The Evolution of Man Volume Ii Part 16

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 s.e.x. 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 ca.n.a.l, 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 r.e.c.t.u.m). 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 ca.n.a.l 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 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 ca.n.a.l 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 ca.n.a.l 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 cla.s.ses of Vertebrates, and is first found in the fishes. This organ is also an outgrowth from the gut.

The last section of the alimentary ca.n.a.l, the large intestine, is at first in the embryo a very simple, short, and straight tube, which opens behind by the a.n.u.s. 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 r.e.c.t.u.m. 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 caec.u.m (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 caec.u.m 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 part.i.tion is afterwards formed, and this separates the urogenital aperture in front from the a.n.u.s-opening behind. (Cf. Chapters 2.22 and 2.29.)

CHAPTER 2.28. 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 princ.i.p.al phylogenetic features. We should be quite unable to explain these if comparative anatomy and ontogeny did not come to our a.s.sistance.

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 a.s.sociated 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 princ.i.p.al 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 ca.n.a.l 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 ca.n.a.ls 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 in the complicated life of the higher animal body as the circulation of money is to the commerce of a civilised community. Just as the citizens meet their needs most conveniently by means of a financial circulation, so the various tissue-cells, the microscopic citizens of the multicellular human body, have their food conveyed to them best by the circulating cells in the blood. These blood cells (haemocytes) are of two kinds in man and all the other Craniotes--red cells (rhodocytes or erythrocytes) and colourless or lymph cells (leucocytes). The red colour of the blood is caused by the great acc.u.mulation of the former, the others circulate among them in much smaller quant.i.ty. When the colourless cells increase at the expense of the red we get anaemia (or chlorosis).

(FIGURE 2.360. Transverse section of the trunk of a chick-embryo, forty-five hours old. (From Balfour.) A ectoderm (h.o.r.n.y-plate), Mc medullary tube, ch chorda, C entoderm (gut-gland layer), Pv primitive segment (episomite), Wd prorenal duct, pp coeloma (secondary body-cavity). So skin-fibre layer, Sp gut-fibre layer, v blood-vessels in latter, ao primitive aortas, containing red blood-cells.)

The lymph-cells (leucocytes), commonly called the "white corpuscles"

of the blood, are phylogenetically older and more widely distributed in the animal world than the red. The great majority of the Invertebrates that have acquired an independent vascular system have only colourless lymph-cells in the circulating fluid. There is an exception in the Nemertines (Figure 2.358) and some groups of Annelids. When we examine the colourless blood of a cray-fish or a snail (Figure 2.358) under a high power of the microscope, we find in each drop numbers of mobile leucocytes, which behave just like independent Amoebae (Figure 1.17). Like these unicellular Protozoa, the colourless blood-cells creep slowly about, their unshapely plasma-body constantly changing its form, and stretching out finger-like processes first in one direction, then another. Like the Amoebae, they take particles into their cell-body. On account of this feature these amoeboid plastids are called "eating cells"

(phagocytes), and on account of their motions "travelling cells"

(planocytes). It has been shown by the discoveries of the last few decades that these leucocytes are of the greatest physiological and pathological consequence to the organism. They can absorb either solid or dissolved particles from the wall of the gut, and convey them to the blood in the chyle; they can absorb and remove unusable matter from the tissues. When they pa.s.s in large quant.i.ties through the fine pores of the capillaries and acc.u.mulate at irritated spots, they cause inflammation. They can consume and destroy bacteria, the dreaded vehicles of infectious diseases; but they can also transport these injurious Monera to fresh regions, and so extend the sphere of infection. It is probable that the sensitive and travelling leucocytes of our invertebrate ancestors have powerfully co-operated for millions of years in the phylogenesis of the advancing animal organisation.

The red blood-cells have a much more restricted sphere of distribution and activity. But they also are very important in connection with certain functions of the craniote-organism, especially the exchange of gases or respiration. The cells of the dark red, carbonised or venous, blood, which have absorbed carbonic acid from the animal tissues, give this off in the respiratory organs; they receive instead of it fresh oxygen, and thus bring about the bright red colour that distinguishes oxydised or arterial blood. The red colouring matter of the blood (haemoglobin) is regularly distributed in the pores of their protoplasm. The red cells of most of the Vertebrates are elliptical flat disks, and enclose a nucleus of the same shape; they differ a good deal in size (Figure 2.358). The mammals are distinguished from the other Vertebrates by the circular form of their biconcave red cells and by the absence of a nucleus (Figure 1.1); only a few genera still have the elliptic form inherited from the reptiles (Figure 1.2).

