Book - Text-Book of the Embryology of Man and Mammals

From Embryology
Text-Book of the Embryology of Man and Mammals

by Dr Oscar Hertwig (1892)

Professor extraordinarius of Anatomy and Comparative Anatomy, Director of the II. Anatomical Institute of the University of Berlin




Hersey Professor of Anatomy in Harvard University

with 339 figures in the text and 2 lithographic plates


  • Introduction
  • manuals and text-books
  1. Description of the Sexual Products
    1. the egg-cell
    2. the seminal filaments
  2. The Phenomena of the Maturation of the Egg and the Process of Fertilisation
    1. the phenomena of maturation
    2. the process of fertilisation
  3. The Process of Cleavage
  4. General Discussion of the Principles of Development
  5. The Development of the Two Primary Germ-Layers (Gastraea-Theory)
  6. The Development of the Two Middle Germ-Layers (Coelom-Theory)
  7. History of the Germ-Layer Theory
  8. Development of the primitive segments
  9. Development of connective substance and blood (the parablast- and mesenchyme-theories)
  10. Establishment of the external form of the body
  11. The foetal membranes of reptiles and birds
  12. The foetal membranes of mammals
  13. The foetal membranes of man
  14. The organs of the inner germ-layer. the alimentary tube with its appended organs
    1. the formation of the mouth, the throat-, gill-, or visceral clefts, and the anus
    2. the differentiation of the alimentary tube into separate regions, and formation of the mesenteries
    3. the development of the separate organs of the alimentary tube
      1. the organs of the oral cavity : tongue, salivary glands, and teeth
      2. the organs arising from the pharynx (1) the thymus (2) thyroid gland (3) lungs and larynx
      3. the glands of the small intestine (1) the liver (2) pancreas
  15. The organs of the middle germ-layer
    1. the development of the voluntary musculature
      1. the primitive segments of the trunk
      2. head-segments
    2. the development of the urinary and sexual organs
      1. the pronephros and the mesonephric duct
      2. mesonephros (wolffian body)
      3. metanephros (kidney)
      4. Mullerian duct
      5. germinal epithelium
      6. ovary
      7. testis
      8. metamorphosis of the different fundaments of the urogenital system into their adult condition
        1. in the male (descemus testiculorum)
        2. female (descemus ovariorum)
      9. the development of the external sexual parts
    3. the development of the suprarenal bodies
  16. the organs of the outer germ-layer
    1. the development of the nervous system
      1. the development of the central nervous system
        1. the development of the spinal cord
        2. the development of the brain (1) metamorphosis of the fifth brain- vesicle (2) fourth (3) third (4) second development of the pineal gland (epiphysis cerebri)., hypophysis (pituitary body) (5) fore-brain vesicle
      2. the development of the peripheral nervous system
        1. the development of the spinal ganglia
        2. the development of the peripheral nerves
        3. the development of the sympathetic system
    2. the development of the sensory organs
      1. the development of the eye
        1. the development of the lens
        2. vitreous body
        3. secondary optic cup and the coats of the eye
        4. optic nerve
        5. accessory apparatus of the eye
      2. the development of the organ of hearing
        1. the development of the otocyst into the labyrinth
        2. membranous ear-capsule into the bony labyrinth and the perilymphatic spaces
        3. middle and external ear
      3. the development of the organ of smell
    3. the development op the skin and its accessory organs
      1. the skin
      2. hair
      3. nails
      4. glands of the skin
  17. the organs of the intermediate layer or mesenchyme
    1. the development of the blood-vessel system
      1. the first developmental conditions of the vascular system
        1. of the heart
        2. vitelline circulation, allantoic and placental circulation
      2. the further development of the vascular system up to the mature condition
        1. the metamorphosis of the tubular heart into a heart with chambers
        2. the development of the pericardia! sac and the diaphragm
        3. metamorphoses of the arterial system
        4. venous
    2. the development of the skeleton
      1. the development of the axial skeleton
        1. the development of the vertebral column
        2. head -skeleton i. bones of the cranial capsule ii. ., visceral skeleton
        3. concerning the relation of the head-skeleton to the trunk-skeleton
      2. the development of the skeleton of the extremities
        1. pectoral and pelvic girdles
        2. skeleton of the free extremity
        3. development of the joints

appendix to literature

Translator's Preface

THE rapidly increasing recognition of the importance of Embryology in all morphological studies makes it desirable that the most valuable text-books upon the subject, in whatever language, be made available for those who are beginning its study. Although the English-reading student already has at command a number of text-books upon this subject, it is evident to any one familiar with HERTWIG'S Lehrbuch d?r Entwicklunysgeschichte des Me?ischen und der Wirbelthiere that thi> work covers the field of Vertebrate Embryology in a more complete and satisfactory way than any book heretofore published in English.

