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pericardium, but with the folding over they come to lie in the dorsal wall and gradually bulge into the cavity as they coalesce to form the heart, so that the heart drops into the dorsal side of the pericardium and draws down a fold of the membrane called the dorsal mesocardium. In mammals A. Robinson (Jour. Anat. and Phys., xxxvii. 1) has shown that no ventral mesocardium exists, though in more lowly vertebrates it is present. Laterally the pericardial cavity communicates with the general cavity of the coelom, but with the growth of the Cuvierian ducts (see development of veins) these communications disappear. Originally the mesocardium runs the whole length of the pericardium from before backward, but later on the middle part becomes obliterated, and so the two separate reflections from the parietal to the visceral layer, already noticed, are accounted for.

Just behind the pericardium and in front of the umbilicus, which at first are close together, the mesoderm forms a mass which is called the septum transversum, and into this the developing lungs push bag-like protrusions of the coelom, consisting of visceral and parietal layers, and these eventually lose their connexion with the rest of the coelom, as the diaphragm develops, and become the pleural cavities. After the pericardium and pleurae have been separated off the remainder of the coelom becomes the peritoneum. At first the stomach and intestine form a straight tube, which is connected to the dorsum of the embryo by a dorsal mesentery and to the mid-ventral wall in front of the umbilicus by a ventral mesentery. Into the ventral mesentery the liver grows as diverticula from the duodenum, so that some of the mesentery remains as the falciform ligament of the liver and some as the lesser omentum. Into the dorsal mesentery the pancreas grows, also as diverticula, from the duodenum, while the spleen is developed from the mesoderm contained in the same fold. As the stomach turns over so that its left side becomes ventral, the dorsal mesentery attached to it becomes pulled out, in such a way that part of it forms the great omentum and part the gastro-splenic omentum. After the caecum is formed as a diverticulum from the intestine it is situated close to the liver and gradually travels down into the right iliac fossa. This passage to the right is accompanied by a throwing over of the duodenal loop to the right, so that the right side of its mesentery becomes pressed against the dorsal wall of the abdomen and obliterated. This accounts for the fact that the pancreas and duodenum are only covered by peritoneum on their anterior surfaces in man. The formation of the lesser sac is due to the turning over of the stomach to the right, with the result that a cave, known sometimes as the bursa omentalis, is formed behind it. Originally, of course, the whole colon had a dorsal mesocolon continuous with the mesentery, but in the region of the ascending and descending colon this usually disappears and these parts of the gut are uncovered by peritoneum posteriorly. The transverse mesocolon persists and at first is quite free from the great omentum, but later, in man, the two structures fuse[1] and the fourth layer of the great omentum becomes continuous with the posterior layer of the transverse mesocolon.

For further details see Quain’s Anatomy (London, 1908).

Comparative Anatomy.—In the Amphioxus the coelom is developed in the embryo as a series of bilateral pouches, called enterocoeles, from the sides of the alimentary canal; these are therefore entodermal in their origin, as in Sagitta and the Echinodermata among the invertebrates. In the adult the development of the atrium causes a considerable reduction of the coelom, represented by two dorsal coelomic canals communicating with a ventral canal by means of branchial canals which run down the outer side of the primary gill bars. Into the dorsal canals the nephridia open. In the intestinal region the coelom is only present on the left side.

In the higher vertebrates (Craniata) the coelom is developed by a splitting of the mesoderm into two layers, and a pericardium is constricted off from the general cavity. In all cases the ova burst into the coelom before making their way to the exterior, and in some cases, e.g. amphioxus, lamprey (Cyclostomata), eels and mud-fish (Dipnoi), the sperm cells do so too. The Cyclostomata have a pair of genital pores which lead from the coelom into the urino-genital sinus, and so to the exterior.

In the Elasmobranch fish there is a pericardio-peritoneal canal forming a communication between these two parts of the coelom; also a large common opening for the two oviducts in the region of the liver, and two openings, called abdominal pores, on to the surface close to the cloacal aperture. In the Teleostomi (Teleostean and Ganoid fish) abdominal pores are rare, but in most Teleostei (bony fish) the ova pass directly down oviducts, as they do in Arthropods, without entering the peritoneal cavity; there is little doubt, however, that these oviducts are originally coelomic in origin. In the Dipnoi (mud-fish) abdominal pores are found, and probably serve as a passage for the sperm cells, since there are no vasa deferentia. In fishes a complete dorsal mesentery is seldom found in the adult; in many cases it only remains as a tube surrounding the vessels passing to the alimentary canal.

In the Amphibia, Reptilia and Aves, one cavity acts as pleura and peritoneum, though in the latter the lungs are not completely surrounded by a serous membrane. In many lizards the comparatively straight intestine, with its continuous dorsal mesentery and ventral mesentery in the anterior part of the abdomen, is very like a stage in the development of the human and other mammalian embryos. In the mammalia the diaphragm is complete (see Diaphragm) and divides the pleuro-peritoneal cavity into its two constituent parts. In the lower mammals the derivatives of the original dorsal mesentery do not undergo as much fusion and obliteration as they do in adult man; the ascending and descending mesocolon is retained, and the transverse mesocolon contracts no adhesion to the great omentum. It is a common thing, however, to find a fenestrated arrangement of the great omentum which shows that its layers have been completely obliterated in many places.

In those animals, such as the rabbit, in which the tests are sometimes in the scrotum and sometimes in the abdomen, the communication between the peritoneum and the tunica vaginalis remains throughout life.

For further details and literature up to 1902, see R. Wiedersheim’s Vergleichende Anatomie der Wirbeltiere (Jena, 1902).  (F. G. P.) 

COEN, JAN PIETERSZOON (1587–1630), fourth governor-general of the Dutch East Indies, was born at Hoorn, and spent his youth at Rome in the house of the famous merchants the Piscatori. In 1607 he sailed from Amsterdam to the Indies as second commercial agent, and remained away four years. He had proved so capable that in 1612 he was sent out a second time at the head of a trading expedition. In the following year he was made a councillor and director-general of the East Indian trade. Afterwards he became president at Bantam, and on the 31st of October 1617 he was promoted in succession to Laurens Reaal to the post of governor-general. To his vigour and intrepidity the Dutch in no small measure owed the preservation and establishment of their empire in the East. He took and destroyed Jacatra, and founded on its ruins the capital of the Dutch East Indies, to which he gave the name of Batavia. In 1622 Coen obtained leave to resign his post and return to Holland, but in his absence great difficulties had arisen with the English at Amboina (the so-called massacre of Amboina), and in 1627 under pressure from the directors of the East India Company he again returned as governor-general to Batavia. In 1629 he was able to beat off a formidable attack of the sultan of Mataram, sometimes styled emperor of Java, upon Batavia. He died the following year.

COENACULUM, the term applied to the eating-room of a Roman house in which the supper (coena) or latest meal was taken. It was sometimes placed in an upper storey and reached by an external staircase. The Last Supper in the New Testament was taken in the Coenaculum, the “large upper room” cited in St Mark (xiv. 15) and St Luke (xxii. 12).

  1. Some authorities hold that this alteration is not brought about by fusion, but by a dragging away of the posterior layer of the great omentum from the dorsal wall of the abdomen.