In the embryos of the mammals the red cells have a nucleus and the power of increasing by cleavage (Figure 1.10).

The origin of the blood-cells and vessels in the embryo, and their relation to the germinal layers and tissues, are among the most difficult problems of ontogeny--those obscure questions on which the most divergent opinions are still advanced by the most competent scientists. In general, it is certain that the greater part of the cells that compose the vessels and their contents come from the mesoderm--in fact, from the gut-fibre layer; it was on this account that Baer gave the name of "vascular layer" to this visceral layer of the coeloma. But other important observers say that a part of these cells come from other germinal layers, especially from the gut-gland layer. It seems to be true that blood-cells may be formed from the cells of the entoderm before the development of the mesoderm. If we examine sections of chickens, the earliest and most familiar subjects of embryology, we find at an early stage the "primitive-aortas" we have already described (Figure 2.360 ao) in the ventral angle between the episoma (Pv) and hyposoma (Sp). The thin wall of these first vessels of the amniote embryo consists of flat cells (endothelia or vascular epithelia); the fluid within already contains numbers of red blood-cells; both have been developed from the gut-fibre layer. It is the same with the vessels of the germinative area (Figure 2.361 v), which lie on the entodermic membrane of the yelk-sac (c). These features are seen still more clearly in the transverse section of the duck-embryo in Figure 1.152. In this we see clearly how a number of stellate cells proceed from the "vascular layer" and spread in all directions in the "primary body-cavity"--i.e. in the s.p.a.ces between the germinal layers. A part of these travelling cells come together and line the wall of the larger s.p.a.ces, and thus form the first vessels; others enter into the cavity, live in the fluid that fills it, and multiply by cleavage--the first blood-cells.

But, besides these mesodermic cells of the "vascular layer" proper, other travelling cells, of which the origin and purport are still obscure, take part in the formation of blood in the meroblastic Vertebrates (especially fishes). The chief of these are those that Ruckert has most aptly denominated "merocytes." These "eating yelk-cells" are found in large numbers in the food-yelk of the Selachii, especially in the yelk-wall--the border zone of the germinal disk in which the embryonic vascular net is first developed. The nuclei of the merocytes become ten times as large as the ordinary cell-nucleus, and are distinguished by their strong capacity for taking colour, or their special richness in chromatin. Their protoplasmic body resembles the stellate cells of osseous tissue (astrocytes), and behaves just like a rhizopod (such as Gromia); it sends out numbers of stellate processes all round, which ramify and stretch into the surrounding food-yelk. These variable and very mobile processes, the pseudopodia of the merocytes, serve both for locomotion and for getting food; as in the real rhizopods, they surround the solid particles of food (granules and plates of yelk), and acc.u.mulate round their nucleus the food they have received and digested. Hence we may regard them both as eating-cells (phagocytes) and travelling-cells (planocytes). Their lively nucleus divides quickly and often repeatedly, so that a number of new nuclei are formed in a short time; as each fresh nucleus surrounds itself with a mantle of protoplasm, it provides a new cell for the construction of the embryo. Their origin is still much disputed.

(FIGURE 2.361. Merocytes of a shark-embryo, rhizopod-like yelk-cells underneath the embryonic cavity (B). (From Ruckert.) z two embryonic cells, k nuclei of the merocytes, which wander about in the yelk and eat small yelk-plates (d), k smaller, more superficial, lighter nuclei, k apostrophe a deeper nucleus, in the act of cleavage, k asterisk chromatin-filled border-nucleus, freed from the surrounding yelk in order to show the numerous pseudopodia of the protoplasmic cell-body.)

Half of the twelve stems of the animal world have no blood-vessels.

They make their first appearance in the Vermalia. Their earliest source is the primary body-cavity, the simple s.p.a.ce between the two primary germinal layers, which is either a relic of the segmentation-cavity, or is a subsequent formation. Amoeboid planocytes, which migrate from the entoderm and reach this fluid-filled primary cavity, live and multiply there, and form the first colourless blood-cells. We find the vascular system in this very simple form to-day in the Bryozoa, Rotatoria, Nematoda, and other lower Vermalia.