Two important objects to be accomplished in a text-book are : first, a clear and methodical exposition of the well-established facts of the science; and, secondly, such a presentation of unsettled questions as shall stimulate the reader to further inquiry and research. I believe it is far too common for the second of these aims to be overlooked. The present work fulfils both requirements in an eminent degree, and in its historical surveys exhibits an exceptional fairness of treatment, notwithstanding the author has been one of the foremost contestants in several of the fields reviewed. The summaries which follow the discussions of the several topics serve a useful purpose in directing attention to the more important conclusions drawn from each subject.

I have aimed to give a clear and accurate reproduction of the author's ideas ; while I have endeavored not always successfully to avoid awkward renderings and German idioms, I have preferred to err on the side of a too literal rather than a too liberal translation. There are a few points that demand a brief explanation. The German word Anlacje has heretofore been variously rendered into English by rudiment, origin, beginning, basis, foundation, etc., while some writers, recognising the inadequacy of any of these words to express the idea, have incorporated the German word itself in their English.

The Anlaye of a structure is its beginning or its undifferentiated state the object in a simple condition which is destined to be followed by a more complicated one. The use of rudiment in this sense is undesirable, because, in the interest of scientific accuracy, it is important to restrict its meaning, as in German, to a structure which is not destined to become more complicated, but which may have been, either ontogenetically or phylogenetically, even more highly developed than it now is. Origin and beginning are abstract terms, whereas A nlage is more frequently used in the concrete; basis and foundation (Grundlage) convey a wrong impression that of the substratum upon which the structure is erected. The need of a new word, which shall be used in the sense of Anlage, is evident. I suggest the adoption of an already existing word, -fundament, used at present only in a sense with which the proposed usage will not produce confusion. This word has been uniformly employed in the present translation, and the reader will see how readily and naturally it lends itself to this use. Fundament would thus bear the same relation to foundation that Anlage does to Grundlage.

I have also departed from authorised usage by sometimes employing for Bindeyeivebe and Stutzgewebe the term sustentative (in a mechanical sense) tissue, instead of connective tissue. My reason for this is the narrower meaning of connective as compared with sustentative.

In deference to a custom still followed in Human Anatomy, the author, in describing the relative positions of parts, has very generally used anterior and posterior for dorsal and ventral, etc. Instead of converting these expressions into terms which are independent of the temporary position of the organism, as I should have preferred, it has seemed better to indicate the direction by a bracketed word in those cases where a misunderstanding was most likely to occur. It has of course not been necessary to repeat this after each term of direction, but only after the first one of a series, the reader's attention being thus sufficiently directed to the matter to prevent any misconception.

The rapid advances in Embryology make it impossible for a book two years old to be a faithful reflection of the science of to-day in all its branches ; there are some topics in which even radical changes must be recognised. I have thought best, however, to reproduce the book as it left the hands of its author, and to content myself with calling the reader's attention to some of the topics in which the most important advances have been made, such as the metamerism of the head, and the plan and metamorphoses of the vessels of the visceral arches.

I am under very great obligations to my colleague, Dr. C. B. Davenport, for kind assistance and valuable criticism, but for which many defects of the translation would have been overlooked. I am also indebted to Drs. T. G. Lee, H. B. Ward, and W. McM. Woodworth for aid in reading portions of the proof.



Authors Preface to the First Edition

" Die Entwickelungsgeschichte 1st cler wahre Lichttrager fur Untersuchungen liber organische Kb'rper." C. E. v. BAER, "Ueber Entwickelungsgeschichte der Thiere " (Bd. L, S. 231).

THE Embryology of Animals, although one of the youngest shoots of morphological research, has, nevertheless, grown up in the course of sixty years, along with the cell-doctrine and that of the tissues, to a vigorous and stately tree. The comprehension of the structure of organisms has been extended in a high degree by numerous developmental investigations. The study of the human body has also derived great advantage from the same. In the newer anatomical textbooks (GrEGENBAUR, ScHWALBE) Embryology is receiving more and more attention in the description of the separate systems of organs. To what extent many things may be more clearly and attractively described in this manner is best shown by a comparison of the cles criptions of brain, eye, heart, etc., in the older and the more recent anatomical text-books.

