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Written by

Charles R, Bardeen, Madison, Wis.; Herbert M. Evans, Baltimore, Md.; Walter Felix, Zurich; Otto Grosser, Prague; Franz Keibel, Freiburg i. Br.; Frederic T. Lewis, Boston, Mass.; Warren H. Lewis, Baltimore, Md.; J. Playfair McMurrich, Toronto; Franklin P. Mall, Baltimore, Md. ; Charles S. Minot, Boston, Mass. ; Felix Pinkus, Berlin ; Florence R. Sabin, Baltimore, Md ; George L. Streeter, Ann Arbor, Mich.; Julius Tandler, Vienna; Emil Zuckerkandl, Vienna.



With 423 Illustrations




Printed by J. B. Lippincott Company The Washington Square Press, Philadelphia, U. S. A.
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The circumstances that led to the publication of this Manual of Human Embryology, of which the present is the first volume, have been recorded in the Introductory Chapter, where the aims of the work are also set forth. A number of German and American embryologists have collaborated in its production, and the work appears simultaneously in Germany and America. The translation of the chapters originally written in German has been made by Professor McMurrich, and the publishers desire to express their thanks to him for his careful work. The English chapters have been translated for the German edition by one of the editors, Professor Keibel. Valuable assistance has been rendered in the correction of the proofs by Dr. C. Elze, and to him also the thanks of the publishers are due. The editors desire to express their indebtedness to the publishers, Messrs. S. Hirzel, of Leipzig, and J. B. Lippincott Company, of Philadelphia, for their generosity in making it possible for the collaborators to enrich the text with numerous excellent illustrations and for the aid they have rendered in bringing the work to a successful completion. The second volume will appear at an early date.

Franz Keibel, Franklin P. Mall.

   Manual of Human Embryology I 1910: The Germ Cells | Fertilization | Segmentation | First Primitive Segment | Gastrulation | External Form | Placenta | Human Embryo and Fetus Age | Ovum Pathology | Integument | Skeleton and Connective Tissues | Muscular System | Coelom and Diaphragm | Figures | Manual of Human Embryology 1 | Manual of Human Embryology 2 | Franz Keibel | Franklin Mall | Embryology History

   Manual of Human Embryology II 1912: Nervous System | Chromaffin Organs and Suprarenal Bodies | Sense-Organs | Digestive Tract and Respiration | Vascular System | Urinogenital Organs | Figures 2 | Manual of Human Embryology 1 | Figures 1 | Manual of Human Embryology 2 | Figures 2 | Franz Keibel | Franklin Mall | Embryology History

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By FRANZ KEIBEL, Freiburg i. Br.

Vesal has rightly been regarded as the founder of human anatomy. He emancipated anatomy from the dogmas of Galenism and showed that Galen's anatomical observations had really been made upon apes and that he had attributed to the human body the structure observed in these forms. Vesal studied the human body and based his immortal work upon that study. Embryology , was, however, only incidentally considered by him, and in his studies in this field he was false to his principles in that, like Galen in anatomy, he attributed to man what he observed in the embryos of animals.[1]

However, the path which Vesal opened in anatomy was also followed by his suCoessors in embryology. In this connection Gabriele Fallopia (1523-1562) deserves mention as the first who gave a correct description of the placenta and of the chorion and its vessels, and also, on the basis of his own observations, denied the oCourrence of an allantois in man. Bartholomeo Eustachi (d. 1574) studied the development of the teeth in human embryos, and Julio CaBsare Aranzi (1530-1588) expressly noted that differences exist between the early stages of development of man and those of the lower animals. Vesal's suCoessor and pupil, Matteo Realdo Colombo (d. 1559), endeavored to do for himian embryology what Vesal had done for human anatomy, promising a consideration of human embryology and not animal embryology, since Nature had formed man and animals upon different plans. His efforts, however, to establish an embryology on the basis of observations on human material must, as Bloch has pointed out, be regarded as a failure. And the reasons why Colombo was able to record only exceedingly incomplete observations are not far to seek. The number of human embryos aCoessible to a single investigator in his day must have been very small ; in addition they were, in part, exceedingly altered pathologically, and, what was still more important, the phenomena of greatest significance for embryology oCour at such an early period of development that they could not possibly be observed with the methods of investigation available at that time. Even in the case of animals these phenomena were still altogether obscure, and almost three hundred years were to elapse before the necessary preliminary work in this field was aCoomplished. Ulisse Aldrovandi (1526-1605) must first lay a foundation for modern embryology. He was the first to trace systematically the development of the chick up to the time of hatching and to give a consecutive aCoount of the development. After Aldrovandi followed his pupil Volcher Koyter (1534-1600), and then Hieronymus Fabricius ab Aquapendente (1537-1619). Harvey and Malpighi need be only mentioned here, but a few words may be devoted to the less known Adrian van den Spieghel (Spigelius) (1578-1625).

