The Evolution of Man, V.2 - Ernst Haeckel (little bear else holmelund minarik TXT) 📗

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, in the middle, is the conical heart; to the right and left of it are the two small lungs.)

In the human embryo and that of all the other Amniotes the lungs develop from the hind part of the ventral wall of the head-gut (Figure 1.149). Immediately behind the single structure of the thyroid gland a median groove, the rudiment of the trachea, is detached from the gullet. From its hinder end a couple of vesicles develop--the simple tubular rudiments of the right and left lungs. They afterwards increase considerably in size, fill the greater part of the thoracic cavity, and take the heart between them. Even in the frogs we find that the simple sac has developed into a spongy body of peculiar froth-like tissue. The originally short connection of the pulmonary sacs with the head-gut extends into a long, thin tube. This is the wind-pipe (trachea); it opens into the gullet above, and divides below into two branches which go to the two lungs. In the wall of the trachea circular cartilages develop, and these keep it open. At its upper end, underneath its pharyngeal opening, the larynx is formed--the organ of voice and speech. The larynx is found at various stages of development in the Amphibia, and comparative anatomists are in a position to trace the progressive growth of this important organ from the rudimentary structure of the lower Amphibia up to the elaborate and delicate vocal apparatus that we have in the larynx of man and of the birds.

We must refer here to an interesting rudimentary organ of the respiratory gut, the thyroid gland, the large gland in front of the larynx, that lies below the "Adam's apple," and is often especially developed in the male sex. It has a certain function--not yet fully understood--in the nutrition of the body, and arises in the embryo by constriction from the lower wall of the pharynx. In many mining districts the thyroid gland is peculiarly liable to morbid enlargement, and then forms goitre, a growth that hangs at the front of the neck. But it is much more interesting phylogenetically. As Wilhelm Muller, of Jena, has shown, this rudimentary organ is the last relic of the hypobranchial groove, which we considered in a previous chapter, and which runs in the middle line of the gill-crate in the Ascidia and Amphioxus, and conveys food to the stomach. (Cf.

Chapter 2.

16, Figure 2.246). We still find it in its original character in the larvae of the Cyclostomes (Figures 2.355 and 2.356).

The second section of the alimentary canal, the trunk or hepatic gut, undergoes not less important modifications among our vertebrate ancestors than the first section. In tracing the further development of this digestive part of the gut, we find that most complex and elaborate organs originate from a very rudimentary original structure. For clearness we may divide the digestive gut into three sections: the fore gut (with oesophagus and stomach), the middle gut (duodenum, with liver, pancreas, jejunum, and ileum, and the hind gut (colon and rectum). Here again we find vesicular growths or appendages of the originally simple gut developing into a variety of organs. Two of these embryonic structures, the yelk-sac and allantois, are already known to us. The two large glands that open into the duodenum, the liver and pancreas, are growths from the middle and most important part of the trunk-gut.

Immediately behind the vesicular rudiments of the lungs comes the section of the alimentary canal that forms the stomach (Figures 2.353 d and 2.354 b). This sac-shaped organ, which is chiefly responsible for the solution and digestion of the food, has not in the lower Vertebrates the great physiological importance and the complex character that it has in the higher. In the Acrania and Cyclostomes and the earlier fishes we can scarcely distinguish a real stomach; it is represented merely by the short piece from the branchial to the hepatic gut. In some of the other fishes also the stomach is only a very simple spindle-shaped enlargement at the beginning of the digestive section of the gut, running straight from front to back in the median plane of the body, underneath the vertebral column. In the mammals its first structure is just as rudimentary as it is permanently in the preceding. But its various parts soon begin to develop. As the left side of the spindle-shaped sac grows much more quickly than the right, and as it turns considerably on its axis at the same