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=THE ANATOMICAL RECORD=
 
=THE ANATOMICAL RECORD=
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EDITORIAL BOARD
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Irving Hardesty
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Tulane University
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Clarence M. Jackson
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University of Minnesota
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Thomas G. Lee
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University of Minnesota
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Frederic T. Lewis
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Harvard University
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Warren H. Lewis
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Johns Hopkins University
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Charles F. W. McClure
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Princeton University
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William S. Miller
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University of Wisconsin
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Florence R. Sarin
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Johns Hopkins University
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George L. Streeter
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University of Michigan
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G. Carl Huber, Managing Editor
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1330 Hill Street, Ann Arbor, Michigan
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VOLUME 11 AUGUST, 1916-JANUARY, 1917
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PHILADELPHIA
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THE WISTAR INSTITUTE OF ANATOMY AND BIOLOGY
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COMPOSED AND PRINTED AT THE
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WAVERLY PRESS
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By THE Williams & Wilkins Company
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Baltimore, Mi>., U. S. A.
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==Contents==
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===No. 1. AUGUST===
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Eliot R. Clauk. A study of the reaction of mesenchyme cells in the tad-pole's tail toward injected oil globules. Five figures
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Louis H. Kornder. An anomalous urinogenital system in a dog. Two figures 10
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W. SoHiER Bryant. Sensory elements in the human cerebral hypophysis. One figure. 25
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Emily Ray Gregory. A method for micro-injection. One figure 29
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===No. 2. SEPTEMBER===
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William B. Kirkham. The prolonged gestation period in suckling mice 81
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Helen Dean King. On the postnatal growth of the body and of the central nervous system in albino rats that are undersized at birth 41
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E. L. JuDAH. Mounting specimens under Petri dishes and clock glasses 5.
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===No. 3. OCTOBER===
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P. E. Smith. The effect of hypophysectomy in the early embryo upon the growth and development of the frog. Ten figures 57
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Carbon Gillaspie, Lewis I. Miller and Morris Baskin. Anomalies in lobation of lungs with review of literature. Five figures Go
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Carbon Gillaspie, Lewis I. Miller and Morris Baskin. Anomalous renal vessels and their surgical significance. Nine figures 77
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Osc.\R Riddle. Size and length relations of the right and left testes of pigeons in health and disease S7
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J. A. Long. Hygienic cages for rats and mice. Two figures 103
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===No. 4. NOVEMBER===
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Alwin M. Pappenheimer. The Golgi apparatus. Personal observations and a review of the literature. Twenty-two figures 107
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G. Carl Huber. On the form and arrangement in fasciculi of striated voluntary muscle fibers. Four figures 149
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G. Carl Huber. A note on the structure of the elastica interna of arteries. One figure 169
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G. Carl Huber. A note on the morphology of the seminiferous tubules of birds. One figure 177
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===No. 5. DECEMBER===
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P. G. Shipley and R. S. Cunningham. The histology of blood and lymphatic vessels during the passage of foreign fluids through their walls. II. Studies on absorption from serous cavities 181
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Wilbur C. Smith. A case of a left superior vena cava without a corresponding vessel on the right side. Two figures 191
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G. S. Hopkins. The innervation of the muscle retractor oculi. One figure 199
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Eben Carey. The anatomy with especial consideration of the embryological significance of the structures of a full-term fetus amorphus. Nineteen figures 207
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Rat Henry Kistler. The thoracic duct in the rabbit. Six figures 233
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Franklin P. Reagan. Some results and possibilities of early embryonic castration. Six figures (four plates) 251
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Helen Dean King. The relation of age to fertility in the rat. Three figures. ....... 269
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Technique Notes. I. The application of Benda's neuroglia stain. II. Some uses of Mallory's connective tissue stain. By H. M. Kingbry. III. The use of the Van Wijhe method for the staining of the cartilaginous skeleton. By Gustave J. Noback. IV. A convenient method of orientation in paraffin imbedding when paper trays or boxes are used. By B. F. Kingsbury 289
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K. Okajima. On the elective staining of the erythrocyte 295
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J. B. Johnston. Neutral red as a cell stain for the central nervous system 297
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===No. 6. JANUARY===
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H. H. Donaldson. Biological Problems and the American Association of Anatomists.
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Address of the President at the Annual Meeting 299
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Proceedings of the American Association of Anatomists, Thirty -third session 311
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Proceedings of the American Association of Anatomists. Abstracts 317
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Proceedings of the American Association of Anatomists. Demonstrations 439
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American Association of Anatomists, Constitution 446
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American Association of Anatomists, List of officers and members 449
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Proceedings of the American Society of Zoologists, Fourteenth Annual Meeting 467
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Proceedings of the American Society of Zoologists, Abstracts 473
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American Society of Zoologists, Constitution 543
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American Society of Zoologists, By-laws 545
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American Society of Zoologists, Historical Review 546
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American Society of Zoologists, List of officers and members 570
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==A study of the reaction of mesenchyme cells in the tad-pole's tail toward injected oil globules==
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ELIOT R. CLARK
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From the Anatuniical LdJtoralonj uf the University of Missouri
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FIVE FIGURES
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In their later studies on the mode of development of the lymphatic system, Huntington and McClure have reiterated the view that the hnnphatics are formed by the transformation of mesenchyme cells in the following manner. They hold that fluid accumulates in the tissue spaces, forming small lakelets; that the mesenchyme cells are pressed upon by the fluid collected: and that, as a result of this mechanical pressure, the mesenchyme cells react by flattening out and by forming membranes around the blisters.
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The main evidence presented in favor of this view consists in the finding, in microscopic sections, of clear spaces in the tissues, unsurrounded by any membrane, of other spaces which have the appearance of being partly surrounded, and of others with a complete covering. Some of these appear to be completely isolated while others are connected with one another. The numerous possibilities of error in the interpretation of the appearances described have been pointed out by Miss Sabin (1), E. L. Clark (2), and mj^self (3), and will not be reviewed at this time.
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The importance laid by these investigators on the part played by the mechanical action of fluid on mesenchyme cells may be seen from the following quotation. Huntington (4) says (page 289):
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^ A preliminary report of these studies was published in the Proceedings of the .American Ass. of Anat., Anat. Rec, vol. 10, no. 3, 1916, p. 191.
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If two embryonal mesenchymal cells are separated from each other by the accumulation of fluid in the resulting intercellular space, then the opposing aspects of the two cells involved will be subjected to the mechanical and hydrostatic influences of accumulated intercellular fluid, which will react upon the surfaces of the cell still held in syncytial relation to the surrounding mesenchytue. The cells whose opposing surfaces have become freed by the development of an intercellular space, and are subjected to fluid pressure, will react as a whole, become flattened, and be transformed into endothelial cells, forming the parietal limit of an originally intercellular mesenchymal space, which is the font and origin of all vertebrate vascular development.
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This should be supplemented by a quotation from McClure (5), for no explanation is given here for the accumulation of fluid in the tissues.
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As soon, however, as the haemal vessels begin to function, lymph begins to collect in the intercellular spaces of the embryo and, as we know, is subseciuently collected by a set of newly formed vessels, the l>nnphatics, which convey it to the venous circulation.
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Those who maintain that the lymphatics sprout centrifugally and continuously from the veins, would necessarily hold that the lymph in the intercellular spaces patiently awaits the arrival of close and hollow outgrowths from the veins, the lymphatics, before it can be received into any portion of the lymphatic system.
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The combined picture, then, is as follows: as soon as the blood-vessels function, "lymph begins to collect in the intercellular spaces." It collects apparently because it cannot get back into the blood-vessels. The mesenchyme cells are subjected to "mechanical and hydrostatic pressure" exerted by the accumulated lymph, and respond by flattening out, and becoming endothelial cells. This comprises the "font and origin of all vertebrate vascular development."
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Some of the theoretical objections to this series of assumptions may be pointed out.
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First, McClure states that "as soon as the haemal vessels begin to function, lymph begins to collect in the intercellular spaces of the embryo;" to the pressure exerted by this accumulated lymph is assigned the role of acting as the formative stimulus for t lie t r;iiisl()ruuiti(<ii of iiicsoucliyuK' colls into lyiuphatics. Ill allot li(Mi)Iace, McC 'liii('arfi;ues that blood-vessels and lymphatit's differ(Mitiat(> in the same manner. Now if the collection of int(M'c(^llular lyiii])li l)ep;ins only after the blood-vessels be^iii to fiiiictioii, how are we to exi)lain the collections of fluid wJiich ai-e su])])osed to have served as a fonnative stimulus for the (liflereutiatioii of blood-vessels? Afi;ain, if both bloodvessels and lymi)liatics ai'e determined solely by the action of mechanical pressure, we are left with the same puzzling problem which confronted Goette (6) in 1873, namely, wliy the two sets of vessels do not everywhere form comnumications with one anotlu^r. The ])uzzle ditfers only in that (Joette conceived of the circulation fi'om blood to lymph capillary as beinj;- intracellular, and tried to explain why the same mesenchyme cell was not sometimes demanded by both the blood-vessel and the lymphatic, while Huntington and McClure conceive of the fluid as being extra-cellular and must explain why it does not happen that the same lakelet does not connect with a blood capillary on the one side, and with a lymphatic capillary on the other. Goette's puzzle has been eliminated by the proof that in the tail of amphibian larvae, mesenchyme cells take no part in the growth of blood or lymphatic vessels.
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Again, it is hardly necessary that "those who maintain that the lymphatic sprout centrifugally and continuously from the veins, would necessarily hold that the lymph in the intercellular spaces patiently awaits the arrival of closed and hollow outgrowths from the veins, the lymphatics," since it was demonstrated clearly by Magendie, over a hundred years ago, and has been proven so many times since that it cannot be questioned, that absorption of substances may take place through the bloodvessels as well as through the lymphatics.
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Another assumption which is quite unwarranted on the basis of facts is that with an increase in intercellular fluid, this fluid will collect in definite lakelets, in the intercellular spaces. All our knowledge of intercellular spaces indicates that they form an irregular, intercommunicating net-work of spaces filled with fluid. It would be expected that, if this fluid were increased, the increase in the intercellular spaces would be a general one, the distention, or edema, being regulated in its intensity only by the relative amount of resistance offered by the tissue. This resistance would, of course, be greater in dense tissues, in which the cells are bound firmly to one another, and less in tissues where cells are less firmly attached to one another. In any region, however, where the tissue is uniform the pressure and separation of cells must be uniform, and no accumulation of tissue fluid in the form of lakelets would seem to be possible. Microscopic examination of embryos bears out this theoretical consideration; there are regions in which the intercellular fluid is large in amount, proportionately, and others where it is small. "VMiere large, the mesenchyme or other cells are uniformly separated from one another, as for example in the umbilical cord, and the ventral body wall.
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It is unjustifiable from our knowledge, or better, lack of knowledge, to speak positively about the stimuli which are responsible for the primary differentiation of organs or tissues, since with, perhaps, the exception of the formation of the lens as the result of the contact of the optic cup with the epidermis (7), there is hardly an instance in embryology of the satisfactory demonstration of the stimulus responsible for the primary differentiation of any organ or tissue (8). In this connection, results obtained in recent studies made on tissues, grown by the 'tissue culture' method of Harrison, are interesting. Harrison (9) found that primitive nerve cells send out processes into a medium consisting of coagulated lymph. The Lewises (10) obtained similar results with sympathetic nerve cells, in a medium of Locke's solution. Shipley (11) has found that undifferentiated heart muscle cells differentiate and start rhythmic contractions, in a medium of coagulated plasma. In all of these cases the cells were removed from their normal environment. Their continued development makes one sceptical of hypotheses as to the nature of formative stimuli, when such hypotheses are not supported by fact.
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The only experimental evidence which has been proposed in support of the hypothesis that collections of lymph furnish
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MESENCHYME TELLS IN TAD-POLE S 'I'AIL O
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the sliiiiuhis to the lonnatioii of lyin])hati('s, consists of tiie results of studies oil 'ex])('i'iiii('iital niosotlieliiini,' l)y W. C. Clurk (12). This iiivestigiitor found tlint if solid blocks of celloidin are ])la('ed in tlie suheutaneous tissue of dogs, they became surrounded i)y flattened cells, which show, when treated with silver salts, black intei'celhdar lines tj^pical of flattened mesotlielial or (>])ithelial tissues. Solid globules of hard ])araflPin, injected into the cornea of rabbits, were surrounded l)y a layer of flattened cells. Flattened cells were also found to line 'dead spaces' in the tissue and artificial channels, such as may be induced by the ligation of the cystic duct, with formation of a mucous fistula.
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Interesting as are these studies, in spite of the absence of evidence as to the source of origin of the cells which formed the flattened lining membrane, there appears to be no justification for the conclusion that we have thereby gained any information as to the mode of differentiation or growth of blood or lymphatic vessels. And 3'et, W. C. Clark concludes (page 316) that "Therefore the second hypothesis, premised in this article, is tenable, namely that the flat cells of serous surfaces and those lining blood vessels may regenerate from deep connective tissue cells, and do not necessarily arise from adjacent intact mesothelial or endothelial cells." Again ^McClure (13) refers to the results of W. C. Clark as bearing out in a most decisive manner" the view that "the gradual increase in the amount of lymph received by the subocular sacs (in the trout) during the stage of their independence, results in the application of a constant and continuous pressure to the mesenchyme cells forming their walls, which in itself must be a positive factor in causing these cells to flatten out and gradually assume an endothelial form."
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It is difficult to conceive how the results obtained by W. C. Clark can have anj^ bearing on the problem which confronts McClure, unless the assumption is made that all mesothelial, endothelial, and epithelial membranes which line spaces or ducts have the same properties — an assumption for which facts furnish no justification. Surely blood-vessel endothelium has
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6 ELIOT R. CLARK
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properties which differ from synovial membranes, peritoneal membranes, or the lining membrane of the urethra, the bile ducts, or the gall-bladder.
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There is, perhaps, a suggestion of support for the hypothesis in question in some of the results obtained in tissue cultures. Several observers — Harrison (14), M. R. and W. H. Lewis (15), Lambert (16), W. C. Clark (17) and others — have found that in bits of tissues, explanted to plasma or Locke's solution, membranes may be formed around solid bodies — such as along the cover-slip and around solid threads, as the threads of spider web, used by Harrison, and around droplets of fluid — such as may be formed, occasionally, in plasma preparations, by the retraction of the fibrous threads. The explanation of the formation of membranes around droplets of fluid is associated with the apparent inability of cells to grow into a purely fluid medium, without mechanical support, or as expressed by Harrison, their dependence on 'stereotropism.' The formation of such a membrane in tissue cultures is not to be interpreted as a reaction by flattening out, on the part of the culture cells, but rather, in all probability, as due to the fact that cells grow around the periphery of such a droplet. Moreover, it has not yet been possible to determine the origin of the cells which form membranes in tissue cultures. At present there appears to be no ground for claiming that the formation of membranes in this manner, furnishes us any information as to the mode of differentiation of blood-vessel or lymphatic endothelium.
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In order to put to the test the hypothesis that the differentiation of blood or lymph vessel endothelium may be stimulated by the mechanical pressure exerted on mesenchyme cells by accumulations of fluid, and to plan the test in such a way as to make it approach as nearly as possible the actual conditions supposed to exist, the present studies were started. The aim was to inject an inert fluid, in globules of size sufficient to press against the mesenchyme cells, into a region of embryonic tissue, in which the reaction of the cells could be watched in the living animal; to see whether a membrane were formed, and, if so, by what type of cell, and what would be the properties of such a
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MESENC'in.Mi; CELLS IN TAD-POLES TAIL 7
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monihrano, osperially its roaction towanl hlood-vosscl and lyin])liati(' (Mulothcliuin.
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In ordci- to simulate* as nearly as ])()ssU)l(' tiic (liiid wliose })resen('e is thought to excite the traiisfoi'niatioii of inesenchyme cells into lyin])hatics, and at the same thne to have a fluid which would he inert, wliicli would not be absorbed, which w(.uld n.erely exert a iniH'hanical pressure, paraffin oil was selected for injection. The ol)ject chosen was the trans])arent fin expansion of the tail of youn^" fi'oji and toad tad-poles wliere it is possible to see the indi\idual mesenchyme cells, as well as blood-vessels, lym]ihatics and leucocytes, and to watch their reactions in the living- larvae, from qIsly to day. The tad-poles were anaesthetized with chloretone (1 : 4000 to 1 : 5000) and small globules of oil injected into both fins, through fine glass cannulae, under the binocular microscope. The oil was sterilized by heating. In some cases the tadpole was washed in several changes of sterile w^ater, but the results did not difTer materially from those obtained when the only antiseptic precaution consisted in sterilizing the oil. The observations were made by a method previously described in detail (18) — the larva, anesthetized, was placed in a micro-aciuariiun, in chloretone of the proper strength, and the tube of the microscope tilted to the horizontal, to enable the tadpole to retain its normal upright position. The oil globules w^ere of varying sizes, from 20 to 100 micra in diameter.
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The larvae used were those of Rana catesbiana (bull-frog) and of Fowler's toad (Bufo lentiginosus Fowleri). The latter have beautifully clear tails, with very few pigment cells at the stages used, while the mesenchyme cells are far apart and stand out most clearly.
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Since the oil could not be injected without a certain amount of injury, there were alwaj^s some temporary effects of the injection, not attributable to the presence of the oil. These consisted principalh' of a more or less intense leucocytosis, probably caused by greater or milder degrees of infection. In some cases large numbers of leucocytes gathered about the globules, many of the leucocytes containing pigment. Several of the globules, around which the leucocytosis was most intense
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ELIOT R. CLARK
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.mi:si;n(Iiv.mi: cklls in tad-i'olk's tail 9
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wore oxtnidcd. In raso tliis did not occur, tho loiicocytosis firadiially subsided luitil in most cases tlu'cc or four days after the injection, the i-ej2;ioii surrouii(lin},j; tlie globule contained no more leucocyt(>s than the other ])arts of tlie tail. In some mstances, a sH^litly increased number of leucocytes near the p;lobul(^ continued for several days longer. From now on the globules remained ai)i)arently inert, so far as could be judged from the beluuior of leucocytes in their vicinity. The longest time over which a globule was watched was 12 days.
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In a])])(\ii'ance the oil globules, wh(>n "[iresent in the tail, form sphei'es with the central portion clear and transparent and a dark periphery, with a sharp outline. Structures over the central portion can be seen most clearly. Thus it is possible, in case the diameter of the globule is sufficient to distend the skin slightly, to see the nuclei of the cells of the epidermis, and the details of other structures most distinctly. Many of the globules w^ere oval in shape, immediately after injection. Later, after a day or tw^o, they usually rounded up to a spherical shape though sometimes remaining slightly oval.
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The beha^'ior of the mesench\nne cells will now be described. In order to follow them with accuracy camera lucida records were made of all mesenchyme cells in the neighborhood of the globules, and their changes from day to day were noted. When a mesenchyme cell happened to be in the outer dark zone of the globule, it was difficult to make out its outlines. Occasionally only one or two of its processes could be clearly seen. The following of such cells, however, was made possible by the fact
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Fig. 1 From larva of Fowler's toad. 9 nun. long. Four globules of paraffin oil injected into fin expansion of tail, on Aug. 19. .Much leucocytosis about three of them, and all three extruded within forty-eight hours". There was very little leucocytosis aljout the fourth globule, of which three drawings are shown, two, four, and six daj-s after injection. This globule was in the ventral fin. The mesenchj-me cells in the immediate vicinity of the globule are shown. The letters, a, b, etc., indicate the same cells. * leucocytes against the globule; pig.L., pigmented leucocyte against the globule. The mesenchyme cells were in three different planes; those nearest the observer are represented in solid black, those furthest away are dotted, while those in the midst are cross-hatched. Enlarged 267 times. Drawn with camera lucida.
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10 ELIOT R. CLARK
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that the globules from time to time shifted their position slightly, so that a cell, at one time not clearly seen, later could be clearly outlined. This applied to only a very few cells, particularly in the toad larvae, because of the relative rarity of their mesenchyme cells. In view of the descriptions in the literature of the reaction of connective tissue cells to the pressure exerted by foreign substances, the behavior of the mesenchyme cells was a great surprise. It was expected that the cells near the globule would flatten out on its surface and form a membrane. On the contrary, the mesenchyme cells apparently paid no particular attention to the globules. They maintained their identity as 'star-shaped' cells, with thickened central portion and branched processes, and their property of slow progression, described in an earlier paper. That the mesenchyme cells are not influenced by the pressure exerted by the globules was brought out quite strikingly in one instance in which the globule shifted its position in such a way as to come to lie against a mesenchyme cell which had been at a slight distance from the globule. For a day or two it was rather difficult to make out the outlines of the cell. The globule then shifted its position in the opposite direction, and the mesenchyme cell could now be seen clearly, apparently unchanged, at a slight distance from the globule. Occasionally there are to be seen, over the clear part of the globule, if the globule is of sufficient size to distend the skin slightly, one or two flattened cells which, at first glance, might be interpreted as cells flattened out by the pressure of the globule. Such cells were seen over only a few of the globules, and their explanation was obvious on studying other parts of the tail, at a distance from the globules. Such flattened out mesenchyme cells appear more or ess evenly distributed, lying just below the epidermis, and are not particularly associated with the oil globules. That they are associated with the skin and not with the oil globules, is shown by the fact that none are present over the majority of the globules, and also by the fact that, if the globules shift, these cells maintain the same position with reference to the skin, while they are left behind by the oil globules (fig. 4). There are no other appearances on
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MESENCllVMK CKLLS IX TAI)-1'()LK S TAIL
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11
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Aug. 4
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Fig. 2 From larva of Rana catesbiana, 10.5 mm. long. Two globules of paraffin oil injected into the ventral, and a small globule into the dorsal fin; all three remained. ]\Iuch leucocytosis around each. The first drawing (Aug. 3) was made immediatel}- after the injection. Small mass of cellular debris is shown, in the path of the injection. Mesenchyme nearest the globule lettered as in fig. 1. In drawings Aug. 4, 5, and 7, some of the leucocytes about the globule are shown. In drawing of Aug. 5 are shown the successive positions taken by a leucocyte as it moved to the globule, moved along the surface of the globule a short distance, and then moved away. The shifting of the globule is clearh" seen, by comparing its relation to the blood vessel. The cells fZ and e, which lie close to the globule Aug. 3 and 4, are left behind, while the cells b and c are approached by the shifting of the globule.
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Enlarged 267 times. Drawn with camera lucida.
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12 ELIOT R. CLARK
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the part of the mesenchyme cells which even remotely suggested a flattening out. The 'act that some of the globules watched shifted their position would also indicate that no surrounding membrane had beeii formed, for a membrane would prevent such movement.