Spieghel, as may be seen from the introduction to his work De formate fetu, published the year after his death (1626), had a clear appreciation of the value of embryology to the physician, and while his predecessors studied especially the fetal aCoessory organs, he turned his attention mainly to the development of the fetus itself and that of its organs. Since his object was a history of the development of the human embryo, he was met by the diflSculties mentioned above, but where these diflSculties were less pronounced, as in the history of the development of the bones, he made valuable contributions and materially advanced the investigations of Koyter and Fallopia in this field. In this connection he made the first attempt at a description based on observation of the genesis of a tissue, distinguishing those osseous elements which were formed in preformed cartilage from those which arose in membrane and describing the growth of membrane bones by the apposition of osseous spicules at the margins. What Spieghel had to say concerning the development of the other organs of the body does not compare favorably with his description of the development of the bones, and he also devoted more than half the contents of his book to the consideration of the chorion, the placenta, and the umbilical vessels ; he recognized the occurrence of an allantois in man. And Spieghel also rendered important service by his observations on the fetal circulation. In his time the teaching of Galen that the direction of the blood-stream was the same in both the umbilical arteries and veins was generally aCoepted, and with this false belief it was naturally impossible to obtain a correct xmderstanding of the fetal circulation. Spieghel, for a number of good reasons, concluded that the direction of the flow in the umbilical arteries was centrifugal ; he believed, however, that the vessels carried not blood but Galenas spiritus vitalis, and this prevented him from obtaining a clear idea of the true relations and an aCourate knowledge of the circulation of the blood. The difficulties which stood in the way of this discovery were exceedingly great, and the service rendered by Harvey, as Bloch points out (l. c, p. 333,,^ note), can be rightly appreciated only when, from a study of the literature, an idea is obtained of how confused were the notions of even the most distinguished physiologists of the time concerning the movements of the blood.

Without entering into details the names may be mentioned here of (lualtherus Needham, Nicolaus Hoboken, Nicolaus Steno, and Thomas Wharton, who rendered services in connection with the anatomy and physiology of the egg-membranes. The discoveries of Regner de Graaf, Swammerdam, Hamm, and Leeuwenhoek are generally known and have many times been recounted ; and the much-admired *'Icones Ossiuui Foetus Humani" of Albinus gave a certain amount of completeness to the knowledge of the development of the human skeleton. Albrecht von Haller and Kaspar Friedrich Wolff are worthy of mention for more than the foimulation of the theories of evolution and epigenesis. Haller wrote important works on the development of the osseous system and of the heart, and Kaspar Friedrich Wolff was the founder of the theory of the germ layers, which was further developed by Dollinger's pupil. Christian Pander, and especially by Karl Ernst von Baer and Remak. Karl Ernst von Baer finally discovered the long-sought-for ovum of man and of the mammalia, and is the real founder of comparative embryology. His work, "Ueber Entwicklungsgeschichte der Tiere, Beobachtung und Reflexion" (Konigsberg, 1828 and 1837), must be considered, aCoording to Kolliker ("Entwicklungsgeschichte," 1879, p. 14), the best that embryological literature of all times and all peoples has to show.

