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

same thing happening in the head. This simple primary pericardial cavity has been well called by Gegenbaur the “head-coeloma,” and by Hertwig the “pericardial breast-cavity.” As it now encloses the heart, it may also be called cardiocoel.

The cardiocoel, or head-coelom, is often disproportionately large in the Amniotes, the simple cardiac tube growing considerably and lying in several folds. This causes the ventral wall of the amniote embryo, between the head and the navel, to be pushed outwards as in rupture (cf. Figure 1.180 h). A transverse fold of the ventral wall, which receives all the vein-trunks that open into the heart, grows up from below between the pericardium and the stomach, and forms a transverse partition, which is the first structure of the primary diaphragm (Figure 2.380 d). This important muscular partition, which completely separates the thoracic and abdominal cavities in the mammals alone, is still very imperfect here; the two cavities still communicate for a time by two narrow canals. These canals, which belong to the dorsal part of the head-coelom, and which we may call briefly pleural ducts, receive the two pulmonary sacs, which develop from the hind end of the ventral wall of the head-gut; they thus become the two pleural cavities.

The diaphragm makes its first appearance in the class of the Amphibia (in the salamanders) as an insignificant muscular transverse fold of the ventral wall, which rises from the fore end of the transverse abdominal muscle, and grows between the pericardium and the liver. In the reptiles (tortoises and crocodiles) a later dorsal part is joined to this earlier ventral part of the rudimentary diaphragm, a pair of subvertebral muscles rising from the vertebral column and being added as “columns” to the transverse partition. But it was probably in the Permian sauro-mammals that the two originally separate parts were united, and the diaphragm became a complete partition between the thoracic and abdominal cavities in the mammals; as it considerably enlarges the chest-cavity when it contracts, it becomes an important respiratory muscle. The ontogeny of the diaphragm in man and the other mammals reproduces this phylogenetic process to-day, in accordance with the biogenetic law; in all the mammals the diaphragm is formed by the secondary conjunction of the two originally separate structures, the earlier ventral part and the later dorsal part.

Sometimes the blending of the two diaphragmatic structures, and consequently the severance of the one pleural duct from the abdominal cavity, is not completed in man. This leads to a diaphragmatic rupture (hernia diaphragmatica). The two cavities then remain in communication by an open pleural duct, and loops of the intestine may penetrate by this “rupture opening” into the chest-cavity. This is one of those fatal mis-growths that show the great part that blind chance has in organic development.

(FIGURE 2.381. Transverse section of the head of a chick-embryo, thirty-six hours old. Underneath the medullary tube the two primitive aortas (pa) can be seen in the head-plates (s) at each side of the chorda. Underneath the gullet (d) we see the aorta-end of the heart (ae), hh cervical cavity or head coelom, hk top of heart, ks head-sheath, amniotic fold, h horny plate. (From Remak.)

(FIGURE 2.382. Transverse section of the cardiac region of the same chick-embryo (behind the preceding). In the cervical cavity (hh) the heart (h) is still connected by a mesocard (hg) with the gut-fibre layer (pf). d gut-gland layer, up provertebral plates, jb rudimentary auditory vesicle in the horny plate, hp first rise of the amniotic fold. (From Remak.))

Thus the thoracic cavity of the mammals, with its important contents, the heart and lungs, belongs originally to the HEAD-PART of the vertebrate body, and its inclusion in the trunk is secondary. This instructive and very interesting fact is entirely proved by the concordant evidence of comparative anatomy and ontogeny. The lungs are outgrowths of the head-gut; the heart develops from its inner wall. The pleural sacs that enclose the lungs are dorsal parts of the head-coelom, originating from the pleuroducts; the pericardium in which the heart afterwards lies is also double originally, being formed from ventral halves of the head-coelom, which only combine at a later stage. When the lung of the air-breathing Vertebrates issues from the head-cavity and enters the trunk-cavity, it follows the example of the floating bladder of the fishes, which also originates from the pharyngeal wall in the shape of a small pouch-like outgrowth, but soon grows so large that, in order to find room, it has to pass far behind into the trunk-cavity. To put it more precisely, the lung of the quadrupeds retains this hereditary growth-process of the fishes; for the hydrostatic floating bladder of the latter is the air-filled organ from which the air-breathing organ of the former has been evolved.

There is an interesting cenogenetic phenomenon in the formation of the heart of the higher Vertebrates that deserves special notice. In its earliest form the heart is DOUBLE, as recent observation has shown, in all the Amniotes, and the simple spindle-shaped cardiac tube, which we took as our starting-point, is only formed at a later stage, when the two lateral tubes move backwards, touch each other, and at last combine in the middle line. In man, as in the rabbit, the two embryonic hearts are still far apart at the stage when there are already eight primitive segments (Figure 1.134 h). So also the two coelom-pouches of the head in which they lie are still separated by a broad space. It is not until the permanent body of the embryo develops and detaches from the embryonic vesicle that the separate lateral structures join together, and finally combine in the middle line. As the median partition between the right and left cardiocoel disappears, the two cervical cavities freely communicate (Figure 2.381), and form, on the ventral side of the amniote head, a horseshoe-shaped arch, the points of which advance backwards into the pleuroducts or pleural cavities, and from there into the two peritoneal sacs of the trunk. But even after the conjunction of the cervical cavities (Figure 2.381) the two cardiac tubes remain separate at first; and even after they have united a delicate partition in the middle of the simple endothelial tube (Figures 2.379 s and 2.382 h) indicates the original separation. This CENOGENETIC “primary cardiac septum” presently disappears, and has no relation to the subsequent permanent partition between the halves of the heart, which, as a heritage from the reptiles, has a great PALINGENETIC importance.