The first step in the improvement of this primitive vascular system is the formation of larger ca.n.a.ls or blood-conducting tubes. The s.p.a.ces filled with blood, the relics of the primary body-cavity, receive a special wall. "Blood-vessels" of this kind (in the narrower sense) are found among the higher worms in various forms, sometimes very simple, at other times very complex. The form that was probably the incipient structure of the elaborate vascular system of the Vertebrates (and of the Articulates) is found in two primordial princ.i.p.al vessels--a dorsal vessel in the middle line of the dorsal wall of the gut, and a ventral vessel that runs from front to rear in the middle line of its ventral wall. From the dorsal vessel is evolved the aorta (or princ.i.p.al artery), from the ventral vessel the princ.i.p.al or subintestinal vein. The two vessels are connected in front and behind by a loop that runs round the gut. The blood contained in the two tubes is propelled by their peristaltic contractions.

(FIGURE 2.362. Vascular system of an Annelid (Saenuris), foremost section. d dorsal vessel, v ventral vessel, c transverse connection of two (enlarged in shape of heart). The arrows indicate the direction of the flow of blood. (From Gegenbaur.)

The earliest Vermalia in which we first find this independent vascular system are the Nemertina (Figure 2.244). As a rule, they have three parallel longitudinal vessels connected by loops, a single dorsal vessel above the gut and a pair of lateral vessels to the right and left. In some of the Nemertina the blood is already coloured, and the red colouring matter is real haemoglobin, connected with elliptical discoid cells, as in the Vertebrates. The further evolution of this rudimentary vascular system can be gathered from the cla.s.s of the Annelids in which we find it at various stages of development. First, a number of transverse connections are formed between the dorsal and ventral vessels, which pa.s.s round the gut ring-wise (Figure 2.362).

Other vessels grow into the body-wall and ramify in order to convey blood to it. In addition to the two large vessels of the middle plane there are often two lateral vessels, one to the right and one to the left; as, for instance, in the leech. There are four of these parallel longitudinal vessels in the Enteropneusts (Balanoglossus, Figure 2.245). In these important Vermalia the foremost section of the gut has already been converted into a gill-crate, and the vascular arches that rise in the wall of this from the ventral to the dorsal vessel have become branchial vessels.

We have a further important advance in the Tunicates, which we have recognised as the nearest blood-relatives of our early vertebrate ancestors. Here we find for the first time a real heart--i.e. a central organ of circulation, driving the blood into the vessels by the regular contractions of its muscular wall, it is of a very rudimentary character, a spindle-shaped tube, pa.s.sing at both ends into a princ.i.p.al vessel (Figure 2.221). By its original position behind the gill-crate, on ventral side of the Tunicates (sometimes more, sometimes less, forward), the head shows clearly that it has been formed by the local enlargement of a section of the ventral vessel. We have already noticed the remarkable alternation of the direction of the blood stream, the heart driving it first from one end, then from the other (Chapter 2.16). This is very instructive, because in most of the worms (even the Enteropneust) the blood in the dorsal vessel travels from back to front, but in the Vertebrates in the opposite direction. As the Ascidia-heart alternates steadily from one direction to the other, it shows us permanently, in a sense, the phylogenetic transition from the earlier forward direction of the dorsal current (in the worms) to the new backward direction (in the Vertebrates).

(FIGURE 2.363. Head of a fish-embryo, with rudimentary vascular system, from the left. dc Cuvier's duct (juncture of the anterior and posterior princ.i.p.al veins), sv venous sinus (enlarged end of Cuvier's duct), a auricle, v ventricle, abr trunk of branchial artery, s gill-clefts (arterial arches between), ad aorta, c carotid artery, n nasal pit. (From Gegenbaur.)

FIGURE 2.364. The five arterial arches of the Craniotes (1 to 5) in their original disposition, a arterial cone or bulb, a double apostrophe aorta-trunk, c carotid artery (foremost continuation of the roots of the aorta). (From Rathke.)

FIGURE 2.365. The five arterial arches of the birds; the lighter parts of the structure disappear; only the shaded parts remain. Letters as in Figure 2.364. s subclavian arteries, p pulmonary artery, p apostrophe branches of same, c apostrophe outer carotid, c double apostrophe inner carotid. (From Rathke.)

