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

the tympanic cavity of the ear (Figure 2.320 e). The pharynx is continued in a long, narrow tube, the oesophagus (sr). By this the food passes into the stomach when masticated and swallowed. Into the gullet also opens, right above, the trachea (lr), that leads to the lungs. The entrance to it is covered by the epiglottis, over which the food slides. The cartilaginous epiglottis is found only in the mammals, and has developed from the fourth branchial arch of the fishes and amphibia. The lungs are found, in man and all the mammals, to the right and left in the pectoral cavity, with the heart between them. At the upper end of the trachea there is, under the epiglottis, a specially differentiated part, strengthened by a cartilaginous skeleton, the larynx. This important organ of human speech also develops from a part of the alimentary canal. In front of the larynx is the thyroid gland, which sometimes enlarges and forms goitre.

The oesophagus descends into the pectoral cavity along the vertebral column, behind the lungs and the heart, pierces the diaphragm, and enters the visceral cavity. The diaphragm is a membrano-muscular partition that completely separates the thoracic from the abdominal cavity in all the mammals (and these alone). This separation is not found in the beginning; there is at first a common breast-belly cavity, the coeloma or pleuro-peritoneal cavity. The diaphragm is formed later on as a muscular horizontal partition between the thoracic and abdominal cavities. It then completely separates the two cavities, and is only pierced by several organs that pass from the one to the other. One of the chief of these organs is the oesophagus. After this has passed through the diaphragm, it expands into the gastric sac in which digestion chiefly takes place. The stomach of the adult man (Figure 2.349) is a long, somewhat oblique sac, expanding on the left into a blind sac, the fundus of the stomach (b apostrophe), but narrowing on the right, and passing at the pylorus (e) into the small intestine. At this point there is a valve, the pyloric valve (d), between the two sections of the canal; it opens only when the pulpy food passes from the stomach into the intestine. In man and the higher Vertebrates the stomach itself is the chief organ of digestion, and is especially occupied with the solution of the food; this is not the case in many of the lower Vertebrates, which have no stomach, and discharge its function by a part of the gut farther on. The muscular wall of the stomach is comparatively thick; it has externally strong muscles that accomplish the digestive movements, and internally a large quantity of small glands, the peptic glands, which secrete the gastric juice.

(FIGURE 2.349. Human stomach and duodenum, longitudinal section. a cardiac (end of oesophagus), b fundus (blind sac of the left side), c pylorus-fold, d pylorus-valves, e pylorus-cavity, fgh duodenum, i entrance of the gall-duct and the pancreatic duct. (From Meyer.)

FIGURE 2.350. Median section of the head of a hare-embryo, one-fourth of an inch in length. (From Mihalcovics.) The deep mouth-cleft (hp) is separated by the membrane of the throat (rh) from the blind cavity of the head-gut (kd). hz heart, ch chorda, hp the point at which the hypophysis develops from the mouth-cleft, vh ventricle of the cerebrum, v3, third ventricle (intermediate brain), v4 fourth ventricle (hind brain), ck spinal canal.)

Next to the stomach comes the longest section of the alimentary canal, the middle gut or small intestine. Its chief function is to absorb the peptonised fluid mass of food, or the chyle, and it is subdivided into several sections, of which the first (next to the stomach) is called the duodenum (Figure 2.349 fgh). It is a short, horseshoe-shaped loop of the gut. The largest glands of the alimentary canal open into it—the liver, the chief digestive gland, that secretes the gall, and the pancreas, which secretes the pancreatic juice. The two glands pour their secretions, the bile and pancreatic juice, close together into the duodenum (i). The opening of the gall-duct is of particular phylogenetic importance, as it is the same in all the Vertebrates, and indicates the principal point of the hepatic or trunk-gut (Gegenbaur). The liver, phylogenetically older than the 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 underside 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