Thorough opponents of the biogenetic law have laid great stress on these and similar cenogenetic phenomena, and endeavoured to urge them as striking disproofs of the law. As in every other instance, careful, discriminating, comparative-morphological examination converts these supposed disproofs of evolution into strong arguments in its favour. In his excellent work, On the structure of the Heart in the Amphibia (1886), Carl Rabl has shown how easily these curious cenogenetic facts can be explained by the secondary adaptation of the embryonic structure to the great extension of the food-yelk.

The embryology of all the other parts of the vascular system also gives us abundant and valuable data for the purposes of phylogeny. But as one needs a thorough knowledge of the intricate structure of the whole vascular system in man and the other Vertebrates in order to follow this with profit, we cannot go into it further here. Moreover, many important features in the ontogeny of the vascular system are still very obscure and controverted. The characters of the embryonic circulation of the Amniotes, which we have previously considered (Chapter 1.15), are late acquisitions and entirely cenogenetic. (Cf. Chapter 1.15 and Figures 1.198 to 1.202.)

 

In the Selachii also we find a longitudinal row of segmental canals on each side, which open outwards into the primitive renal ducts (nephrotomes, Chapter 1.14). The segmental canals (a pair in each segment of the middle part of the body) open internally by a ciliated funnel into the body-cavity. From the posterior group of these organs a compact primitive kidney is formed, the anterior group taking part in the construction of the sexual organs.

In the same simple form that remains throughout life in the Myxinoides and partly in the Selachii we find the primitive kidney first developing in the embryo of man and the higher Craniotes (Figures 2.386 and 2.387). Of the two parts that compose the comb-shaped primitive kidney the longitudinal channel, or nephroduct, is always the first to appear; afterwards the transverse “canals,” the excreting nephridia, are formed in the mesoderm; and after this again the Malpighian capsules with their arterial coils are associated with these as coelous outgrowths. The primitive renal duct, which appears first, is found in all craniote embryos at the early stage in which the differentiation of the medullary tube takes place in the ectoderm, the severance of the chorda from the visceral layer in the entoderm, and the first trace of the coelom-pouches arises between the limiting layers (Figure 2.385). The nephroduct (ung) is seen on each side, directly under the horny plate, in the shape of a long, thin, thread-like string of cells. It presently hollows out and becomes a canal, running straight from front to back, and clearly showing in the transverse section of the embryo its original position in the space between horny plate (h), primitive segments (uw), and lateral plates (hpl). As the originally very short urinary canals lengthen and multiply, each of the two primitive kidneys assumes the form of a half-feathered leaf (Figure 2.387). The lines of the leaf are represented by the urinary canals (u), and the rib by the outlying nephroduct (w). At the inner edge of the primitive kidneys the rudiment of the ventral sexual gland (g) can now be seen as a body of some size. The hindermost end of the nephroduct opens right behind into the last section of the rectum, thus making a cloaca of it. However, this opening of the nephroducts into the intestine must be regarded as a secondary formation. Originally they open, as the Cyclostomes clearly show, quite independently of the gut, in the external skin of the abdomen.

(FIGURE 2.395. Primitive kidneys and germinal glands of a human embryo, three inches in length (beginning of the sixth week), magnified fifteen times. k germinal gland, u primitive kidney, z diaphragmatic ligament of same, w Wolffian duct (opened on the right), g directing ligament (gubernaculum), a allantoic duct. (From Kollmann.))

In the Myxinoides the primitive kidneys retain this simple comb-shaped structure, and a part of it is preserved in the Selachii; but in all the other Craniotes it is only found for a short time in the embryo, as an ontogenetic reproduction of the earlier phylogenetic structure. In these the primitive kidney soon assumes the form (by the rapid growth, lengthening, increase, and serpentining of the urinary canals) of a large compact gland, of a long, oval or spindle-shaped character, which passes through the greater part of the embryonic body-cavity (Figures 1.183 m, 1.184 m, 2.388 n). It lies near the middle line, directly under the primitive vertebral column, and reaches from the cardiac region to the cloaca. The right and left kidneys are parallel to each other, quite close together, and only separated by the mesentery—the thin narrow layer that attaches the middle gut to the under surface of the vertebral column. The passage of each primitive kidney, the nephroduct, runs towards the back on the lower and outer side of the gland, and opens in the cloaca, close to the starting-point of the allantois; it afterwards opens into the allantois itself.

(FIGURES 2.396 TO 2.398. Urinary and sexual organs of ox-embryos. Figure 2.396, female embryo one and a half inches long; Figure 2.397, male embryo, one and a half inches long. Figure 2.398 female embryo two and a half inches long. w primitive kidney, wg Wolffian duct, m Mullerian duct, m apostrophe upper end of same (opened at t), i lower and thicker part