The Evolution of Man, vol 2 by Ernst Haeckel (fun books to read for adults TXT) 📕

the Vertebrates, and that proves irrefragably the affinity of the Ascidiae to the Vertebrates. From the epidermis of the gastrula a medullary tube is formed on the dorsal side, and, between this and the primitive gut, a chorda; these are the organs that are otherwise only found in Vertebrates. The formation of these very important organs takes place in the Ascidia-gastrula in precisely the same way as in that of the Amphioxus. In the Ascidia (as in the other case) the oval gastrula is first flattened on one side—the subsequent dorsal side. A groove or furrow (the medullary groove) is sunk in the middle line of the flat surface, and two parallel longitudinal swellings arise on either side from the skin layer. These medullary swellings join together over the furrow, and form a tube; in this case, again, the neural or medullary tube is at first open in front, and connected with the primitive gut behind by the neurenteric canal. Further, in the Ascidia-larva also the two permanent apertures of the alimentary canal only appear later, as independent and new formations. The permanent mouth does not develop from the primitive mouth of the gastrula; this primitive mouth closes up, and the later anus is formed near it by invagination from without, on the hinder end of the body, opposite to the aperture of the medullary tube.

During these important processes, that take place in just the same way in the Amphioxus, a tail-like projection grows out of the posterior end of the larva-body, and the larva folds itself up within the round ovolemma in such a way that the dorsal side is curved and the tail is forced on to the ventral side. In this tail is developed—starting from the primitive gut—a cylindrical string of cells, the fore end of which pushes into the body of the larva, between the alimentary canal and the neural canal, and is no other than the chorda dorsalis. This important organ had hitherto been found only in the Vertebrates, not a single trace of it being discoverable in the Invertebrates. At first the chorda only consists of a single row of large entodermic cells. It is afterwards composed of several rows of cells. In the Ascidia-larva, also, the chorda develops from the dorsal middle part of the primitive gut, while the two coelom-pouches detach themselves from it on both sides. The simple body-cavity is formed by the coalescence of the two.

When the Ascidia-larva has attained this stage of development it begins to move about in the ovolemma. This causes the membrane to burst. The larva emerges from it, and swims about in the sea by means of its oar-like tail. These free-swimming larvae of the Ascidia have been known for a long time. They were first observed by Darwin during his voyage round the world in 1833. They resemble tadpoles in outward appearance, and use their tails as oars, as the tadpoles do. However, this lively and highly-developed condition does not last long. At first there is a progressive development; the foremost part of the medullary tube enlarges into a brain, and inside this two single sense-organs are developed, a dorsal auditory vesicle and a ventral eye. Then a heart is formed on the ventral side of the animal, or the lower wall of the gut, in the same simple form and at the same spot at which the heart is developed in man and all the other vertebrates. In the lower muscular wall of the gut we find a weal-like thickening, a solid, spindle-shaped string of cells, which becomes hollow in the centre; it begins to contract in different directions, now forward and now backward, as is the case with the adult Ascidia. In this way the sanguineous fluid accumulated in the hollow muscular tube is driven in alternate directions into the blood-vessels, which develop at both ends of the cardiac tube. One principal vessel runs along the dorsal side of the gut, another along its ventral side. The former corresponds to the aorta and the dorsal vessel in the worms. The other corresponds to the subintestinal vein and the ventral vessel of the worms.

With the formation of these organs the progressive development of the Ascidia comes to an end, and degeneration sets in. The free-swimming larva sinks to the floor of the sea, abandons its locomotive habits, and attaches itself to stones, marine plants, mussel-shells, corals, and other objects; this is done with the part of the body that was foremost in movement. The attachment is effected by a number of outgrowths, usually three, which can be seen even in the free-swimming larva. The tail is lost, as there is no further use for it. It undergoes a fatty degeneration, and disappears with the chorda dorsalis. The tailless body changes into an unshapely tube, and, by the atrophy of some parts and the modification of others, gradually assumes the appearance we have already described.

(FIGURE 2.225. An Appendicaria (Copelata), seen from the left. m mouth, k branchial gut, o gullet, v stomach, a anus, n brain (ganglion above the gullet), g auditory vesicle, f ciliated groove under the gills, h heart, t testicles, e ovary, c chorda, s tail.)

Among the living Tunicates there is a very interesting group of small animals that remain throughout life at the stage of development of the tailed, free Ascidia-larva, and swim about briskly in the sea by means of their broad oar-tail. These are the remarkable Copelata (Appendicaria and Vexillaria, Figure 2.225). They are the only living Vertebrates that have throughout life a chorda dorsalis and a neural string above it; the latter must be regarded as the prolongation of the cerebral ganglion and the equivalent of the medullary tube. Their branchial gut also opens directly outwards by a pair of branchial clefts. These instructive Copelata, comparable to permanent Ascidia-larvae, come next to the extinct Prochordonia, those ancient worms which we must regard as the common ancestors of the Tunicates and Vertebrates. The chorda of the Appendicaria is a long, cylindrical string (Figure 2.225 c), and serves as an attachment for the muscles that work the flat oar-tail.

