaria larva of the Enteropneusta (fig. 11), which recalls Echinoderms in the possession of two ciliary bands, the one preoral and the other postoral and partly longitudinal, and in the presence of gut diverticula which give rise to the coelom; but, like the trochosphere, it possesses an apical plate with sensory organs on the preoral lobe. The resemblance of the tornaria to the bipinnaria is so close that, taking into consideration certain additional resemblances in the arrangement
