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- The Evolution of Man, V.2 - 5/63 -


in the rear through the respiratory pore (Figure 2.210 c).

On the inner surface of these mantle-folds (M1), in the ventral half of the wide mantle cavity (atrium), we find the sex-organs of the Amphioxus. At each side of the branchial gut there are between twenty and thirty roundish four-cornered sacs, which can clearly be seen from without with the naked eye, as they shine through the thin transparent body-wall. These sacs are the sexual glands they are the same size and shape in both sexes, only differing in contents. In the female they contain a quantity of simple ova (Figure 2.219 g); in the male a number of much smaller cells that change into mobile ciliated cells (sperm-cells). Both sacs lie on the inner wall of the atrium, and have no special outlets. When the ova of the female and the sperm of the male are ripe, they fall into the atrium, pass through the gill-clefts into the fore-gut, and are ejected through the mouth.

(FIGURE 2.216. Transverse section of the lancelet, in the fore half. (From Ralph.) The outer covering is the simple cell-layer of the epidermis (E). Under this is the thin corium, the subcutaneous tissue of which is thickened; it sends connective-tissue partitions between the muscles (M1) and to the chorda-sheath. N medullary tube, Ch chorda, Lh body-cavity, A atrium, L upper wall of same, E1 inner wall, E2 outer wall, Lh1 ventral remnant of same, Kst gill-reds, M ventral muscles, R seam of the joining of the ventral folds (gill-covers), G sexual glands.)

Above the sexual glands, at the dorsal angle of the atrium, we find the kidneys. These important excretory organs could not be found in the Amphioxus for a long time, on account of their remote position and their smallness; they were discovered in 1890 by Theodor Boveri (Figure 2.217 x). They are short segmented canals; corresponding to the primitive kidneys of the other vertebrates (Figure 2.218 B). Their internal aperture (Figure 2.217 B) opens into the body-cavity; their outer aperture into the atrium (C). The prorenal canals lie in the middle of the line of the head, outwards from the uppermost section of the gill-arches, and have important relations to the branchial vessels (H). For this reason, and in their whole arrangement, the primitive kidneys of the Amphioxus show clearly that they are equivalent to the prorenal canals of the Craniotes (Figure 2.218 B). The prorenal duct of the latter (Figure 2.218 C) corresponds to the branchial cavity or atrium of the former (Figure 2.217 C).

(FIGURE 2.217. Transverse section through the middle of the Amphioxus. (From Boveri.) On the left a gill-rod has been struck, and on the right a gill-cleft; consequently on the left we see the whole of a prorenal canal (x), on the right only the section of its fore-leg. A genital chamber (ventral section of the gonocoel), x pronephridium, B its coelom-aperture, C atrium, D body-cavity, E visceral cavity, F subintestinal vein, G aorta (the left branch connected by a branchial vessel with the subintestinal vein), H renal vessel.

FIGURE 2.218. Transverse section of a primitive fish embryo (Selachii-embryo, from Boveri.). To the left pronephridia (B), the right primitive kidneys (A). The dotted lines on the right indicate the later opening of the primitive kidney canals (A) into the prorenal duct (C). D body-cavity, E visceral cavity, F subintestinal vein, G aorta, H renal vessel.)

If we sum up the results of our anatomic study of the Amphioxus, and compare them with the familiar organisation of man, we shall find an immense distance between the two. As a fact, the highest summit of the vertebrate organisation which man represents is in every respect so far above the lowest stage, at which the lancelet remains, that one would at first scarcely believe it possible to class both animals in the same division of the animal kingdom. Nevertheless, this classification is indisputably just. Man is only a more advanced stage of the vertebral type that we find unmistakably in the Amphioxus in its characteristic features. We need only recall the picture of the ideal Primitive Vertebrate given in a former chapter, and compare it with the lower stages of human embryonic development, to convince ourselves of our close relationship to the lancelet. (Cf. Chapter 1.11.)

It is true that the Amphioxus is far below all other living vertebrates. It is true that it has no separate head, no developed brain or skull, the characteristic feature of the other vertebrates. It is (probably as a result of degeneration) without the auscultory organ and the centralised heart that all the others have; and it has no fully-formed kidneys. Every single organ in it is simpler and less advanced than in any of the others. Yet the characteristic connection and arrangement of all the organs is just the same as in the other vertebrates. All these, moreover, pass, during their embryonic development, through a stage in which their whole organisation is no higher than that of the Amphioxus, but is substantially identical with it.

(FIGURE 2.219. Transverse section of the head of the Amphioxus (at the limit of the first and second third of the body). (From Boveri) a aorta (here double), b atrium, c chorda, co umlaut coeloma (body-cavity), e endostyl (hypobranchial groove), g gonads (ovaries), kb gill-arches, kd branchial gut, l liver-tube (on the right, one-sided), m muscles, n renal canals, r spinal cord, sn spinal nerves, sp gill-clefts.)

