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time, it soon comes to lie obliquely. The upper end is more to the left, and the lower end more to the right. The foremost end draws up into the longer and narrower canal of the oesophagus. Underneath this on the left the blind sac (fundus) of the stomach bulges out, and thus the later form gradually develops (Figures 2.349 and 1.184 e). The original longitudinal axis becomes oblique, sinking below to the left and rising to the right, and approaches nearer and nearer to a transverse position. In the outer layer of the stomach-wall the powerful muscles that accomplish the digestive movements develop from the gut-fibre layer. In the inner layer a number of small glandular tubes are formed from the gut-gland layer; these are the peptic glands that secrete the gastric juice. At the lower end of the gastric sac is developed the valve that separates it from the duodenum (the pylorus, Figure 2.349 d).

Underneath the stomach there now develops the disproportionately long stretch of the small intestine. The development of this section is very simple, and consists essentially in an extremely rapid and considerable growth lengthways. It is at first very short, quite straight, and simple. But immediately behind the stomach we find at an early stage a horseshoe-shaped bend and loop of the gut, in connection with the severance of the alimentary canal from the yelk-sac and the development of the first mesentery. The thin delicate membrane that fastens this loop to the ventral side of the vertebral column, and fills the inner bend of the horseshoe formation, is the first rudiment of the mesentery (Figure 1.147 g). We find at an early stage a considerable growth of the small intestine; it is thus forced to coil itself in a number of loops. The various sections that we have to distinguish in it are differentiated in a very simple way--the duodenum (next to the stomach), the succeeding long jejunum, and the last section of the small intestine, the ileum.

From the duodenum are developed the two large glands that we have already mentioned--the liver and pancreas. The liver appears first in the shape of two small sacs, that are found to the right and left immediately behind the stomach (Figures 2.353 f, and 2.354 c). In many of the lower Vertebrates they remain separate for a long time (in the Myxinoides throughout life), or are only imperfectly joined. In the higher Vertebrates they soon blend more or less completely to form a single large organ. The growth of the liver is very brisk at first. In the human embryo it grows so much in the second month of development that in the third it occupies by far the greater part of the body-cavity (Figure 2.357). At first the two halves develop equally; afterwards the left falls far behind the right. In consequence of the unsymmetrical development and turning of the stomach and other abdominal viscera, the whole liver is now pushed to the right side. Although the liver does not afterwards grow so disproportionately, it is comparatively larger in the embryo at the end of pregnancy than in the adult. Its weight relatively to that of the whole body is 1 : 36 in the adult, and 1 : 18 in the embryo. Hence it is very important physiologically during embryonic life; it is chiefly concerned in the formation of blood, not so much in the secretion of bile.

Immediately behind the liver a second large visceral gland develops from the duodenum, the pancreas or sweetbread. It is wanting in most of the lowest classes of Vertebrates, and is first found in the fishes. This organ is also an outgrowth from the gut.

The last section of the alimentary canal, the large intestine, is at first in the embryo a very simple, short, and straight tube, which opens behind by the anus. It remains thus throughout life in the lower Vertebrates. But it grows considerably in the mammals, coils into various folds, and divides into two sections, the first and longer of which is the colon, and the second the rectum. At the beginning of the colon there is a valve (valvula Bauhini) that separates it from the small intestine. Immediately behind this there is a sac-like growth, which enlarges into the caecum (Figure 2.357 v). In the plant-eating mammals this is very large, but it is very small or completely atrophied in the flesh-eaters. In man, and most of the apes, only the first portion of the caecum is wide; the blind end-part of it is very narrow, and seems later to be merely a useless appendage of the former. This "vermiform appendage" is very interesting as a rudimentary organ. The only significance of it in man is that not infrequently a cherry-stone or some other hard and indigestible matter penetrates into its narrow cavity, and by setting up inflammation and suppuration causes the death of otherwise sound men. Teleology has great difficulty in giving a rational explanation of, and attributing to a beneficent Providence, this dreaded appendicitis. In our plant-eating ancestors this rudimentary organ was much larger and had a useful function.