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The behavior of wandering cells towards the globules was watched with interest. As already stated there was a leucocytosis of greater or less intensity following the introduction of the globules, for the first two or three days. Leucocytes, most of them small and clear, others larger, and containing pigment, collected around the globules. Many of them flattened themselves out on the surface of the globule, or formed irregular humps on the profile. Occasionally such a flattened leucocyte formed a thin, circular structure, with nucleus visible over the clearest, central portion of the globule. Such a cell, coupled with the irregular humps on the profile, if seen only at one stage, and not followed, might well give the impression of membrane formation by wandering cells. When, however, such cells were followed, it was seen that they gradually moved away (fig. 2). The humps on the profile changed shape, with each drawing, even when the records were made several times daily, while the flattened cells on the clearer part moved away. After four or five days most of the globules were quite free from the presence of leucocytes. In order to be sure of the behavior of the wandering cells, some were watched intensively. They were seen to move, w th the typical amoeboid type of progression, up to the oil globule, to flatten themselves out on its surface and again move away. The impression was gained, that, had sections been made at a time when the leucocytes were flattened on the globule, they might have been interpreted as forming a membrane, an interpretation which the study of the living shows would be quite unjustifiable.
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Chromatophores which are present in large numbers in the dorsal fins of bull-frog larvae and to a somewhat less extent in the ventral fins of the same larvae, sometimes wrapped around the globules with their long branched processes, when the glob
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MESENCHYME CELLS IN TAD-POLE's TAIL 13
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ulcs wvw iiij(>('t(Ml Ileal' llicm, hut did iiol I'onii ;i dcfinito UKMiibruiie.
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Since the inosonchyiiio colls fnilod to form a ineinbnme around the ^l()i)ulcs, the second ]xirt of tlu^ inciuiry, namely, the reaction of such a membrane, if foinuMl, toward the lymphatics or bloodvessels in their vicinity, could not be followed. It was of interest, however, to observe the reaction of blood-vessel and lymphatic endothelium to the oil globules. In one case a particularly faA'oi'able o])])ortiuiity was afforded to study the reaction of the bl()od-ca])illaiy, since the sl<^^^ule pressed against a blood-capillary, forcing it to make a bend in its course (fig. 4).
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Fig. 3 From same Rana catesbiana larva from which figure 2 was taken. To show relation of pigment cells to globule. This small oil globule was injected into the dorsal fin, on Aug. 3; the drawing was made Aug. 5.
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Enlarged 267 times. Camera lucida.
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The capillary then appeared to wrap around a part of the globule. In this position the capillary showed no tendency to give ofT cells which might grow around the globule, but instead remained as a distinct vessel. The circulation of blood cells through it, which was at first interrupted, was later resumed. Blood-capillaries and lymphatics near the oil globules showed no tendency to grow toward it, or to send out sprouts to it.
 +
 +
The results of this study, then, indicate that, aside from the temporaiy inflammatory reaction, due probably to the injury and to the bacteria introduced at the time of injection, the presence in the fin of the tad-pole's tail of injected globules of paraffin oil, of sufficient size to cause a distension of the tissues, fails
 +
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14
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ELIOT R. CLARK
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to stimulate the formation of membranes about the globules, on the part of mesenchyme cells, wandering cells, or of bloodvessel or lymphatic endothelium.
 +
 +
These results are in disagreement with those of W. C. Clark (19), already mentioned. They are, however, in agreement with the older findings of E. Juckuff (20) who found that soft paraffin, injected subcutaneously, travelled long distances from the injection site, showing that no membrane was formed about
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Aug. 20
 +
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Fig. 4 Globule of paraffin oil injected into ventral fin of tail of Rana catesl)iana larva on Aug. 13. The globule rested against a blood-capillary, forcing it to bend slight!}'. The two sketches shown, made four and seven days respectively after the injection, show the relation of the globule to the blood capillary, to a nearljy lymphatic, {lijm) and to two mesenchyme cells which happened to be under the epidermis immediately over the globule. Note that the globule has shifted to the right, so that the two mesenchyme cells, which on Aug. 17 are over the right part of the glol)ule, are over the left central portion Aug. 20. Other mesenchyme cells not shown.
 +
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Enlarged 267 times. Drawn with camera lucida.
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Fig. 5 Same larva as figure 4. Small globule injected in dorsal fin on Aug. 13. Drawing made nine days later — on Aug. 22, d.c, pigment cell. Enlarged 267 times. Drawn with camera lucida.
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MKSHXCIIVMK CKM.rt IN TAD-I'OLe's TAIL 15
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it — a liinliii^ \('i-iru'(l hy M;ic( 'alhiiii CJI). Il will he rciiiciiiboretl tluit the uttoin])! was made, in selecting tJ»e substance to be injected, to find sonietiiinfi; which would sinuilate tlie supposed lak(>lets of tissue fhiid wliose presence are lield by Huntinji;ton and Ab'("hire to stimulate, mei-ely l)y the mechanical ])r(;ssure whicli tliey are su])])osed to exert, the mesencliyme cells to form membranes. It is oljvious that in point of size and consistency the small globules of oil nmch more nearly reproduce the supposed conditions than the relatively enormous pus ])ockets, or tlie relatively huge solid blocks of celloidin or the globules of hard ])arafHn. It is also obvious that in a transparent object, like the tad-pole's tail, where the individual cells may be seen with great clearness and the reaction process watched in the living animal, the conditions for observing what happens are much more favorable than in the case of the other experiments referred to.
 +
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Since, then, the action of pressure alone fails to stimulate the formation of membranes on the part of mesenchyme cells, an important link in the argument used in favor of the origin of lymphatics from mesenchyme cells, as presented by Huntington and McClure, drops out.
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 +
A certain feeling of disappointment must be confessed, that the mesenchyme cells failed to respond to the presence of the oil globules by the formation around them of membranes, as it was expected they would, because of the desire to see w^hat would be the reaction of such membranes toward blood-vessels and lymphatics.
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It is true that cells derived from the middle layer or mesoderm differentiate at various stages into pavement epithelium, or endothelium, other than that which lines the blood and l^^llph vascular systems. Among such may be mentioned the lining of the large cavities pleural, peritoneal, and pericardial, the lining of bursae and sj^novial membranes, and the outer layers of tendons and fasciae. It should also be remembered that the same middle embryonic layer differentiates into smooth and striated muscle, into cartilage and bone, into blood cells, and other types of tissues. Each of these tissues has certain
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16 ELIOT R. CLARK
 +
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modes of reaction, a specific life history, the property of responding each in its own individual way, to various stimuli. To transfer the modes of reaction of one set of, tissues derived from the mesoderm to another set is quite unjustifiable. To be more specific, it is not justifiable to claim that, if connective tissues, in adult animals, are capable of forming membranes about solid foreign bodies, or large accumulations of fluid, or if membranes form around liquid vesicles in the midst of coagulated lymph in tissue culture preparations, then lymphatics arise as the result of the pressure exerted by accumulated lakelets of Ij^mph. It is even unjustifiable to transfer to lymphatics the properties of structures which resemble them morphologically so nearly as blood-vessels, for, while there are many points of similarity between the modes of reaction of the two, there are also striking differences.
 +
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It is conceivable that future studies may reveal the various stimuli which are responsible for the primary differentiation of tissues and organs. For the lymphatic endothelium it would seem a more hopeful field to investigate the chemical nature of the intercellular fluid, to see whether any evidence can be gained as to the collection there of especial chemical substances which stimulate its differentiation. To propose such an hypothesis at the present time, however, would be pure speculation.
 +
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Much confusion has arisen because quite different structures have been grouped together under the name of endothelium or mesothelium. It would seem that the time is ripe to separate these different forms of flattened lining cells under different names. If, for example, we could speak of blood-vessel endothelium as Haem-angiothelium, and of lymphatic endothelium as Lymph-angiothelium , or some equally specific names, and if distinctive names could be selected for the other forms of pavement epithelium, much of the confusion would disappear.
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In conclusion, it is a pleasure to express my gratitude to the Marine Biological Laboratory at Wood's Hole, where these studies were made, for generously granted laboratory facilities.
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MKSKXCIIYMK CELLS LV TAI)-1'0LE'.S TAIL 17
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liii:ka'iii{|': ciii;!)
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(1) Sauin, F. R. lOL'J JoliMs Hopkins llosp. U.-p,)ils. Mononiaphs. New
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Series, no. 5.
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(2) Clark, E. L. 11)12 Anal. Wi-c, vol. t>.
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(3) Clauk, K. R. 1911 -Vnal. I{cc., vol. f..
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(4) Huntington, C. S. HU I .\iii. .lour. .\iial., vol. 10.
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(5) McCluue, C. F. \V. I'.llf) Aiiat. ]{rc., vol. <J, no. 4, p. 281.
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(6) CiOKTTE 1875 Die KntAvickclunfisfrcschichte der Unke. Leipzig.
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(7) Lkwis, \V. H. 1907 Am. Jour. .\iia(., vol C, p. 473.
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1913 Storkard Am. Jour. .Vii.it., vol. !.'>, p. 253. 1910 vol. 10, pp. 393-423.
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(8) IIkrbst 1901 Fornuitive Reize in der 'I'hierischen Ontogcnese, Leipzig.
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(9) Hahrison, a. G. 1910 Jour. Exp. Zoo!., vol 9.
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(10) Lewis and Lewis 1912 .\nat. Rcc, vol. 6, j). 7.
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(11) Shipley, P. G. 1910 .Vnat. Rec, vol. 10, p. 347.
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(12) Clark, \V. C. 1914 Anat. Rec, vol. 8.
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1910 Anat. Rec. vol. 10.
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(13) .McCluhe, C. F. W. 1915 Mem. of Wistar Inst., no. 4, p. 29.
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(14) Harrison, R. G. 1910 Jour. E.xp. Zool., vol. 9.
 +
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1911 Science, vol. 34.
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1914 Jour. Exp. Zool., vol. 17.
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(15) Lewis, M. R. and W. H. 1911 Anat. Rec, vol. 5.
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(16) Lambert, R. A. 1912 Anat. Rec, vol. 6.
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(17) Clark, W. C. 1910 loc cit., p. 313.
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(18) Clark, E. R. 1912 Am. Jour. Anat., vol. 13.
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(19) Clark, W. C. 1910 loc cit.
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(20) JucKUFF, E. 1893 Archiv. f. Pathol, u. Physiol., Bd. 32, p. 124.
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(21) Mac C.a.lltjm, W. G. 1903 Johns Hopkins Hospital Bulletin vol. 14. pp.
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5 and 0.
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THE ANATOMIC.M, RECORD, VOL. 11, NO. 1
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AX AXO.MALors rHIXOGENITAL SYSTEM LX
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A DOC
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i>()ris II. K()K\i)i:i{
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From the Ano!o>/iical Lnboralori/ of the A'urthwcslcrn Unu'crsilij Medical School^
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TWO FIGl'KES
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Anomalies of the iiriiiogenital system are frequent and have ceased to attract nmch attention. Few, however, present features of such embryological interest as the following case. Because of this and on account of its value, as illustrating the physiological adaptability of one system to the needs of another, this case seems worthy of mention, ^ly acknowledgment is due Dr. L. B. Arey for valuable suggestions regarding the embryological considerations.
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On opening the abdomen of a dog it is commonly observed that the bladder is large and lies almost entirely in the abdominal cavity. In a medium sized mature female, selected at random for the purpose of obtaining certain tissues used in a research problem, the bladder lay deep in the pelvic cavity and was rather small, the size and shape being that of a walnut. On palpation it felt extremely firm, much as though it were a solid mass of tissue. A longitudinal incision through its wall revealed but a very small lumen, less than 1 cm. in diameter and 2 cm. in length.
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Two broad ligaments passed from this bladder over the rectum and gained attachment to the front of the sacrum. One ligament was considerably longer than the other, due to the bladder lying ventral and to the left of the uterus instead of directly ventral as is normally the case. Except for these two ligaments and a slightly shortened urethra which merged into the left wall of the urinogenital sinus, other connections with the bladder could not be established.
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1 Contribution Xo. 40, :May 15, 191(3.
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19
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20 LOUIS H. KORNDER
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These findings led to an examination of the kidneys, which were found to be normal in shape, size and position. Each possessed one short ureter, the right being 6.5 cm. and the left 7.2 cm. in length.
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Originating in the pelvis of the kidneys the ureters coursed downward over the psoas muscles and passed one on each side into the horns of the bicornuate uterus. This union occurred about a centimeter below the place where the short Fallopian tubes merge into the uterine horns. The uterine horns and the uterus were not soft and pliable as is usual but were hard and rigid and on making a longitudinal incision through their walls, were found to be filled with debris, composed mainly of desquamated epithelial cells.
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The structures in this region were surrounded and some deeply imbedded in a mass of fibrous and adipose tissue. This appeared to form a common capsule which covered the union of the ureters with the uterine horns and extended over the Fallopian tubes including the ovaries, becoming at this connection part of the ovarian bursa.
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The ovarian bursa exists normalh' in the dog as a separate fold of peritoneum covering each ovary. This is usually covered by adipose tissue but opens through a small slit-like opening into the abdominal cavity.
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It has been mentioned that the urethra was sUghtly shorter than normal and passed from the left into the wall of the urinegenital sinus. This relation of the urethra to the urinogenital sinus and the original location of the bladder explains why in the accompanying figure (fig. 1) the bladder is shown as lying between the uterus and rectum, instead of ventral to the uterus as is normal.
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Histological preparations of the bladder show a slightly changed epithelial lining, consisting in two to three layers of low cuboidal epithelium. The uterine surface epithelium instead of being high columnar in type is pseudo-stratified. The deeper glandular epithelium, however, is the same as in a normal dog's uterus. The ovaries which were deeply imbedded in their ovarian bursae on sectioning showed nothing atypical, with the exception of
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ANO.MAI.ors IKIXOdKM TAL SYSTEM IN A DOG 21
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ureter
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\ XjOvarian bursa.
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entranceop ureter
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liter ine hornopened
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opt . uterus ,
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Lateral/ tiaaynent- y of the Ucbdder, .^ bUdde.r..jjf- -^j|^\
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cervix.. ■
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uretkrou.
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recuum ji_.jr__»a_
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vcuamcL. i* |fl^
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^^ IT
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Fig. 1 \'entral view. Ureters shown as they enter both horns of uterus. Ovarian bursae and upper end of uterine horns opened. Bladder small and abnormal in location. 22 LOUIS H. KORNDER
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a slightly more fibrous stroma than is usual. Several large Graffian follicles present indicated a normal functional activity of the ovaries. Histologically, then, practically nothing unusual exists, the anomaly being one of gross anatomy, this consisting in a union of the urinary with the reproductive tract, the fusion of ureters and uterine horns leaving the bladder as a cul de sac which leads through the urethra into the urinogenital sinus.
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E.AIBRYOLOGICAL CONSIDERATIONS
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The structures in^'olved here are embryological derivatives of the mesonephric ducts, metanephros, jNIuellerian ducts and cloaca. That the cloaca developed normally is indicated by the presence of a rectum, bladder, urethra and urinogenital sinus. The presence of the uterus, tubes and vagina likewise indicate the normal development of the Muellerian ducts. The anomaly then must be due to a defective embryological growth of the mesonephric ducts and their derivatives, with a deficiency in the development of the nephrogenic cord as a possible causal stimulus.
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In pig embryos of approximately 5 mm. length the mesonephric ducts give rise to the ureteric anlage of the metanephros where the ducts bend to join the cloaca. But that the ureteric anlages do not always originate at this bend is indicated by the frequency of double or triple ureters. In these instances the first ureteric bend develops usually into the ureter most normal, while the rest show evidence of slowed or mal-development. From these cases it may be assumed that the ureteric anlage need not necessarily arise at a definite location but can occur at any point along the ducts.
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Figure 2, a very diagrammatic sketch, shows both the Muellerian and Wolffian ducts leading into the urinogenital sinus. The approximate position where normally a single metanephric anlage arises from the Wolffian duct is indicated by (A). However, in man as many as six such anlages have been observed. While in these instances the anlage corresponding to (A) de^'elops into the adult ureter the possibility exists that a more
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ANOMALors ri{INOGENITAL SVSTKM IN A l)()(i
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23
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cranial aiilajio. Toi- iiistaii('(> (/>'), may hccdiiic llic functional urct(>r.
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Kct'crcncc to figure 2 will show tlic lowci- i)art of the Wolffian duct not cross-hatched. Tiiis ])()rtion which extends from the normal ureteric anlaj>;e on downwards is durijij;' further development drawn into the urin()f2;enital sinus. Thi'ouf^h this fusion the uieteis ivceive theii' normal connection with tJie definitive bladder which develo])s ])ai1ly out of this portion of the sinus.
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Fig. 2 Diagrammatic sketch. M, Muellerian duct; W, Wolffian duct; A, location of normal metanephric anlage; B, possible upwardly displaced metanephric anlage; D, extended ureter merging into the Muellerian duct at [/; N, portion of Wolffian duct taken into wall of urogenital sinus; UgS., cross-hatched portion of Wolffian duct degenerates.
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If in this particular case, however, the ureteric anlage did not develop low enough to be included in that lower portion of the mesonephric duct then just as soon as the normal degeneration of the upper part of the Wolffian duct occurred, the upwardly displaced anlage (B) which developed into the ureter was without connection with the urinogenital sinus. Being thus isolated it seems probable that the ureter (D) extended to the nearby Muellerian duct and merged into it at (U). This established an outlet into the urinogenital sinus.
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The above is ofTered as one possibility to which the present anomaly may be due. It is entirely hypothetical as any con
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24 LOUIS H. KORNDER
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sideration of this case must be. Because of this a further explanation may possibly be found in the following. In addition to those known embryological facts mentioned above it should be recalled that in embryos of 8 to 11 mm. length the Muellerian ducts develop caudalward beneath the epithelium of the mesonephric fold. Reference to dissections of the pig embryo show the Muellerian ducts lying very close to the Wolffian ducts so that the occurrence of a more or less complete longitudinal fusion of these two ducts seems not impossible.
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The establishment of this anomalous union of ureters and uterine horns presumably occurred early in the development of the animal. Since in the female, during the normal development the mesonephric ducts degenerate and disappear almost entirely, it may be assumed that in this case these ducts gradually fused with the Muellerian ducts. It may be that in this way an early connection occurred on either side between the Muellerian duct and an upwardly displaced ureter.
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The question may be raised why a metanephros thus displaced should have abandoned the mesonephric duct and appropriated a new outlet by way of the Muellerian ducts? It may be that the functional need of maintaining the patency of the mesonephric duct was ineffectual compared with the tendency toward atrophy and consequent occlusion which the cranial portion normally shows. This query becomes all the more pertinent in view of the recent report by Bremer (Jour. Anat., vol. 19, '16) that in the cat the mesonephros maintains its activity until the permanent kidney assumes the excretory function. This being true of the cat it is more than probable that it also exists in the dog since both belong to the Carnivora.
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SENSORY ELEMENTS IN THE HUMAN CEREBRAL
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HYPOPHYSIS
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w. S()hii:r hhvaxt
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ONE FIGURE
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111 the i)ast few years, the ^rt^ater i)art of the work relating to the cerebral hy])()])]i>'sis has been of a therapeutic, a clinical or a surgically experimental nature, and the interest aroused in these aspects of the pituitary has tended to obscure the fact that certain histological elements in the structure of the organ still remain a mystery. The following report re-introduces the subject of the sensoiy elements of the hypophysial cavity, of which I have made a careful examination in human specimens : These sensory elements occur in maculae, which, in sagittal sections of the pituitary are seen situated on the posterior wall of the cavity, and sometimes, apparently, on the anterior wall. The maculae are composed of tall columnar ciliated sensory cells interspersed with bipolar cells, which have their nuclei towards the periphery; whereas in the ciliated cells, the nuclei are near the base which terminates in a caudal prolongation. Between these caudal processes of the ciliated cells, there is a layer of round cells, resting on a thin basement membrane. An area of ciliated cuboidal cells occurs at the margins of the maculae. I have found these sensory cells in all the freshly hardened human hypophyses that I have examined, except those in which the parenchyma had been almost completely replaced by connective-tissue; the sensory cells are moreover encountered even in pituitaries which have undergone very extensive pathological change. In their gross arrangement, the sensory elements of the hypophyseal cavity are suggestive of the sensory elements of the maculae acousticae.
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Gentes (3), in his examination of the hypophysis of cats and dogs, found in the juxta-nervous layer, a stratified cylinder
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25
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26
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W. SOHIER BRYANT
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Fig. 1 Sensory epithelium of the hypophysial cavity of a human adult. Stained with hematoxylin-eosin.
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This material was procured through the kindness of Dr. William Mabon and the assistance of Dx. Clarence O. Cheney, Pathologist of the Manhattan State Hospital; the work was done in the New York Psychiatric Institute. Special thanks are due Dr. Charles Bates Dunlap for his technical assistance and supervision of the work.
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SENsoKv i:lk.mi;n rs i.\ ckukiucai, ii vi-onivsis 2/
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opitliclimii i-('S('ml)liii (lay and ni^ht. hut much more commonly at night or, according to Long and Mark ('1 1), in tlu^ early morning.
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1 day })()st-/>(tiii(ni. 11" the female white mouse comes in heat, as is the rule during the warmer part of the year (April to October) and the new born j'oung are not suckled, within twenty-four hours after parturition ovulation and pairing (provided a male is present) will occur.
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2 days post-partum. The eggs have been fertilized in the ui)per third of the Fallopian tubes and the first cleavage has occurred.
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3 days post-partum. The eggs are still in the two-cell stage.
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4 days post-partimi. Cleavage is again in progress and morulas of 8 to 10 blastomeres are found at this time.
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5 days post-partum. Morulas of 12 to 16 blastomeres. At the close of this day the morulas develop a central cavity, thus becoming blastulas, and at the same time they pass from the Fallopian tubes into the horns of the uterus.
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6 days post-partum. Blastodermic vesicles lie free in the horns of the uterus.
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7 days post-partum. The blastodermic vesicles are now implanted in proliferated masses of uterine cells which completely obstruct the lumen. The embryos themselves are in the 'eggcylinder' stage, the 'cylinder' almost filling the vesicle and possessing a single, undivided cavity.
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8 days post-partum. The egg-cylinder in embryos of this age has its lumen divided into three cavities.
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9 days post-partuin. The embryo now possesses a medullary groove, which is open except at the extreme anterior end.
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10 days post-partum. Embryos of this age have the medullary groove closed in its anterior half, and for the first time show a heart.
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11 days post-partum. The medullary groove is now closed for more than half its length; the optic vesicles are budding off from the brain; and the auditory vesicles appear as cup-shaped depressions in the ectoderm.
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12 days post-partum. The medullary groove has closed except at the extreme posterior end; the auditory vesicles are almost or
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34 WILLIAM B. KIRKHAM
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entirely closed. The fore limb buds are present, together with the first nephric tubules and the anlage of the liver.
