The Evolution of Man — Volume 2

by Ernst Haeckel

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 hypobranchial groove which we have previously found at the same spot in the Amphioxus. The food of the Ascidia also consists of tiny organisms, infusoria, diatoms, parts of decomposed marine plants and animals; etc. These pass with the water into the gill-crate and the digestive part of the gut at the end of it, at first into an enlargement of it that represents the stomach. The adjoining small intestine usually forms a loop, bends forward, and opens by an anus (Figure 2.220 a), not directly outwards, but first into the mantle cavity; from this the excrements are ejected by a common outlet (a apostrophe) together with the used-up water and the sexual products. The outlet is sometimes called the branchial pore, and sometimes the cloaca or ejection-aperture. In many of the Ascidiae a glandular mass opens into the gut, and this represents the liver. In some there is another gland besides the liver, and this is taken to represent the kidneys. The body-cavity proper, or coeloma, which is filled with blood and encloses the hepatic gut, is very narrow in the Ascidia, as in the Amphioxus, and is here also usually confounded with the wide atrium, or peribranchial cavity, full of water.

There is no trace in the fully-developed Ascidia of a chorda dorsalis, or internal axial skeleton. It is the more interesting that the young animal that emerges from the ovum HAS a chorda, and that there is a rudimentary medullary tube above it. The latter is wholly atrophied in the developed Ascidia, and looks like a small nerve-ganglion in front above the gill-crate. It corresponds to the upper "gullet-ganglion" or "primitive brain" in other vermalia. Special sense-organs are either wanting altogether or are only found in a very rudimentary form, as simple optic spots and touch-corpuscles or tentacles that surround the mouth. The muscular system is very slightly and irregularly developed. Immediately under the thin corium, and closely connected with it, we find a thin muscle tube, as in the worms. On the other hand, the Ascidia has a centralised heart, and in this respect it seems to be more advanced than the Amphioxus. On the ventral side of the gut, some distance behind the gill-crate, there is a spindle-shaped heart. It retains permanently the simple tubular form that we find temporarily as the first structure of the heart in the vertebrates. This simple heart of the Ascidia has, however, a remarkable peculiarity. It contracts in alternate directions. In all other animals the beat of the heart is always in the same direction (generally from rear to front); it changes in the Ascidia to the reverse direction. The heart contracts first from the rear to the front, stands still for a minute, and then begins to beat the opposite way, now driving the blood from front to rear; the two large vessels that start from either end of the heart act alternately as arteries and veins. This feature is found in the Tunicates alone.

Of the other chief organs we have still to mention the sexual glands, which lie right behind in the body-cavity. All the Ascidiae are hermaphrodites. Each individual has a male and a female gland, and so is able to fertilise itself. The ripe ova (Figure 2.221 o apostrophe) fall directly from the ovary (o) into the mantle-cavity. The male sperm is conducted into this cavity from the testicle (t) by a special duct (vd). Fertilisation is accomplished here, and in many of the Ascidiae developed embryos are found. These are then ejected with the breathing-water through the cloaca (q), and so "born alive."

If we now glance at the entire structure of the simple Ascidia (especially Phallusia, Cynthia, etc.) and compare it with that of the Amphioxus, we shall find that the two have few points of contact. It is true that the fully-developed Ascidia resembles the Amphioxus in several important features of its internal structure, and especially in the peculiar character of the gill-crate and gut. But in most other features of organisation it is so far removed from it, and is so unlike it in external appearance, that the really close relationship of the two was not discovered until their embryology was studied. We will now compare the embryonic development of the two animals, and find to our great astonishment that the same embryonic form develops from the ovum of the Amphioxus as from that of the Ascidia—a typical chordula.

CHAPTER 2.17. EMBRYOLOGY OF THE LANCELET AND THE SEA-SQUIRT.

The structural features that distinguish the vertebrates from the invertebrates are so prominent that there was the greatest difficulty in the earlier stages of classification in determining the affinity of these two great groups. When scientists began to speak of the affinity of the various animal groups in more than a figurative—in a genealogical—sense, this question came at once to the front, and seemed to constitute one of the chief obstacles to the carrying-out of the evolutionary theory. Even earlier, when they had studied the relations of the chief groups, without any idea of real genealogical connection, they believed they had found here and there among the invertebrates points of contact with the vertebrates: some of the worms, especially, seemed to approach the vertebrates in structure, such as the marine arrow-worm (Sagitta). But on closer study the analogies proved untenable. When Darwin gave an impulse to the construction of a real stem-history of the animal kingdom by his reform of the theory of evolution, the solution of this problem was found to be particularly difficult. When I made the first attempt in my General Morphology (1866) to work out the theory and apply it to classification, I found no problem of phylogeny that gave me so much trouble as the linking of the vertebrates with the invertebrates.

But just at this time the true link was discovered, and at a point where it was least expected. Towards the end of 1866 two works of the Russian zoologist, Kowalevsky, who had lived for some time at Naples, and studied the embryology of the lower animals, were issued in the publications of the St. Petersburg Academy. A fortunate accident had directed the attention of this able observer almost simultaneously to the embryology of the lowest vertebrate, the Amphioxus, and that of an invertebrate, the close affinity of which to the Amphioxus had been least suspected, the Ascidia. To the extreme astonishment of all zoologists who were interested in this important question, there turned out to be the utmost resemblance in structure from the commencement of development between these two very different animals—the lowest vertebrate and the mis-shaped, sessile invertebrate. With this undeniable identity of ontogenesis, which can be demonstrated to an astounding extent, we had, in virtue of the biogenetic law, discovered the long-sought genealogical link, and definitely identified the invertebrate group that represents the nearest blood-relatives of the vertebrates. The discovery was confirmed by other zoologists, and there can no longer be any doubt that of all the classes of invertebrates that of the Tunicates is most closely related to the vertebrates, and of the Tunicates the nearest are the Ascidiae. We cannot say that the vertebrates are descended from the Ascidiae—and still less the reverse—but we can say that of all the invertebrates it is the Tunicates, and, within this group, the Ascidiae, that are the nearest blood-relatives of the ancient stem-form of the vertebrates. We must assume as the common ancestral group of both stems an extinct family of the extensive vermalia-stem, the Prochordonia or Prochordata ("primitive chorda-animals").