Page:EB1911 - Volume 18.djvu/703

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MOLLUSCA
673


in later and special lines of descent—show approximately the following history. By division, of the egg-cell a mulberry-mass of embryonic-cells is formed (morula), which dilates, forming a one-cell-layered sac (blastula). By invagination one portion of this sphere becomes tucked into the other—as in the preparation of a woven night-cap for the head. The orifice of invagination (blastopore) narrows, and we now have a two-cell-layered sac—the gastrula. The invaginated layer is the enteric cell-layer or endoderm; the outer cell-layer is the dermic cell-layer or ectoderm. The cavity communicating with the blastopore and lined by the endoderm is the arch-enteron. The blastopore, together with the whole embryo, now elongates. The blastopore then closes along the middle portion of its extent, which corresponds with the later developed foot. At the same time the stomodaeum, or oral invagination, forms around the anterior remnant of the blastopore, and the proctodaeum, or anal invagination, forms around the posterior remnant of the blastopore. There are, however, variations in regard to the relation of the blastopore to the mouth and to the anus which are probably modifications of the original process described above.

(After Lankester, 15.)
Fig. 2.—Development of the Pond-Snail, Limnaeus stagnalis.
r, Directive corpuscle.
bl, Blastopore.
en, Endoderm or enteric cell layer.
ec, Ectoderm or deric cell-layer.
v, Velum.
m, Mouth.
f, Foot.
t, Tentacles.
fp, Pore in the foot (belonging to the pedal gland?).
mf, The mantle-flap or limbus pallialis.
sh, The shell.
l, The sub-pallial space, here destined to become the lung.
A, First four cells resulting from the cleavage of the original egg-cell.
B, Side-view of the same.
C, Diblastula stage showing the two cell-layers and the blastopore.
D, E, F, Trochosphere stage, D older than E or F.
G, Three-quarter view of a Diblastula, to show the orifice of invagination of the endoderm or blastopore, (bl).
H, I, Veliger stage later than D.

In eggs which contain a larger quantity of food-yolk, the process by which the endoderm is enveloped by the ectoderm is somewhat different. Segmentation in these is very unequal, and results in the formation of small cells called micromeres and large cells called megameres, as in fig. 4. As the micromeres become more numerous they gradually envelop the megameres until the latter are completely enclosed. The gastrula is in these cases said to be formed by epibole. Between ectoderm and endoderm a third intermediate cell-layer is formed, which is called the mesoderm, and gives rise to the muscular and connective tissues to the vascular system, and to the excretory and generative organs. The mesoderm arises for the most part from the endoderm. When the segmentation is unequal one of the megameres gives rise by successive divisions to two primary mesoderm cells called mesomeres; these divide to form two masses of cells called mesoblastic bands. The coelom is formed as a cavity or cavities in the interior of these cell-masses. In some cases the coelom is formed as a single cavity, and renal and generative organs are formed from its walls. This is the primitive method, but in other cases the organs mentioned may be formed separately in the mesoderm. The renal organs are tubular outgrowths of the pericardial parts of the coelom; the reproductive cells are derived from cells lining the generative portion.

(After Lankester, 17.)
Fig. 3.—Development of the River-Snail, Paludina vivipara.
dc, Directive corpuscle (outcast cell).
ae, Arch-enteron or cavity lined by the enteric cell-layer or endoderm.
bl, Blastopore.
vr, Velum or circlet of ciliated cells.
dv, Velar area or cephalic dome.
sm, Site of the as yet unformed mouth.
f, Foot.
mes, Rudiments of the skeletotrophic tissues.
pi, The pedicle of invagination, the future rectum.
shgl The primitive shell-sac or shell-gland.
m, Mouth. an, Anus.
A, Gastrula phase (optical section).
B, The Gastrula has become a Trochosphere by the development of the ciliated ring vr (optical section).
C, Side view of the Trochosphere with commencing formation of the foot.
D, Further advanced Trochosphere (optical section).
E, The Trochosphere passing to the Veliger stage, dorsal view showing the formation of the primitive shell-sac.
F, Side view of the same, showing foot, shell-sac (shgl), velum (vr), mouth and anus.

N.B.—In this development the blastopore is not elongated; it persists as the anus. The mouth and stomodaeum form independently of the blastopore.

The external form of the embryo meanwhile passes through highly characteristic changes, which are on the whole fairly constant throughout the Mollusca. A circlet of cilia forms when the embryo is still nearly spherical in an equatorial position. As growth proceeds, one hemisphere remains relatively small, the other elongates and enlarges. Both mouth and anus are placed in the larger area; the smaller area is the prostomium simply; the ciliated band is therefore in front of the mouth. The larval form thus produced is known as the trochosphere. It exactly agrees with the larval form of many Chaetopod worms and other Coelomata. Most remarkable is its resemblance to the adult form of the Wheel animalcules, or Rotifera, which retain the prae-oral ciliated band as their chief organ of locomotion and prehension throughout life. So far the young mollusc has not reached a definitely molluscan stage of