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13 days post-partum. At this age the embryo is a decidedly complex organism, and from this time on the daily changes are rather matters of detail than the appearance of entirely new structures. The characteristic features of this particular stage of development are these: a well developed cranial flexure; optic vesicles completely separated from the brain, invaginated, and showing the beginnings of lens formation. The nasal capsules are visible; the liver has developed into a distinct organ; hind limb buds are present. The embryo has the anlagen of the lungs, and a few pancreatic tubules; the auditory vesicles have withdrawn from the surface and are connected by nerve fibers with the brain. Along the free border of the kidneys, especially at the posterior end appears the genital ridge with a few large cells with large round nuclei, the primordial germ cells, scattered through a much larger number of smaller, epithelial cells.
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14 days post-partum. The eyes have developed to the stage where the lenses have a sohd, clear core. Dense masses of connective tissue foreshadow the future location of the bones of the limbs, girdles, and ribs. Whisker follicles are present; also the semi-circular canals of the ears. The kidneys possess definite boundaries. The nuclei of the red blood corpuscles are smaller and stain less deeply than in earlier stages, while their cytoplasm shows a faint indication of haemoglobin. The first indications of teeth follicles are found in embryos of this age; also the anlagen of the thymus and thyroid glands. The gonads differ from the preceding stage merely in having more of the primordial germ cells in the genital ridges.
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15 days post-partum. Embryonic cartilage cells constitute the most striking characteristic of this stage, clearly differentiating embryos of this age from all younger specimens. Other features are the fewer and smaller blood spaces in the liver, as compared with fourteen-day embryos; the deeper straw yellow color in the cytoplasm of the red blood corpuscles, together with a few which are non-nucleated; the well-developed anlagen of
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GESTATION PERIOD IN SUCKLING MICE 35
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the tooth; aiul tlio prosonce of cartilage cells in the floor of the crauiiiin. Sexual difforoiitiation is present in (embryos fifteen days post-partum, male specimens showing gonads in which follicle formation has already started, wliile the female gonads preserve the earlier condition of prunordial germ cells scattered through a mass of epithoiial colls.
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16 days -post-partum. Differential characteristics now become still more matters of detail and of direct comparison with earlier stages, however, embryos of this age differ from all younger ones in having a decidedly transparent cornea. The heart has assumed its final shape. The pancreas is a clearly defined organ. The nasal capsules open into the front part of the mouth, while the naso-pharynx is connected with both the nasal capsules and the back part of the mouth. The gonads show no marked change from those of fifteen days embryos.
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17 days post-partum. The chief characteristic of embryos of this age is the commencement of ossification around the rib cartilages. Nucleated red blood corpuscles are very scarce. The tongue possesses conspicuous striated muscle cells, stratified epithelium, and at least one circumvallate papilla. The nasal capsules have lost their direct connection with the mouth, but the nasopharynx opens into both the anterior and posterior regions of the mouth. The anlagen of the cartilaginous rings of the trachea are present. The testes have tubules with a peripheral layer of small cells while the larger primordial germ cells occupy the lumen. In the ovaries an ingrowth of connective tissue is noticeable.
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18 days post-partum. Ossification of the cartilaginous skeleton is now the striking feature, and the membrane bones of the upper jaw, hard palate and roof of the skull are also being formed. The testes show a considerable amount of connective tissue between the tubules, while the ovaries differ from those of the preceding stage only in that they project further into the abdominal cavity.
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 +
19 days post-partum. The ribs of embrj^os of this age have an outer shell of bone, and the underlying cartilage is being torn down to make a marrow cavity. There is a marked spongy
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36 WILLIAM B. KIRKHAM
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structure in the lungs; the eye balls show a differential curvature in the cornea and sclerotic; the naso-pharynx no longer opens into the front of the mouth. The testes show no change, but the ovaries are more spherical, and the ingrowth of connective tissue has forced most of the germ cells toward the periphery.
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 +
20 days post-partum. The iris and choroid of the eyes first show pigmentation at this time, and lymphoid tissue appears in the tonsils. One litter of four animals was born twenty days after the birth of a preceding litter and grew to maturity.
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21 days post-partum. The feature of this stage of development is the ossification of the metacarpals and metatarsals. The testes have a well organized tunica albuginea and show less space between the individual tubules than in twenty-day specimens, while in the ovaries the primordial germ cells, or oogonia, are of varjdng sizes, the largest of them beginning to form follicles about themselves.
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22 days post-partum. The embryos in non-suckling white mice have now completed their intra-uterine development, and parturition occurs.
 +
 +
Such, in brief, is the record of development from day to day of white mouse embryos carried by females not suckling young. More extended study and more material would certainly yield many more details, but the above data are sufficient basis for estimating the age of embryos of unknown history, provided onlj'they come from non-suckling females. Also the evidence collected is adequate for comparison with the facts noted below concerning the development of embryos carried by suckling mothers.
 +
 +
THE DEVELOPMENTAL CYCLE IN SUCKLING FEMALES
 +
 +
White mice are able to become pregnant while lactating but when suckling a litter only about one female in five undergoes a complete pregnancy; those which do not complete a pregnancy either failing to ovulate (the majority of cases) or the fertilized eggs developing normally until shortly after implantation in the uterus, when they die and are absorbed (the minority of
 +
 +
 +
 +
GESTATION PEUIOD IN SUCKLINC; MICE
 +
 +
 +
 +
37
 +
 +
 +
 +
cases). In some instances we find auotlier state of alTairs, certain ones of a set of implanted (Mnhryos undergoing normal development wliile others die and are absorbed, a very rare condition in non-suckling females.
 +
 +
Suckling females which arc not going to skip an o\ulation cycle shed their eggs within twenty-four hours of parturition as do non-suckling females. These eggs are then fertilized and divide according to the same time scheme as given above for eggs in non-suckling females, being in the two-cell stage on the second and third days following parturition, morulas on the fourth and fifth days, and blastulas lying free in the uterus on the sixth day post-partum.
 +
 +
Now comes the point of greatest interest in this investigation. The fertilized eggs in non-suckling females, as stated earlier in this paper, become implanted in the uterus at the close of the sixth daj^ post-partum, the fertilized eggs in suckling white mice, on the contrar}'^, lie free in the lumen of the uterus from the sixth to the end of the fourteenth day following parturition. The material on which this statement is based comprises serial sections of the entire uterus of ten females, suckling from three to eight young, killed at various times from the sixth to the fourteenth day post-partum, all of w^hich show normal blastulas lying free in the uterus, with no sign of any reaction in the adjacent cells of the uterine epithelium (table 1).
 +
 +
TABLE 1
 +
 +
Data regarding all suckling white mice from which embryos were obtained from the sixth to the fourteenth day following 'parturition
 +
 +
 +
 +
AGE IN DATS POST-PARTUM
 +
 +
 +
STAGE OF DEVELOPMENT
 +
 +
 +
NUMBER
 +
 +
 +
OF SUCKLING TOUNQ
 +
 +
 +
6
 +
 +
 +
Free blastulas in uterus
 +
 +
 +
 +
 +
6
 +
 +
 +
7
 +
 +
 +
Free blastulas in uterus
 +
 +
 +
 +
 +
4
 +
 +
 +
8
 +
 +
 +
Free blastulas in uterus
 +
 +
 +
 +
 +
5
 +
 +
 +
9
 +
 +
 +
Free blastulas in uterus
 +
 +
 +
 +
 +
3
 +
 +
 +
10
 +
 +
 +
Free blastulas in uterus
 +
 +
 +
 +
 +
5
 +
 +
 +
10
 +
 +
 +
Free blastulas in uterus
 +
 +
 +
 +
 +
8
 +
 +
 +
11
 +
 +
 +
Free blastulas in uterus
 +
 +
 +
 +
 +
6
 +
 +
 +
12
 +
 +
 +
Free blastulas in uterus
 +
 +
 +
 +
 +
4
 +
 +
 +
13
 +
 +
 +
Free blastulas in uterus
 +
 +
 +
 +
 +
4
 +
 +
 +
14
 +
 +
 +
Free blastulas in uterus
 +
 +
 +
 +
 +
5
 +
 +
 +
 +
38 WILLIAM B. KIRKHAM
 +
 +
Here evidently is one, and perhaps the main cause of the prolonged gestation in suckling white mice. Ovulation, fertilization and early cleavage stages occur at the same time intervals in both suckling '• and non-suckling animals, but in the sue ling females the uterine cells will not react to the eggs and enable the latter to become implanted while the mammary glands are withdrawing the surplus nourishment from the parent organism. In support of this statement it should be said that young mice derive all of their nourishment from the mother for the first ten or eleven days after they are born, and thereafter appear to nurse as much and as often as the female will let them, a fact which undoubtedly accounts for the precise number of suckling young exerting a definite influence on the rate of growth of intrauterine embryos.
 +
 +
If only one or two young mice are suckling the development of eggs and embryos proceeds as though none were suckling, but if three or more young nurse the gestation period is lengthened, according to both Daniel ('10) and to the small amount of evidence on this point possessed by the present writer, approximately one day for each animal suckled. This observed fact is, however, very difficult to correlate with the series of embryos obtained from females suckling three to eight young and killed from fifteen to twenty-four days after parturition (table 2) . These embryos show a variation in state of development which appears to vary neither with the number of young suckled nor with the number of embryos carried; in fact it would seem as though in one instance (female killed eighteen days post partum table 2) that the embryos, being as fully developed as in a non-suckling female of the same age post-partum, would have come to birth on the twenty-second day following the previous parturition in spite of there being three young suckling. Even supposing this had happened, were all the facts known such an exception would probably be explainable, since occasional females may reasonably be expected to possess amounts of nourishment far in excess of the average, and some litters of young may start eating grain at an earlier age than usual. More difficult of explanation are such cases as the embryos of a female killed seventeen days
 +
 +
 +
 +
GESTATION I'KKIUl) IN SUCKLING MICE
 +
 +
 +
 +
39
 +
 +
 +
 +
TABLE 2
 +
 +
 +
 +
Data regarding alt suckling white mice from which embryos were obtained from the fifteenth to the tievnty-fourth day following parturition. Embryos labeled {small) tvould probably all have been absorbed
 +
 +
 +
 +
AGK IN DAYS P08T-P\RTUM
 +
 +
 +
DEVELOPMENT EQUAL
 +
 +
TO EMURYOB OP NON-8DCKLINO 9 8 OF
 +
 +
 +
NUMBER OP SUCKLING YOUNG
 +
 +
 +
NUMBER OF EMBRYOS
 +
 +
 +
 +
 +
days p. p. 15
 +
 +
 +
8
 +
 +
 +
8
 +
 +
 +
11
 +
 +
 +
16
 +
 +
 +
12
 +
 +
 +
4
 +
 +
 +
8
 +
 +
 +
17
 +
 +
 +
14
 +
 +
 +
5
 +
 +
 +
2 large + 6 small
 +
 +
 +
18
 +
 +
 +
18
 +
 +
 +
3
 +
 +
 +
9
 +
 +
 +
19
 +
 +
 +
12
 +
 +
 +
7
 +
 +
 +
2 large + 4 small
 +
 +
 +
20
 +
 +
 +
8
 +
 +
 +
6
 +
 +
 +
5
 +
 +
 +
21
 +
 +
 +
8
 +
 +
 +
5
 +
 +
 +
9
 +
 +
 +
21
 +
 +
 +
12
 +
 +
 +
8
 +
 +
 +
9
 +
 +
 +
22
 +
 +
 +
8
 +
 +
 +
?
 +
 +
 +
10
 +
 +
 +
23
 +
 +
 +
12
 +
 +
 +
7
 +
 +
 +
9
 +
 +
 +
24
 +
 +
 +
14
 +
 +
 +
4
 +
 +
 +
6 large + 1 small
 +
 +
 +
 +
post-partum (table 2) which, if, as we have every reason to beheve, they became implanted at the close of the fourteenth day post-partum must in the course of the three days following have undergone a development which in embryos in non-suckling females requires eight days to complete, and this in spite of there being five suckling young.
 +
 +
At the present time work is in progress with a view to explaining, if possible, these apparent contradictions and until that work is completed, which may not be for some time, it does not seem desirable to attempt any further analysis of the facts presented in table 2.
 +
 +
 +
 +
SUMMARY
 +
 +
1. The present work has brought together sufficient data with which to determine, within the possible error of one day, the age of all embryos obtained from non-suckling white mice.
 +
 +
2. Ox-ulation, fertilization, and the early cleavage of the eggs bear the same time relations to parturition and to one another in both suckling and non-suckling white mice except the former are much more apt to skip an ovulation period.
 +
 +
 +
 +
40 WILLIAM B. KIRKHAM
 +
 +
3. Implantation of embryos in the uterus occurs in non-suckling white mice on the fifth day following parturition (provided the female did not skip an ovulation cycle).
 +
 +
4. Implantatioii of embryos in the uterus occurs in suckling white mice, with 3 or more young, on the fourteenth day following parturition (provided the female did not skip an ovulation cycle). In these lactating females the blastulas lie free in the lumen of the uterus from the sixth to the fourteenth day postpartum due supposedly to the activity of the mammary glands.
 +
 +
5. The available material of stages following implantation in suckling females shows no evident correlation with either the number of nursing young or the number of embryos being carried. It also is impossible at present to reconcile the development of these embryos with the observed facts regarding the time of parturition in suckling mice.
 +
 +
6. The conflicting evidence from post-implantation stages in suckling females is at present being subjected to further study.
 +
 +
JUNE, 1916
 +
 +
LITERATURE CITED
 +
 +
Daniel, J. F. 1910 Observations on the period of gestation in white mice.
 +
 +
Jour. Exp. ZooL, vol. 9. King, H. D. 1913 Some anomalies in the gestation of the albino rat (Mus
 +
 +
norvegicus albinus). Biol. Bull., vol. 24. KiRKHAM, W. B. 1907 Maturation of the egg of the white mouse. Trans. Conn.
 +
 +
Acad., vol. 13.
 +
 +
1910 Ovulation in mammals, with special reference to the mouse and
 +
 +
rat. Biol. Bull., vol. 18. KiRKHAM, W. B., AND BuRR, H. S. 1913 The breeding habits, maturation of
 +
 +
eggs, and ovulation of the albino rat. Am. Jour. Anat., vol. 15.
 +
 +
1916 The prolonged gestation period in nursing mice. Anat. Rec,
 +
 +
vol. 11. Long, J. A. and Mark, E. L. 1911 The maturation of the egg of the mouse.
 +
 +
Pub. Carnegie Inst., Washington, D. C.
 +
 +
 +
==On the postnatal growth of the body and of the central nervous system in albino rats that are undersized at birth==
 +
 +
HELEN DEAN KING
 +
 +
The Wistar Institute of Anatomy and Biology
 +
 +
Among the newborn young of various species of mammals there occasionally appear individuals that are undersized and have a very small weight at birth, although they are apparently normal in all other respects. Such individuals are very generally called 'runts' and they are usually discarded by breeders at birth, since it is the popular belief that they never attain normal size and that they are always sterile. Little is known of the true status of these animals, as few attempts have been made to rear them for the purpose of studying their growth processes and reproductive capacity.
 +
 +
In the course of an extensive series of breeding experiments with the albino rat a number of litters have been obtained in which one or more of the rats was very small at birth and weighed much less than the average birth weight for the young of the species, which is about 4.5 grams for the male and 4.3 grams for the female rat (King, '15 a). An examination of these litters some twenty-four hours after birth has shown, as a rule, that the very small individuals were dead, although all of the other members of the litter were alive and vigorous. Thus even under very favorable environmental conditions rats that are much below the average size at birth have little chance, apparently, of surviving even the first few hours of postnatal life; in a state of nature probably very few of them ever live to reach maturity.
 +
 +
In three of the htters examined the undersized young survived the seemingly crucial twenty-four period, and it was found possible to rear them with the other members of the litter and to study their growth in bodj^ weight. In order that the small
 +
 +
41
 +
 +
 +
 +
42 HELEN DEAN KING
 +
 +
individuals might have every possible chance for subsequent growth, the young that were not of the same sex as the smallest member were discarded from each litter. This gave two series of female rats, on6 containing three and the other four individuals; and one series of four males. The rats in the first and those in the third series belonged to the twentieth generation of an inbred strain of albinos in which matings had been made only between brother and sister of the same litter in each generation; the rats in the second series were the offspring of an inbred female (nineteenth generation) and a stock male. For convenience litters with a parentage like that of the second series are designated as 'half -inbred' litters.
 +
 +
The rats in each series were weighed for the first time before they had suckled and then daily for one week. Thereafter the weighings were made at intervals varying from two to fifteen days until the rats were 150 days old w^hen the experiment was ended. In an investigation of this kind it is impossible to obtain the exact body weights owing to the varying amounts of food in the digestive tract at the time that the animals are weighed. To minimize this source of error as much as possible all of the weight records were taken in the morning before the rats had received their daily ration.
 +
 +
Table 1 gives the birth weights of the rats belonging to the three series and also their later body weights at the different ages for which records were taken.
 +
 +
There was considerable variation in the birth weights of the rats in each of the three litters, as is shown in table 1, but in no series was the range of variation as great as that known to occur within the species (King, '15 a). In the first series the heaviest rat (no. 3) weighed but slightly more than the average weight for the female albino rat at birth, yet this weight is 77 per cent greater than the weight of rat no. 1 which had the smallest birth weight that has been found, as yet, in any female rat in our colony. The range of variation in the birth weights of the rats in the second series is much less than that in either of the other series, yet the weight of the heaviest member (no. 4) is 46 per cent greater than that of the smallest member. In the
 +
 +
 +
 +
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 +
 +
a 2
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 +
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93 142 111 162 119 180 125 182
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lO lO o t
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Scries 1 1 (Females) |
 +
 +
 +
Series 2 j (Females) ]
 +
 +
 +
Series 3 J
 +
 +
(Males) 1 1
 +
 +
 +
 +
43
 +
 +
 +
 +
44 HELEN DEAN KING
 +
 +
third series the birth weights ranged from 2.7 grams to 5.3 grams, the largest individual being nearly twice as heavy as the smallest.
 +
 +
All of these rats followed the normal com'se of the growth in body weight, as already determined by Donaldson ('06), in spite of the very great differences in their birth weights. Increase in body weight was very rapid during the early days of postnatal life. The rate of growth dropped off somewhat abruptly at about thirty days, and again showed a marked decline at seventyfive days. After the rats reached ninety days of age the period of rapid growth was ended, and the rats gained relatively little in body weight up to 150 days of age when the weighings were discontinued. With few exceptions the body weights of all individuals fall within the range of variation in the body weights of stock albino rats of like age (King, '15 b, table 3). None of the rats that were unusually small at birth, therefore, could properly be considered as 'runts' when they became mature.
 +
 +
The rats that were undersized at birth never succeeded in attaining a body weight equal to that of the other individuals in the same series at any period of their growth, and at successive weighings the actual weight differences between the individuals that were small and those that were heavy at birth tended to increase. This was true for the individuals in each of the three series, irrespective of sex, as is shown by the data in table 1. At the end of 150 days the rats in each series still maintained the same order with respect to body weight that they had at birth, with the exception of rat no. 3 in the third series. This rat overtook its brother, which had a heavier birth weight, when it was ninety days of age and it subsequently kept the lead in body weight until the end of the experiment.
 +
 +
According to the 'standard' tables for the relation of age to body weight and to body length in the albino rat as given by Donaldson ('15), breeding females should have a body length of 193 mm. and a body weight of 186.1 grams when they are 150 days of age; the body length of a male albino rat of the same age should be 207 mm. and the body weight 218.7 grams. Records for the growth in body weight of a selected series of stock, albino rats reared in The Wistar Institute annual colony under
 +
 +
 +
 +
nUOWTll OF 1U)1)V AND NERVOUS SYSTEM IN RATS 45
 +
 +
environmental conditions similar to those under which the rats used in the present experiment lived (King, '15 b) show that for this group the average body weight of breeding females at 151 days of age is the same as that given by Donaldson, namely 186.1 grams; the average weight of the males of the same age is 244.8 grams, which is 26.1 grams above the computed weight for the male as given in Donaldson's tables.
 +
 +
The a\'erage body weight of all of the females used in this experiment was 146.7 grams, and that of the males was 258.7 grams, when the animals were 150 days old. The females, as a group, are too light in weight for their age, while the males are much too heavy, wliichever of the above series of records is taken as a standard for comparison.
 +
 +
The fact that the females were either strictly inbred or halfinbred does not account for their small size, since in the strain of albinos from which these rats were taken inbreeding has increased rather than diminished the average body weight of both males and females (Popenoe, '16). Investigations made by Watson ('05) have shown that female rats that are allowed to breed are heavier at a given age than non-breeding females. It is probable that the low weight of these females is due, in some measure at least, to the fact that the rats were never mated (the stock females whose weights were given for comparison were all breeding animals). The relatively large size of the males in the third series can be attributed to the fact that the animals were from a selected inbred strain.
 +
 +
The average daily percentage gain of these rats in body weight during the period covered by the experiment is shown in table 2.
 +
 +
The percentage values for weight increase, unhke the weight data, show no definite order with respect to the birth weights of the individuals concerned. At some periods the rat which had the smallest birth weight shows a greater daily percentage gain in weight than smy other member of the same series; at other periods the weight excess is in favor of the individual with the heavier birth weight (table 2).
 +
 +
If, instead of considering all of the percentage values in a given series, only the data for the two individuals that had the extreme H D3
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t; HOW I'll OF llODV AND NERVOUS SYSTEM IN HATS 47
 +
 +
birth weights are coinpared, the results ()])taiiietl seem consistent enough to be significant. In the total of 01) records for the three series, 42, or nearly- two-thirds, show that the daily percentage weight increase is greater for the individuals with a low birth weight than for those with a lieavy birth weight. In other words, regardless of sex, rats that are very small at birth tend to grow more rapidly than do rats that have a heavy birth weight, although their actual body weights are always less at any given period.
 +
 +
On computing the percentage increase in body weight at 150 days over the birth weight for the various individuals it was found that the rats that were small at birth had gained a greater amount than had the rats that were heavy at birth. For each series, as shown in the last column in table 2, these percentage values stand in inverse order to that of the birth weights, with the one exception in the second series (rat no. 4).
 +
 +
The results of this investigation are in accord with those obtained by Dunn ('08) in her study of the weight increase in a 'group' of seven albino rats (three males and four females) w^hich had very unUke body weights when they were fourteen days old. Dunn found that, with, one exception, the order relation of the weight at fourteen days of age was maintained until the end of the experiment ; the rats having the heaviest initial weights were also the heaviest at sixty-sLx days of age when the weighings were discontinued. The lighter rats, on the other hand, while putting on less absolute weight, had gained at the end a greater percentage of their original weight than had the individuals with the heavier initial weight.