The origin of the germ layers and the origin of the organs from these were now subjected to most careful investigation on all sides and in all classes of animals. Here Remakes work, which exercised a very deep and lasting influence, need only be mentioned. The progress in human embryology did not, however, at first keep pace with that in comparative embryology. William Hunter's "Anatomia Uteri Gravidi" (Birmingham, 1774) gave splendid representations of the egg-membranes and of the gravid uterus, but did not advance our actual knowledge of embryology, the condition of which at the close of the eighteenth century can be well understood from the book by D. Ferdinand Georg Danz ("Grundriss der Zergliederungskimde des ungebomen Kindes", vol. i, EVankfurt and Leipzig, 1792; vol. ii, Giessen, 1793), which was accompanied by annotations by Sommerring, the first authority of that period. Sommerring himself published in 1799 his "Icones Embryonum Humanomm", which does not, however, contain satisfactory representations of even the later stages of human development, although it is interesting as citing all important earlier observations. How small was the amount of useful material of the early stages of human development accessible even to Karl E. von Baer can readily be appreciated if one examines the concluding part of his great work, edited by Stieda in 1888.

In addition the methods of satisfactory fixation, of sectioning, to say nothing of serial sections and of reconstruction, were still lacking. Consequently, the most important early stages in the development of man remained unknown, and progress even in the case of animals, of which abundant material was available and could be studied under the lens while still fresh and transparent, was slow and difficult, even when the method of fixing the embryos in alcohol and dissecting them under the lens with fine needles and knives was learned. Indeed, it is remarkable what was accomplished in spite of these arduous and uncertain methods, but results could be obtained only by oft-repeated observations, and, in the case of man, the material for these was wanting. Thus, we read in Friedrich Tiedemann's "Anatomie und Bildungsgeschichte des Gehirns im Fetus des Menschen'^ (Niimberg, 1816) that in embryos of the first month the place of the spinal cord and brain is oCoupied by a clear fluid. His observations begin with an embryo which had been preserved for some time in alcohol and measured seven lines in length (from the figure it is to be concluded that Ehenish lines of 2.18 mm. are meant; the Parisian line corresponds to 2.25 mm.), that is to say, about 15 mm. Johannes Miiller in his Bildungsgeschichte der Genitalien (Diisseldorf, 1830) begins his observations on human embryos with an embiyo of 8 lines[2] in length (measurement of the figure gives a length of 20 mm.). What the younger stages were like could only be conjectured from observations on animals, and so matters practically remained until His published his "Anatomie menschlicher Embryonen."

It is true that human embryos and the egg membranes were extensively investigated and to a certain extent very well figured at this time — as in Coste's Embryogenie (Paris, 1837), in the works of Pockel (Isis, 1825), Seller ("Die (Jebarmutter und das Ei des Menschen," Dresden, 1831), Breschet ("Etudes anatomiques sur PcBuf humain," Paris, 1832), Th. L. W. Bischoff ("Beitrage zur Lehre von den Eihiillen des menschlichen Fetus," Bonn, 1831), and in the briefer publications of E. H. Weber, Joh. Miiller, R. Wagner, Von Baer, Wharton Jones, Allen Thomson, and Eschricht — but the observations remained isolated, they were not suflScient to round out the whole story, and frequently the results failed to attain completeness owing to a disinclination to destroy a valuable specimen. The technic of investigation made little progress, and it is in this particular that His made a change. He had prepared himself by a study of the embryology of the chick and in connection with this study had worked out a successful technic; he fixed his embryos, he had constructed an apparatus for section cutting which, primitive as it now seems, led the way to our modem microtomes, and he had thought out a method of reconstruction which has been gradually improved, especially by Born, into a method that is now absolutely indispensable in embryologicaL investigation. With this equipment His began the study of human embryology. Although he was fortunate in obtaining material, yet this was still scanty, and he was far from imagining that with it he could study the complete development of man ; but with the help of the method of reconstruction he first thoroughly worked out the anatomv of the individual embryos so far as his technic permitted, and the results so obtained formed a sure foundation for human embryology. His models, as well as those he had constructed from the chick, were reproduced by Dr. Ziegler in Freiburg, and, since these came to be regarded as indispensable by every anatomical institute, they served, more than anything else, to increase the knowledge and interest in human embryology. Gradually the gaps which existed were filled in by His's own continued observations and by those of others who studied their material by similar methods, and in this connection there may be mentioned the work of Fol, of Phisalix, of Eternod, and, especially, of Graf Spee. Monographs of individual embryos were contributed by Piper, Mall, Frederic T. Lewis, Susanna Phelps Gage, and Bremer, and in connection with the "Normentaf el zur Entwicklungsgeschichte des Menschen" of Keibel and Elze, to be considered later, by Thompson, Ingalls, Elze, and Low. In the meantime the investigation of the individual organ-systems was not standing still. It was very natural that the foundation stones upon which certain systems of organs are built up should be brought together sooner than those of other systems, and, this happening, His directed his attention to these systems, although the development of the organism as a whole still remained the chief object of his thoughts. Thus were achieved the results recorded in the third part of his '^Anatomie menschlicher Embryonen,'of which those treating of the general differentiation of the digestive tract and those concerning the heart may be especially mentioned. He also published other larger works on the development of the nervous system, one of which appeared shortly before his death.