Although it is generally recognised that Embryology constitutes " a foundation-stone of our comprehension of organic forms," nevertheless the attention which its importance warrants is not yet given to it ; it is especially true that it has not become as extensively as it should be a component of well-rounded medical and natural-history instruction, to which it is indispensable. The cause of this is perhaps in part to be sought in the fact that in student-circles the study of Embryology is often held to be especially difficult and a comprehension of it to be laborious. And thus many do not venture into this apparently obscure realm.

But ought the development of an organism to be really more difficult to comprehend than the complicated finished structure ? To a certain extent this was the case at a time when the most divergent and contradictory opinions prevailed concerning many of the most important processes of development, such as the formation of the germ-layers, the proto vertebrae, etc., which the lecturer had to take into account, and when many processes were not yet understood in their essence and their significance. But, thanks to the results of Comparative Embryology, the number of the unintelligible processes has been every year diminished, and in the same ratio the study of Embryology even for the beginner has been rendered easier.

At least, it is not in any way an essential feature of the process of development that it should be more difficult to understand than the structure of the completed form. For every development begins with a very simple condition, from which the more complicated is gradually derived and by which it is explained.

Inasmuch as I have for twelve years pursued the study of Embryology with especial interest, both in annually recurring academic lectures and in a series of scientific investigations, the desire has been awakened in me to acquire for Embryology a broader and more secure foundation in education, and to procure for it admission into larger circles of medical men and well-educated naturalists. As the result of this there has come into existence the book which is before us, in which the especial problem has been to make the complicated structure of the human body more intelligible through the knowledge of its development.

For the solution of this problem I have in the present text-book placed the comparative method of investigation in the foreground. I do not thereby find myself in any way in opposition to another direction of embryological research, which places the objective point in the physiological or mechanical explanation of the form of the animal body. Such a direction I hold to be fully warranted, and I believe that, instead of being opposed to a comparative-morphological direction, it can be of the most permanent value to it in the solution of its problems. One will find that I have here given full attention to the mechanico-physiological explanation of forms. Compare the sections on cell-division and Chapter IV., " General Discussion of the Principles of Development," in which the laws of unlike growth and the processes of the formation of folds and evaginations are treated.

In the presentation of the separate processes of development, in the main the important things only have been selected, the subsidiary left out, in order thus to make the introduction into embryological study easier. In the case of fundamental theories I have gone into their history extensively, because it is of great interest, and under certain circumstances operates as a stimulus, for one to see in what way the state of a scientific question for the time being has been attained. In pending controversial questions.

I have, it is true, employed chiefly as the foundation of my presentation the views which appear to me the most entitled to acceptance, but have not left immentioned opposing conceptions.

Numerous figures in the text, as well as some colored plates, will contribute materially to the easier comprehension of the various developmental processes.

I submit, then, this text-book to physicians and to students of medicine and the natural sciences, with the desire that it may promote and facilitate the study of Embryology in wider circles, and that it may thereby contribute to a deeper insight into the structure of our own bodies.

OSCAK HERTWIG. JENA, October 1886.

Authors Preface to the Second Edition

THE friendly reception which the " Text-book of the Embryology of Man and Mammals" has found, is an indication of the increased interest which this branch of Morphology now meets with.

Even more than a year ago, after the first part of the text-book appeared and while the second part was in the press, the necessity of preparing a second edition became evident.

In this edition fundamental changes have not been undertaken ; the text has, however, undergone an expansion in some places, owing to the attention given to several works which have recently appeared. This has been the case with the section on the first developmental processes of the egg (WEISMANX, BLOCHMANN) ; that on the origin of the vascular s}^stem (RABL, RUCKERT) ; that on the development of the foetal membranes (DuvAL, OSBORX) ; and that on the human placenta (KASTSCHEXKO, WALDEYER, HUGE).

As the second part of the text-book has just appeared, it has been possible to incorporate it in the second edition without alteration.

It has, furthermore, seemed to me expedient in the second edition to distribute at the ends of the several chapters the synopses of the literature, which in the first edition were brought together at the close of the whole work. Finally, there has been added an index of subjects, by which a more rapid orientation concerning the separate topics will be facilitated ; this will increase the usefulness of the work.

May the book in this form make for itself new friends, not only among students of medicine and the natural sciences, but also with all those who have a fondness for and a comprehension of studies in natural science.