 +
 +
In addition to the undersized young which, as shown above, are capable of developing into adults which are only slightly below normal, a litter sometimes contains individuals of a much lower grade to which the term 'runt' properly applies. In these individuals, which apparently are indistinguishable from the other young at birth, the normal action of the growth factors is inhibited from the very beginning of postnatal life by unknown constitutional causes, not by environmental conditions. When the young rats are old enough to leave the nest the runts can
 +
 +
THE ANATOMICAL RECORD, VOL. 11, N'O. 2
 +
 +
 +
 +
48 HELEN DEAN KING
 +
 +
easily be distinguished from the other members of the Utter, not only because of their very small size, but also because of their slower movements and apparent lack of normal vitality. Runts grow slowly for a certain time, but no matter how favorable the external conditions, they never exhibit normal vigor and they are always dwarfed and stunted in their body growth. In such animals growth is not merely retarded, as it is in the case of rats experimentally stunted (Hatai, '07 ; Osborne and Mendel, '14) , but it is permanently checked at an early age. Several attempts have been made in our colony to increase the size of runts by special feeding, and to breed them for the production of a dwarfed race of rats. Only a few Jitters could be obtained from such stock, and these contained a very small number of young which were puny from birth and which died at an early age. The reproductive powers of these animals are apparently never developed in a normal way, as the males rarely mate and most of the females are sterile.
 +
 +
All of the rats used in this study were killed at the end of 150 days, the body weights and body lengths determined, and t-he brains and spinal cords removed and weighed. This was done in order to ascertain whether the central nervous system in adult rats that were small at birth bears the same relation to body weight and to body length as that found in adult Individ uals that were of average size, or above, at birth.
 +
 +
Table 3 gives the body lengths of the individuals, the observed weights of the brains and of the spinal cords, and the brain and cord weights corrected according to table 68 in 'The rat: data and reference tables' (Donaldson, '15) which gives the computations for the 'standard' weights of the central nervous system in albino rats of various body lengths. In addition to the above data, table 3 shows the percentage deviations of the observed weights of the brains and of the spinal cords from the corresponding standard weights.
 +
 +
It is evident, from the percentage values given in the fifth and in the eighth columns of table 3, that 'the observed weights of the brain and of the spinal cord in all of the rats are considerably below the standard weights for these organs in animals
 +
 +
 +
 +
GRO^VTH OF 1U)I)V AM) NERVOUS SYSTEM IN RATS
 +
 +
 +
 +
49
 +
 +
 +
 +
Showing the bodij lengths, wilh the observed and the 'standard' weights for the brain and for the spinal cord, of the albino rats whose weight data are given in table 1
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«. 1
 +
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183
 +
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1.461
 +
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1.773
 +
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-17.5
 +
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0.455
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0,524
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 +
-13.1
 +
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 +
Scries 1
 +
 +
 +
(Females)
 +
 +
 +
2
 +
 +
 +
185
 +
 +
 +
1.611
 +
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 +
1.782
 +
 +
 +
- 9.5
 +
 +
 +
0.473
 +
 +
 +
0.532
 +
 +
 +
-11.0 3
 +
 +
 +
182
 +
 +
 +
1.679
 +
 +
 +
1.768
 +
 +
 +
- 5.0
 +
 +
 +
0.499
 +
 +
 +
0.520
 +
 +
 +
- 4.0 1
 +
 +
 +
175
 +
 +
 +
1.511
 +
 +
 +
1.735
 +
 +
 +
-12.9
 +
 +
 +
0.418
 +
 +
 +
0.492
 +
 +
 +
-15.0 2
 +
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 +
184
 +
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 +
1.616
 +
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 +
1.778
 +
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- 9.1
 +
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0.467
 +
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0.528
 +
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 +
-11.5
 +
 +
 +
Series 2
 +
 +
 +
(Females) <
 +
 +
 +
3
 +
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 +
192
 +
 +
 +
1.653
 +
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 +
1.814
 +
 +
 +
- 8.8
 +
 +
 +
0.485
 +
 +
 +
0.560
 +
 +
 +
-13.4 4
 +
 +
 +
188
 +
 +
 +
1.643
 +
 +
 +
1.796
 +
 +
 +
- 8.5
 +
 +
 +
0.475
 +
 +
 +
0.544
 +
 +
 +
-12.6
 +
 +
 +
 +
 +
c
 +
 +
 +
1
 +
 +
 +
198
 +
 +
 +
1.697
 +
 +
 +
1.849
 +
 +
 +
- 8.1
 +
 +
 +
0.529
 +
 +
 +
0.556
 +
 +
 +
- 4.8 2
 +
 +
 +
210
 +
 +
 +
1.765
 +
 +
 +
1.903
 +
 +
 +
- 7.2
 +
 +
 +
0.579
 +
 +
 +
0.603
 +
 +
 +
- 3.9
 +
 +
 +
Series 3
 +
 +
 +
(Males) <
 +
 +
 +
3
 +
 +
 +
217
 +
 +
 +
1.955
 +
 +
 +
1.933
 +
 +
 +
+ 1.1
 +
 +
 +
0.631
 +
 +
 +
0.630
 +
 +
 +
+ 0.2 4
 +
 +
 +
212
 +
 +
 +
1.999
 +
 +
 +
1.911
 +
 +
 +
+ 4.6
 +
 +
 +
0.669
 +
 +
 +
0.611
 +
 +
 +
+ 9.4
 +
 +
 +
 +
of like bod}^ length, except in the case of the two largest males where the observed weights slightly exceed the standard weights. That the weights of the central nervous system in all individuals of a given litter, having the same sex and about the same body weight, should deviate from the standard weights in the same direction was not an unexpected result. In the rat variation within the litter unit is usually in the same direction and much less than that in the general population as regards body weight (Jackson, '13; King, '15 b), and doubtless this rule holds for the central nervous system and other organs as well. In every series, as shown in table 3, the rats with the smallest birth weights are the ones whose brain and cord weights show the most marked deviations from the standard weights, regardless of their body length and body weight with which the weight of the central nervous system is, as a rule, closely correlated. Thus, in the first series, female no. 1 had a body length of 183 mm. and a body weight of 145 grams while female no. 3 was shorter and heavier,
 +
 +
 +
 +
50 HELEN DEAN KING
 +
 +
yet in the former individual the brain was 17.5 per cent and the cord 13.1 per cent below the corresponding standard weights; in the latter individual the brain and cord weights were only about 5 per cent less tl^ian the standard weights. All of the rats in the second series had cord weights that showed relatively greater deviations from the standards than did the brain weights; the lowest weight for both brain and cord being found in the rat that had the shortest body length and the smallest body weight (no. 1). The most interesting result is found on comparing the records for the rats belonging to the third series. The brains of the two rats that had birth weights much lower than the average birth weight (nos. 1 and 2) were each about 8 per cent less than the standard and the cord weights showed a minus deviation of some 4 per cent from the standard, yet in body measurements rat no. 1 was below and rat no. 2. was above the average for stock males of like age. The brain and cord weights of their brothers, each of which was unusually heavy at birth, were above the computed standard weights for the central nervous system in animals of like body length.
 +
 +
That the postnatal growth of the central nervous system is influenced to some extent by the factors that determine the size of a rat at birth seems to be a conclusion warranted by the analysis of data given above.
 +
 +
SUMMARY AND CONCLUSIONS
 +
 +
The results obtained in this investigation seem to indicate that the undersized individuals which are sometimes found in a newborn litter of rats are not necessarily 'runts' in the generally accepted use of that term. Some of these small individuals, as shown above, attain an adult size that not only is within the normal limits of variation in the body weights of standard stock rats of like age, but may even exceed the average body weight of a large number of stock animals (rat no. 2, series 3).
 +
 +
All rats in a litter are not born with a like capacity for growth, as the data in table 1 indicate, and even when environmental conditions are as favorable and as uniform as it is possible to make them, individuals having unlike birth weights show marked differences in their rates of growth from birth to the adult state.
 +
 +
 +
 +
(;i{()\\Tll OF BODY AND NERVOUS SYSTEM IN RATS 51
 +
 +
In the cases stiidietl the individuals liaving a small birth weight seemed to possess a 'very great capacity for growth from the very beginning of postnatal life. Female no. 1 of the first series increased 2 per cent more in body weight during the first twentyfour hours after birth than did either of her sisters; for the same period the gain in body weight of the two smallest males in the third series was over G per cent more than that of their brothers. In the second series, howe\'er, the weight increase for the first day was greater for the rats that were heavy than for those that were small at -birth.
 +
 +
With an early acceleration in the rate of growth there is seemingly correlated an early cessation in the extent of growth, as in the adult state individuals that were undersized at birth are alwa>'s smaller than the other members of the same litter. On the other hand, rats with a hea\'y birth weight tend to grow more slowly at first than do the smaller individuals, but they continue to grow for a longer time and eventually reach a greater size. This rule seems to apply to females as w^ell as to males.
 +
 +
Not only does body weight at birth indicate the probable capacity of the individual for subsequent growth, but it also indicates the probable size of the central nervous system, since rats that are undersized at birth tend to have a much smaller central nervous system when they become mature than do other rats. The factors, whatever their nature, that determine the body size of a rat at birth seem to have a marked effect on the subsequent postnatal development of the individual, influencing the ultimate body weight as well as the size of the central nervous system.
 +
 +
A very small weight at birth indicates that a rat has a handicap in its organization that environment, however favorable, cannot overcome. Such animals, although they appear vigorous and healthy during their growth period and after reaching the adult state, are unquestionabty sub-normal in regard to the size of the body and of the central nervous system. If allowed to breed these rats would probabty produce young having a weaker constitution than their ow-n, and from such stock one W'Ould ultimately get 'runts' and an increasing tendency towards sterility that w^ould soon bring disaster to the colony.
 +
 +
 +
 +
52 HELEN DEAN KING
 +
 +
Judging from the results obtained in this study a newborn litter of rats may contain individuals of three kinds as regards their inherent capacity for body growth. As a rule, only young rats having a normal birth weight and a normal capacity for growth are found in a small or medium sized litter produced by a female rat in good physical condition. Occasionally rats are born which have a very small birth weight, and in these individuals, if they are able to survive, the growth capacity is lessened to some extent but not sufficiently to prevent them from being classed as 'normal' after they have reached maturity. If a litter is very large, or if the mother is not in good physical condition during the gestation period, some of her young may be born with their growth capacity so impaired that it is impossible for them to grow beyond a certain stage. These individuals are true 'runts' and, fortunately, they are lacking in reproductive vigor as well as in growth capacity so that they are usually unable to reproduce their kind and so prove a menace to the colony in which they live.
 +
 +
LITERATURE CITED
 +
 +
Donaldson, H. H. 1906 A comparison of the white rate with man in respect
 +
 +
to the growth of the entire body. Boas anniversary volume, New
 +
 +
York.
 +
 +
1915 The rat. Data and reference tables. Memoirs of The Wistar
 +
 +
Institute of Anatomy and Biology, no. 6. Philadelphia. Dunn, Elizabeth 1908 A study of the gain in weight for the light and heavy
 +
 +
individuals of a single group of albino rats. Anat. Rec, vol. 2. Hatai, S. 1907 Effects of partial starvation followed by a return to normal
 +
 +
diet, on the growth of the body and central nervous system of albino
 +
 +
rats. Amer. Jour. Phys., vol. 18. Jackson, C. M. 1913 Postnatal growth and variability of the body and of
 +
 +
the various organs in the albino rat. Am. Jour. Anat., vol. 15. King, Helen Dean 1915 a . On the weight of the albino rat at birth and the
 +
 +
factors that influence it. Anat. Rec, vol. 9.
 +
 +
1915 b The growth and variability in the body weight of the albino
 +
 +
rat. Anat. Rec, vol. 9. Osborne, T. B., and Mendel, L. B. 1914 The suppression of growth and the
 +
 +
capacity to grow. Jour. Biol. Chem., vol. 18. PoPENOE, Paul 1916 Experimental inbreeding. Jour. Heredity, vol. 7. Watson, J. B. 1905 The effects of bearing young upon the body-weight and
 +
 +
the weight of the central nervous system of the female white rat. Jour.
 +
 +
Comp. Neur. and Psychol., vol. 15.
 +
 +
 +
 +
MOUxTixc spp:cimens under PF/nn dishes and
 +
 +
CLOCK GLASSES
 +
 +
E. L. JUDAH McGill University, Monlrcnl
 +
 +
Since obtainiii};- a suitable cement for the sealing of square museum jars, the mounting of thin sections of pathological and anatomical specimens under Petri dishes and clock glasses has been made both easy and cheap. Several years ago these mounts attracted a great deal of attention, lacing put on the market as paper weights, etc., but were gradually adopted for the exhibition and display of museum specimens. The method, however, being patented was expensive and beyond the reach of the average museum; besides it was necessary to send your material for mounting to the manufacturer. In 1906 Dr. Hutchinson of the Roj^al Victoria Hospital, Montreal, read a paper before the British Medical Association at Toronto on this method; but unfortunately he did not have a suitable cement and the process was slow and laborious.
 +
 +
METHOD
 +
 +
The fluid used for mounting should be brought to a boil in the same dish that is to be used for mounting and allowed to stand over night, or until cool, to get rid of as much air as possible. The dish should be deep enough to come well up over the mount to allow of easy manipulation of both Petri dish and specimen. Get a Petri dish of suitable size to hold the specimen so that when the sheet of glass which is to form the cover of the mount is in position it will not quite touch it. Great care must be taken that the Petri dish or clock glass fits perfectl}'^ on the base and does not rock.
 +
 +
In placing the Petri dish in the mounting fluid, do so without causing any air bubbles. When the dish and fluid are read}^, wash the specimen in several changes of the same fluid that you are mounting in, to get rid of any loose particles or dirt. In the last change of fluid, work out all air and remove quickly to the Petri dish, face downwards, again eliminating air bubbles. A small piece of looking-glass in the bottom of the mounting dish is very convenient, as by tipping the Petri dish over and on its side shghtly, any bubbles under the specimen may be seen.
 +
 +
The specimen now being in position, put on the base or cover and remove from the mounting-dish, holding it firmly so that it cannot slip
 +
 +
53
 +
 +
 +
 +
54 E. L. JUDAH
 +
 +
and admit air while turning the mount right side up. Over the junction of Petri dish and cover pour hot cement^ to the thickness of about one quarter of an inch, and allow the mount to stand on a flat surface for a few days, or until the cement is perfectly adherent to both Petri dish and cover. If after several days it is desired to finish the mount, remove any bubWes of fluid between the cover and the cement with a very hot knife, working the knife to the outer edge of the base. This must be carefully done, and the knife kept hot enough so that the cement will be kept liquid and not be drawn away from the Petri dish. All the fluid must be removed from under the edge of the Petri dish in this way.
 +
 +
The best results are obtained, however, by allowing the mount to stand for several weeks when most of the excess fluid will work out by itself. I usually mount several dozen specimens at a time and let them stand until they are ready to finish. If any air bubbles should happen to get under the edge of the Petri dish they will have to be worked into the mount in the same way that fluid is removed, only work your knife inwards instead of outwards.
 +
 +
Even with the greatest care air is often retained in the specimen itself, and only detected after the mount has stood for several days. Should the bubbles be very small they will quite frequently be absorbed by themselves; if not, shake the mount until they are all in one large bubble, and resubmerge in mounting fluid which must be heated as hot as you can comfortably stand your hand in. When the cement has become pliable enough so that it is possible to move the Petri dish with the fingers, insert the point of a sharp knife between the dish and base, gently forcing them apart and allowing a little fluid to enter. The mount must then be tilted on its side to bring the air bubble to the opening, where it will escape when you remove the point of the knife; then press the Petri dish up against the base, expelling all superfluous fluid. Close the hole by pressing the soft cement together with the fingers.
 +
 +
Air bubbles may have to be removed several times depending on the specimen. Sections of lung give the most trouble. When sure that there are no more bubbles in the mount and that all fluid has been removed from between the cement base and the Petri dish, coat the cement over to the thickness of about one-eighth of an inch with refined asphalt, being careful that it does not burn; I usually melt it in a tea-spoon over a Bunsen burner. When the asphalt has been all apphed, reheat the whole with a very hot knife and apply a bezel ring which must be heated red hot, and pressed down into the cement so that the lower edge will rest upon the base. Clean with gasoHn and polish with bon ami soap.
 +
 +
The Petri dishes used are manufactured specially from 2 to 3 mm. thick, as the ordinary ones sometimes break with the expansion and
 +
 +
Sec Muir and Judah, Sealing of museum jars. Bulletin .5, Inter. Assoc. Med.
 +
Mus., page 87.
 +
 +
 +
 +
MOUNTING SPECIMENS UNDER I'ETRI DISHES
 +
 +
 +
 +
OO
 +
 +
 +
 +
contraction of tlu> Ihiid. 'I'hc l)t'zcl iin;;,s which arc nscd to f>;iv(! a tinishcil appearance to the ceniont can In* made out of any in(;tal tliat will stand heinj? heated red hot. They arc no't, howevci-, absohitely necessary, as you can use a sheet of cardboartl with a hole cut in the centre and passe partout the edges. Dr. Higgins of the Experimental Farm, Ottawa, has a very convenient cardboard case into which he slips the mount. If a bezel ring is to be used, tjic base or cover should be made out of plate glass with the edges bevelled.
 +
 +
Clock glasses are inferior to Petri dishes for this method because they magnify and distort the spochncn.
 +
 +
The following list of sizes have been found to answer all requirements. While 36 in number they onl}' require ten different sized bezel rings and plate glass bases, a desired advantage when the glass ware is made to order.
 +
 +
Sizes (Outside measurements)
 +
 +
 +
 +
SIZE NO.
 +
 +
 +
WIDTH
 +
 +
 +
HEIGHT
 +
 +
 +
SIZE NO.
 +
 +
 +
WIDTH
 +
 +
 +
HEIGHT
 +
 +
 +
 +
 +
cm.
 +
 +
 +
cm.
 +
 +
 +
 +
 +
C7n.
 +
 +
 +
cm.
 +
 +
 +
1
 +
 +
 +
18
 +
 +
 +
3
 +
 +
 +
19
 +
 +
 +
8
 +
 +
 +
2.5
 +
 +
 +
2
 +
 +
 +
18
 +
 +
 +
2
 +
 +
 +
20
 +
 +
 +
8
 +
 +
 +
2
 +
 +
 +
3
 +
 +
 +
18
 +
 +
 +
7
 +
 +
 +
21
 +
 +
 +
8
 +
 +
 +
1.5
 +
 +
 +
4
 +
 +
 +
16
 +
 +
 +
3
 +
 +
 +
22
 +
 +
 +
8
 +
 +
 +
1
 +
 +
 +
5
 +
 +
 +
16
 +
 +
 +
2
 +
 +
 +
23
 +
 +
 +
6
 +
 +
 +
2.5
 +
 +
 +
6
 +
 +
 +
16
 +
 +
 +
1.5
 +
 +
 +
24
 +
 +
 +
6
 +
 +
 +
2
 +
 +
 +
7
 +
 +
 +
14
 +
 +
 +
3
 +
 +
 +
25
 +
 +
 +
6
 +
 +
 +
1.5
 +
 +
 +
8
 +
 +
 +
14
 +
 +
 +
2.5
 +
 +
 +
26
 +
 +
 +
6
 +
 +
 +
1
 +
 +
 +
9
 +
 +
 +
14
 +
 +
 +
2
 +
 +
 +
27
 +
 +
 +
4
 +
 +
 +
2.5
 +
 +
 +
10
 +
 +
 +
14
 +
 +
 +
1.5
 +
 +
 +
28
 +
 +
 +
4
 +
 +
 +
2
 +
 +
 +
11
 +
 +
 +
12
 +
 +
 +
3
 +
 +
 +
29
 +
 +
 +
4
 +
 +
 +
1.5
 +
 +
 +
12
 +
 +
 +
12
 +
 +
 +
2.5
 +
 +
 +
30
 +
 +
 +
4
 +
 +
 +
1
 +
 +
 +
13
 +
 +
 +
12
 +
 +
 +
2
 +
 +
 +
31
 +
 +
 +
3
 +
 +
 +
2
 +
 +
 +
14
 +
 +
 +
12
 +
 +
 +
1
 +
 +
 +
32
 +
 +
 +
3
 +
 +
 +
1.5
 +
 +
 +
15
 +
 +
 +
10
 +
 +
 +
2.5
 +
 +
 +
33
 +
 +
 +
3
 +
 +
 +
1
 +
 +
 +
16
 +
 +
 +
10
 +
 +
 +
2
 +
 +
 +
34
 +
 +
 +
2
 +
 +
 +
2
 +
 +
 +
17
 +
 +
 +
10
 +
 +
 +
1.5
 +
 +
 +
35
 +
 +
 +
2
 +
 +
 +
1.5
 +
 +
 +
18 .
 +
 +
 +
10
 +
 +
 +
1
 +
 +
 +
36
 +
 +
 +
2
 +
 +
 +
1
 +
 +
 +
 +
ANNOUNCEMENT
 +
 +
 +
 +
An Authors' Index has been prepared and printed for each of the following journals:
 +
 +
ft
 +
 +
JOURNAL OF MORPHOLOGY
 +
 +
25 volumes — 1887-1914 Price per copy, 50 cents
 +
 +
THE AMERICAN JOURNAL OF ANATOMY
 +
 +
18 volumes — 1901-1915 Price per copy, 50 cents
 +
 +
THE ANATOMICAL RECORD
 +
 +
10 volumes — 1906-(July) 1916 Price per copy, 50 cents
 +
 +
Sent post-paid to any address
 +
 +
THE WISTAR INSTITUTE OF ANATOMY AND BIOLOGY
 +
 +
Philadelphia
 +
 +
 +
 +
56
 +
 +
 +
 +
yi
 +
 +
 +
 +
THE EFFECT OF IIYPOPIIYSIX'TOMY IX THE EARLY
 +
 +
EMiuno UPON THi: (;uo^vT^ and develop:\ient of the frog
 +
 +
A PRELIMINARY REPORT
 +
 +
p. E. SMITH From Ihc Anatomical Laboratory, University of California
 +
 +
TEX FIGURES
 +
 +
The extirpation of the hjpopliysis in the adult frog lias not given uniform results. Caselli ('00) and Gaglio ('02) who reported no changes following M^pophysectomies were followed by Boteano ('06) \\'ho reported a neuromuscular asthenia in the operated animals. Houssay ('10) came to the conclusion that the removal of the gland was followed by death. Adler ('14) bm-ned out the hypophysis of a 20 mm. Rana temporaria larvae ^^■ith the electric cautery. Out of the 1200 operated animals thi'ee were found to have been h>T3oplwsectomized, not, however, without great injury^ to the smTounding soft parts, particularly the brain. In not one of those three animals did hind legs develop beyond a small bud, and transformation did not take place, the specimens remaining as neotonic tadpoles.