There should also be mentioned in this connection the work of A. von KoUiker (eye and olfactory organ) ; of Hammar (the branchial arch region) ; of Broman, Mall, and Swaen (coelom and diaphragm) ; of Hochstetter, Tandler, Erik Miiller, and de Vriese (vascular system; also Elze) ; of Bardeen (skeleton); of W. H. Lewis (musculature) ; of Nagel, Keibel, and Bayer (the urogenital system) ; of the younger His, Hammar, and Streeter (ear) ; and of the younger His and Romberg (the sympathetic system).

The above enumeration makes no pretence of completeness, but is merely intended to show how human embryology was approached from all sides and how the material for a full exposition of the subject was gradually accumulated.

As has been stated, the idea of working out a complete aCoount of the development of the human body was always before the mind of His, but as time went on the hope of aCoomplishing the task single-handed failed him; and so he suggested that I should collaborate with him in writing a text-book on human embryology. Unfortunately, the plan was never carried out; I have, however, gone further with it and prepared the way for its fulfilment by another work. In the pages of my "Normentaf el zur Entwicklungsgeschichte der Wirbeltiere" I have published, in conjunction with Curt Elze, a Normentafel for the development of man. In this I have had the assistance of a large number of other investigators, some of whom, such as Hammar and Tandler, have made personal contributions, while others have contributed valuable material, frequently material obtained by operation and most admirably preserved. Thus I have been able to follow the development of the body in an almost perfect series of embryos from about the twelfth day[3] up to the end of the second month and to obtain many data concerning the first appearance and degree of development of the organs. An explanation of the stages of development earlier than those that I examined had been furnished by the admirable investigations of Graf Spee and Hubert Peters,[4] already mentioned, and the time seemed to me to be propitious for giving an account of the development of the human body, based throughout on human material. We have already, it is true, a whole series of text-books on human embryology, some of them excellent ; but they are based, for the most part, on other than human material. They have been written from the comparative embryological stand-point, and, in the interests of continuity, an endeavor to conceal the gaps in our knowledge is frequently evident. This will not be the case in the present book. On the contrary, its endeavor will be to indicate clearly and precisely these gaps, for thus only can they be filled in; and frequently, as I know, the material for filling them in is already available. I do not propose, of course, to dispense with the assistance of comparative embryology and anatomy, but when a gap has been indicated the manner in which it is probably to be bridged will be pointed out, and, similarly, attention will be given to those facts of comparative embryology and anatomy which serve to render intelligible to us special processes of development in man. As a rule, however, such considerations will be printed in smaller type, in order that there may be a clear distinction between facts and deductions. So far as I can see, all the material necessary for a human embryology, with the exception of the earlier stages which concern the development of the germ layers and the first stages of placentation, is already available or at least is comparatively readily obtainable. It would be possible, therefore, to describe the development of all the organs exclusively from human material if it were possible to bring together all the material which is now available. If my colleagues and myself have not suCoeeded in this, nevertheless we have been able to approach the goal, and we may hope that it will fare better with us than with Eealdo Colombo, and even if the first attempt has not been perfectly suCoessful, the second, whether made by us or by others, will come so much nearer the goal. The material is in hand and to be obtained ; it will be brought forward if we only point out clearly what is lacking.