OSCAR HERTWIG. JEXA, February 1888.

Authors Preface to the Third Edition

IN the two years which have elapsed since the appearance of the second edition of this text-book, our knowledge of the embryology of Vertebrates has experienced many important enrichments, thanks to the numerous investigations which are annually published. Therefore, as the problem of preparing a third edition of the text-book confronted me, I was compelled to make extensive changes in many places. Thus the second and third chapters, concerning the processes of fertilisation and cleavage of the egg, have undergone expansion, owing to the presentation of the important discoveries which have been made on the the egg of Ascaris megalocephala. I have given an entirely new wording to the ninth chapter on the development of connective substance and blood, also to the sections on the origin of the urinary organs and the development of the peripheral nervous system, and, finally, to the account of the development of the heart and the venous system. Also at other places one will often recognise the hand of improvement.

The third edition has been essentially improved by the addition of thirty new figures, which I have taken from the investigations of VAN BENEDEN, BOVERI, DUVAL, FLEMMING, HERMANN, His, BORN, GEGENBAUR, NAGEL, VAN WIJHE, GRAF SPEE, BONNET, and IVETBEL. Through the friendliness of Professor VAN BENEDEN I was also put in a position to employ for my text-book three figures out of his hitherto unpublished extensive work on the development of the germinal layers of the Rabbit. By means of the increase in the number of figures I hope that I have been able to render still easier the comprehension of many of the processes of development.

And so I close the preface to the third edition by expressing my thanks to all those who have rendered me friendly aid, and especially to the publisher, who in the further equipment of the text-book has met my wishes with the greatest willingness.



THE history of the development of the individual, or Ontogeny (Embryology), is the science of the growth of an organism ; it describes the morphological changes which an organism passes through from its origin in the ovum up to its complete maturity, and presents these in their natural connection. We can regard the fertilisation of the egg-cell as the beginning of the process of development for Vertebrates, as it also is for all the rest of the higher animals.

In giving an account of the changes of the egg-cell, which begin with fertilisation, one may choose between two different methods.

According to one method a particular organism is made the basis .of the account, and one describes the changes which its germ undergoes from the moment of fertilisation onward, from hour to hour, and from day to day. It is in this way that the embryology of the Chick has been worked out by C. E. VON BAER in his classical paper, and by FOSTER AND BALFOUR in their " Elements of Embryology." This method has the advantage that the reader acquires a view of the total condition of an organism in the separate stages of its development.

A book of that kind is especially suitable for such persons as desire to acquaint themselves, by their own observation, with the embryology of a single animal, as, for example, the Chick, by repeating the investigations of others. It is, on the contrary, less adapted to those who wish to acquire a connected view of the development of the separate organs, as the eye, the heart, the brain, etc. For the formation of these will of course be treated of at different places in describing younger and older embryos. In order to procure a general survey of the course of development of an organ, the reader must consult various places in the text-book, and collect for himself w r hat relates to the subject.

For beginners, and for the needs of theoretical instruction in Embryology, the second method commends itself, in which the separate organs are considered in succession, each for itself, and the changes which a single organ has to pass through during development are set forth connectedly from beginning to end. It is in this way that KOLLIKER'S " Embryology of Man and the Higher Animals" is written. The second method is, moreover, the only one applicable when the problem is to investigate in a comparative way the development of several organisms, and to fill up the gaps which exist in our knowledge of one by that which we know concerning nearly related animals. But it is precisely in this position that we find ourselves, when we wish to acquire a survey of the development of the human body. An account which should limit itself to that which we know about Man would exhibit numerous and extensive gaps. For up to the present the eye of man has not seen how the human ovum is fertilised, how it divides, how the germ-layers are formed, or how the establishment of the most important organs is effected. It is especially the period of the first three weeks, during which the greatest variety of fundamental processes of development take place, concerning which we know next to nothing ; there is also little prospect that a change will soon occur in this regard. The time will therefore perhaps never come when a complete embryology of Man in the strict sense of the word will be possible.

However, the existing gaps can be filled out in another manner, and one which is entirely satisfactory. The study of the most widely differing Vertebrates teaches us that they are developed according to a common plan, that the first processes of development agree in all really important points, and that the differences which we encounter here and there are produced by causes of a subordinate kind, as, e.g., by the egg's possessing a greater or less amount of yolk.