 +
 +
This work was commenced in the Spring of 1914, repeated in 1915, and again in 1916, Diemyctylus torosus, Rana pipiens, and Rana boylei being successively used. In this paper the results obtained with the Cahfornia yellow-legged frog, R. boylei are reported. Shortly after the closure of the medullary plate, Kopsch's stages d-e, was found to be the size in which the hj^ophysial invagination could be most successfully removed. About 200 larvae of this stage were operated upon. In specimens of this size the h^^Dophysis was successfully removed in over 60 per cent of the operated animals. Approximately 30 per cent of those animals in which the gland was extirpated did
 +
 +
57
 +
 +
THE AXATOMICAL RECORD, VOL. 11, NO. 3 OCTOBER, 1916
 +
 +
 +
 +
58 p. E. SMITH
 +
 +
not give reliable results in the rate of growth as the mouth was wholly or partially removed thus interfering with feeding. Unoperated animals and those in which the ablation of the gland was unsuccessfully attempted were available for checks.
 +
 +
The operation is a simple procedure. The hyj^ophysial invagination can be accurately determined from the pit that it early forms or from its location between the protuberance of the forebrain and the stomadeum, which is just forming. This epithelial ingrowth was removed with some neighboring epithelium. The wound healed within three hours in most cases, less than 1 per cent of the larvae disintegrating after the operation. The operated animals and checks were kept in boiled water for five days and then transferred to' a frog tank where they were in an essentially normal environment.
 +
 +
The rate of growth in the hypophysis-free animals has been slower than in the checks. The larger hypophysectomized animals averaged smaller in size than the larger checks, the averages of the two showing a noticeable difference. On June 6 the operated but not hypophysectomized animals had an average length of 40 to 43 mm., the hypophysis-free animals averaging 33 to 35 mm., a ratio constant throughout their growth. The ratio of body to tail length is the same in the two classes, the difTerence in size being uniform for all parts of the animal. The tail fin did not show an increased width or pleating in the hypophysectomized animals as reported by Adler ('14).
 +
 +
In activity the two classes of animals showed no marked differences. The hypophysectomized specimens were perhaps slightly more alert, darted more quickly, and consequently were more difficult to capture with the pipette than were the checks.
 +
 +
The resistance of the hypophysectomized animals was greater than that of the checks. Towards the close of the experiment the animals were attacked by disease, none reaching the adult stage. The normal specimens succumbed more rapidly to this infection than did the hypophysectomized ones. Some of the intrinsic factors which induce growth of legs and transformation were lacking in the abnormal specimens as will be shown later. The absence of these factors may well be conducive to a greater
 +
 +
 +
 +
EKFKc'i' or in i'(M'iivsK( ro.Mv rrox thk fuog 5!)
 +
 +
liariliiioss in an animal wJicn (■()ni])ar('(l to tlic iiornial tadjiole in wliic'li tlio usual ra])i(l ('lianf:;(>s arc takiiij;- ])la('('.
 +
 +
nirtci'ciiccs ill color l)c<>;aii to be not iccahic hcforc a i('nfi;tli of IT) nun. was jvaciuHi, and IVoni tlicn on the contrast in ])if2;nicntation between the hypopliysectoniized animals and the cliccks was strikhig. Those animals without hypophyses were characterized by a light grayish appearance; however, the dorsal side was more pigmented tlian the ventral (figs. 7, 10). These are referred to as albinos. The checks were a brown-black color often showing a mottling (figs. 8, 9). This color difference was more noticeable over tlie body tlian on the tail, but was evident in both regions and was the most striking feature up to the time of the appearance of the hind legs in the checks. Sections show that these pigment differences are referable chiefly, if not solely, to the condition of the epidermis. Counts of the melanophores of corresponding areas in the albinos and in the checks show that the number of these cells, in the epidermis, are reduced in the former. Further the melanophores of the albino specimens contain fewer pigment granules than do those of the checks and thus have a distinctly lighter appearance. The melanophores are equally expanded in the two types, consequently, the lighter color of the albinos cannot be due to the contracted condition of the chromatophores but must be referred, in part, to the reduced number of melanin granules in the pigment cells of the epidermis. In addition to this the free pigment granules which form a distinct zone in the superficial layer of the epidermis in the normal checks are much reduced in number in the albino specimens (figs. 5, 6). It is surprising that in the albinos the deeper or subcutaneous pigment is present in as great a quantity as in the normal animals, if not greater. The amount and distribution of the retinal pigment seem to be identical in the two.
 +
 +
Another important feature was the inhibition in growth of the hind legs of the operated animals. There was only a slight retardation in the time of appearance of the hind leg buds, normalh^, appearing when the tadpole has reached a length of 25 to 27 mm. In the albino, averages show that the hind limb buds appear when the larvae are from 26 to 28 mm. in length.
 +
 +
 +
 +
{
 +
 +
 +
 +
60
 +
 +
 +
 +
p. E. SMITH
 +
 +
 +
 +
From this state on, however, the hind Umbs in an h^i^ophysectoniized animal grew but httle if at all, although the animal's length increased at a rate but slightly under the normal. The accompanying tabte shows the increase in length of the hind legs in relation to total length for the albinos and for the checks. (See also figs. 7, 8).
 +
 +
Average rate of growth in millimeters in terms of total length, of the hind legs of the checks and the albinos
 +
 +
 +
 +
H1TOPHYSECTOMIZED AXIMALS
 +
 +
 +
CHECKS
 +
 +
 +
Total length
 +
 +
 +
Hind le? length
 +
 +
 +
Total length
 +
 +
 +
Hind leg length'
 +
 +
 +
26
 +
 +
 +
barelj' visible
 +
 +
 +
25
 +
 +
 +
barely visible
 +
 +
 +
28
 +
 +
 +
0.1
 +
 +
 +
28
 +
 +
 +
1.0
 +
 +
 +
30
 +
 +
 +
0.1
 +
 +
 +
30
 +
 +
 +
2.0
 +
 +
 +
35
 +
 +
 +
0.1
 +
 +
 +
35
 +
 +
 +
3.0
 +
 +
 +
37
 +
 +
 +
0.12
 +
 +
 +
38
 +
 +
 +
4.0 40
 +
 +
 +
5.0 45
 +
 +
 +
9.0
 +
 +
 +
 +
Only one exception to the rule that no hind legs grew on albinos was found. A 36 mm. albino had hind legs 4.2 mm. long when killed. The above is in accord with Adler ('14) who found that removal of the hypophysis in a 20 mm. stage inhibited the growth of the hind legs.
 +
 +
Examination of sections of albino and normal animals shows striking differences in the endocrine glands. The sectioned hypophysectoniized animals show no trace of the anterior lobe of the h>T)ophysis. That part of the floor of the diencephalon which normally abuts against the hypophysis, rests upon the floor of the cranium (fig. 2). This apparently demonstrates conclusively that the entoderm has not the intrinsic power to form a hypophysis. If it enters into the formation of the gland at all it must be considered as a tissue inclusion which became changed through its adaptabihty into glandular parenchyma, a conclusion previously drawn by the ^^Titer, Smith ('14). The infundibulum shows some structural modifications when compared to the checks, although the saccus vasculosus, as far as determined, appears to be normal. In the checks that region of the diencephalon which rests against the pars glandularis is
 +
 +
 +
 +
EKFKCT OF ll'* I'til'HYSECTOMY UPON TIIK FUOG
 +
 +
 +
 +
(>L
 +
 +
 +
 +
 +
"■"^v
 +
 +
 +
 +
 +
Fig. 1 A section throufih tlic hypophysial region of a 3S mm. normal tadpole. X 100.
 +
 +
Fig. 2 A section through the hypophysial region of a 87 mm. albino. Note the much reduced pars nervosa. X 100.
 +
 +
 +
 +
 +
Fig. 3 A sagittal section through a lobe of the thyroid of a 38 mm. check. X 100.
 +
 +
Fig. 4 A sagittal section through a lobe of the thyroid of a 37 mm. albino. X 100.
 +
 +
5 6
 +
 +
Fig. 5 A section through the epidermis, in the mid-brain region, of a normal
 +
 +
39 mm. check. The pigment granules are indicated by dots. X 200.
 +
 +
Fig. 6 A section through the epidermis, in the mid-brain region, of a 38
 +
 +
mm. albino. A faint melanophore in the left part of the figure. X 200.
 +
 +
 +
 +
62
 +
 +
 +
 +
p. E. SMITH
 +
 +
 +
 +
of considerable thickness, that is, in addition to the ependyma there is a rudimentary pars nervosa. Caudad to this the wall is formed almost entirely of ependyma. The pars nervosa is reduced throughout most of its extent to an ependymal layer in the hypoi^hysectomized animals. There may be a small localized thickening but nothing to correspond to the normal animal (figs. 1, 2).
 +
 +
The thyroid shows marked modifications in the albinos. In the accompanying table the size of one lobe of the thyroid of a normal 38 mm. tadpole with 4.0 hind legs and of a 37.0 mm. albino with 0.1 mm, hind legs is given.
 +
 +
Size in millimeters oj one lobe of the thyroid SS mm. check 37 mm. albino
 +
 +
Length 0.6 Length 0.21
 +
 +
Width 0.3 Width 0.15
 +
 +
Thickness 0.16 Thickness 0.04
 +
 +
 +
 +
 +
-#■
 +
 +
 +
 +
8 *»
 +
 +
 +
 +
 +
Fig. 7 Photograph of an albino. X 2. Note the very small hind limb bud. Fig. 8 Photograph of a normal tadpole. Figures 7 and 8 were photograi)hed on the same plate' X 2.
 +
 +
Fig. 9 Photograph of a normal tadpole. X 2. Fig. 10 Photograph of an albino. X 2.
 +
 +
 +
 +
EFFECT OK m I't >l'in SK( I'c ).M V Ul'ON 'I'lIK FKOCJ G3
 +
 +
Tlic al)^)^■(' tahlc sliows tlial the thyroid of the all)iiio is ap])roximat('ly oiic-thii-d normal size. Tlic contrast is oven more strikinji wlicii the compactness and character of the parenchyiTm is noted. A sagittal section thronfi;li the thyroid of a :iS mm. cliecU sliows on an averago 12 to lo vesicles, many of which are larjicly distended witli colloid, the parenchyma of the whole gland Ixnng compactcMl togetliei'. A sagittal section through the thyroid of a hyp()])hysectomized 37 nnn. s])ecimen shows 6 to 8 atro})hied vesicles containing but a slight amount, or no colloid, and with large spaces between the vesicles. The cells making u]) tlie vesicles of the former are cuboidal and protoplasmicricli, in the latter little but the nuclei remain (figs. 3, 4). The results from exj^erimental feeding of thyi-oid by Gudernatsch and other workers suggests that the non-development of the hind legs in the albinos is due not to the hj^^ophysis but rather to the failure of the thyroid. In this connection the 36 mm. albino with 4.2 mm. hind legs, mentioned above, is of interest. Sections of this specimen sh(nv that the hypophysis was completely ablated but that the thyroid is normal. This specimen thus gives adcUtional evidence that the retarded development of the hind legs must be referred to the thyroid and not to the hypophysis. Also the reduction in pigment is not due to the atrophy of the thyroids. The modifications of the thyroid obtained by Adler ('14) were similar but less striking.
 +
 +
An examination of a large number of male and female albinos and checks has, as yet, failed to show any constant variation from the noraial in the sex glands of the hypophysectomized animals. The sex glands of the albinos although varying considerably apparently do not exceed the limit of variation met with constantly in the normal animals. This conclusion stands in contradiction to the results previously adduced by the author and to the results of Adler ('14) in the hypophysectomized tadpole and to the conclusions of Hahn ('12) in the tadpole with hypertrophied hypophysis as well to the results obtained in mammals by pituitary feeding, notably that of Goetsch ('16).
 +
 +
The WTiter wishes to express his appreciation to Dr. H. M. Evans for his generous aid.
 +
 +
 +
 +
64 p. E, SMITH
 +
 +
LITERATURE CITED
 +
 +
Adler, L. 191-1 jMetamorphosestudien an Betrachierlarven. I. Extirpation
 +
 +
endokriner Drlisen. A. Extirpation der Hypophyse. Arch. f. Entw.
 +
 +
mech. d. Or^anis., Bd. 39. BiEDL, A. 1913 Innere Sekretion. Zweite Aufi. Berlin. BoTEANO, E. R. 1906 Contr. la physiol. glandei pituitare la brosca. These,
 +
 +
Bucarest, zeit. n. Paulesco, cit. f. Biedl. Caselli, a. 1900 Influence de la fonction de I'hypophyse suz le development
 +
 +
de I'organisme. Riv. sper. di fren., vol. 37, cit. f. Biedl. Gaglio, G. 1902 Recherches zur la fonction de I'hypophyse de cerveau chez
 +
 +
les grenouilles. Arch. ital. d. Biol., vol. 38, cit. f. Biedl. GoETSCH, E. 1916 The influence of pituitary feeding upon growth and sexual
 +
 +
development. Bull. Johns Hopk. Hosp., vol. 27. Hahn, a. 1912 Einige Beobachtungen an Riesenlarven von Rana esculenta.
 +
 +
Arch. f. mikr. Anat., vol. ,80. HoussAY 1910 La hypofisio de la rana. Trabajos de Labor de Univ. Nacional
 +
 +
de Buenos Aires, cit. f. Biedl. Smith, P. E. 191-1 The development of the hypophysis of Amia calva. Anat.
 +
 +
Rec, vol. 8.
 +
 +
 +
 +
ANO.MALIES IN LOBATION OF LUNGS W 11 H REVIEW OF IJTKRATrRE
 +
 +
CARBON' (ill.LASPIE, LEWIS I. MILLER, .VXD MORRIS BASKIX Anatoniiail Department, University of Colorado
 +
 +
FIVK FIGURES
 +
 +
Judging from tlio literature, abnormal lobation of the lungs is relatively rare.
 +
 +
Lindsay ('10) di^'ided the abnormalities in the lobation of the lungs into two classes: those in which the normal number is decreased and those in which the number of lobes is increased. The former is due either to a deficiency of the lobes themselves, or to a deficiencj' of the fissures, which normally separate the lobes. The latter is due to an increase of lobes, or to an increase in fissures. Complete absence of lobation in definitely formed lungs is rarely if ever found, though its homologue is to be found in the Orang, with two lungs each existing as single lobes.
 +
 +
Rokitansky ('61) showed that arrests of development may occur and that this may lead to complete absence or great deficiency of one or both lobes. This arrest may be so early that the lungs can scarcely be observed as small round bodies situated at the ends of the bronchi. This condition is generally due to contraction of the volume of the thorax.
 +
 +
Pontif ('60) in 'Virchows Ai'chives' recorded a case in which the right bronchus was connected wdth an ovoid body, which was imbedded in gelatinous tissue and filled the right half of the thorax.
 +
 +
According to Lindsay, cases of class two, that is those which have an excessive number of fissures, are occasionally encountered, and are probably the most common form of abnormahty. They do not appear to present any regularity, and lack the interest which is attached to accessory lobes. There are two
 +
 +
65
 +
 +
 +
 +
66 CARBON GILLASPIE, L. I. MILLER, AND M. BASKIN
 +
 +
groups of accessory lobes. One which is of considerable developmental interest, is composed of completely isolated masses of pulmonary tissue formed between the diaphragm and the base of the left lung; occasionally on the right side such masses are found even in the abdominal cavity. Other masses containing arteries, veins, nerves, and bronchial tissue, but devoid of bronchi, are attached to the oesophagus, aorta or other mediastinal structures by a pedicle. These masses are apparently quite functionless. Two such cases are recorded by Vogel ('99), in each of which he found a deficiency in the bronchial tree. Simpson ('99) described a case of a deficient bronchial tree found in a foetus.
 +
 +
In the cases with additional fissures there is a normally placed lung presenting an excessive number of lobes. These abnormalities are usually very definite in their position, and occur more commonly on the right side.
 +
 +
Wrisberg ('77), who was the first to notice an accessory lobe in the human lung, recorded a most interesting and unique case of an accessory lobe on the left side produced by the left azygous vein; i.e., the superior intercostal vein which preserved its foetal condition and opened into the left innominate vein.
 +
 +
Chiene ('76), described a pear-shaped supernumerary lobe, lying between the upper lobe of the right lung and the bodies of the dorsal vertebrae, having its origin from the angle formed by the junction of the upper lobe with the root of the lung. The supernumerary lobe was separated from the upper lobe of the lung by a double fold of the plenral membrane, which descended vertically for seven centimeters from the apex of the thoracic cavity where it was continuous with the pleura costalis. It enclosed in its free border the vena azygous, and formed the outer wall of the cul-de-sac, in which the supernumerary lobe was contained. The left side of the chest was normal; both sides were healthy.
 +
 +
E. W. Collins ('88), recorded a case of an accessory lobe immediately above the posterior part of the root in the angle between it and the upper portion of the right lung. This accessory lobe was somewhat pyriform in shape, with a broad pedun
 +
 +
 +
.\\(>M \I.IKS IN I.OHA'rio.N OK I.INCS 67
 +
 +
cular at tac'liinciit . In all. ( (tlliiis was al)lc to collect st'X'cii cuses of {icccssory lohcs in lniiiiaii hui^s.
 +
 +
A. I'. .Mai> land ('90) descrilxul ahiioiintditios in lohcs of three liiiijj;s. 'Vhv first was a rif»;]it lun^ with no iiuhcations of ;t middle lobe, l)ut a (h^Nclopnient of ;i tliird, or accessoi-y one, on the inner side of the lun^". Tlie second was :l left Inn^- with a sulxlixision of the iii)i)er lobe. The third was a right lunj; with an iiiconiplete separation of a normal middle lobe. ^Maryland states: "Cases of more than four right lobes and three left lobes are exceedingly rare. '
 +
 +
Patterson ('09-' 10) described a condition of two additional lobes. One of these was above the root, and separated from th(> n])i)er lobe by a fissure. This accessory lobe was enclosed within a pleural pouch, w^hich contained the vena azygous. The second additional lobe was below the root, and between the upper and middle lobes.
 +
 +
Case I of the present specimens presents features entirely different from those hitherto described. It w^as obtained in the dissecting room during the term of 1915-1916 from a male subject, aged sixty-one. The cause of death as given on the death certificate was dementia. No clinical history was obtainable. Besides the anomalous condition of the lungs, there was a persistent thymus, and a number of anomalous arteries. On gross inspection the lungs presented no pathologic lesions.
 +
 +
The left lung apex-base measured 22 cm., dorso-ventrally 20 cm. The normal fissure (FS), which separated the superior from the inferior lobe, was in its normal position, starting at the junction of the antero-inferior border 25 cm. from the apex, and running obliquely upward and backward, dividing the superior and inferior portions completely.
 +
 +
The superior portion, however, presented two other fissures, as shown in the diagram, thus dividing this portion into three more or less distinct lobes.
 +
 +
Fissure number one (Fl) started at the anterior border 7 cm. below the apex, and ran horizontally backward on the anterolateral surface of the lung for 6 cm. The depth of the fissure was on an average 1.5 cm.
 +
 +
 +
 +
68 CARBON GILLASPIE, L. I. MILLER, AND M. BASKIN
 +
 +
Fissure number two {F2) started at the antero-median border 17 cm. below the apex and ran upward and backward for 9 cm. This fissure extended through the lung tissue, completely separating the middle from the inferior division of the superior lobe. Neither of the two mentioned fissures extended as far as the main fissure, which separated the superior from the inferior lobe.
 +
 +
Three distinct divisions of the superior lobe were evident from an antero-lateral view. The upper division (L7) was pyramidal in
 +
 +
 +
 +
F "
 +
 +
 +
1 F,3 , •U
 +
 +
 +
^
 +
 +
 +
 +
Fig. 1 Anterior aspect
 +
 +
shape, forming the apex. It measured 8 cm. antero-posteriorly, and 7 cm. from apex to base.
 +
 +
The middle portion {L2) was wedge-shaped, wider on the anterior border than on the posterior. It measured 9 cm. on the anterior border between the first and second fissures, and 17 cm. antero-posteriorly.
 +
 +
The lower division {L3) is hngual in shape. This division measured 6 cm. on the anterior border and 21 cm. anteroposteriorly.
 +
 +
The inferior lobe presented two more supernumerary fissures as did the superior lobe.
 +
 +
 +
 +
.\.\(tM AI.IKS IN I.oHAIlo.N oi' UNCiS 09
 +
 +
Mssurc iiuiiilxM' I'diir (A'./) stiirlcd ,") cm. uiitcro-inlci'jorl^' I'roin tlic sii])(M()-iiitVriur (issiiic ;m(l i':ni ]);ii'all('l wilh (he above fissure tor a (lis(anc(> of 10 ciii. Tliis fissure also exten(l('(l llii-ou^h tile lunji' tissue se])ai'at iii^ the u])j)('i' IVoiii llic ])ostero-iiif('i-i()r divisions.
 +
 +
Fissure iiuiiil)er lix'e (Fo) Avas on the lateral surface, extending' ohli(iuely, su])erioi'ly and inferiorly, and incompletely dividing' \ho ])()st(M-o-lat(M-al fi'oin tlie i)()stero-inferior divisions.
 +
 +
 +
 +
 +
2
 +
 +
Fig. 2 Posterior asjiect
 +
 +
The inferior lobe, which was triangular, also presented three fairly distinct di\dsions.
 +
 +
The upper division {L4), which is oblong in shape, having a small tongue-like projection on the anterior border, ran obliquely upward, and backward, following the direction of the superoinferior, or great fissure. This division measured 5 cm. anteroinferiorly, and 19 cm. antero-posteriorly.
 +
 +
The postero-inferior division (Ld) was more or less quadrangular in shape. It measured 8 cm. on the superior border and 12 cm. on the inferior border, i.e., at the base.
 +
 +
The postero-lateral division {L6) was irregular in outline. The upper border was made by the supero-inferior fissure, and
 +
 +
 +
 +
70 CARBON GILLASPIE, L. I. MILLER, AND M. BASKIN
 +
 +
the lower by a separate fissure between this division and the postero-inferior division.
 +
 +
The right lung apex-base measured 19 em,, and dorso-ventrally 18 cm. In this lung the fissures were deei:)er, and went through the lung tissue, completely dividing the lung into distinct divisions. The normal fissure (F'S), which divided the apex lobe and middle lobe from the inferior, or base lobe, was a httle more irregular than the corresponding fissure on the left lung,
 +
 +
 +
 +
 +
Fig. 3 Right lung: lateral view
 +
 +
and ran obliquely upward and backward, dividing the lung into an upper and lower division.
 +
 +
The upper di\'ision presented an ir egular outline from a lateral view. There wei'e two large fissures; one which normally divided the apex lobe from the middle lobe, and the other a supernumerary fissure dividing the middle lobe into two. There was also a small fissure in the apex lobe.
