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

stomach, is a large gland, rich in blood, in the adult man, immediately under the diaphragm on the left side, and separated by it from the lungs. The pancreas lies a little further back and more to the left. The remaining part of the small intestine is so long that it has to coil itself in many folds in order to find room in the narrow space of the abdominal cavity. It is divided into the jejunum above and the ileum below. In the last section of it is the part of the small intestine at which in the embryo the yelk-sac opens into the gut. This long and thin intestine then passes into the large intestine, from which it is cut off by a special valve. Immediately behind this "Bauhin-valve" the first part of the large intestine forms a wide, pouch-like structure, the caecum. The atrophied end of the caecum is the famous rudimentary organ, the vermiform appendix. The large intestine (colon) consists of three parts--an ascending part on the right, a transverse middle part, and a descending part on the left. The latter finally passes through an S-shaped bend into the last section of the alimentary canal, the rectum, which opens behind by the anus. Both the large and small intestines are equipped with numbers of small glands, which secrete mucous and other fluids.

For the greater part of its length the alimentary canal is attached to the inner dorsal surface of the abdominal cavity, or to the lower surface of the vertebral column. The fixing is accomplished by means of the thin membranous plate that we call the mesentery.

Although the fully-formed alimentary canal is thus a very elaborate organ, and although in detail it has a quantity of complex structural features into which we cannot enter here, nevertheless the whole complicated structure has been historically evolved from the very simple form of the primitive gut that we find in our gastraead-ancestors, and that every gastrula brings before us to-day. We have already pointed out (

Chapter 1.

9) how the epigastrula of the mammals (Figure 1.67) can be reduced to the original type of the bell-gastrula, which is now preserved by the amphioxus alone (Figure 1.35). Like the latter, the human gastrula and that of all other mammals must be regarded as the ontogenetic reproduction of the phylogenetic form that we call the Gastraea, in which the whole body is nothing but a double-walled gastric sac.

We already know from embryology the manner in which the gut develops in the embryo of man and the other mammals. From the gastrula is first formed the spherical embryonic vesicle filled with fluid (gastrocystis, Figure 1.106). In the dorsal wall of this the sole-shaped embryonic shield is developed, and on the under-side of this a shallow groove appears in the middle line, the first trace of the later, secondary alimentary tube. The gut-groove becomes deeper and deeper, and its edges bend towards each other, and finally form a tube.

As we have seen, this simple cylindrical gut-tube is at first completely closed before and behind in man and in the Vertebrates generally (Figure 1.148); the permanent openings of the alimentary canal, the mouth and anus, are only formed later on, and from the outer skin. A mouth-pit appears in the skin in front (Figure 2.350 hp), and this grows towards the blind fore-end of the cavity of the head-gut (kd), and at length breaks into it. In the same way a shallow anus-pit is formed in the skin behind, which grows deeper and deeper, advances towards the blind hinder end of the pelvic gut, and at last connects with it. There is at first, both before and behind, a thin partition between the external cutaneous pit and the blind end of the gut--the throat-membrane in front and the anus-membrane behind; these disappear when the connection takes place.

Directly in front of the anus-opening the allantois develops from the hind gut; this is the important embryonic structure that forms into the placenta in the Placentals (including man). In this more advanced form the human alimentary canal (and that of all the other mammals) is a slightly bent, cylindrical tube, with an opening at each end, and two appendages growing from its lower wall: the anterior one is the umbilical vesicle or yelk-sac, and the posterior the allantois or urinary sac (Figure 1.195).

The thin wall of this simple alimentary tube and its ventral appendages is found, on microscopic examination, to consist of two strata of cells. The inner stratum, lining the entire cavity, consists of larger and darker cells, and is the gut-gland layer. The outer stratum consists of smaller and lighter cells, and is the gut-fibre layer. The only exception is in the cavities of the mouth and anus, because these originate from the skin. The inner coat of the mouth-cavity is not provided by the gut-gland layer, but by the skin-sense layer; and its muscular substratum is provided, not by the gut-fibre, but the skin-fibre, layer. It is the same with the wall of the small anus-cavity.

If it is asked how these constituent layers of the primitive gut-wall are related to the various tissues and organs that we find afterwards in the fully-developed system, the answer is very simple. It can be put in a single sentence. The epithelium of the gut--that is to say, the internal soft stratum of cells that lines the cavity of the alimentary canal and all its appendages, and is immediately occupied with the processes of nutrition--is formed solely from the gut-gland layer; all other tissues and organs that belong to the alimentary canal and its appendages originate from the gut-fibre layer. From the latter is also developed the whole of the outer envelope of the gut and its appendages; the fibrous connective tissue and the smooth muscles that compose its muscular layer, the cartilages that support it (such as the cartilages of the larynx and the trachea), the blood-vessels and lymph-vessels that absorb the nutritive fluid from the intestines--in a word, all that there is in the alimentary system besides the epithelium of the gut. From the same layer we also get the whole of the mesentery, with all the organs embedded in it--the heart, the large blood-vessels of the body, etc.

(FIGURE 2.351. Scales or cutaneous teeth of a shark (Centrophorus calceus). A three-pointed tooth rises obliquely on each of the quadrangular bony plates that lie in the corium. (From Gegenbaur.))

Let us now leave this original structure of the mammal gut for a moment, in order to compare it with the alimentary canal of the lower Vertebrates, and of those Invertebrates that we have recognised as man's ancestors. We find, first of all, in the lowest Metazoa, the Gastraeads, that the gut remains permanently in the very simple form in which we find it transitorily in the palingenetic gastrula of the other animals; it is thus in the Gastremaria (Pemmatodiscus), the Physemaria (Prophysema), the simplest Sponges (Olynthus), the freshwater Polyps (Hydra), and the ascula-embryos of many other Coelenteria (Figures 2.233 to 2.238). Even in the simplest forms of the Platodes, the Rhabdocoela (Figure 2.240), the gut is still a simple straight tube, lined with the entoderm; but with the important difference that in this case its single opening, the primitive mouth (m), has formed a muscular gullet (sd) by invagination of the skin.

(FIGURE 2.352. Gut of a human embryo, one-sixth of an inch long, magnified fifteen times. (From His. Showing: Epiglottis, Tongue, Hypophysis, Hepatic duct, Tail, Allantoic duct, Tail-gut, Umbilical cord, Larynx, Rudimentary lungs, Stomach, Pancreas, Bladder, Wolffian duct, Rudimentary kidneys.))

We have the same simple form in the gut of the lowest 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