The Evolution of Man, vol 2 - Ernst Haeckel (sight word books .TXT) 📗

one whip-like hair each, and so are called flagellate (whip) cells (in contrast with the ciliated cells, which have a number of short lashes or cilia).

In the further course of its rapid development the roundish bell-gastrula becomes elongated, and begins to flatten on one side, parallel to the long axis. The flattened side is the subsequent dorsal side; the opposite or ventral side remains curved. The latter grows more quickly than the former, with the result that the primitive mouth is forced to the dorsal side (Figure 1.39). In the middle of the dorsal surface a shallow longitudinal groove or furrow is formed (Figure 1.79), and the edges of the body rise up on each side of this groove in the shape of two parallel swellings. This groove is, of course, the dorsal furrow, and the swellings are the dorsal or medullary swellings; they form the first structure of the central nervous system, the medullary tube. The medullary swellings now rise higher; the groove between them becomes deeper and deeper. The edges of the parallel swellings curve towards each other, and at last unite, and the medullary tube is formed (Figures 1.83 m and 1.84 m). Hence the formation of a medullary tube out of the outer skin takes place in the naked dorsal surface of the free-swimming larva of the Amphioxus in just the same way as we have found in the embryo of man and the higher animals within the foetal membranes.

Simultaneously with the construction of the medullary tube we have in the Amphioxus-embryo the formation of the chorda, the coelom-pouches, and the mesoderm proceeding from their wall. These processes also take place with characteristic simplicity and clearness, so that they are very instructive to compare with the vermalia on the one hand and with the higher vertebrates on the other. While the medullary groove is sinking in the middle line of the flat dorsal side of the oval embryo, and its parallel edges unite to form the ectodermic neural tube, the single chorda is formed directly underneath them, and on each side of this a parallel longitudinal fold, from the dorsal wall of the primitive gut. These longitudinal folds of the entoderm proceed from the primitive mouth, or from its lower and hinder edge. Here we see at an early stage a couple of large entodermic cells, which are distinguished from all the others by their great size, round form, and fine-grained protoplasm; they are the two promesoblasts, or polar cells of the mesoderm (Figure 1.83 p). They indicate the original starting-point of the two coelom-pouches, which grow from this spot between the inner and outer germinal layers, sever themselves from the primitive gut, and provide the cellular material for the middle layer.

Immediately after their formation the two coelom-pouches of the Amphioxus are divided into several parts by longitudinal and transverse folds. Each of the primary pouches is divided into an upper dorsal and a lower ventral section by a couple of lateral longitudinal folds (Figure 1.82). But these are again divided by several parallel transverse folds into a number of successive sacs, the primitive segments or somites (formerly called by the unsuitable name of “primitive vertebrae”). They have a different future above and below. The upper or dorsal segments, the episomites, lose their cavity later on, and form with their cells the muscular plates of the trunk. The lower or ventral segments, the hyposomites, corresponding to the lateral plates of the craniote-embryo, fuse together in the upper part owing to the disappearance of their lateral walls, and thus form the later body-cavity (metacoel); in the lower part they remain separate, and afterwards form the segmental gonads.

In the middle, between the two lateral coelom-folds of the primitive gut, a single central organ detaches from this at an early stage in the middle line of its dorsal wall. This is the dorsal chorda (Figures 1.83 and 1.84 ch). This axial rod, which is the first foundation of the later vertebral column in all the vertebrates, and is the only representative of it in the Amphioxus, originates from the entoderm.

In consequence of these important folding-processes in the primitive gut, the simple entodermic tube divides into four different sections:—

1. underneath, at the ventral side, the permanent alimentary canal or permanent gut;

2. above, at the dorsal side, the axial rod or chorda; and

3. the two coelom-sacs, which immediately subdivide into two structures:—

3A. above, on the dorsal side, the episomites, the double row of primitive or muscular segments; and

3B. below, on each side of the gut, the hyposomites, the two lateral plates that give rise to the sex-glands, and the cavities of which partly unite to form the body-cavity. At the same time, the neural or medullary tube is formed above the chorda, on the dorsal surface, by the closing of the parallel medullary swellings.

All these processes, which outline the typical structure of the vertebrate, take place with astonishing rapidity in the embryo of the Amphioxus; in the afternoon of the first day, or twenty-four hours after fertilisation, the young vertebrate, the typical embryo, is formed; it then has, as a rule, six to eight somites.

The chief occurrence on the second day of development is the construction of the two permanent openings of the gut—the mouth and anus. In the earlier stages the alimentary tube is found to be entirely closed, after the closing of the primitive mouth; it only communicates behind by the neurenteric canal with the medullary tube. The permanent mouth is a secondary formation, at the opposite end. Here, at the end of the second day, we find a pit-like depression in the outer skin, which penetrates inwards into the closed gut. The anus is formed behind in the same way a few hours later (in the vicinity of the additional gastrula-mouth). In man and the higher vertebrates also the mouth and anus are formed, as we have seen, as flat pits in the outer skin; they then penetrate inwards, gradually becoming connected with the blind ends of the closed gut-tube. During the second day the Amphioxus-embryo undergoes few other changes. The number of primitive segments increases, and generally amounts to fourteen, some forty-eight to fifty hours after impregnation.