FIGURE 2.366. The five arterial arches of mammals; letters as in Figure 2.365. v vertebral artery, b Botall's duct (open in the embryo, closed afterwards). (From Rathke.))

As the new direction became permanent in the earlier Prochordonia, which gave rise to the Vertebrate stem, the two vessels that proceed from either end of the tubular heart acquired a fixed function. The foremost section of the ventral vessel henceforth always conveys blood from the heart, and so acts as an artery; the hind section of the same vessel brings the blood from the body to the heart, and so becomes a vein. In view of their relation to the two sections of the gut, we may call the latter the intestinal vein and the former the branchial artery. The blood contained in both vessels, and also in the heart, is venous or carbonised blood--i.e. rich in carbonic acid; on the other hand, the blood that pa.s.ses from the gills into the dorsal vessel is provided with fresh oxygen--arterial or oxydised blood. The finest branches of the arteries and veins pa.s.s into each other in the tissues by means of a network of very fine, ventral, hair-like vessels, or capillaries (Figure 2.359).

When we turn from the Tunicates to the closely-related Amphioxus we are astonished at first to find an apparent retrogression in the formation of the vascular system. As we have seen, the Amphioxus has no real heart; its colourless blood is driven along in its vascular system by the princ.i.p.al vessel itself, which contracts regularly in its whole length (cf. Figure 2.210). A dorsal vessel that lies above the gut (aorta) receives the arterial blood from the gills and drives it into the body. Returning from here, the venous blood gathers in a ventral vessel under the gut (intestinal vein), and goes back to the gills. A number of branchial vascular arches, which effect respiration and rise in the wall of the branchial gut from belly to back, absorb oxygen from the water and give off carbonic acid; they connect the ventral with the dorsal vessel. As the same section of the ventral vessel, which also forms the heart in the Craniotes, has developed in the Ascidia into a simple tubular heart, we may regard the absence of this in the Amphioxus as a result of degeneration, a return in this case to the earlier form of the vascular system, as we find it in many of the worms. We may a.s.sume that the Acrania that really belong to our ancestral series did not share this retrogression, but inherited the one-chambered heart of the Prochordonia, and transmitted it directly to the earliest Craniotes (cf. the ideal Primitive Vertebrate, Prospondylus, Figures 1.98 to 1.102).

(FIGURES 2.367 TO 2.370. Metamorphosis of the five arterial arches in the human embryo (diagram from Rathke). la arterial cone, 1, 2, 3, 4, 5 first to fifth pair of arteries, ad trunk of aorta, aw roots of aorta. In Figure 2.367 only three, in Figure 2.368 all five, of the aortic arches are given (the dotted ones only are developed). In Figure 2.369 the first two pairs have disappeared again. In Figure 2.370 the permanent trunks of the artery are shown; the dotted parts disappear, s subclavian artery, v vertebral, ax axillary, c carotid (c apostrophe outer, c double apostrophe inner carotid), p pulmonary.)

The further phylogenetic evolution of the vascular system is revealed to us by the comparative anatomy of the Craniotes. At the lowest stage of this group, in the Cyclostomes, we find for the first time the differentiation of the vasorium into two sections: a system of blood-vessels proper, which convey the RED blood about the body, and a system of lymphatic vessels, which absorb the colourless lymph from the tissues and convey it to the blood. The lymphatics that absorb from the gut and pour into the blood-stream the milky food-fluid formed by digestion are distinguished by the special name of "chyle-vessels." While the chyle is white on account of its high proportion of fatty particles, the lymph proper is colourless. Both chyle and lymph contain the colourless amoeboid cells (leucocytes, Figure 1.12) that we also find distributed in the blood as colourless blood-cells (or "white corpuscles"); but the blood also contains a much larger quant.i.ty of red cells, and these give its characteristic colour to the blood of the Craniotes (rhodocytes, Figure 2.358). The distinction between lymph, chyle, and blood-vessels which is found in all the Craniotes may be regarded as an outcome of division of labour between various sections of our originally simple vascular system. In the Gnathostomes the spleen makes its first appearance, an organ rich in blood, the chief function of which is the extensive formation of new colourless and red cells. It is not found in the Acrania and Cyclostomes, or any of the Invertebrates. It has been transmitted from the earliest fishes to all the Craniotes.

