Book - Vertebrate Embryology (1949)

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A personal message from Dr Mark Hill (May 2020)  
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I have decided to take early retirement in September 2020. During the many years online I have received wonderful feedback from many readers, researchers and students interested in human embryology. I especially thank my research collaborators and contributors to the site. The good news is Embryology will remain online and I will continue my association with UNSW Australia. I look forward to updating and including the many exciting new discoveries in Embryology!

McEwen RS. Vertebrate Embryology. (1949) IBH Publishing Co., New Delhi.
1949 McEwen: 1 Germ Cells and Amphioxus | 2 Frog | 3 Teleosts and Gymnophiona | 4 Chick | 5 Mammal | 1949 Vertebrate Embryology

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This 1949 textbook by McEwen describes embryonic development.

1923 Edition

Modern Notes:

Historic Disclaimer - information about historic embryology pages 
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Pages where the terms "Historic" (textbooks, papers, people, recommendations) appear on this site, and sections within pages where this disclaimer appears, indicate that the content and scientific understanding are specific to the time of publication. This means that while some scientific descriptions are still accurate, the terminology and interpretation of the developmental mechanisms reflect the understanding at the time of original publication and those of the preceding periods, these terms, interpretations and recommendations may not reflect our current scientific understanding.     (More? Embryology History | Historic Embryology Papers)

Vertebrate Embryology

Robert S. McEwen

Professor Emeritus of Zoology, Oberlin College


Part I The Germ Cells and Early Development of Amphioxus

  • 1. Introduction
  • 2. Fertilization and Early Stages in Development
  • The Early Development of Amphioxus

Part II The Development of the Frog

  • The Frog: from the Production of the Germ Cells through Gastrulation
  • The Frog: Early or Embryonic Development Subsequent to Gastrulation
  • The Frog: Later or Larval Development

Part III The Teleosts and Gymnophiona

  • The Teleosts and Gymnophiona: their Segmentation and Gastrulation

Part IV The Development of the Chick

  • The Chick: the Adult Reproductive Organs, and the Development of the Egg Previous to Gastrulation
  • Gastrulation and Development through the First Day of Incubation
  • The Chick: Development during the Second Day of Incubation
  • The Chick: Development during the Third Day of Incubation
  • The Chick: Development during the Fourth Day of Incubation
  • The Chick: Development during the Fifth and Subsequent Days

Part V The Development of the Mammal

  • The Early Development of the Mammal and its Embryonic Appenclages
  • Development of the Pig to the Ten Millimeter Stage
  • The Later Development of the Pig
  • The Skeleton, Teeth, Hair, Hoofs and Horns


Preface to the Fourth Edition

As in previous revisions, the fourth edition of this text does not purport to be a new book. It again frankly retains the fundamental plan and character of the older editions, in that it is primarily descriptive. but with enough experimental results interwoven with the descriptive material to stimulate interest, and to elucidate such principles of development as have been firmly established.

Though not radically altered, the older book has nevertheless been carefully gone over page by page, and, as before, changes have been made. wherever it was thought desirable in order to bring the subject matter up to date, to clarify statements, or to correct errors. In some cases. whole pages have been entirely rewritten, and in certain instances, as in the section on maturation of the germ cells, this has involved several successive pages. Mistakes in figures have also been corrected, and in a few cases. as in the diagram of frog gastrulation, the figure has been completely modified and, the writer believes, greatly improved.

Thanks are due to various colleagues who have made suggestions and pointed out errors. Especial gratitude is felt by the author to Dr. Roland Walker for his meticulous notations of errors both large and small, and for his constructive eiiorts to aid in their correction.

R. S. MCE. Oberlin College,

September, 1956.

Preface to the First Edition

This book is designed as an introductory text in Vertebrate Embryology, a work which seems to be justified on the following grounds: The older texts upon this subject, though in many cases excellent, do not cover exactly the field which is now covered in many colleges; these texts, moreover, are becoming somewhat out of date in various details. Among the newer books the best ones tend to do one of two things. Either, in the interest of thoroughness, they confine their attention entirely tn one form, e.g., the Chick, or else, for the sake of a broader viewpoint, they deal with a considerable number of animals, but in doing so touch only upon the earlier developmental stages of each. Now it is obvious that there is great value for the student, both in the accuracy gained by the careful intensive study of a single type, and also in the possession of less detailed knowledge of the history of other forms which are nearly related to it. Hence, what has seemed to be needed was a book which would, so far as is possible, make available both these advantages. To meet this need, the major part of the present text comprises a mo-,leratcl}‘ complete account of the development of two typical forms. i.e., the Frog and the Chick, each of which, in the writer’s opinion, has special features which justify such treatment. These relatively detailed discussions are then supplemented by chapters which present brief comparisons, not only with the Mammal, but also with certain other significant members of the Vertebrate group. Furthermore, the essentially embryological portion of the book is preceded by an optional introductory chapter dealing with the elements of cytology. Upon this basis the effort throughout the work has been to produce something adapted to the requirements of the general student of Zoology. us well as to the individual particularly interested in premedical preparation.