Among the various modifications which the Ascidia-larva undergoes after its establishment at the sea-floor, the most interesting (after the loss of the axial rod) is the atrophy of one of its chief organs, the medullary tube. In the Amphioxus the spinal marrow continues to develop, but in the Ascidia the tube soon shrinks into a small and insignificant nervous ganglion that lies above the mouth and the gill-crate, and is in accord with the extremely slight mental power of the animal. This insignificant relic of the medullary tube seems to be quite beyond comparison with the nervous centre of the vertebrate, yet it started from the same structure as the spinal cord of the Amphioxus. The sense-organs that had been developed in the fore part of the neural tube are also lost; no trace of which can be found in the adult Ascidia. On the other hand, the alimentary canal becomes a most extensive organ. It divides presently into two sections—a wide fore or branchial gut that serves for respiration, and a narrower hind or hepatic gut that accomplishes digestion. The branchial or head-gut of the Ascidia is small at first, and opens directly outwards only by a couple of lateral ducts or gill-clefts—a permanent arrangement in the Copelata. The gill-clefts are developed in the same way as in the Amphioxus. As their number greatly increases we get a large gill-crate, pierced like lattice work. In the middle line of its ventral side we find the hypobranchial groove. The mantle or cloaca-cavity (the atrium) that surrounds the gill-crate is also formed in the same way in the Ascidia as in the Amphioxus. The ejection-opening of this peribranchial cavity corresponds to the branchial pore of the Amphioxus. In the adult Ascidia the branchial gut and the heart on its ventral side are almost the only organs that recall the original affinity with the vertebrates.

The further development of the Ascidia in detail has no particular interest for us, and we will not go into it. The chief result that we obtain from its embryology is the complete agreement with that of the Amphioxus in the earliest and most important embryonic stages. They do not begin to diverge until after the medullary tube and alimentary canal, and the axial rod with the muscles between the two, have been formed. The Amphioxus continues to advance, and resembles the embryonic forms of the higher vertebrates; the Ascidia degenerates more and more, and at last, in its adult condition, has the appearance of a very imperfect invertebrate.

If we now look back on all the remarkable features we have encountered in the structure and the embryonic development of the Amphioxus and the Ascidia, and compare them with the features of man’s embryonic development which we have previously studied, it will be clear that I have not exaggerated the importance of these very interesting animals. It is evident that the Amphioxus from the vertebrate side and the Ascidia from the invertebrate form the bridge by which we can span the deep gulf that separates the two great divisions of the animal kingdom. The radical agreement of the lancelet and the sea-squirt in the first and most important stages of development shows something more than their close anatomic affinity and their proximity in classification; it shows also their real blood-relationship and their common origin from one and the same stem-form. In this way, it throws considerable light on the oldest roots of man’s genealogical tree.

 

CHAPTER 2.18. DURATION OF THE HISTORY OF OUR STEM.

Our comparative investigation of the anatomy and ontogeny of the Amphioxus and Ascidia has given us invaluable assistance. We have, in the first place, bridged the wide gulf that has existed up to the present between the Vertebrates and Invertebrates; and, in the second place, we have discovered in the embryology of the Amphioxus a number of ancient evolutionary stages that have long since disappeared from human embryology, and have been lost, in virtue of the law of curtailed heredity. The chief of these stages are the spherical blastula (in its simplest primary form), and the succeeding archigastrula, the pure, original form of the gastrula which the Amphioxus has preserved to this day, and which we find in the same form in a number of Invertebrates of various classes. Not less important are the later embryonic forms of the coelomula, the chordula, etc.

Thus the embryology of the Amphioxus and the Ascidia has so much increased our knowledge of man’s stem-history that, although our empirical information is still very incomplete, there is now no defect of any great consequence in it. We may now, therefore, approach our proper task, and reconstruct the phylogeny of man in its chief lines with the aid of this evidence of comparative anatomy and ontogeny. In this the reader will soon see the immense importance of the direct application of the biogenetic law. But before we enter upon the work it will be useful to make a few general observations that are necessary to understand the processes aright.

We must say a few words with regard to the period in which the human race was evolved from the animal kingdom. The first thought that occurs to one in this connection is the vast difference between the duration of man’s ontogeny and phylogeny. The individual man needs only nine months for his complete development, from the fecundation of the ovum to the moment when he leaves the maternal womb. The human embryo runs its whole course in the brief space of forty weeks (as a rule, 280 days). In many other mammals the time of the embryonic development is much the same as in man—for instance, in the cow. In the horse and ass it takes a little longer, forty-three to forty-five weeks; in the camel, thirteen months. In the largest mammals,