When working at my Normentafel I had very striking evidence of the conununity of purpose which to-day inspires our scientific world. A considerable number of investigators deprived themselves for long periods of time of valuable material which they themselves had not yet had opportunity to study thoroughly, in order that a larger undertaking might be completed. In the time of Eealdo Colombo such scientific co-operation did not exist; we can with pride regard this as a great step in advance. Without it the completion of such a work as the Normentafel would have been unthinkable; to-day the post and the telegraph, railways and steamboats, are at our service and may be turned to the service of science. And so it has been possible to bring to completion such a "Handbook of Human Embryology" as His had planned with me in Germany with the assistance of one of his most loyal pupils on the far side of the ocean, Franklin P. Mall, of the Johns Hopkins Medical School in Baltimore. It has seemed proper to Mall and myself to enlist the services of a considerable number of collaborators, the necessary similarity of treatment being secured by the common purpose and by editorial supervision ; in this way the book would the sooner be brought to completion and, what is more important, the various chapters would be written with a complete mastery of the subject. Considered purely objectively and scientifically the embryology of man has no more interest than that of any other mammal or vertebrate, and from this stand-point special endeavors to work out human embryology by itself may seem to be misplaced. Our interest in the development of the human body is justified, however, first, by the fact that we are human beings, and, secondly, because, as Spieghel long ago pointed out, it is of importance to physicians. That we do not undervalue the information to be obtained from comparative embryology and anatomy has already been stated.

  1. Those interested in the history of Embryology are referred to the paper of Bruno Bloch, Die geschichtlichen Grundlagen der Embryologie bis auf Harvey, Nova Acta Leop.-Carol. Akademie, No. 3, 1904, p. 295 et seq., a work written under the stimulus and influence of Rudolf Burckhardt.
  2. A wonderful human embryo, 3 J lines in length measured along its curvature, with a long-stalked umbilical vesicle and traces of branchial clefts," he could not sacrifice for investigation; he described it in Meckel's Archiv, 1830. He also refers to a similar embryo in Meckel's possession (Beitrage zur vergl. Anat., vol. i, pp. 71 and 72).
  3. According to the age estimate usually employed; if the estimates of Bryce and Teacher, to which I shall refer later, are correct, the above statement should be "from about the eighteenth day." (Compare Chapter VIII, The Age of Human Embryos and Fetuses.)
  4. Quite recently the observations of Spee and Peters have been confirmed and extended by Ph. Jung ("Beitrag zur friihesten Eieinbettung beim menschlichen Weibe," Berlin, 1908) ; Bryce, Teacher, and Kerr ("Contribution to the Study of the Early Development and Imbedding of the Human Ovum," Glasgow, 1908) ; and Frassi ("Ueber ein junges menschliches Ei in Situ," Arch. f. mikr. Anat., vol. Ixx, 1907, and Weitere Ergebnisse des Studiums eines jungen menschliches Eies in Situ, ibid,, vol. Ixxi, 1908).

   Manual of Human Embryology I 1910: The Germ Cells | Fertilization | Segmentation | First Primitive Segment | Gastrulation | External Form | Placenta | Human Embryo and Fetus Age | Ovum Pathology | Integument | Skeleton and Connective Tissues | Muscular System | Coelom and Diaphragm | Figures | Manual of Human Embryology 1 | Manual of Human Embryology 2 | Franz Keibel | Franklin Mall | Embryology History

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