As i'crgzvx'tls certain details concerning the method of handling the topics involved, the following remains to be said. Because of the character of the book, the chapter upon cytology places special emphasis upon the structure, development, and function of the germ cells, with particular reference to nuclear phenomena and their genetic significance. The strictly cnibryological subject tn:-ttter is then introduced by a short general discussion of the more lundaixiierxtal and universal proc of Vertebrate development from the comparative standpoint. This includes a description of the various types of segmentation, gastrulation, and the formation of the rudiments of the nervous system and the main mesodermal structures. Following these introductory chapters,

‘Amphioxus is the first particular type to be considered lI(’('£lUSt‘. of the

relatively primitive character of most of its early history. The later development of this animal, i.e,, that following the fnrnizuion of the mt'.s'ndermal somites. is, however, quite highly distinguish it from the vast majority of Clionlates are without great significance for the general student. tliey are mniuml.

The Frog, as suggested above, is one of the two forms which have been treated at some length. The reasons for suvli extencled mnsirl<~ration in this instance and in that of the Chick are presunmbly olwious to every Zoiilogist. For the sake of the student. however. the uzlim uf these animals as subjects of enibryologitral study is lt\[llt‘txil,’il in tinparagraphs of the text which introduce them. ln the case ui lhv "I":-u;_». its early history has been presented under the head of c-ertuin fairly. well recognized stages which lend themselves well to corre-l;1tion with work in the laboratory. In further pursuance of this method the-. internal changes have been noted in alternation with those or-currin;__r cxtc-rnall_\ . This was done in order that the reader might obtain. so far as pm-s_<il»le. a correct idea of the really simultaneous character of tliese processes. It did not seem feasible, however, in a work of this St'(}pt.' to continue this plan throughout the entire course of development in this animal. The later external changes. therefore, are included under one lieading. while the more advanced details of organogeny are described in terms of particular systems.

Following the treatment of the Frog, there has been introduced a very brief account of segmentation and gastrulation in the Teleosts and the Gymnophiona. This has been done despite the realization that in the case of the latter group laboratory consideration will in most cziscs be impossible. The reason for this is the authors opinion that segnu-xi1;+ tion and gastrulation in these two classes of animals are extrem:-ly valuable in assisting the student to relate these processes in the Frog In those which he is about to study in the Bird. Experience, xnoremm‘, has seemed to indicate that the relation of avian and mammalian gztstrulzb tion to that in more primitive forms is always particularly clillicult for

i the beginner to grasp, and it is believed, therefore. that any legitinmte aid to this end is worth while.

In treating the early stages of the Chick a good deal of stress has been placed upon the method of segmentation and gastrulation. The latter especially has been emphasized because of its peculiar character, and the desirability of making clear its relationship to that in the forms already studied. The later history of this animal is then presented in daily periods, according to the well-known plan of Foster and Balfour. This has been done because it seems to the writer that at least in a beginning course, this method has certain marked advantages over that of stuclying the complete embryology of one system at a time. In the first place the Bird lends itself particularly well to treatment by periods, and secondly, the simultaneous development of all the systems is what is actually seen to occur in any animal. This latter fact it would seem well to impress upon the student when possible by the method of presentation. Finally it has appeared not only possible but easier to conduct the class work in correlation with the laboratory when development is studied by periods rather than by systems. It should be noted, nevertheless, that in this book the material has been so arranged that the student can readily follow through the complete growth of any one system if the instructor so desires.

As regards the Mamxnals, it is felt that the detailed differences between the organogeny of this group and that of the Birds are not, on the. whole, of great general biological significance. Of very considerable significance, however, are those unique characteristics of both mother and embryo connected with mammalian gestation. For this reason the discussion in this portion of the text is confined chiefly to the earlier developinental stages, which are treated largely from the comparative standpoint. The subject is introduced by a description of the structure and functions of the adult reproduetige organs in the,same manner as in the case of preceding forms. This involves the process of ovulation, and in that connection it has seemed worth while to describe briefly the peculiar cyclic phenomena which accompany this process in the mammalian female. Following this, the comparative idea is pursued with particular reference to the development of the extra-embryonic z1ppt’ll(l£lgC.‘.‘~. This is believed to be especially important from an evolutionary viewpoint because it shows how these appendages, already observed in the Chick. have been modified in the various Mammals. This discussion is naturally accompanied by a description of the structure and probable evolution of the placenta. For the general plan of treatmom of these latter topics the author frankly acknowledges his indebtedness to Professor Jenl<inson’s excellent book, Vertebrate Embryology.