 +
 +
Fissure number one (F'l), which divided the apex lobe from the middle on , started 10 cm. from the apex laterally, midway between the apex and the base.
 +
 +
 +
 +
ANOMAMKS IN l.( )HA'n( ).\ OF IJ'NCS
 +
 +
 +
 +
71
 +
 +
 +
 +
Plssiirc lumihci' two (/'"J), which (hxidcd the middle lohc into two (h\'isioiis, i-;iii \ crt ically for ;i (Hstaiicc of S cm., slartiiif^ from tJie base and nmiiiiif;" towurd tlie iipvx.
 +
 +
The ai)ox lobe (/>'/) was ([uadrilateral in slia])e, Juiving' the upper border narrower than the base and nieasiirinjij 9 cm. ai)exbase, and 8 cm. antero-posteriorly.
 +
 +
The inferior di\isi()n (L'2) of tlie n]i])er lobe was lingual in shape, extcMidiiig obli(iuely upward and backward 13 cm. in its long direction, and measuring 4 cm. a])ex-base.
 +
 +
 +
 +
■VVSI«*«M»»
 +
 +
 +
 +
4
 +
 +
 +
 +
Fig. 4 Left lung: lateral view
 +
 +
The inferior division (L'S) of the upper half of the lung was elongated in outline measuring 4 cm. in antero-posterior direction, and 13 cm. apex-base.
 +
 +
The lower half of the right lung was triangular in shape, having two clearly visible fissures which ran through the lung tissue and divided this portion of the lung into three distinct lobes. There are also two smaller fissures in the inferior border of the lung, merely forming small tongue-like lobules.
 +
 +
The fourth fissure {F'4) started 10 cm. from the apex midw^ay between apex-base, running downward and backward for a dis
 +
 +
 +
rz
 +
 +
 +
 +
CARBON GILLASPIE, L. I. MILLER, AND M. BASKIN
 +
 +
 +
 +
tance of 14 cm., and completely separating the upper from the lower division of this half of the right lung.
 +
 +
The fifth fissure {F'5) started from the base, or inferior border, and ran vertically upward for a distance of 10 cm. It extended through the lung tissue, dividing this inferior half into two divisions, a posterior and an antero-median.
 +
 +
The superior division {L%) of this inferior half of the right lung was triangular in shape, measuring 13 cm. on its inferior
 +
 +
 +
 +
 +
border, 8 cm. on its superior border, and 11 cm. on its posterior border.
 +
 +
The postero-inferior division {L'5) was quadrilateral in outline, measuring 8 cm. on its shortest border and 14 cm. on its longest, or posterior border.
 +
 +
The antero-median division {L'6) had the form of an elongated triangle, measuring 11 cm. on the anterior border, 6 cm. on the median posterior border.
 +
 +
At the base of the right lung was a very irregularly shaped division distinctly separate from the rest of the lung as shown
 +
 +
 +
 +
ANOMALIES IN LOHATION OK LUNGS 73
 +
 +
in tJie diagram. 'I'liis basal clivlsion (L 7) iiicasunHi 10 cin. in the long direction and 6 cm. in the short direction. This division was separated from the rest of the lung by two distinct fissures {F'6-7) which started at the postero-inforioi- border, and ran for 9 cm. anteriorly and backwards,
 +
 +
Case 11 was obtained in the dissecting room during the term 1915 -191(), from a female subject, aged thirty-eight.
 +
 +
The cause of tleatli as given on the death certificate was 'septic meningitis.' No clinical history was obtainable.
 +
 +
The left lung apex-base measured 20 cm., dorso-ven trail y 18 cm. The right lung apex-base measured 18 cm., dorso- ventral ly 17 cm.
 +
 +
The left lung presented upon examination one accessory lobe, one accessor}^ lobule, and three accessory fissures in the inferior portion of the left lung.
 +
 +
The right lung presented upon examination two accessory lobes and two accessory fissures. The first accessory lobe was in the upper portion of the inferior division of the right lung. The other accessory lobe was a basal lobe and was identical with the basal lobe seen in the right lung in Case I.
 +
 +
X-Ray^ pictures of Case II show separate bronchi going to each of the main lobes including the accessory lobes, as may be seen by referring to the plates.
 +
 +
X-Ray views of Case I were unsuccessful on account of a poor bismuth injection.
 +
 +
From the foregoing description it is evident that these specimens show no distinct azygous lobe, which is the common accessory lobe described. These lungs retain their normal shape; but they present, besides the normal fissures, a number of accessory fissures which divide the lungs into distinct accessory lobes.
 +
 +
Complete absence or deficiency of one or both lobes may be due as Rokitansky points out to arrests of development as contraction of the volume of the thorax. Supernumerary lobes have been variously explained. Lindsay ascribes a slight adhesion of the lungs to the thoracic wall as the cause of the super 1 Stereoscopic X-ray plates show the separate bronchi as described. Reduced prints fail to show the necessary details and, therefore, are not reproduced here.
 +
 +
THE ANATOMICAL RECORD, VOL. 11, NO. 3
 +
 +
 +
 +
74 CARBON GILLASPIE. L. I. MILLER, AND M. BASKIN
 +
 +
numerary lobe, or possibly an undue curvature of the embryo, so that the Venae Cavae, as it bent down to a position at right angles to its original position, instead of slipping behind the pleura and lung, dragged down a fold of the former and deeply notched the latter.
 +
 +
Fischer remarks that it is difficult to conceive of the azygous lobe without an already preexisting anomalous course of the azygous, and makes no attempt to explain the phenomenon.
 +
 +
Collins explains the azygous lobe as a persistent foetal condition of the left azygous vein.
 +
 +
All previous attempts to explain the origin of the supernumerary lobes apply only to cases in which there is a definite azygous lobe. The anomalous course of the azygous vein has constricted off a portion of the lung tissue, thereby forming a new lobe. This explanation does not apply to cases in which there are no azygous lobes and in which, however, there are other supernumerary lobes, as the azygous lobes are not in reality true lobes. A true lobe as applied to the lung indicates a separate bronchus. Whether the azygous lobe has a separate bronchus, or not, has not been stated in their descriptions.
 +
 +
Considered from the viewpoint of their origin, a constricted portion — such as the azygous lobe — is not a separate entity, but a part of the mother lobe from which it has separated.
 +
 +
The formation of lobes of the lungs has first been studied by Aeby and was worked out by Narath. Aeby defines lung lobe as follows: A true lobe is never supported by more than a single bronchus and therefore includes no portion of the stem bronchus." Soon after the formation of the first lateral bud in the embryo, each bud becomes marked out upon the surface of the mesodermal anlage of the lung. Before development of lateral bronchi, the surface of this anlage is smooth. Later it becomes almost mulberry-shaped, and secondary elevations are then formed by the budding of the bronchial bud which it contains. The process goes on until the surface becomes covered with fine granules. These disappear with further growth of the Jung — only the first formed furrows persisting normally.
 +
 +
 +
 +
AXO.MALIKri 1\ LDHAIION oi' LUNCJS /O
 +
 +
Accessory lobes may then be tliie to a retention of tlie foetal condition in which not only the primary divisions persist, but on account of the rather slow growth of the bronchi, the secondary conditions persist — that is to say, accessory lobes are not new formations. Init represent a stage in the normal development of the lung.
 +
 +
Accessory fissures dividing the lobe into a number of lobules — as exists in the present specimens — may have been due to folds of splanchnopleura which have been carried down and caused the formation of a groove, lined with pleura, or they may have been caused by incomplete obliteration of the secondary divisions, which are ])roduced in the mulberry stage and which normally disappear.
 +
 +
BIBLIOGRAPHY
 +
 +
Chiene 1876 Jour. Anat. and Phys., vol. 4. Clel.\nd 1861 Jour. Anat. and Phys., vol. 43. Collins, E. W. 1888 Royal Irish Academy. Fischer 1898-1899 Anat. Anzeig. Keib.\l and jSIall Human Embryology. LiNDS.\Y, M. A. 1910 ^Maritime Med. News, vol. 22. Maryland, A. E. 1890 Jour, of Anat. and Phys. McAllister Text Book of Anatomy, p. 340. Paterson 1909-1910 Jour. Anat. and Phys., vol. 44. PoNTiF 1860 Virch. Arch., vol. 50. Simpson 1899 Jour. Anat. and Phys., vol. 42. Wrisberg 1874 Trans, for the Royal Irish Academy.
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ANOMALOUS RENAL VESSELS AND THEIR SURGICAL
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 +
SIGNIFICANCE
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CARBON GILLASPIE, LEWIS I. MILLER AND MORRIS RASKIN
 +
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Anatomical Deparlmcnt, University of Colorado
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NINE FIGURES
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Abnormalities of the renal arteries occur more frequently, perhaps, than anomalies of any of the other larger vessels. In view of the enormous number of investigations of the different structures of the kidneys recorded in the literature on the subject, it seems strange that only scanty information exists concerning the actual course of the larger blood vessels, and their relations to the pelvis of the kidney. The normal, as well as the abnormal, arrangement of the renal vessels at the hilum is known; the microscopic picture of the vessels in the cortex and pyramids are likewise thoroughly familiar to every student; but as to the form of the pelvis, and the actual course and distribution of the larger vessels around its walls, very vague ideas still prevail.
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Professor Thane states that irregularities of the renal arteries are met with in about 25 per cent of cases, and that the most comimon irregularity is the presence of an additional vessel in about 20 per cent.
 +
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Young and Thompson report four cases of anomalous renal arteries: the first is that of multiple renal arteries and malposition of the right kidney; the second that of multiple renal arteries, malposition, and malformation of both kidneys; the third that of a horseshoe kidney with multiple renal arteries; the fourth that of multiple renal arteries, two of which were on the left side, two on the right. In the last case there were also multiple spermatic arteries, and two renal veins on the right side, while the left side was normal.
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77
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/8 CARBON GILLASPIE, L. I. MILLER, AND M. BASKIN
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Irregularities in renal vessels have also been mentioned by McAllister ('83) who found anomalous renal arteries in 43 per cent of the cases examined. Levings ('12) reported two cases of anomalous renal vessels going to the lower poles. Harvey ('14) described a case of multiple renal arteries.
 +
 +
The specimens of the present description were obtained in the dissecting room through the help of Dr. E. B. Trovillion, Instructor in Anatomy, University of Colorado; and from autopsies through the courtesy of Dr. R. C. Whitman, Professor of Pathology, University of Colorado. We were able to examine, in all, 33 cases of which 22, or 73 per cent possessed anomalous renal vessels.
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Case I. The kidneys were normally placed in the abdominal cavity. Both were somewhat smaller than normal. The right showed two small cysts on its anterior surface. It lacked the normal large renal artery, but possessed instead two renal arteries of equal size, which were almost as large as the normal vessel should have been. The lower artery arose from the ventro-lateral portion of the aorta, 6 cm. above its bifurcation, and passed to the lower pole where it divided into two vessels, 2 cm. before it entered the kidney substance. . The second artery arose from the aorta 6 cm. above the lower one, and split into two vessels 2 cm. before it reached the kidney substance. There were also two large renal veins, closely accompanying the arteries; one arising from the upper, the other from the lower pole of the kidney. Both veins emptied into the vena cava.
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The left kidney possessed three renal arteries and one renal vein. The lowest renal artery arose from the left ventral portion of the aorta 4 cm. above the birufcation of the aorta, and supplied the lower pole of the kidney. The second artery arose from the aorta 6 cm. above the first, and supplied the upper pole. The third arose 1 cm. above the second, crossed it and entered the kidney at the hilum. One large renal vein arose from the kidney at the hilum.
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Case II. Both kidneys were larger than normal, and in the proper position. The right possessed one renal artery and one renal vein. The left kidney, however, possessed three main renal arteries, and one renal vein. The first renal artery, which seemed to be the normal vessel, arose from the ventro-lateral aspect of the aorta just opposite the origin of the superior mesenteric artery. The second renal artery arose 1 cm. above the first. This was a long slender artery having a tortuous direction, and entered the kidney 4 cm. above the inferior border. The third artery was the smallest, and arose from the aorta just lateral to the second. This artery broke up into thi-ee smaller branches, all of which supplied the upper polo of the kidney. The; veins were normal.
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woMAi.ors I {i:\.\L \ KssELs 7'.)
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('.•ISC III. rilis picscritcd two normally placed Uidiicys, (lie ri^litIxMiip; sniallcr than the left. The riu,lit kidiicN' presented upon examination two renal aiteiies and one renal vein. The lower icnal ail(uy arosj' 'J cm. al)ov(> the hifincation of the aorta from its ventral aspect, and entei'cd the kidney at the lower pole I cm. al)o\c the inferior border-. The iip|)er icnal arlei_\-, which was the normal one, arose; from the dorsal aspect of the aorta 1 1 cm. above the bifurcation of the latter, and, aftei' runnin<2; a tortuous coui'se, (Mitered the kidney at \ho. hilum. The left kidney was normal. Two spermatic xcins emptied into the left renal \-ein.
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Case I\'. This presented upon examination two normally placed kidneys. The li^ht [jossessed two arteries; the normal one arose from the aoi'ta, and entertMJ the kidney at the hilum; the anomalous one arose from the ventro-latcM-al aspcH't of th(> aorta 5 cm. above the normal renal vessel, and entered the upper jiole after it had si)lit into two branches.
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The left kidney also possessed two arteries; the normal one coming from the ventro-lateral aspect of the aorta just opposite the normal rifiht I'cnal arteiy. and enterin^j; the kidney at the hilum; the accessory renal ai'tery branchin<;' from the superior mesenteric, and entering; the upper pole of th(> left kidney 3 cm. l)elow the superior l)order. The veiiis in both kidneys were normal.
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Case y. This sul)ject presented two normally placed kidneys of normal size. The right kidney contained a small anomalous artery arising from the aorta, and supplying the upper pole. The left kidney was normal, as were also the veins.
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Case VI. This presented two normally placed kidneys. The right contained four lai-ge renal arteries and two large renal veins. The first renal artery aros efrom the ventro-lateral aspect of the aorta 3 cm. above the bifurcation, and entered the kidney on the posterior surface of the lower pole 3 cm. above the inferior border. The second and third arteries took origin from the aorta at the ventral surface as a connnon large branch 1 cm. above the bifurcation but immediately divided into two rather large branches, which entered the kidney at the hilum. The fourth artery was a long slender one coming from the coeliac axis, and entering the kidney at the upper pole 4 cm. below the superior border. There were two renal veins present ; a large one entering the vena cava, and another somewhat smaller one also entering the posterior aspect of the vena cava. The left kidney possessed one accessory artery arising directly from the aorta and entering the kidney substance at the lower pole.
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Case VH. This was a case of a right kidney with a small accessory artery arising just above the normal renal artery, and entering the kidney at the upper pole 3 cm. below the superior border.
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Case VIII. In this case the left kidney was somewhat larger than normal, and contained three large renal arteries, one of which divided into three smaller branches. The first artery arose from the aorta, and entered the kidney at the lower pole, 4 cm. above the inferior
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80 CARBON GILLASPIE, L. I. MILLER, AND M. BASKIN
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bordei'. The second artery was the largest of the three. It arose from the aorta above the first, and entered the kidney at the hihim. The third artery arose from the aorta above the second, and divided into three smaller branches all of which supplied the upper pole. Two large renal veins were present which arose from the hilum and entered the vena cava.
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 +
Case IX. Both kidneys were normal in size and position. On the right side one large arterial trunk sprang from the aorta. This divided into two rather long slender branches after it had continued its course for 2 cm. Each of the two branches further divided into two other branches, thus four arteries entered the kidne.v: two at the upj^er pole 3 cm. below the superior border; the other two at the lower pole 4 cm. above the inferior border. Connecting the main renal trunk with the aorta was a plexus of vessels of varying sizes. This plexus, covering the walls of the aorta, appeared to be a persistence of the embryonic periaortic plexus. There was one renal vein which came from the hilum and entered the vena cava.
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 +
The left kidney possessed two large and two small arteries. The first, which seemed to be the normal one, arose directly from the aorta and entei'ed the kidney at the hilum. The second artery, whidi was a lorig sleilder vessel, came from the aorta and entered the anterior surface of the kidney 4 cm. below the superior border. On the left side two small arteries came from a plexus which resembled the periaortic plexus. On this side, these small arteries entered the kidney substance, while on the right side the plexus merely connected the large renal vessels with the aorta.
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 +
There were also two large renal veins, two smaller veins and a plexus of still smaller veins. This plexus connected the larger veins with the vena cava. The largest vein, which seemed to be the noi-mal one, came from the hilum of the kidney and entered the vena cava. The second large renal vein, which was a long slender one, connected the left spermatic vein with the anterior surface of the kidney. The plexus not only connected the vena cava with the large renal veins, but also formed an anastomosis between this venous plexus and the arterial plexus, so that the venous and arterial blood had a chance to mix.
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Cases X, XI, XII, and XIII. These four subjects had kidneys which were similar in most respects. Thei'e were two renal arteries arising from the aorta and entering the hila of the right and left kidneys in each case. The veins were normal.
 +
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Cases XIV, XV and XVI. These three cases possessed small accessory arteries arising from the aorta and entering the poles of the kidneys. In cases fourteen and fifteen the accessory arteries entered the lower poles, and in case sixteen the small arteries entered the upper pole.
 +
 +
Case XVII. This case possessed four arteries to the right kidney and thi-ee to the left, all of which arose dii'cctly from the aorta. There were also two renal veins from the left kidney, both of which arose from the hilum and cntcj-ed the vena cava.
 +
 +
 +
 +
AN'OMALOl'S UKN'M. \ KSSELS 81
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 +
Case X\'IIT. The lifjht kidney possessed two snudl renal arteries eoniinji; directly- fioin the aorta and enteiiiiK tlie kidney at the hilum. The left kichiey as well as the veins were normal.
 +
 +
Case XIX. Both kidneys posse.ssed three renal arteries rather small in size, whieh entered the kidney at the hilum. The veins were normal.
 +
 +
Case XX. The rijiht kidney was normal. The left po.sscsscd two renal arteries, both of which were branches of the aorta, and entered the kidney at the hilum. The veins were normal.
 +
 +
Case XXI. The right kidney possessed two renal arteries, one of which entered the supiMior, the other the inferior pole. The left kidney possessed three renal arteries, which arose fiom the aorta as separate branches, and entered the kidney at the hilum. The veins were normal.
 +
 +
Case XXII. The right kidney possessed three renal arteries, arising from the aorta and entering the kidney from the hilum. The left kidney possessed two renal arteries, one arising from the aorta and entering the kidney at the hilum, the other arising from the coeliac axis and entering the kidney at the superior pole 2 cm. from the upper border. The veins wove normal.
 +
 +
The embryological origin of the renal arteries has not yet been satisfactorily explained. The explanation as here offered is after Keibel and Mall.
 +
 +
His ('80) first observed multiple branches of the aorta supplying the mesonephi'os in 7 mm. embryos. A more extended account of them has been given by Broman. At first, when the Wolffian bodies are relatively small, the mesonephric vessels are correspondingly small. They come from the middle portion of the aorta (2nd to 8th thoracic). At the end of the first month the mesonephros reaches its greatest development. It receives many direct branches from the aorta at levels cranial as well as caudal to the original ones. In 8 mm. embryos there are twenty mesonephric arteries on each side (8 cervical to 12th thoracic segments). The last vessels to appear grow out from the region between the first and second lumbar segments. These are destined to persist as the remainder atrophy. There are then on each side^ — in maximo — thirty vessels distributed thi'oughout the entire mesonephric area. At first these are entirely distributed to the mesonephros, but later also supply the reproductive glands, suprarenal bodies, metanephi'oi, and diaphragm. These new regions of distribution prevent their com
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82
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CARBON GILLASPIE, L. I. MILLER, AND M. BASKIN
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 +
plete degeneration when the mesonephros disappears. A variable number of them persist as phrenic, suprarenal, renal, accessory renal, internal spermatic, accessory spermatic arteries, and as the rami ad lympho-glandulas and ad sympathetic um. The first mesonephric arteries found in embryos 5.3 mm. arise from the lateral surface of the aorta, and pass horizontally to the urogenital fold, reaching the malpighian corpuscles, and terminating in them with an enlargement which usually assumes a spherical shape. Later a network of vessels occupies the place of the enlargement. This network is also connected with the
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Fig. 1 Mesonephric arteries in human embryo of 18 mm. greatest length. Circles on anterior surface of aorta indicate origin of coeliac, superior and inferior mesenteric arteries (after Keibel and Mall).
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 +
 +
posterior cardinal vein. Case IX is an example of the persistence of this anastomosis. With increasing age the arteries continually recede into the lumbar segments disappearing from the thoracic ones.
 +
 +
The arteries are di\T.ded into three groups bj'^ the suprarenal body: the cranial group, which is dorsal to the suprarenal {1-2); the middle group, whose vessels pass through the suprarenal {R. 3-~4, L. 3-5); and the caudal group whose vessels pass over the ventral side of the suprarenal body {R. 5-6, L. 6-9). The mesonephric arteries {5-9) situated in the angle formed by the reproductive gland ventrally, the mesonephros laterally, and the
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ANOMALOUS HHXAL VESSELS 83
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 +
niotano])lin)s dorsally, form ;i network: tlio rote iirtoriosum urogiMiitalo. 'I'lio ines()ii('])liros, rei)r()tluctivo gland, and the nietanephros are siip])lie(l witli aiterial branches from this network, thus makinfi; tliese ()r}i;ans indei)endent of shigle branches for their blood sui)i)ly. Should one or several roots degenerate, neighboring arteries can take their places. In the above diagram, for instance, the second mesonephric artery has divided into an ascending and descending branch; the ascending one sui)plies tlie entire ii])per lialf of the mesonephros and the reproductive gland, a region that in the young embryo receives its blood from several mesonephric arteries belonging to more cranial segments. Tlie occurrence of this network at once explains why all persistent arteries that arise from the roots of this network show, within certain Umits a variability in the points of their origin from the aorta. Each of the nine to eleven remaining mesonephric arteries may become an internal spermatic artery, since all supply the reproductive glands. This explains the frequently observed multiplicity of these arteries, and the not infrequent difference in the place of origin of the right and left ones.
