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The Development of the Lymphatics in the Small Intestine of the Pig

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The Development of the Lymphatics in the Small Intestine of the Pig THE DEVELOPMENT OF THE LYMPHATICS IN THE SMALL INTESTINE OF THE PIG. BY GEORGE HEUER. From the dnatoinical Laboratory of the Johns Hopkins Uiaiversity. WITH 17 FIGUEES. The recent American work on the lymphatic system has given us a new conception of the morphology of the system as a whole. The primitive lymphatic system consists of a number of sacs which are derived from the veins and which become united into a system for the most part by the thoracic duct. A further essential of this primitive system is that all of the sacs give up their connection with the veins, and only the two in the neck rejoin the vein to form the permanent opening. Thns far three sets of paired sacs and two unpaired ones have been described: the jugular sacs, the subclavian which in human embryos are an extension of the jugular, and the sciatic are the paired, the retroperitoneal sac and the cisterna chyli are the unpaired. It is the retroperitoneal sac which especially concerns us in this paper, for it is the source of the lym- phatics of the intestine. This primitive system is complete and can be injected in pig embryos 2.7 cm. long. The idea that the sacs form a primitive lymphatic system is not the most fundamental conception of this new theory, but rather that these sacs arise from the veins and in turn give rise to the lym- phatic vessels, so that we may say that the lymphatic system as a whole is derived from the blood vascular system, that lymphatics are modified veins, and that the growth of lymphatics is alwayvs from center to periphery. There is now a general agreement in regard to the origin of the primitive sacs, but in regard to the second THB AMERICANJOURNAL OF ANATOMY.-VOL. Ix’, NO. 1. 94 George EIeuer. exceedingly important point, namely, that the lymphatic vessels grow from the sacs, there are still differences. It was to test this point that the present study was undertaken at the suggestion of Dr. Sabin. I believe that this work, showing that the vessels of the intestine grow from the retroperitoneal sac, strengthens her position that the lymphatic system is derived from the sacs. Cer- tainly the theory is a fruitful one, for injections of the retro- peritoneal sac give the key for tracing the growth of the lymphatic vessels to the viscera. The literature and general relations of the problem are given by Dr. Sabin in the article on the development of the lymphatics in human embryos in this same journal and therefore only the work on the retroperitoneal sac will be mentioned here. This sac was discovered by F. T. Lewis' who described it as a part of the lymphatic system in 1906. It has been thoroughly worked out by Mr. W. Bsetjer,2 in this laboratory. Mr. Bdjer has s1iowIi tliaL in pig einbryos 17-19 ~m.lung there are small branches of the large renal anastornosing vein in the root of the mesentery. These small veins are ventral to the renal vein and run in an antero-posterior direction. In embryos 19 and 20 mm. long these veins increase markedly in size and number, and by the time the embryo is 21 mm. long shorn sac-like dilatations which are still readily injected from the renal vein. From this time on the sac formation goes on rapidly; in embryos 22-23 mm. long, these small sacs have been completely transformed into a large median sac entirely cut off from the veins and likewise independent of the cisterna chyli. By the time the pig is 2.7 em. long, this sac is abundantly connected with the cisterna chyli, which forms from the veins dorsal to the aorta, and an injection into the thoracic duct will flow through the cisterna chyli into the retroperitoneal sac. Methods and Material. This paper begins with the stage at which Mr. Bztjer left off, namely, where the retroperitoneal sac 'Lewis, Amer. Jour. it€ Anat., Vol. T., 1943106. 'BLetjer, Amer. Jour. of Anat., Vol. VIII. Lymphatics in Small Intestine of the Pig. 95 is connected with the thoracic duct through the cisterna chyli. Such an injection is shown in Fig. 1 from an embryo 3 cm. long. The earliest stage in which the thoracic duct and sac have been injected is 2.7 em., where the appearances do not differ from the specimen shown in Fig. 1, except that the sac is a little smaller. FIG.1.