The heart also, the central organ of circulation in all the Craniotes, shows an advance in structure in the Cyclostomes. The simple, spindle-shaped heart-tube, found in the same form in the embryo of all the Craniotes, is divided into two sections or chambers in the Cyclostomes, and these are separated by a pair of valves. The hind section, the auricle, receives the venous blood from the body and pa.s.ses it on to the anterior section, the ventricle. From this it is driven through the trunk of the branchial artery (the foremost section of the ventral vessel or princ.i.p.al vein) into the gills.

In the Selachii an arterial cone is developed from the foremost end of the ventricle, as a special division, cut off by valves. It pa.s.ses into the enlarged base of the trunk of the branchial artery (Figure 2.363 abr). On each side 5 to 7 arteries proceed from it. These rise between the gill-clefts (s) on the gill-arches, surround the gullet, and unite above into a common trunk-aorta, the continuation of which over the gut corresponds to the dorsal vessel of the worms. As the curved arteries on the gill-arches spread into a network of respiratory capillaries, they contain venous blood in their lower part (as arches of the branchial artery) and arterial blood in the upper part (as arches of the aorta). The junctures of the various aortic arches on the right and left are called the roots of the aorta. Of an originally large number of aortic arches there remain at first six, then (owing to degeneration of the fifth arch) only five, pairs; and from these five pairs (Figure 2.364) the chief parts of the arterial system develop in all the higher Vertebrates.

(FIGURE 2.371. Heart of a rabbit-embryo, from behind, a vitelline veins, b auricles of the heart, c atrium, d ventricle, e arterial bulb, f base of the three pairs of arterial arches. (From Bischoff.)

FIGURE 2.372. Heart of the same embryo (Figure 2.371), from the front.

v vitelline veins, a auricle, ca auricular ca.n.a.l, l left ventricle, r right ventricle, ta arterial bulb. (From Bischoff.))

The appearance of the lungs and the atmospheric respiration connected therewith, which we first meet in the Dipneusts, is the next important step in vascular evolution. In the Dipneusts the auricle of the heart is divided by an incomplete part.i.tion into two halves. Only the right auricle now receives the venous blood from the veins of the body. The left auricle receives the arterial blood from the pulmonary veins. The two auricles have a common opening into the simple ventricle, where the two kinds of blood mix, and are driven through the arterial cone or bulb into the arterial arches. From the last arterial arches the pulmonary arteries arise (Figure 2.365 p). These force a part of the mixed blood into the lungs, the other part of it going through the aorta into the body.

From the Dipneusts upwards we now trace a progressive development of the vascular system, which ends finally with the loss of branchial respiration and a complete separation of the two halves of the circulation. In the Amphibia the part.i.tion between the two auricles is complete. In their earlier stages, as tadpoles (Figure 2.262), they have still the branchial respiration and the circulation of the fishes, and their heart contains venous blood alone. Afterwards the lungs and pulmonary vessels are developed, and henceforth the ventricle of the heart contains mixed blood. In the reptiles the ventricle and its arterial cone begin to divide into two halves by a longitudinal part.i.tion, and this part.i.tion becomes complete in the higher reptiles and birds on the one hand, and the stem-forms of the mammals on the other. Henceforth, the right half of the heart contains only venous, and the left half only arterial, blood, as we find in all birds and mammals. The right auricle receives its carbonised or venous blood from the veins of the body, and the right ventricle drives it through the pulmonary arteries into the lungs. From here the blood returns, as oxydised or arterial blood, through the pulmonary veins to the left auricle, and is forced by the left ventricle into the arteries of the body. Between the pulmonary arteries and veins is the capillary system of the small or pulmonary circulation. Between the body-arteries and veins is the capillary system of the large or body-circulation. It is only in the two highest cla.s.ses of Vertebrates--the birds and mammals--that we find a complete division of the circulations. Moreover, this complete separation has been developed quite independently in the two cla.s.ses, as the dissimilar formation of the aortas shows of itself. In the birds the RIGHT half of the fourth arterial arch has become the permanent arch (Figure 2.365). In the mammals this has been developed from the LEFT half of the same fourth arch (Figure 2.366).

(FIGURE 2.373. Heart and head of a dog-embryo, from the front, a fore brain, b eyes, c middle brain, d primitive lower jaw, e primitive upper jaw, f gill-arches, g right auricle, h left auricle, i left ventricle, k right ventricle. (From Bischoff.)