 +
 +
The renal arteries are not new formations, as some have claimed, but each is formed from a mesonephric artery. The kidney climbs upward to the mesonephric artery and as soon as sufficient blood supply is assured cranially, the caudal branches separate from it. ^\Tien the kidney has acquired its definitive position it possesses several arteries, and of these one becomes greath^ enlarged to form the definitive arterj-, while the others either degenerate, or persist as accessory renals. The definitive renal arterj'^ is either the last vessel of the second group, or first of the thkd group. The relations of both groups, i.e., of the second to the suprarenal artery, and of the third to the internal spermatic, explain the variation in which the renal artery arises from a suprarenal or from an internal spermatic. In the first case it may be the principal stem; in the latter, only an accessory renal. The relations between the urogenital rete and metanephros show how the accessory renal arteries ma^^ develop, and explain their varied relations to the kidney. Accessory renal
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84 CARBON GILLASPIE, L. I. MILLER AND M. BASKIN
 +
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arteries from the fii'st group will be branches of the superior suprarenal, and must pass over the dorsal surface of the kidneys. These consequently first reach the kidney on its dorsal surface and there penetrate its cortex. Those from the second group will be branches of either the middle or inferior suprarenal, and will reach either the hilus or the kidney or the medial edge above this. Those from the third group may enter the hilus at the medial edge below it, or on the ventral surface of the caudal half of the organ. Should a caudal branch be retained, it will be drawn upwards by the migration of the kidney. This condition explains why such an artery may cross the principal stem or another accessory.
 +
 +
Anomalous renal vessels are not only interesting from a purely scientific point of view, but are also of very great significance from a clinical and surgical standpoint.
 +
 +
The Mayos in 20 out of 27 cases operated upon by them for hydronephrosis, found anomalous blood vessels. The obstruction in each case was caused by the blood vessel crossing the uretero-pelvic juncture. The vessels passed to the lower pole of the kidney, and varied from the size of a knitting needle to that of the radial artery.
 +
 +
Levings, in a report of four cases on which operations were performed, found anomalous arteries which crossed the ureter, and to which condition he attributed the clinical symptoms of the patients. He also reported post-operative improvement in every case.
 +
 +
Rupert in 1913 found anomalous renal vessels in 35 out of 50 cadavers studied. In every case the kidneys were normally placed, and of normal size and shape. He concluded that the usual percentage given is too low, and that anomalous renal veins, while not as common as the arteries, do occur, and that on account of the thinness of their walls and lack of pulsations they increase the hazards of kidney operations.
 +
 +
It is very evident from the specimens at hand that a number of these arteries might be overlooked if it were necessary to remove or examine any of these kidneys. In view of the fact that anomalous renal arteries are important it is rather strange
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ANOMALorS i{i;nai, \ ksskls
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85 i. 4 86 CARBON GILLASPIE, L. I. MILLER, AND M. BASKIN
 +
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that their existence has been neglected in surgical anatomy teaching. As Rupert points out, the most common text books of anatomy and surgery make but brief mention of this condition, and some do not even refer to it at all.
 +
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The accepted percentage of 20 to 25 as given by Quaine and Gerrish is evidently too low. Senator as recently as 1905 made the statement that RedupUcations of one or both renal arteries is a rare condition and may be dismissed." Rupert found 35 cases or 70 per cent of anomalous renal vessels, while in this present investigation there are 22 cases out of 33, or 73 per cent. In all of these cases the kidneys were normally placed and the anomalous arteries were so , arranged that they would easily complicate surgical procedures.
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BIBLIOGRAPHY
 +
 +
Brodel, M. 1901 Johns Hopkins Hosp. Bull., Baltimore.
 +
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Broman, F. 1906 Ergeb. der Anat. U. Entw., Bd. 16.
 +
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Bremer, J. L. 1912 Am. Jour. Anat., vol. 13, no. 2.
 +
 +
Clark, E. R. 1912 Am. Jour. Anat., vol. 13.
 +
 +
Harvey, R. W. 1914 Anat. Rec, vol. 8.
 +
 +
Hill, E. C. 1905-1906 Johns Hopkins Hosp. Bull., vol. 16-17.
 +
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Jeidell, H. 1914 Anat. Rec, vol. 5.
 +
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Keibal and Mall Human Embryology.
 +
 +
Lewls, F. T. 1902 .\m. Jour. Anat., vol. 13.
 +
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Lewis, Papez 1914 Abstracts Proceedings, Am. Jour. Anat.
 +
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Levings, a. H. 1912 Wis. Med. Jour., March.
 +
 +
PoHLMAN, A. G. 1905 Johns Hopkins Hosp. Bull., vol. 16.
 +
 +
Rupert, R. R. 1913 Jour, of Gyn. and Obst., vol. 17.
 +
 +
Wilson, L. B. 1912 Collection of Papers, St. Mary's Hosp.
 +
 +
YoxjNG and Thompson 1903 Jour. Anat. and Phys., vol. 38.
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 +
 +
 +
SIZE AM) LENGTH RELATIONS OL THE RIGHT AND
 +
 +
LEFT TESTES OF PIGEONS IN HEALTH
 +
 +
AND DISEASE
 +
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OSCAR RIDDLE
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Carnegie Station for Experimental Evolution
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The right ovary luulergoes an early and more or less complete atroplw in most species of birds. Etzold ('91) has shown that in the sparrow the left testis is larger than the right. Firket ('14) and Swift ('15) have shown that in the chick embryo there were more primordial germ cells in the left gonad, and that this gonad is there also distinct^ larger than the right. Allen ('07) found that the sex cells were unequally distributed to the two gonads of the turtle, the left receiving most. In this form only 24-70 per cent of the sex cells ever enter the gonads. Our own accumulation of data on the size and length relations of the two testes of young and adult pigeons show a very decided predominate number of larger right testes; and also a distinct difference in shape of the two glands — the left though actually smaller in size is usually absolutely longer than the right. Changes in the size relation in birds dead of certain diseases — particularly tuberculosis — and in hybrids are also suggested by our data
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The meaning of this pronounced inequahty in the distribution of the primordial germ cells which is plainly associated with a larger left embryonic gonad, and the finding in adults of two groups of birds of a marked and nearly constant larger gonad, but this a different gonad in the two cases, is by no means clear. But, whatever this may mean, it is probably a situation of importance to the theory^ of sex. We present our present data then with the confession that on the main points the meaning is not clear, but with the conviction that they are not less valuable because of our present inability to clarify the puzzling situation, and hopeful that the data may stimulate the further
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87
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88 OSCAR RIDDLE
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accumulation of facts from enough forms, and of such varied kinds, as may lead to a better understanding of the embryonic and adult inequalities of the sex glands of birds.
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An examination of our data has shown that the measurements of glands of healthy birds should be grouped apart from those dead of disease; and those of pure species should be separated from hybrids. The justification of these separate groupings will appear later.
 +
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The relative size of the sex glands in healthy common pigeons
 +
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The weights of 31 pairs of testes from healthy common pigeons are recorded in table 1. In 27 of these pairs the left testis was the smaller; in 4 the left was the larger. In two or three of these latter cases — 12, 15 (22?) — the disparity of the two glands is so great as to make it clear that the smaller gland was wholly abnormal. In healthy common pigeons the right testis is larger than the left in a high proportion of cases.
 +
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Size relations of the testes of common pigeons dead of disease
 +
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In tables 2 and 6 the weights of 9 pairs of testes are given. In 7 of these the left gland was the smaller. It was larger in two instances; in one of these irregular cases, the smaller gland was again quite abnormally proportioned in reference to its larger associate
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Size relations of testes of pure species, healthy and dead of disease
 +
 +
The testes of only 5 healthy birds of pure species (dead of cold, exposure, accident) are included in table 6. In all of these cases the right testis was the larger.
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 +
The data for 46 individuals of pure species dead of disease are available. In table 2, 9 of the 10 individuals listed had larger right testes; the tenth had the two glands of equal size. Eleven further comparisons are supplied in table 3. Of these, 7 right testes are larger, 2 are smaller, and 2 are the size-equivalents of the left. Table 6 gives the data for 30 additional pairs. Of these, 7 of St. risoria all had larger right testes; 2 of T. orientalis both had larger right testes; 13 of Spil. tigrina — mostly not
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 +
 +
 +
TESTES OF llGEONS IN HEALTH AND DISEASE
 +
 +
 +
 +
89
 +
 +
 +
 +
mature birds luul, ."> larger, 5 siiiall(>r, and '.] ('([uivalont right testes. Four inisccllaiicous birds here had 2 larger and 2 smaller' rig! it testes.
 +
 +
Weight of right unit left testes of heallhj/ common pigeons
 +
 +
 +
 +
3 4 5 6
 +
 +
7 8 9 10 11 12 13 14 15 16
 +
 +
 +
 +
April 5. April 5. April 5. April 5. April 7. April 7. April 7. April 7. April 7. April 9. April 9. April 9. April 9. April 9. April 9. April 9.
 +
 +
 +
0.885
 +
0.515 1.475 0.845 1.185 0.990 1.055 0.820 1.410 0.765 1.190 0.975 1.260 1.000 1.235 0.945 1.280 1.225 1.075 0.900 1.025 0.710 0.025 0.715 1.460 1.390 1.010 0.720 0.275 1.425 1.125 0.500
 +
 +
 +
 +
PER CENT OP DIFF.
 +
 +
 +
 +
-71.8* -74.6 -19.7 -28.7 -84.3 -22.1 -26.0 -30.7
 +
 +
- 4.5 -19.4 -44.4
 +
 +
+2760.0
 +
 +
- 5.0 -40.3
 +
 +
+418.2 -125.0
 +
 +
 +
 +
April 9. July 5. . July 5. . July 5. . July 5.. July 5. . July 9.. July 9.. July 13. July 13. July 16. July 18. July 20. July 20.
 +
 +
 +
 +
July 21 (juv.; WEIGHT
 +
 +
 +
PER CENT OF DIFF.
 +
 +
 +
R
 +
 +
 +
= 1.115
 +
 +
 +
 +
 +
L
 +
 +
 +
= 1.015
 +
 +
 +
-9.9
 +
 +
 +
R
 +
 +
 +
= 1.010
 +
 +
 +
 +
 +
L
 +
 +
 +
= 0.750
 +
 +
 +
-34.7
 +
 +
 +
R
 +
 +
 +
= 1.220
 +
 +
 +
 +
 +
L
 +
 +
 +
= 1.375
 +
 +
 +
+ 12.7
 +
 +
 +
R
 +
 +
 +
= 1.140
 +
 +
 +
 +
 +
L
 +
 +
 +
= 0.970
 +
 +
 +
-17.5
 +
 +
 +
R
 +
 +
 +
= 1.158
 +
 +
 +
 +
 +
L
 +
 +
 +
= 0.765
 +
 +
 +
-51.4
 +
 +
 +
R
 +
 +
 +
= 0.370
 +
 +
 +
 +
 +
L
 +
 +
 +
= 0.720
 +
 +
 +
+94.6
 +
 +
 +
R
 +
 +
 +
= 0.855
 +
 +
 +
 +
 +
L
 +
 +
 +
= 0.800
 +
 +
 +
- 6.9
 +
 +
 +
R
 +
 +
 +
= 1.643
 +
 +
 +
 +
 +
L
 +
 +
 +
= 1.030
 +
 +
 +
-59.5
 +
 +
 +
R
 +
 +
 +
= 0.571
 +
 +
 +
 +
 +
L
 +
 +
 +
= 0.540
 +
 +
 +
- 5.7
 +
 +
 +
R
 +
 +
 +
= 0.820
 +
 +
 +
 +
 +
L
 +
 +
 +
= 0.631
 +
 +
 +
-29.9
 +
 +
 +
R
 +
 +
 +
= 1.820
 +
 +
 +
 +
 +
L
 +
 +
 +
= 1.500
 +
 +
 +
-21.3
 +
 +
 +
R
 +
 +
 +
= 1.600
 +
 +
 +
 +
 +
L
 +
 +
 +
= 0.536
 +
 +
 +
-198.5
 +
 +
 +
R
 +
 +
 +
= 0.051
 +
 +
 +
 +
 +
L
 +
 +
 +
= 0.040
 +
 +
 +
-27.5
 +
 +
 +
R
 +
 +
 +
= 1.390
 +
 +
 +
 +
 +
L
 +
 +
 +
= 1.085
 +
 +
 +
-28.1
 +
 +
 +
R
 +
 +
 +
= 0.0004
 +
 +
 +
 +
 +
L
 +
 +
 +
= 0.0003
 +
 +
 +
-33.3
 +
 +
 +
 +
Left smaller in 27; larger in 4.
 +
 +
 +
In calculating percentage differences in these tables the smaller gland is
 +
considered as equal to 100 per cent.
 +
 +
' In both of these cases where the right testes weighed less than the left it will be seen that both testes were quite small — so small as perhaps to raise a question as to the reliability of the weights.
 +
 +
 +
 +
90
 +
 +
 +
 +
OSCAR RIDDLE
 +
 +
 +
 +
Size of testes in healthy specific hybrids
 +
 +
In tables 4 and 5 the data for 30 healthy young hybrids are given. The very, small gonad size of most of these young birds
 +
 +
TABLE 2 Weights of right and left testes of various pigeons (classified) dead of disease
 +
 +
 +
 +
PER CENT OF DIFF.
 +
 +
 +
 +
PER CENT OF DIFF.
 +
 +
 +
 +
1. Common pigeons
 +
 +
 +
 +
32 33^
 +
 +
 +
 +
July 28
 +
 +
September 1 ,
 +
 +
 +
 +
R = 0.122
 +
 +
L = 0.460
 +
 +
R = 0.580
 +
 +
L = 0.415
 +
 +
 +
 +
+277.0 -39.8
 +
 +
 +
 +
September 28 December 14.
 +
 +
 +
 +
= 0.037 = 0.034 = 1.300
 +
 +
= 0.850 (?)
 +
 +
 +
 +
-52.9
 +
 +
 +
 +
2. Blond and white wing doves and their hybrids
 +
 +
 +
 +
Hybrids, (specific)
 +
 +
 +
 +
Pure
 +
 +
 +
 +
36*
 +
 +
37*
 +
 +
38*
 +
 +
39
 +
 +
40
 +
 +
41*
 +
 +
 +
 +
June 10
 +
 +
June 27
 +
 +
July 23
 +
 +
September 12 October 17.. October 31 . . .
 +
 +
 +
 +
0.417
 +
 +
0.305
 +
 +
0.050
 +
 +
0.035
 +
 +
0.032
 +
 +
0.046
 +
 +
0.765
 +
 +
0.705
 +
 +
0.0053
 +
 +
0.0067
 +
 +
0.090
 +
 +
0.090
 +
 +
 +
 +
-36.7
 +
 +
 +
42*
 +
 +
 +
-42.9
 +
 +
 +
43*
 +
 +
 +
+43.8
 +
 +
 +
44*
 +
 +
 +
-8 5
 +
 +
 +
45*
 +
 +
 +
+26.4
 +
 +
 +
46
 +
 +
 +
= 0.0
 +
 +
 +
47*
 +
 +
 +
 +
April 9
 +
 +
April 17
 +
 +
September 26 October 25... October 25 . . October 28...
 +
 +
 +
 +
0.045 0.025 0.060 0.050 0.055 0.055 0.068 0.056 0.046 0.033 0.030 0.027
 +
 +
 +
 +
-80.0 -20.0 0.0 -21.4 -39.3 -11.1
 +
 +
c
 +
 +
 +
. Other hybr
 +
 +
 +
ds (spec
 +
 +
 +
ific,
 +
 +
 +
ex. 51, 52 = gen.)
 +
 +
 +
 +
 +
48 49 50
 +
 +
 +
August 28.. June 10.... August 28..
 +
 +
 +
■{ ■{ ■{
 +
 +
 +
R = 0.845 L = 0.600 R = 0.190 L = 0.160 R = 0.133 L = 0.115
 +
 +
 +
-40.8 -18.8 -15.7
 +
 +
 +
51*
 +
 +
52
 +
 +
53*
 +
 +
 +
September 18. I September 23. | September 27. <
 +
 +
 +
R = 0.580 L = 0.510 R = 0.040 L = 0.031 R = 0.010 L = 0.012
 +
 +
 +
-13.7 -29.0 +20.0
 +
 +
 +
 +
4. Other pure species
 +
 +
 +
 +
54^ 55^
 +
 +
 +
 +
August 26
 +
 +
November 10
 +
 +
 +
 +
R = 0.040
 +
 +
L = 0.022
 +
 +
R = 0.033
 +
 +
L = 0.023
 +
 +
 +
 +
-81.8
 +
 +
 +
 +
-43.5
 +
 +
 +
 +
56
 +
 +
 +
 +
57^
 +
 +
 +
 +
October 31.
 +
 +
 +
 +
November 9.
 +
 +
 +
 +
0.465 0.445 0.015 0.013
 +
 +
 +
 +
4.5 15.4
 +
 +
 +
Tuberculosis found.
 +
 +
 +
TESTKS OK PIGEONS IN HH.VLTH .\N1) DISEASE
 +
 +
 +
 +
91
 +
 +
 +
 +
T.\ni.i:3. Weights of testes of doves (classified) dead of disease March 29 to November 25, 1915
 +
 +
 +
 +
PER CENT or DIFF.
 +
 +
 +
.NO.
 +
 +
 +
 +
PER CENT OV DIPF. Pure species Specific hybrids
 +
 +
 +
 +
 +
59*
 +
 +
60*
 +
 +
61*
 +
 +
62*
 +
 +
63
 +
 +
64*
 +
 +
65
 +
 +
66*
 +
 +
67*
 +
 +
68*
 +
 +
69*
 +
 +
 +
March •_".». ... |
 +
 +
April 1 I
 +
 +
April 2 \
 +
 +
April 16 <
 +
 +
September 2. <
 +
 +
September 5. . <
 +
 +
September 11 f (Juv.) \
 +
 +
September 14 < September 29 < October 3. . . . < November 11. <
 +
 +
 +
R = 0.012 L = 0.018 R = 0.008 L = 0.005 R = 0.105 L = 0.105 R = 0.022' L = 0.060 R = 0.1582 L = 0.146 R = 0.032 L = 0.020 R = 0.003 L = 0.002 R = 0.023 L = 0.023 R = 0.138 L = 0.116 R"= 0.045 L = 0.030 R = 0.082 L = 0.067
 +
 +
 +
+50.0
 +
 +
-60.0
 +
 +
= 0.0
 +
 +
+ 172.7
 +
 +
- 8.2 • -60.0
 +
 +
-50.0 = 0.0 -18.9 -50.0
 +
 +
— 22.4
 +
 +
 +
76*
 +
 +
77* 78* 79* 80
 +
 +
81*
 +
 +
82*
 +
 +
83* 84*
 +
 +
85* 86*
 +
 +
87* 88
 +
 +
 +
May 1 1
 +
 +
July 23 I
 +
 +
July 30 1
 +
 +
August 22 — \ August 29.... \ September 19 < September 28 \
 +
 +
October 7. . . . {
 +
 +
October 19 f (Juv.) 1
 +
 +
October 24 . . . <
 +
 +
November 21 f (Juv.) 1
 +
 +
November 22. < November 25. <
 +
 +
 +
R = 0.030 L = 0.031 R = 0.262 L = 0.212 R = 0.040 L = 0.032 R = 0.076 L = 0.077 R = 0.6052 L = 0.454 R = 0.188 L = 0.152 R = 0.026 L = 0.026 R = 0.025 L = 0.021 R = 0.007 L = 0.006 R = 0.0.30 L = 0.030 R = 0.015 L = 0.020 R = 0.014 L = 0.016 R = 0.018 L = 0.015
 +
 +
 +
+ 3.3 -23.6 -25.0 + 1.3 -33.3 -23.7 = 0.0 -19.0 -16.6 - 0.0 +.33.3
 +
 +
 +
Generic hybrids
 +
 +
 +
+ 14.3
 +
 +
 +
 +
 +
April 2 1
 +
 +
May 30 |
 +
 +
June 20 <
 +
 +
July 16 1
 +
 +
August 2 <
 +
 +
August 17. . . . <
 +
 +
 +
R = 0.140 L = 0.125 R = 0.320 L = 0.280 R = 0.082 L = 0.076 R = 0.300 L = 0.298 R = 0.012 L = 0.010 R = 0.040 L = 0.037
 +
 +
 +
-12.0 -14.3
 +
 +
- 7.9
 +
 +
- 0.7 -20.0
 +
 +
- 8.1
 +
 +
 +
-20.0
 +
 +
 +
70*
 +
 +
 +
Common pigeons
 +
 +
 +
71*
 +
 +
72 73* 74* 75*
 +
 +
 +
89
 +
 +
90*
 +
 +
91
 +
 +
 +
April 28 1
 +
 +
August 25 ... . < April 5 \
 +
 +
 +
R = 1.260 L = 1.035 R = 0.098 L = 0.087 R = 1.105' L = 0.990
 +
 +
 +
-21.7 -12.6 -11.6
 +
 +
 +
Tuberculosis found.
 +
'The left suprarenal wholly involved in a tubercle nodule weighing nearly 1.0 gr.
 +
 +
2 Healthy.
 +
 +
' A hybrid from a family cross.
 +
 +
 +
 +
92
 +
 +
 +
 +
OSCAR RIDDLE
 +
 +
 +
 +
TABLE 4 Weight, length and ividth of testes of young Ring dove (specific) hybrids — killed
 +
 +
PER CENT
 +
 +
 +
 +
 +
PER CENT
 +
 +
 +
so.
 +
 +
 +
DATE
 +
 +
 +
'•
 +
 +
 +
 +
 +
WEIGHT
 +
 +
 +
OP DIFFERENCE
 +
 +
 +
LENGTH AND WIDTH
 +
 +
 +
OF DIFFERENCE
 +
 +
 +
88a
 +
 +
 +
December 4,
 +
 +
 +
1915.
 +
 +
 +
/ I
 +
 +
 +
R = 0.035 L =0.030
 +
 +
 +
-16.7
 +
 +
 +
8.0x3.8
 +
 +
 +
 +
 +
S9a
 +
 +
 +
December 4,
 +
 +
 +
1915.
 +
 +
 +
/ I
 +
 +
 +
R = 0.007(?) L = 0.005 (?)
 +
 +
 +
-40.0
 +
 +
 +
5.1 xl.6 .6x 1.4
 +
 +
 +
-10.9
 +
 +
 +
90a
 +
 +
 +
December 4,
 +
 +
 +
1915.
 +
 +
 +
 +
 +
R = 0.004(?) L = 0.005(?) R = less than
 +
 +
 +
+25.0
 +
 +
 +
4.1 xO.9 4.9x0.9
 +
 +
 +
+ 19.5
 +
 +
 +
91a
 +
 +
 +
December 4,
 +
 +
 +
1915.
 +
 +
 +
<
 +
 +
 +
0.005(?)
 +
 +
 +
 +
 +
5.1 xO.8 L = 0.005(?)
 +
 +
 +
+ ?
 +
 +
 +
5.1 xl.5
 +
 +
 +
= 0.0
 +
 +
 +
92
 +
 +
 +
December 4,
 +
 +
 +
1915.
 +
 +
 +
{
 +
 +
 +
R = 0.005(?) L = 0.005(?)
