The Evolution of Man, vol 2, Ernst Haeckel [online e reader txt] 📗

Vermalia (Gastrotricha, Figure 2.242, Nematodes, Sagitta, etc.). But in these a second important opening of the gut has been formed at the opposite end to the mouth, the anus (Figure 2.242 a).

We see a great advance in the structure of the vermalian gut in the remarkable Balanoglossus (Figure 2.245), the sole survivor of the Enteropneust class. Here we have the first appearance of the division of the alimentary tube into two sections that characterises the Chordonia. The fore half, the head-gut (cephalogaster), becomes the organ of respiration (branchial gut, Figure 2.245 k); the hind half, the trunk-gut (truncogaster), alone acts as digestive organ (hepatic gut, d). The differentiation of these two parts of the gut in the Enteropneust is just the same as in all the Tunicates and Vertebrates.

It is particularly interesting and instructive in this connection to compare the Enteropneusts with the Ascidia and the Amphioxus (Figures 2.220 and 2.210)—the remarkable animals that form the connecting link between the Invertebrates and the Vertebrates. In both forms the gut is of substantially the same construction; the anterior section forms the respiratory branchial gut, the posterior the digestive hepatic gut. In both it develops palingenetically from the primitive gut of the gastrula, and in both the hinder end of the medullary tube covers the primitive mouth to such an extent that the remarkable medullary intestinal duct is formed, the passing communication between the neural and intestinal tubes (canalis neurentericus, Figures 1.83 and 1.85 ne). In the vicinity of the closed primitive mouth, possibly in its place, the later anus is developed. In the same way the mouth is a fresh formation in the Amphioxus and the Ascidia. It is the same with the human mouth and that of the Craniotes generally. The secondary formation of the mouth in the Chordonia is probably connected with the development of the gill-clefts which are formed in the gut-wall immediately behind the mouth. In this way the anterior section of the gut is converted into a respiratory organ. I have already pointed out that this modification is distinctive of the Vertebrates and Tunicates. The phylogenetic appearance of the gill-clefts indicates the commencement of a new epoch in the stem-history of the Vertebrates.

In the further ontogenetic development of the alimentary canal in the human embryo the appearance of the gill-clefts is the most important process. At a very early stage the gullet-wall joins with the external body-wall in the head of the human embryo, and this is followed by the formation of four clefts, which lead directly into the gullet from without, on the right and left sides of the neck, behind the mouth. These are the gill or gullet clefts, and the partitions that separate them are the gill or gullet-arches (Figure 1.171). These are most interesting embryonic structures. They show us that all the higher Vertebrates reproduce in their earlier stages, in harmony with the biogenetic law, the process that had so important a part in the rise of the whole Chordonia-stem. This process was the differentiation of the gut into two sections—an anterior respiratory section, the branchial gut, that was restricted to breathing, and a posterior digestive section, the hepatic gut. As we find this highly characteristic differentiation of the gut into two different sections in all the Vertebrates and all the Tunicates, we may conclude that it was also found in their common ancestors, the Prochordonia—especially as even the Enteropneusts have it. (Cf. Chapters 1.12, 1.14 and 2.20, and Figures 2.210, 2.220, 2.245.) It is entirely wanting in all the other Invertebrates.

(FIGURE 2.353. Gut of a dog-embryo (shown in Figure 1.202, from Bischoff), seen from the ventral side, a gill-arches (four pairs), b rudiments of pharynx and larynx, c lungs, d stomach, f liver, g walls of the open yelk-sac (into which the middle gut opens with a wide aperture), h rectum.

FIGURE 2.354. The same gut seen from the right. a lungs, b stomach, c liver, d yelk-sac, e rectum.)

There is at first only one pair of gill-clefts in the Amphioxus, as in the Ascidia and Enteropneusts; and the Copelata (Figure 2.225) have only one pair throughout life. But the number presently increases in the former. In the Craniotes, however, it decreases still further. The Cyclostomes have six to eight pairs (Figure 2.247); some of the Selachii six or seven pairs, most of the fishes only four or five pairs. In the embryo of man, and the higher Vertebrates generally, where they make an appearance at an early stage, only three or four pairs are developed. In the fishes they remain throughout life, and form an exit for the water taken in at the mouth (Figures 2.249 to 2.251). But they are partly lost in the amphibia, and entirely in the higher Vertebrates. In these nothing is left but a relic of the first gill-cleft. This is formed into a part of the organ of hearing; from it are developed the external meatus, the tympanic cavity, and the Eustachian tube. We have already considered these remarkable structures, and need only point here to the interesting fact that our middle and external ear is a modified inheritance from the fishes. The branchial arches also, which separate the clefts, develop into very different parts. In the fishes they remain gill-arches, supporting the respiratory gill-leaves. It is the same with the lowest amphibia, but in the higher amphibia they undergo various modifications; and in the three higher classes of Vertebrates (including man) the hyoid bone and the ossicles of the ear develop from them. (Cf. Chapter 2.25.)

(FIGURE 2.355. Median section of the head of a Petromyzon-larva. (From Gegenbaur,) h hypobranchial groove (above it in the gullet we see the internal openings of the seven gill-clefts), v velum, o mouth, c heart, a auditory vesicle, n neural tube, ch chorda.)