Almost simultaneously with the formation of the mouth the first gill-cleft breaks through in the fore section of the Amphioxus-embryo (generally forty hours after the commencement of development). It now begins to nourish itself independently, as the food material stored up in the ovum is completely used up. The further development of the free larvae takes place very slowly, and extends over several months. The body becomes much longer, and is compressed at the sides, the head-end being broadened in a sort of triangle. Two rudimentary sense-organs are developed in it. Inside we find the first blood-vessels, an upper or dorsal vessel, corresponding to the aorta, between the gut and the dorsal cord, and a lower or ventral vessel, corresponding to the subintestinal vein, at the lower border of the gut. Now, the gills or respiratory organs also are formed at the fore-end of the alimentary canal. The whole of the anterior or respiratory section of the gut is converted into a gill-crate, which is pierced trellis-wise by numbers of branchial-holes, as in the ascidia. This is done by the foremost part of the gut-wall joining star-wise with the outer skin, and the formation of clefts at the point of connection, piercing the wall and leading into the gut from without. At first there are very few of these branchial clefts; but there are soon a number of them—first in one, then in two, rows. The foremost gill-cleft is the oldest. In the end we have a sort of lattice work of fine gill-clefts, supported on a number of stiff branchial rods; these are connected in pairs by transverse rods.

(FIGURES 2.222 TO 2.224. Transverse sections of young Amphioxus-larvae (diagrammatic, from Ralph.) (Cf. also Figure 2.216.) In Figure 2.222 there is free communication from without with the gut-cavity (D) through the gill-clefts (K). In Figure 2.223 the lateral folds of the body-wall, or the gill-covers, which grow downwards, are formed. In Figure 2.224 these lateral folds have united underneath and joined their edges in the middle line of the ventral side (R seam). The respiratory water now passes from the gut-cavity (D) into the mantle-cavity (A). The letters have the same meaning throughout: N medullary tube, Ch chorda, M lateral muscles, Lh body-cavity, G part of the body-cavity in which the sexual organs are subsequently formed. D gut-cavity, clothed with the gut-gland layer (a). A mantle-cavity, K gill-clefts, b = E epidermis, E1 the same as visceral epithelium of the mantle-cavity, E2 as parietal epithelium of the mantle-cavity.)

At an early stage of embryonic development the structure of the Amphioxus-larva is substantially the same as the ideal picture we have previously formed of the “Primitive Vertebrate” (Figures 1.98 to 1.102). But the body afterwards undergoes various modifications, especially in the fore-part. These modifications do not concern us, as they depend on special adaptations, and do not affect the hereditary vertebrate type. When the free-swimming Amphioxus-larva is three months old, it abandons its pelagic habits and changes into the young animal that lives in the sand. In spite of its smallness (one-eighth of an inch), it has substantially the same structure as the adult. As regards the remaining organs of the Amphioxus, we need only mention that the gonads or sexual glands are developed very late, immediately out of the inner cell-layer of the body-cavity. Although we can find afterwards no continuation of the body-cavity (Figure 2.216 U) in the lateral walls of the mantle-cavity, in the gill-covers or mantle-folds (Figure 2.224 U), there is one present in the beginning (Figure 2.224 Lh). The sexual cells are formed below, at the bottom of this continuation (Figure 2.224 S). For the rest, the subsequent development into the adult Amphioxus of the larva we have followed is so simple that we need not go further into it here.

We may now turn to the embryology of the Ascidia, an animal that seems to stand so much lower and to be so much more simply organised, remaining for the greater part of its life attached to the bottom of the sea like a shapeless lump. It was a fortunate accident that Kowalevsky first examined just those larger specimens of the Ascidiae that show most clearly the relationship of the vertebrates to the invertebrates, and the larvae of which behave exactly like those of the Amphioxus in the first stages of development. This resemblance is so close in the main features that we have only to repeat what we have already said of the ontogenesis of the Amphioxus.

The ovum of the larger Ascidia (Phallusia, Cynthia, etc.) is a simple round cell of 1/250 to 1/125 of an inch in diameter. In the thick fine-grained yelk we find a clear round germinal vesicle of about 1/750 of an inch in diameter, and this encloses a small embryonic spot or nucleolus. Inside the membrane that surrounds the ovum, the stem-cell of the Ascidia, after fecundation, passes through just the same metamorphoses as the stem-cell of the Amphioxus. It undergoes total segmentation; it divides into two, four, eight, sixteen, thirty-two cells, and so on. By continued total cleavage the morula, or mulberry-shaped cluster of cells, is formed. Fluid gathers inside it, and thus we get once more a globular vesicle (the blastula); the wall of this is a single stratum of cells, the blastoderm. A real gastrula (a simple bell-gastrula) is formed from the blastula by invagination, in the same way as in the amphioxus.

Up to this there is no definite ground in the embryology of the Ascidiae for bringing them into close relationship with the Vertebrates; the same gastrula is formed in the same way in many other animals of different stems. But we now find an embryonic process that is peculiar to