 +
 +
 +
= 0.0
 +
 +
 +
4.8x 1.5 5.3 X 1.4
 +
 +
 +
+ 10.4
 +
 +
 +
93
 +
 +
 +
December 4,
 +
 +
 +
1915
 +
 +
 +
7
 +
 +
I
 +
 +
 +
R = 0.007 (?) L = 0.005(?)
 +
 +
 +
-40.0
 +
 +
 +
4.8x 1.6 5.3x0.9
 +
 +
 +
+ 10.4
 +
 +
 +
94
 +
 +
 +
December 4,
 +
 +
 +
1915
 +
 +
 +
{
 +
 +
 +
R = 0.015 L = 0.010
 +
 +
 +
-50.0
 +
 +
 +
6.6x 1.9 6.6x 1.5
 +
 +
 +
= 0.0
 +
 +
 +
95*
 +
 +
 +
December 8,
 +
 +
 +
1915.
 +
 +
 +
/
 +
 +
I
 +
 +
 +
R = 0.025 L = 0.025
 +
 +
 +
= 0.0
 +
 +
 +
8.3x2.2 9. Ox 1.9
 +
 +
 +
+8.4
 +
 +
 +
96
 +
 +
 +
December 8,
 +
 +
 +
1915.
 +
 +
 +
{
 +
 +
 +
R = 0.037 L = 0.025
 +
 +
 +
-48.0
 +
 +
 +
. 8.4x2.7 7.2x2.2
 +
 +
 +
-16.7
 +
 +
 +
97
 +
 +
 +
December 8,
 +
 +
 +
1915
 +
 +
 +
{
 +
 +
 +
R = 0.010 L = 0.007
 +
 +
 +
-42.9
 +
 +
 +
5.0x1.8 5. Ox 1.5
 +
 +
 +
= 0.0
 +
 +
 +
98
 +
 +
 +
December 8,
 +
 +
 +
1915.
 +
 +
 +
{
 +
 +
 +
R = 0.320 L = 0.280
 +
 +
 +
-14.3 99
 +
 +
 +
Deceml^er 8,
 +
 +
 +
1915.
 +
 +
 +
{
 +
 +
 +
R = 0.007(?) L = 0.005 (?)
 +
 +
 +
-40.0
 +
 +
 +
4.4x 1.7 5.0x1.0
 +
 +
 +
+ 13.6
 +
 +
 +
100
 +
 +
 +
December 8,
 +
 +
 +
1915
 +
 +
 +
{
 +
 +
 +
R = 0.025 L = 0.020
 +
 +
 +
-25.0
 +
 +
 +
6.8x2.2 6.7x2.1
 +
 +
 +
-1.5
 +
 +
 +
101
 +
 +
 +
December 8,
 +
 +
 +
1915.
 +
 +
 +
/
 +
 +
I
 +
 +
 +
R = 0.010 L = 0.007
 +
 +
 +
-42.9
 +
 +
 +
6.5x 1.9 6.5x 1.5
 +
 +
 +
= 0.0
 +
 +
 +
 +
 +
December 8,
 +
 +
 +
1915.
 +
 +
 +
{
 +
 +
 +
R = 0.012 L = 0.012
 +
 +
 +
= 0.0 102
 +
 +
 +
Summary :
 +
 +
 +
 +
 +
103
 +
 +
 +
December 8,
 +
 +
 +
1915.
 +
 +
 +
{
 +
 +
 +
R = 0.035 L = 0.025
 +
 +
 +
-40.0
 +
 +
 +
Left larger i Left smaller
 +
 +
 +
n 2
 +
 +
in ... . 13
 +
 +
 +
104
 +
 +
 +
December 8,
 +
 +
 +
1915.
 +
 +
 +
/ I
 +
 +
 +
R = 0.100 L = 0.070
 +
 +
 +
-42.9
 +
 +
 +
Two equal ir Left longer i
 +
 +
 +
1 3
 +
 +
n... 5
 +
 +
 +
105
 +
 +
 +
December 8,
 +
 +
 +
1915.
 +
 +
 +
_{
 +
 +
 +
R = 0.017 L = 0.015
 +
 +
 +
-13.3
 +
 +
 +
Left shorter Left equal ii
 +
 +
 +
in ... . 3 1 4
 +
 +
 +
Tuberculosis found.
 +
 +
 +
TESTES OF PIGEONS IN IlKAl/ni AND DISEASE
 +
 +
 +
 +
WA
 +
 +
 +
 +
is probal)!} r('s]M)iisil)l(> for (he f;iilui-(' of our wei^Iiings to (lilTci-eutiuto betwoon llie iiuisscs of s(!Vonil jKiirs of the testes. In the 30 pnirs IS right testes were larger. 5 were smaller, 7 were not differentiated by the weighings.
 +
 +
Size, of testes in hybrids dead of disease
 +
 +
Seven of the 10 .s]^ocific li3^brids of table 2 had larger right testes; 2 had smaller; 1 had the testes of e(iiial size. Of the two generic hybrids (52, 53) represented in this table, one had a larger and one a smaller right testes. Tn table 3 are listed 13 s])orific hybrids; 7 larger right testes, 4 smaller, and 2 equivalents.
 +
 +
TABLE 5 Weight, length and width of testes of young Ring dove (specific) hybrids — killed
 +
 +
PER CE.\T
 +
 +
 +
 +
 +
PER CENT
 +
 +
 +
NO.
 +
 +
 +
DATE WEIGHT
 +
 +
 +
OF DIFFERENCE
 +
 +
 +
LENGTH .\ND WIDTH
 +
 +
 +
OP DIFFERENCE f
 +
 +
 +
R
 +
 +
 +
= too small to
 +
 +
 +
 +
 +
3.2 X 1.5
 +
 +
 +
 +
 +
106
 +
 +
 +
Novembor 29, 191.5
 +
 +
 +
J
 +
 +
 +
 +
 +
weigh
 +
 +
 +
L
 +
 +
 +
= 0.005
 +
 +
 +
+ ?.o
 +
 +
 +
4.7 .X 1.8
 +
 +
 +
+ 46.9
 +
 +
 +
107
 +
 +
 +
November 29, 1915
 +
 +
 +
/ 1
 +
 +
 +
R L
 +
 +
 +
= 0.007 = 0.007
 +
 +
 +
= 0.0
 +
 +
 +
5.8 X 1.6 5.4x2.0
 +
 +
 +
- 7.4
 +
 +
 +
108
 +
 +
 +
November 29, 1915
 +
 +
 +
 +
 +
R L
 +
 +
 +
= 0.009 = 0.010
 +
 +
 +
+11.1
 +
 +
 +
5.8x 1.8 6.4x 1.8
 +
 +
 +
+ 10.3
 +
 +
 +
109
 +
 +
 +
November 29, 1915
 +
 +
 +
1
 +
 +
 +
R L
 +
 +
 +
= 0.020 = 0.019
 +
 +
 +
- 5.3
 +
 +
 +
8.2x1.9 8.2x1.9
 +
 +
 +
= 0.0
 +
 +
 +
110
 +
 +
 +
November 29, 1915
 +
 +
 +
{
 +
 +
 +
R L
 +
 +
 +
= 0.018 = 0.020
 +
 +
 +
+11.1
 +
 +
 +
7.4x2.0 7.4x2.0
 +
 +
 +
= 0.0
 +
 +
 +
111
 +
 +
 +
November 29, 1915
 +
 +
 +
(
 +
 +
 +
R L
 +
 +
 +
= 0.010 = 0.010
 +
 +
 +
= 0.0
 +
 +
 +
5.9x2.2 7.4x1.6
 +
 +
 +
+25.4
 +
 +
 +
112
 +
 +
 +
November 29, 1915
 +
 +
 +
/ \
 +
 +
 +
R L
 +
 +
 +
= 0.190 = 0.170
 +
 +
 +
-11.8
 +
 +
 +
17.0x4.5
 +
 +
15.7x4.5
 +
 +
 +
- 8.3
 +
 +
 +
113
 +
 +
 +
December 3, 1915. .
 +
 +
 +
1
 +
 +
 +
R L
 +
 +
 +
= 0.007 = 0.005
 +
 +
 +
-40
 +
 +
 +
5.7x 1.5 4.9x0.9
 +
 +
 +
-16.3
 +
 +
 +
114
 +
 +
 +
December 3, 1915. . December 3, 1915. . December 3, 1915. . December 3, 1915. .
 +
 +
 +
/
 +
 +
I / I / I / I
 +
 +
 +
R L R L R L R L
 +
 +
 +
= 0.007 = 0.005 = 0.007 = 0.007 = 0.005 = 0.005 = 0.007 = 0.005
 +
 +
 +
-40.0 = = -40.0
 +
 +
 +
5.5x1.4 4.2x1.0
 +
 +
 +
-30.9
 +
 +
 +
11,5 116 117
 +
 +
 +
Summary : Left larger ii Left smaller Two equal ii Left longer i Left shorter
 +
 +
 +
1 3
 +
 +
in 5
 +
 +
1 4
 +
 +
n 3
 +
 +
in 4
 +
 +
 +
Two equal ir
 +
 +
 +
1 2
 +
 +
 +
 +
THE ANATOMIC.VL RECORD, VOL. 11, NO. 3
 +
 +
 +
 +
TABLE 6 Weights and measurements of testes — birds classified as to kind and disease PER CENT
 +
 +
 +
 +
 +
PER
 +
 +
 +
NO.
 +
 +
 +
DATE
 +
 +
 +
DISEASE'
 +
 +
 +
WEIGHTS
 +
 +
 +
OF DIFFERENCE
 +
 +
 +
LENGTH AND WIDTH
 +
 +
 +
CENT OF
 +
 +
DrFFER ENCE
 +
 +
 +
 +
Pure species — Spil. tigrina
 +
 +
 +
 +
118 119 120 121 122 123 124 125 126 127 128 129 130
 +
 +
 +
 +
January 7 Januarj' 9 January 11 January 11 January 13 January 15 January 18 January 25 January 29 March 1 May 17 January 28 January 28
 +
 +
 +
 +
Worms
 +
 +
Worms (juv.)
 +
 +
Liver
 +
 +
Liver and worms. Worms
 +
 +
 +
 +
Worms liver.
 +
 +
 +
 +
Intest. and liver. . .,
 +
 +
Intest
 +
 +
Worms (old)
 +
 +
Intest. (juv.)
 +
 +
Sp. Li. (old)
 +
 +
Liver and spleen . . . Worms
 +
 +
 +
 +
0.050
 +
 +
0.050
 +
 +
0.012
 +
 +
0.014
 +
 +
0.040
 +
 +
0.045
 +
 +
0.055
 +
 +
0.055
 +
 +
0.080
 +
 +
0.080
 +
 +
0.040
 +
 +
0.045
 +
 +
031
 +
 +
0.027
 +
 +
030
 +
 +
0.045
 +
 +
0.030
 +
 +
0.022
 +
 +
0.004t
 +
 +
0.004 0.047
 +
 +
0.035
 +
 +
0.030
 +
 +
0.025
 +
 +
0.027
 +
 +
0.030
 +
 +
 +
 +
= 0.0
 +
 +
 +
+ 16.6
 +
 +
 +
+ 12.5
 +
 +
 +
=
 +
 +
 +
= 0.0
 +
 +
 +
+ 12.5
 +
 +
 +
-14.8
 +
 +
 +
+ 16.6
 +
 +
 +
-36.4
 +
 +
 +
- ?.o
 +
 +
 +
-34 3
 +
 +
 +
-20.0
 +
 +
 +
+ 11.1
 +
 +
 +
 +
10.8 5.3 7.0 7.3 9.0 8.3 8 9
 +
 +
10 6
 +
 +
 +
 +
7
 +
 +
6.8 7.2 6.7
 +
 +
 +
 +
x3.4 x3.0 x2.2 xl.8 x3.0 X 2.8 x3.2 x3.0 X 4.5 x4 3 x3.0 x2.8 x3.4 x2.7 x3.0 x3.1 x2.6 x2.3 x2.1 X 1.9
 +
 +
 +
 +
x3.1 x2.4 x2.5 x2.5
 +
 +
 +
 +
+38.5 +32.1 +23.3 = 0.0 + 13.9 +26.2 +31.4 +21.1 - 8.6 +4.8 -20.8 + 5.9 +31.3
 +
 +
 +
 +
Pure species — T. orientalis
 +
 +
 +
 +
131 132 133 134 135
 +
 +
 +
 +
March 8 March 23 March 23 March 23 March 25
 +
 +
 +
 +
Lu. liver
 +
 +
Cold (juv.)... Cold (juv.). . . Cold (juv.). . . Intest. (juv.).
 +
 +
 +
 +
R = 0.016 L = 0.015 R = 010 L = 0.008 R = 0.010 L = 0.008 R = 0.011 L = 0.009 R = 0.010 L = 0.007
 +
 +
 +
 +
6 6
 +
 +
 +
25.0
 +
 +
 +
25
 +
 +
 +
22 2
 +
 +
 +
42.9
 +
 +
 +
 +
5.4x 1
 +
 +
3.7 X 1 6. Ox 1 4 7 x 1 7 0x1 6.3 X 1
 +
 +
5.8 X 1.9 5.5 X 1 .6
 +
 +
 +
 +
-45.9 -27.6 -11.1 - 5.5
 +
 +
 +
 +
1 Abbreviations of the names of organs to their first two letters, implies that advanced and very evident tuberculosis was found in those organs (lungs, spleen, liver, joints, mesentery, intestine). Where more than one organ was affected the name of the organ (or organs) apparently most affected is written first. When the word is written out it denotes that this organ was abnormal, but not necessarily tubercular; immature birds are designated — (juv.).
 +
 +
94
 +
 +
 +
 +
TESTKS OI' I'KiKONS IN II KA I/I'll \.\|) DISK TABLK O-CoiitiiuuMl
 +
 +
 +
 +
.\SK
 +
 +
 +
 +
<).')
 +
 +
 +
 +
PEXt CENT
 +
 +
OP DIKKBH ENCE
 +
 +
 +
 +
LRNQTH AND WIDTH
 +
 +
 +
 +
PER CENT OP DIPFER BNOB
 +
 +
 +
 +
Pure species — St. risoria
 +
 +
 +
136
 +
 +
 +
Docomhor 4
 +
 +
 +
Sp. li.
 +
 +
 +
lu {
 +
 +
 +
R = 030 L = 025
 +
 +
 +
-20 137
 +
 +
 +
January 29
 +
 +
 +
Sp. lu.
 +
 +
 +
li • ■ 1
 +
 +
 +
R = 053 L = 031
 +
 +
 +
-70 9
 +
 +
 +
9 2x3 3 10. ox 1.9
 +
 +
 +
+ 14.1
 +
 +
 +
138
 +
 +
 +
January 29
 +
 +
 +
Sp. lu.
 +
 +
 +
li 1
 +
 +
 +
R = 0.015 L = 010
 +
 +
 +
-50
 +
 +
 +
6.0x2.3 6.6x2.0
 +
 +
 +
+ 10.0
 +
 +
 +
139
 +
 +
 +
February 4
 +
 +
 +
Tntost.
 +
 +
 +
 +
 +
R = 037 L = 020
 +
 +
 +
-85.0
 +
 +
 +
8.4x2 9
 +
 +
8.8x2.3
 +
 +
 +
+ 4.8
 +
 +
 +
140
 +
 +
 +
^Farch 15
 +
 +
 +
Lu. (?
 +
 +
 +
liver <
 +
 +
 +
R = 0.550 L = 0.401
 +
 +
 +
-37.1 141
 +
 +
 +
May 10
 +
 +
 +
Sp
 +
 +
 +
 +
 +
R = 0.023 L = 0.019
 +
 +
 +
-21.1
 +
 +
 +
SO
 +
 +
S.O
 +
 +
 +
= 0.0
 +
 +
 +
142
 +
 +
 +
May 14
 +
 +
 +
Jo., lu
 +
 +
 +
sp. ; liver ■ ■ ■ i
 +
 +
 +
R = 0.020 L = 0.017
 +
 +
 +
-17.6
 +
 +
 +
6.7 6.9
 +
 +
 +
+ 2.9
 +
 +
 +
 +
Miscellaneous — pure species
 +
 +
 +
 +
143
 +
 +
 +
November 17
 +
 +
 +
Canker, li
 +
 +
 +
, sp. lu .
 +
 +
 +
/
 +
 +
■ 1
 +
 +
 +
R = 010 L = 0.007
 +
 +
 +
-42.9 144
 +
 +
 +
November 17
 +
 +
 +
Fight and
 +
 +
 +
hemorr.
 +
 +
 +
/
 +
 +
 +
R =0.595 L = 0.475
 +
 +
 +
-25.3 145
 +
 +
 +
November 18
 +
 +
 +
I'nknown
 +
 +
 +
(juv.)...
 +
 +
 +
^{
 +
 +
 +
R = 0038 L = 0.0042
 +
 +
 +
+ 10.5 146
 +
 +
 +
November 20
 +
 +
 +
Cold {?)..
 +
 +
 +
 +
 +
^
 +
 +
 +
R = 0.165 L = 0.127
 +
 +
 +
-29.9 147
 +
 +
 +
December 3
 +
 +
 +
Liver
 +
 +
 +
 +
 +
■{
 +
 +
 +
R = 0.015 L = 0.016
 +
 +
 +
+ 6.7
 +
 +
Common pigeons
 +
 +
 +
 +
148
 +
 +
 +
January 3
 +
 +
 +
Unknown (juv.). . .
 +
 +
 +
{
 +
 +
 +
R = 0.005(?) L = 0.005(?)
 +
 +
 +
 +
 +
5.4x 1.5 5.2x1.3
 +
 +
 +
- 3.8
 +
 +
 +
149
 +
 +
 +
January 17
 +
 +
 +
L'nknown (juv.). . .
 +
 +
 +
/ I
 +
 +
 +
R = 0.009(?) L = 0.010(?) R = 0.004?
 +
 +
 +
+ 11.1
 +
 +
 +
5.8x 1.5
 +
 +
6.5x1.7
 +
 +
about 6.0
 +
 +
 +
+ 12.1
 +
 +
 +
150
 +
 +
 +
February 15
 +
 +
 +
Li. pleura
 +
 +
 +
<
 +
 +
 +
L = 0.003?
 +
 +
 +
-33.3?
 +
 +
 +
mm. long about 5.3 mm. long
 +
 +
 +
+ 13.2
 +
 +
 +
151
 +
 +
 +
April 13
 +
 +
 +
Hemorrhage, liver.
 +
 +
 +
/
 +
 +
 +
R = 0.014 L = 0.010
 +
 +
 +
-40.0
 +
 +
 +
6.4 6.4
 +
 +
 +
= 0.0
 +
 +
 +
152
 +
 +
 +
April 17
 +
 +
 +
Appar. healthy. . . .
 +
 +
 +
1 1
 +
 +
 +
R = 1.340 L = 1 315
 +
 +
 +
- 1.1
 +
 +
 +
20.2x12.1 23.9x10.7
 +
 +
 +
+ 18.3
 +
 +
 +
153
 +
 +
 +
May 4
 +
 +
 +
Weakling (juv.). . .
 +
 +
 +
/ I
 +
 +
 +
R = 0.006 L = 0.004
 +
 +
 +
-50.0
 +
 +
 +
5.2 5.2
 +
 +
 +
= 0.0
 +
 +
 +
 +
96
 +
 +
 +
 +
OSCAR RIDDLE
 +
 +
 +
 +
TABLE 6— Continued
 +
 +
 +
 +
WEIGHTS
 +
 +
 +
 +
PER CENT OP DIF FEREXCE
 +
 +
 +
 +
PER LENGTH CEXT OF
 +
 +
AND WIDTH DIFFERENCE
 +
 +
 +
 +
Hybrids — from crosses of species
 +
 +
 +
 +
154 155 156 157 158
 +
 +
159
 +
 +
160
 +
 +
161 162 163 164
 +
 +
165
 +
 +
166
 +
 +
167 168 169 170 171 172 173
 +
 +
 +
 +
January 1
 +
 +
January 4
 +
 +
January 8
 +
 +
January 15
 +
 +
December 3
 +
 +
January 15 January 27
 +
 +
Febi'uary 18
 +
 +
March 6
 +
 +
January 22
 +
 +
January 24
 +
 +
January 27 February 13
 +
 +
March 17 March 18 March 26 March 28 March 29 April 12 April 20
 +
 +
 +
 +
Cold? (juv.). Sp. liver ....
 +
 +
(Juv.)
 +
 +
Lu
 +
 +
Li. (juv.)....
 +
 +
 +
 +
Sp. jo I
 +
 +
Healthy d w a r f ; / killed \
 +
 +
 +
 +
Sp- 111 I
 +
 +
Sp. li., etc i
 +
 +
Li., sp \
 +
 +
Lu., liver, spleen. . .
 +
 +
Liver, lu. (?)
 +
 +
Lu., liver, spleen J (juv.) \
 +
 +
Sp., lu. liver
 +
 +
 +
 +
Sp. li., lu
 +
 +
Intest. (?) (juv.). Sp. me. lungs . . . .
 +
 +
Li. sj)., me
 +
 +
Cold, lungs
 +
 +
(Juv.)
 +
 +
 +
 +
0.000
 +
 +
0.000
 +
 +
014
 +
 +
0.017
 +
 +
0.000
 +
 +
0.000
 +
 +
0.037
 +
 +
0.035
 +
 +
0.003
 +
 +
0.003
 +
 +
0.027
 +
 +
0.020
 +
 +
0.010(?)
 +
 +
0.013(?)
 +
 +
0.015
 +
 +
0.015
 +
 +
0.028
 +
 +
0.025
 +
 +
007
 +
 +
0.007
 +
 +
0.050
 +
 +
0.050
 +
 +
0.052
 +
 +
0.050
 +
 +
0.002?
 +
 +
0.003+
 +
 +
0.022
 +
 +
0.021
 +
 +
0.025
 +
 +
0.025
 +
 +
0.002
 +
 +
0.002
 +
 +
0.048
 +
 +
0.042
 +
 +
0.031
 +
 +
0.035
 +
 +
1.015
 +
 +
0.888
 +
 +
0.002(?)
 +
 +
0.002(?)
 +
 +
 +
 +
+21.4
 +
 +
 +
 +
= 0.0 -35.0 +30,0? = 0.0 -12.0 = 0.0 = 0.0
 +
 +
- 4.0 +50.0
 +
 +
- 4.7 = 0.0 = 0.0 -14.3 + 12.9 -14 3 = 0.0
 +
 +
 +
 +
4.2 xO.9 5.2x 1.2 6.3x2.1 8.6x2.5 3.8x 1.5 5.7x 1.2 8.7x2.7 8.4x2.8
 +
 +
 +
 +
5.0x2.1 5.6x2.0 9.7x2.9
 +
 +
10.4x2.7 9.0x3.7
 +
 +
11.4x2.9 5.7 5.9
 +
 +
 +
 +
7.6
 +
 +
4.4
 +
 +
3.9
 +