From the first gill-arch, from the inner surface of which the muscular tongue proceeds, we get the first structure of the maxillary skeleton—the upper and lower jaws, which surround the mouth and support the teeth. These important parts are wholly wanting in the two lowest classes of Vertebrates, the Acrania and Cyclostoma. They appear first in the earliest Selachii (Figures 2.248 to 2.251), and have been transmitted from this stem-group of the Gnathostomes to the higher Vertebrates. Hence the original formation of the skeleton of the mouth can be traced to these primitive fishes, from which we have inherited it. The teeth are developed from the skin that clothes the jaws. As the whole mouth cavity originates from the outer integument (Figure 2.350), the teeth also must come from it. As a fact, this is found to be the case on microscopic examination of the development and finer structure of the teeth. The scales of the fishes, especially of the shark type (Figure 2.351), are in the same position as their teeth in this respect (Figure 2.252). The osseous matter of the tooth (dentine) develops from the corium; its enamel covering is a secretion of the epidermis that covers the corium. It is the same with the cutaneous teeth or placoid scales of the Selachii. At first the whole of the mouth was armed with these cutaneous teeth in the Selachii and in the earliest amphibia. Afterwards the formation of them was restricted to the edges of the jaws.

Hence our human teeth are, in relation to their original source, modified fish-scales. For the same reason we must regard the salivary glands, which open into the mouth, as epidermic glands, as they are formed, not from the glandular layer of the gut like the rest of the alimentary glands, but from the epidermis, from the horny plate of the outer germinal layer. Naturally, in harmony with this evolution of the mouth, the salivary glands belong genetically to one series with the sudoriferous, sebaceous, and mammary glands.

Thus the human alimentary canal is as simple as the primitive gut of the gastrula in its original structure. Later it resembles the gut of the earliest Vermalia (Gastrotricha). It then divides into two sections, a fore or branchial gut and a hind or hepatic gut, like the alimentary canal of the Balanoglossus, the Ascidia, and the Amphioxus. The formation of the jaws and the branchial arches changes it into a real fish-gut (Selachii). But the branchial gut, the one reminiscence of our fish-ancestors, is afterwards atrophied as such. The parts of it that remain are converted into entirely different structures.

(FIGURE 2.356. Transverse section of the head of a Petromyzon-larva. (From Gegenbaur.) Beneath the pharynx (d) we see the hypobranchial groove; above it the chorda and neural tube. A, B, C stages of constriction.)

But, although the anterior section of our alimentary canal thus entirely loses its original character of branchial gut, it retains the physiological character of respiratory gut. We are now astonished to find that the permanent respiratory organ of the higher Vertebrates, the air-breathing lung, is developed from this first part of the alimentary canal. Our lungs, trachea, and larynx are formed from the ventral wall of the branchial gut. The whole of the respiratory apparatus, which occupies the greater part of the pectoral cavity in the adult man, is at first merely a small pair of vesicles or sacs, which grow out of the floor of the head-gut immediately behind the gills (Figures 2.354 C, 1.147 l). These vesicles are found in all the Vertebrates except the two lowest classes, the Acrania and Cyclostomes. In the lower Vertebrates they do not develop into lungs, but into a large air-filled bladder, which occupies a good deal of the body-cavity and has a quite different purport. It serves, not for breathing, but to effect swimming movements up and down, and so is a sort of hydrostatic apparatus—the floating bladder of the fishes (nectocystis, Chapter 2.21). However, the human lungs, and those of all air-breathing Vertebrates, develop from the same simple vesicular appendage of the head-gut that becomes the floating bladder in the fishes.

At first this bladder has no respiratory function, but merely acts as hydrostatic apparatus for the purpose of increasing or lessening the specific gravity of the body. The fishes, which have a fully-developed floating bladder, can press it together, and thus condense the air it contains. The air also escapes sometimes from the alimentary canal, through an air-duct that connects the floating bladder with the pharynx, and is ejected by the mouth. This lessens the size of the bladder, and so the fish becomes heavier and sinks. When it wishes to rise again, the bladder is expanded by relaxing the pressure. In many of the Crossopterygii the wall of the bladder is covered with bony plates, as in the Triassic Undina (Figure 2.254).

This hydrostatic apparatus begins in the Dipneusts to change into a respiratory organ; the blood-vessels in the wall of the bladder now no longer merely secrete air themselves, but also take in fresh air through the air-duct. This process reaches its full development in the Amphibia. In these the floating bladder has turned into lungs, and the air-passage into a trachea. The lungs of the Amphibia have been transmitted to the three higher classes of Vertebrates. In the lowest Amphibia the lungs on either side are still very simple transparent sacs with thin walls, as in the common water-salamander, the Triton. It still entirely resembles the floating bladder of the fishes. It is true that the Amphibia have two lungs, right and left. But the floating bladder is also double in many of the fishes (such as the early Ganoids), and divides into right and left halves. On the other hand, the lung is single in Ceratodus (Figure 2.257).

(FIGURE 2.357. Thoracic and abdominal viscera of a human embryo of twelve weeks, natural size, (From Kolliker.) The head is omitted. Ventral and pectoral walls are removed. The greater part of the body-cavity is taken up with the liver, from the middle part of which the caecum and the vermiform appendix protrude. Above the diaphragm,