When we see that the establishment of the central nervous system, of the eye, of the spinal column, of the viscera, etc., takes place in Mammals on the whole just as it does in Amphibia, Birds, and Reptiles, the conclusion is near at hand, and justified, that Man also in his development is no exception to this general phenomenon. Thus in the study of Embryology we are naturally led to the comparative method. What, owing to the nature of the difficulties, we cannot learn directly about the development of Man, we seek to deduce by the investigation of other Vertebrates.

In earlier decennia the Hen's egg was the favorite object, and it is upon this that we possess the most numerous and most complete series of observations. During the last twenty years research has also been directed to Mammals, in the investigation of which the greatest difficulties have to be surmounted, as well as to Reptiles, Amphibia, Fishes, etc. Only through the observation of such various objects has insight been acquired into many processes, which in their essence remained unintelligible to us from the study of the Chick alone. For it was thus that one first learned to distinguish the important from the accessory and unimportant, and to understand the laws of development in their generality.

In this text-book, therefore, I shall not confine myself to a single object, such as the egg of the Hen or the Rabbit, but from more general comparative standpoints shall endeavour to present what, through extensive series of investigations, we have thus far recognised as the rule in regard to the real nature of the processes of fertilisation and cleavage, the formation of the germ layers, etc.

However, let no one expect a text-book of comparative Embryology. The purpose and the problem is first of all to learn to comprehend the development and the structure of the human body. What we know about that has been placed before everything else, and the embryology of the remaining Vertebrates has been cited, and, as it were, fully utilised, only in so far as was necessary for the purpose indicated.

In the division of the embryological material proposed by us, according to the separate systems of organs, there is a long series of processes, with which the development begins, which do not permit of an arrangement, because at the beginning the fundaments of definite, afterwards differentiated organs, are not recognisable in the germ. Before there is any formation of organs, the egg is divided into numerous cells, and these then arrange themselves into a few larger complexes, which have been called the germ-layers, or the primitive organs of the embryo. Further, in the higher Vertebrates there are formed certain organs, which are useful only during embryonic life, and are subsequently lost namely, the foetal membranes and foetal appendages. All of the processes of that nature we shall treat of connectedly, and by themselves. In accordance with this, we can divide our theme into two main sections, the first of which will deal with the initial processes of development and the embryonic membranes, the second with the origin of the separate systems of organs. In order to facilitate for the advanced a more thorough study, and a penetration into embryological literature, a survey of the more important original works is given at the close of the separate chapters. On the other hand, text-books of Embryology may be mentioned in this place. [Compare also the larger monographic works cited at the end of the book.]

Manuals and Text-Books

Valentin, G-. Handbuch der Entwicklnngsgeschichte cles Menschen mit vergleichender EUcksicht der Entwicklung der Siiugethiere und Vogel.

Berlin 1845. Bischoff. EntwicklungFgeschichte der Saugethiere uncl cles Menschen.

Leipzig 1842.

Rathke, H. Entwicklungsgeschichte der Wirbelthiere. Leipzig 1861. Kolliker, A. Entwicklungsgeschichte des Menschen und der hoheren Thiere.

Academische Vortrage. Leipzig 1861. 2. ganz umgearbeitete Auflage.

Leipzig 1879. Kolliker, A. Grundriss der Entwicklungsgeschichte des Menschen und der hoheren Thiere. 2. Auflage. Leipzig 1884. Schenk. Lehrbuch der vergleichenden Embryologie der Wirbelthiere. Wien 1874. Haeckel, E. Anthropogenie oder Entwicklungsgeschichte des Menschen.

Leipzig 1874. Dritte Auflage. 1877. Foster, M., and F. M. Balfour. The Elements of Embryology. Part 1.

(Chick.) London 1874. 2nd edit, by Adam Sedgwick and Walter Heape 1883. German translation by Kleinenberg. Leipzig 1876. His, W. Unsere Korperform und das physiologische Problem ihrer Ent stehung. Leipzig 1875. Balfour, F. M. A Treatise on Comparative Embryology. London 1880, -81, 2 vols. German translation by Dr. C. Vetter. Jena 1881. Romiti, G-. Lezioni di embriogenia umana e comparata dei vertebrati. Siena 1881, -82, -88.

Preyer, W. Specielle Physiologic des Embryo. 1883, -84. Hoffmann, C. K. Grondtrekken der vergelijkende Ontwikkelingsgeschie denis van de gewervelde Dieren. Leiden 1884. Duval, M. Atlas d'Ernbryologie. Paris 1888.