Concerning bibliographical material, references to the more important literature of each subject are appended to the chapter which concludes consideration of the topic in question. As intimated, it will be quite obvious that these references make no pretense of being exhaustive. Their object is rather merely to point the way to further study for the reader who desires it. This is done, first, because the present volume is intended primarily as a text rather than as a book of reference, and, secondly, because it is felt that the beginner’s interest may be more effectively aroused in this manner than by presenting to him at once every reference available. The latter, if desired, can be readily obtained in the more advanced books which are cited.

It is recognized that illustrations constitute an extremely important feature in a text of this character, and the writer has spared no pains in the attempt to make the figures adequate both in number and quality. It will be evident, however, that the majority of them are not original. This is due to the fact that through the kindness of the authors and publishers indicated below, there were made available a large number of excellent illustrations, which it seemed hardly worth while to attempt to improve upon. Nevertheless, in every instance where it was felt that such improvement was possible, or where it appeared that a new figure would be profitable, original drawings have been inserted. Lastly. it remains to be. stated in this connection that in the case of all borrowed illustrations, great care has been taken to have the illustration and the terms used in its legend agree with the respective description and terminology in the text. The desirability of this, especially in an clexnemarj.' book, is obvious; yet, according to the writer’s observation, it is a feature which is too frequently overlooked.

In conclusion I desire to express my appreciation of the following favors. To Professor Frank R. Eillie and to Henry Holt and Co., I am indebted for their generous permission to use a large number of figures from Lillie’s Development of the Chick; to Professor T. H. Morgan. his co-authors, arid Henry Holt and Co., for certain illustrations from The Mechanism of Memlelian Heredity; to Henry Holt and Co., for numerous figures from Kellicott’s General Embryology and Chordate Development; and to the Delegates and Secretary of the Clarendon Press for a like favor as regards .lenkinson’s Vertebrate Embryology. It is also a pleasure to acknowledge a similar debt to Professor Morgan and The Columbia University Press fr;-2' figures from Heredity and Sex: to Professor J. Playfair McMurrich and P. Blakiston’s Son and Co. for cliches from McMurrich’s Development of the Human Body‘; to P. Blalcistozfs Son and Co. for further clichés from Minot’s Laboratory Text Book of Embryology; to Messrs. Longmans, Green and Co. for cliches from Quain’s Anatomy; to Messrs. G. P. Putnam and Co., for permission to use again certain figures from Marshall’s Vertebrate Embryology, copied and slightly modified by Kellicott; and to Professor 0. Van der Stricht and Dr. T. W. Todd for allowing the use of photomicrographs made in the Anatomical Department of Western Reserve University Medical School from preparations presented to that department by Professor Van der Stricht. In all cases the illustrations thus borrowed are acknowledged in the legends of the figures concerned.

I wish further to express particular gratitude to Professor T. H. Morgan for reading and criticizing the first half of the manuscript; to Professor J. H. McCregor for performing a similar service for the entire hook; to Professor M. M. Metcalf for suggestions regarding the earlier chapters: to my wife for assistance with the proof; and to Pro.fessor R. C. llarrison for the identification of the frog larvae used in niaking certain of my original drawings. Especial gratitude is also felt for the constant interest and helpfulness shown by my colleagues, Professors R. A. Budington and C. G. Rogers.

R. S. MCE.

Oberlin College, August 15, 1923.

Historic Disclaimer - information about historic embryology pages 
Mark Hill.jpg
Pages where the terms "Historic" (textbooks, papers, people, recommendations) appear on this site, and sections within pages where this disclaimer appears, indicate that the content and scientific understanding are specific to the time of publication. This means that while some scientific descriptions are still accurate, the terminology and interpretation of the developmental mechanisms reflect the understanding at the time of original publication and those of the preceding periods, these terms, interpretations and recommendations may not reflect our current scientific understanding.     (More? Embryology History | Historic Embryology Papers)

1949 McEwen: 1 Germ Cells and Amphioxus | 2 Frog | 3 Teleosts and Gymnophiona | 4 Chick | 5 Mammal | 1949 Vertebrate Embryology

Cite this page: Hill, M.A. (2020, May 28) Embryology Book - Vertebrate Embryology (1949). Retrieved from

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