-An embryo pig 3 cm. long, to sliow iiii injection of the lymphatics made by puncturing the thoracic duct. x 2.5, R. o., reproductive organ; r. s., retroperitoneal sac : s., stomach ; s. b., suprareiial body ; t. d., thoracic duct around the aorta; W. b., Wolffian body. The method of injection is as follows: the embryo is opened bg' cutting away the thoracic wall and turning the left lung far to the right or removing it entirely. For injections of the retroperitoneal sac it is better to remove the liver. The aorta and azygos vein should be thoroughly exposed, and the hypodermic needle plunged down behind the aorta, inserting the needle just at the arch where ?)6 George Hener. it is easy to distinguish the vein and thus avoid it. In Fig. 2 is shown a section of an embryo 20 mm. long, taken at the level of the apex of the lung, to show the thoracic duct in its relation to the aorta and to the azygos veins. It shows why it is necessary to insert the needle behind the aorta. It is best to use the finest possible needles; my injections are made with No. 28. For an injection mass either saturated aqueous Prussian blue or India ink are used ; the Prussian blue gives beautiful total specimens which FIG.2.-Transverse section of an embryo pig, 2 cm. long at the level of the apex of the lung to show the relation of the thoracic duct to the azygos veins and to the aorta. x 50. A, aorta; a. v., azygos vein; E., esophagus; p. c., pleural cavity ; t. d., thoracic duct ; 17. c., vertebral cartilage. can be kept either in formalin or hardened in bichloride acetic and kept in alcohol; the India ink runs farther, and the total specimens can he cleared by the Schultze method. The ink is more useful for determining complete injectioas. For studying the development of the lymphatics to the intestine it was found best at first to inject by means of the thoracic duct in embryos from 3.0 to 12.5 cm. Subsequently after the exact position of the retroperitoncal sac was determined, it proved that it was far better to introduce the needle directly into the sac. This Lymphatics in Sniall 1ntcst.inc of the Pig. 07 is done as follows: The embryo is thoroughly opened and the two Wolffian bodies spread apart as far as possible and the entire in- testine turned to the right. This exposes the sac, as can be seen in Fig. 3. By looking closely one can see three or four small arteries FIG. 3.-Embryo pig 3.5 cm. long to shorn an injection of the retroperitoneal sac made through the thoracic duct. x 2.5. r. s., retroperitoneal sac; r. o., reproductive organ ; s, stomach ; W. b., Wolffian body. to the Wolffian body running perpendicular to the rectum ; the needle must enter just dorsal to these arterics. The retroperitoneal sac is the best point for injecting the viscera. To determine the zones in the intestine, the sac has been filled until it ruptured. Good injections are only to be obtained in embryos in wliich the heart is 98 George Heuer. still beating. For any stage it is easier to obtain complete injections through the sac than through the thoracic duct, for in the later case the injection mass must fill the sac before it runs out into the vessels, and the size of the sac decreases the pressure. Figs. 1 to 4 and 7 show that the thoracic duct is fairly sym- metrical below the heart, that there are two vessels, one on either side connected by many cross channels making a plexus around the aorta. Dorsal to the heart, the right duct crosses over and joins the left. Dorsal to the kidneys, the two ducts unite in a median cisterna chyli. In embryos above 12 em. it was found impossible to obtain in- jections of the intestinal wall through the thoracic duct. This is due to the developing lymph nodes, which at first do not check the flow of the injection mass, but later retard it very much. It has been shown that in human embryos the retroperitoneal sac is changed into the group of preaortic lymph nodes extending from the celiac axis to the bifurcation of the aorta, and this change is being made in an embryo 8 cm. long. A secondary, larger group of lymph nodes forms in the center of the mesentery, along the superior mesenteric artery, and this group is also being formed in the same embryo. The early lymph nodes, however, consist of a great plexus of wide lymphatic ducts with very few follicles and hence injection through them is easy. By the time the pig is 12 cm., however, the injection mass passes to the nodes, and increased pressure results in an extravasation at the node. For these stages, therefore, it was found necessary to inject into the wall of the intestine itself. To get good injections by this method it was found best to pierce both muscle coats, and thus have the needle just enter the sub-mucosa and inject slowly under low pressure.
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