Finally, we have important appendages of the alimentary tube in the bladder and urethra, which belong to the alimentary system. These urinary organs, acting as reservoir and duct for the urine excreted by the kidneys, originate from the innermost part of the allantoic pedicle. In the Dipneusts and Amphibia, in which the allantoic sac first makes its appearance, it remains within the body-cavity, and functions entirely as bladder. But in all the Amniotes it grows far outside of the body-cavity of the embryo, and forms the large embryonic "primitive bladder," from which the placenta develops in the higher mammals. This is lost at birth. But the long stalk or pedicle of the allantois remains, and forms with its upper part the middle vesico-umbilical ligament, a rudimentary organ that goes in the shape of a solid string from the vertex of the bladder to the navel. The lowest part of the allantoic pedicle (or the "urachus") remains hollow, and forms the bladder. At first this opens into the last section of the gut in man as in the lower Vertebrates; thus there is a real cloaca, which takes off both urine and excrements. But among the mammals this cloaca is only permanent in the Monotremes, as it is in all the birds, reptiles, and amphibia. In all the other mammals (marsupials and placentals) a transverse partition is afterwards formed, and this separates the urogenital aperture in front from the anus-opening behind. (Cf.

Chapters

2.22 and 2.29.)

CHAPTER XIII(28. EVOLUTION OF THE VASCULAR SYSTEM.)

 

The use that we have hitherto made of our biogenetic law will give the reader an idea how far we may trust its guidance in phylogenetic investigation. This differs considerably in the various systems of organs; the reason is that heredity and variability have a very different range in these systems. While some of them faithfully preserve the original palingenetic development inherited from earlier animal ancestors, others show little trace of this rigid heredity; they are rather disposed to follow new and divergent CENOGENETIC lines of development in consequence of adaptation. The organs of the first kind represent the CONSERVATIVE element in the multicellular state of the human frame, while the latter represent the PROGRESSIVE element. The course of historic development is a result of the correlation of the two tendencies, and they must be carefully distinguished.

There is perhaps no other system of organs in the human body in which this is more necessary than in that of which we are now going to consider the obscure development--the vascular system, or apparatus of circulation. If we were to draw our conclusions as to the original features in our earlier animal ancestors solely from the phenomena of the development of this system in the embryo of man and the other higher Vertebrates, we should be wholly misled. By a number of important embryonic adaptations, the chief of which is the formation of an extensive food-yelk, the original course of the development of the vascular system has been so much falsified and curtailed in the higher Vertebrates that little or nothing now remains in their embryology of some of the principal phylogenetic features. We should be quite unable to explain these if comparative anatomy and ontogeny did not come to our assistance.

The vascular system in man and all the Craniotes is an elaborate apparatus of cavities filled with juices or cell-containing fluids. These "vessels" (vascula) play an important part in the nutrition of the body. They partly conduct the nutritive red blood to the various parts of the body (blood-vessels); partly absorb from the gut the white chyle formed in digestion (chyle-vessels); and partly collect the used-up juices and convey them away from the tissues (lymphatic vessels). With the latter are connected the large cavities of the body, especially the body-cavity, or coeloma. The lymphatic vessels conduct both the colourless lymph and the white chyle into the venous part of the circulation. The lymphatic glands act as producers of new blood-cells, and with them is associated the spleen. The centre of movement for the circulation of the fluids is the heart, a strong muscular sac, which contracts regularly and is equipped with valves like a pump. This constant and steady circulation of the blood makes possible the complex metabolism of the higher animals.

But, however important the vascular system may be to the more advanced and larger and highly-differentiated animals, it is not at all so indispensable an element of animal life as is commonly supposed. The older science of medicine regarded the blood as the real source of life. Even in the still prevalent confused notions of heredity the blood plays the chief part. People speak generally of full blood, half blood, etc., and imagine that the hereditary transmission of certain characters "lies in the blood." The incorrectness of these ideas is clearly seen from the fact that in the act of generation the blood of the parents is not directly transmitted to the offspring, nor does the embryo possess blood in its early stages. We have already seen that not only the differentiation of the four secondary germinal layers, but also the first structures of the principal organs in the embryo of all the Vertebrates, take place long before there is any trace of the vascular system--the heart and the blood. In accordance with this ontogenetic fact, we must regard the vascular system as one of the latest organs from the phylogenetic point of view; just as we have found the alimentary canal to be one of the earliest. In any case, the vascular system is much later than the alimentary.

(FIGURE 2.358. Red blood-cells of various Vertebrates (equally magnified). 1. of man, 2. camel, 3. dove, 4. proteus, 5. water-salamander (Triton), 6. frog, 7. merlin (Cobitis), 8. lamprey (Petromyzon). a surface-view, b edge-view. (From Wagner.)

FIGURE 2.359. Vascular tissues or endothelium (vasalium). A capillary from the mesentery. a vascular cells, b their nuclei.)

The important nutritive fluid that circulates as blood and lymph in the elaborate canals of our vascular system is not a clear, simple fluid, but a very complex chemical juice with millions of cells floating in it. These blood-cells are just as important

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