The changes which are to be observed in the later stages depend principally upon a peculiar feature characteristic of the development of the dorsal mesogastrium. This feature consists in the extreme redundancy of the membrane which grows out of proportion to the requirements of its visceral connections, and to a certain extent becomes independent of the direct mechanical purpose of carrying blood vessels to the viscera. Hence in a transverse section at this period (Figs. 174 and 175) the mesogastrium no longer passes in a direct line between its points of attachment, viz. the greater curvature of the stomach and the vertebral column, but extends beyond the stomach to the left. We will appreciate the significance of this extensive growth of the mesogastrium especially in considering the development of the spleen and pancreas. For the present it will suffice to note (Figs. 174 and 175) that the growth has carried the mesogastrium well to the left of the stomach, consequently the retrogastric space is now bounded toward the left by the bend which the original right leaf of the primitive sagittal mesogastrium takes in order to reach its gastric attachment. The retrogastric space therefore terminates toward the left in a blind pocket formed by this reduplication of the mesogastrium.
One more factor is to be taken into consideration, namely the tendency, already noted, of peritoneal surfaces to become adherent to each other. Such adhesion involves the apposed surfaces of the mesogastrium and of the primitive parietal peritoneum to the left of the vertebral column. The dorsal (original left) layer of the mesogastrium adheres to the parietal peritoneum covering the left side of the abdominal background and the cephalic portion of the ventral surface of the left kidney up to the end of the blind pouch which forms the extreme left limit of the retrogastric space. Hence, after this process of adhesion is completed, the dorsal wall of the retrogastric space is lined by secondary parietal peritoneum covering the left kidney (original right leaf of primitive mesogastrium) (Fig. 175). We obtain (Fig. 175 at x) an apparent continuity of the parietal peritoneum with that portion of the mesogastrium which, derived from the original left layer of the membrane, appears now to extend, as the ventral one of two layers, between the stomach and the abdominal parietes near the lateral border of the left kidney. (Primitive gastro-splenic omentum.)
It should be remembered that the disposition of the peritoneum just indicated is modified by the development of the pancreas and spleen, both of which organs are intimately associated with the mesogastrium. The foregoing statements and diagrams are therefore merely given for the purpose of affording a general view of the extent, growth and changes of the dorsal mesogastrium before proceeding to consider the development of the pancreas and spleen in and from the membrane itself.
In the view directly from in front the redundancy of the peritoneum forming the mesogastrium is shown in Figs. 176 and 177. Just as the membrane extends further to the left than required by its visceral connection with the stomach, so the downward growth exceeds the demand made by the rotation of the attached border (greater curvature) caudad and to the left. The mesogastrium, forming, as it now does, the great omentum, enlarges in descending toward the transverse colon (Fig. 177). The bag thus formed can be distended with air in a fœtus of from 8 to 9 cm. vertex-coccygeal measure, as shown in the figure. Consequently in sagittal section the membrane is seen to extend caudad beyond the level of the greater curvature, and must turn on itself and pass again cephalad in order to reach the stomach ([Fig. 178]). By reason of this excessive growth the limits of the primitive retrogastric space are enlarged, not only toward the left, but more especially in the caudal direction. The bend made by the mesogastrium in returning to the stomach forms the blind extremity of a pouch which continues the retrogastric space caudad beyond the stomach, and whose dorsal and ventral walls are formed by the reduplicated mesogastrium. This pocket or pouch constitutes the omental or epiploic bursa of the lesser peritoneal cavity, for the great omentum is the direct product of this redundant growth of the mesogastrium caudad. It will be observed that the great omentum is made up of four peritoneal layers, the folding of the double-layered mesogastrium naturally producing this result. The first or ventral and the fourth or dorsal layer are derived from the original left layer of the primitive sagittal mesogastrium; the intermediate second and third layers, separated from each other at this stage by the cavity of the omental bursa, are products of the primitive right leaf of the mesogastrium. Since the entire retrogastric space with its extensions becomes the “lesser cavity” of the human adult peritoneum, it will be seen that its serous membrane is derived from the original right leaf of the mesogastrium (second and third omental layers). After the above-described adhesion of the mesogastrium to the parietal peritoneum overlying the ventral surface of the left kidney, the membrane would be traced in sagittal section ([Fig. 179]) from the dorsal surface of the stomach caudad, lining the interior of the omental bursa (second layer) to the turn or blind end of the pouch; thence cephalad as the third omental layer, forming the dorsal wall of the epiploic bursa, to invest, as secondary parietal peritoneum, the cephalic segment of the ventral surface of the left kidney.
C. Development of Spleen and Pancreas in the Dorsal Mesogastrium and Changes in the Disposition of the Great Omentum.
In order to obtain a correct conception of the adult human conditions it is finally necessary to consider the development of the spleen and pancreas in their connection with the dorsal mesogastrium and to note the changes which are produced by adhesion of portions of the great omentum to adjacent serous surfaces. It will be advisable to discuss these subjects at first separately, and to subsequently combine all the facts in an attempt to gain a correct impression of their share in determining the disposition of the adult human peritoneum.
1. Development of Spleen.—The spleen develops from the mesoderm between the layers of the dorsal mesogastrium, near its point of accession to the greater curvature, in the region of the subsequent fundus. It has therefore, like the stomach, originally free peritoneal surfaces. After rotation of the stomach the organ lies between the two layers of the membrane at the extreme left end of the retrogastric space (Fig. 180).
Fig. 180.—Schematic transverse section of the abdomen, showing early stage of development of spleen from extreme left end of dorsal mesogastric pouch.