In order to see this quite clearly, it is particularly useful to compare the Amphioxus with the youthful forms of those vertebrates that are classified next to it. This is the class of the Cyclostoma. There are to-day only a few species of this once extensive class, and these may be distributed in two groups. One group comprises the hag-fishes or Myxinoides. The other group are the Petromyzontes, or lampreys, which are a familiar delicacy in their marine form. These Cyclostoma are usually classified with the fishes. But they are far below the true fishes, and form a very interesting connecting-group between them and the lancelet. One can see how closely they approach the latter by comparing a young lamprey with the Amphioxus. The chorda is of the same simple character in both; also the medullary tube, that lies above the chorda, and the alimentary canal below it. However, in the lamprey the spinal cord swells in front into a simple pear-shaped cerebral vesicle, and at each side of it there are a very simple eye and a rudimentary auditory vesicle. The nose is a single pit, as in the Amphioxus. The two sections of the gut are also just the same and very rudimentary in the lamprey. On the other hand, we see a great advance in the structure of the heart, which is found underneath the gills in the shape of a centralised muscular tube, and is divided into an auricle and a ventricle. Later on the lamprey advances still further, and gets a skull, five cerebral vesicles, a series of independent gill-pouches, etc. This makes all the more interesting the striking resemblance of its immature larva to the developed and sexually mature Amphioxus.

While the Amphioxus is thus connected through the Cyclostoma with the fishes, and so with the series of the higher vertebrates, it is, on the other hand, very closely related to a lowly invertebrate marine animal, from which it seems to be entirely remote at first glance. This remarkable animal is the sea-squirt or Ascidia, which was formerly thought to be closely related to the mussel, and so classed in the molluscs. But since the remarkable embryology of these animals was discovered in 1866, there can be no question that they have nothing to do with the molluscs. To the great astonishment of zoologists, they were found, in their whole individual development, to be closely related to the vertebrates. When fully developed the Ascidiae are shapeless lumps that would not, at first sight, be taken for animals at all. The oval body, frequently studded with knobs or uneven and lumpy, in which we can discover no special external organs, is attached at one end to marine plants, rocks, or the floor of the sea. Many species look like potatoes, others like melon-cacti, others like prunes. Many of the Ascidiae form transparent crusts or deposits on stones and marine plants. Some of the larger species are eaten like oysters. Fishermen, who know them very well, think they are not animals, but plants. They are sold in the fish markets of many of the Italian coast-towns with other lower marine animals under the name of "sea-fruit" (frutti di mare). There is nothing about them to show that they are animals. When they are taken out of the water with the net the most one can perceive is a slight contraction of the body that causes water to spout out in two places. The bulk of the Ascidiae are very small, at the most a few inches long. A few species are a foot or more in length. There are many species of them, and they are found in every sea. As in the case of the Acrania, we have no fossilised remains of the class, because they have no hard and fossilisable parts. However, they must be of great antiquity, and must go back to the primordial epoch.

The name of "Tunicates" is given to the whole class to which the Ascidiae belong, because the body is enclosed in a thick and stiff covering like a mantle (tunica). This mantle--sometimes soft like jelly, sometimes as tough as leather, and sometimes as stiff as cartilage--has a number of peculiarities. The most remarkable of them is that it consists of a woody matter, cellulose--the same vegetal substance that forms the stiff envelopes of the plant-cells, the substance of the wood. The tunicates are the only class of animals that have a real cellulose or woody coat. Sometimes the cellulose mantle is brightly coloured, at other times colourless. Not infrequently it is set with needles or hairs, like a cactus. Often we find a mass of foreign bodies--stone, sand, fragments of mussel-shells, etc.--worked into the mantle. This has earned for the Ascidia the name of "the microcosm."

(FIGURE 2.220. Organisation of an Ascidia (left view); the dorsal side is turned to the right and the ventral side to the left, the mouth (o) above; the ascidia is attached at the tail end. The branchial gut (br), which is pierced by a number of clefts, continues below in the visceral gut. The rectum opens through the anus (a) into the atrium (cl), from which the excrements are ejected with the respiratory water through the mantle-hole or cloaca (a); m mantle. (From Gegenbaur.)

FIGURE 2.221. Organisation of an Ascidia (as in Figure 2.220, seen from the left). sb branchial sac, v stomach, i small intestine, c heart, t testicle, vd sperm-duct, o ovary, o apostrophe ripe ova in the branchial cavity. The two small arrows indicate the entrance and exit of the water through the openings of the mantle. (From Milne-Edwards.))

The hind end, which corresponds to the tail of the Amphioxus, is usually attached, often by means of regular roots. The dorsal and ventral sides differ a good deal internally, but frequently cannot be distinguished externally. If we open the thick tunic or mantle in order to examine the internal organisation, we first find a spacious cavity filled with water--the mantle-cavity or respiratory cavity (Figure 2.220 cl). It is also called the branchial cavity and the cloaca, because it receives the excrements and sexual products as well as the respiratory water. The greater part of the respiratory cavity is occupied by the large grated branchial sac (br). This is so like the gill-crate of the Amphioxus in its whole arrangement that the resemblance was pointed out by the English naturalist Goodsir, years ago, before anything was known of the relationship of the two animals. As a fact, even in the Ascidia the mouth (o) opens first into this wide branchial sac. The respiratory water passes through the lattice-work of the branchial sac into the branchial cavity, and is ejected from this by the respiratory pore (a apostrophe). Along the ventral side of the branchial sac runs a ciliated groove--the


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