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 DOMESTIC

ESSENTIALS OF
ANIMAL
EMBRYOLOGY
Commissioning Editor: Robert Edwards, Joyce Rodenhuis
Development Editor: Nicola Lally
Project Manager: Nancy Arnott
Designer/Design direction: Charles Gray
Illustration Manager: Merlyn Harvey
Illustrator: Oxford Illustrators
 DOMESTIC
ESSENTIALS OF
ANIMAL
EMBRYOLOGY
By
Poul Hyttel
University of Copenhagen, Denmark

Fred Sinowatz
LMU Munich, Germany

Morten Vejlsted
University of Copenhagen, Denmark

With the Editorial Assistance of

Keith Betteridge
Ontario Veterinary College, University of Guelph, Canada

Foreword by
Eric W. Overström, Ph.D.
Professor and Head
Department of Biology & Biotechnology
Director, Life Sciences & Bioengineering Center
Worcester Polytechnic Institute
Worcester, Massachusetts

Edinburgh London New York Oxford Philadelphia St Louis Sydney Toronto 2010
First published 2010, © Elsevier Limited. All rights reserved.

No part of this publication may be reproduced or transmitted in any form or by any means, electronic or
mechanical, including photocopying, recording, or any information storage and retrieval system, without
permission in writing from the publisher. Permissions may be sought directly from Elsevier’s Rights
Department: phone: (+1) 215 239 3804 (US) or (+44) 1865 843830 (UK); fax: (+44) 1865 853333; e-mail:
healthpermissions@elsevier.com. You may also complete your request online via the Elsevier website at
http://www.elsevier.com/permissions.

ISBN 978-0-7020-2899-1

British Library Cataloguing in Publication Data

A catalogue record for this book is available from the British Library

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A catalog record for this book is available from the Library of Congress

Notice
Neither the Publisher nor the Editors assume any responsibility for any loss or injury and/or damage to
persons or property arising out of or related to any use of the material contained in this book. It is the
responsibility of the treating practitioner, relying on independent expertise and knowledge of the patient,
to determine the best treatment and method of application for the patient.
The Publisher

The
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Printed in China
 CONTENTS

Contributors vii 12. Development of the Blood Cells,

Preface ix Heart and Vascular System . . . . . . . 182
Foreword xi Poul Hyttel
Acknowledgements xiii 13. Development of the Immune
System . . . . . . . . . . . . . . . . . . 208
1. History of Embryology . . . . . . . . . . 1 Morten Vejlsted
Poul Hyttel and Gábor Vajta 14. Development of the
2. Cellular and Molecular Mechanisms Gastro-pulmonary System . . . . . . . . 216
in Embryonic Development . . . . . . . 13 Poul Hyttel
Morten Vejlsted 15. Development of the Urogenital
3. Comparative Reproduction . . . . . . . . 25 System . . . . . . . . . . . . . . . . . . 252
Morten Vejlsted Fred Sinowatz

4. Gametogenesis . . . . . . . . . . . . . . 32 16. Musculo-skeletal System . . . . . . . . 286

Poul Hyttel Fred Sinowatz

5. Fertilization . . . . . . . . . . . . . . . . 56 17. The Integumentary System . . . . . . . 317

Fred Sinowatz Fred Sinowatz

6. Embryo Cleavage and Blastulation . . . 68 18. Comparative Listing of

Developmental Chronology . . . . . . . 330
Morten Vejlsted
Poul Hyttel
7. Gastrulation, Body Folding and
Coelom Formation . . . . . . . . . . . . 79 19. Teratology . . . . . . . . . . . . . . . . . 338
Morten Vejlsted Fred Sinowatz

8. Neurulation . . . . . . . . . . . . . . . . 95 20. The Chicken and Mouse as

Models of Embryology . . . . . . . . . . 383
Fred Sinowatz
Palle Serup (chicken) and Ernst-Martin
9. Comparative Placentation . . . . . . . . 104 Füchtbauer (mouse)
Morten Vejlsted 21. Assisted Reproduction
10. Development of the Central and Technologies . . . . . . . . . . . . . . . 402
Peripheral Nervous System . . . . . . . 120 Gábor Vajta, Henrik Callesen,
Fred Sinowatz Gry Boe-Hansen, Vanessa Hall and
Poul Hyttel
11. Eye and Ear . . . . . . . . . . . . . . . . 163
Fred Sinowatz Index 435
This page intentionally left blank
 CONTRIBUTORS

Several highly qualified scientists have demonstrated their willingness to contribute to the book project.

Keith J. Betteridge BVSc MVSc PhD FRCVS Palle Serup Phd

University Professor Emeritus Director of Research
Department of Biomedical Sciences Department of Developmental Biology
Ontario Veterinary College Hagedorn Research Institute
University of Guelph, Ontario Denmark
Canada
Fred Sinowatz Dr.med vet. Dr.med Dr.habil
Gry Boe-Hansen DVM Phd Professor
Lecturer Institute of Veterinary Anatomy, Histology and
School of Veterinary Science Embryology
University of Queensland, Australia LMU Munich,
Germany
Henrik Callesen DVM PhD DVSc
Research Professor Gábor Vajta MD PhD DVSc
Department of Genetics and Biotechnology Scientific Director
Faculty of Agricultural Sciences, Aarhus University, Cairns Fertility Centre
Denmark Australia
Adjunct Professor, University of Copenhagen,
Ernst-Martin Füchtbauer PhD Dr.habil Denmark
Associate Professor Adjunct Professor, James Cook University, Australia
Department of Molecular Biology
Morten Vejlsted DVM Phd
Aarhus University
Denmark Assistant Professor
Department of Large Animal Sciences
Vanessa Hall PhD Faculty of Life Sciences, University of Copenhagen,
Post Doc Denmark
Department of Basic Animal and Veterinary Sciences
Faculty of Life Sciences, University of Copenhagen
Denmark

Poul Hyttel DVM Phd DVSc

Professor
Department of Basic Animal and Veterinary Sciences
Faculty of Life Sciences, University of Copenhagen
Denmark

vii
This page intentionally left blank

For me, some of the most exciting and glorious
moments in teaching gross anatomy of the domestic
animals have occurred during days shared with
open-minded students surrounded by steel tables
full of pregnant uteri and fetuses in the dissection
room. To examine those fetal specimens is to open
an anatomy book; a book in which each organ and
structure is perfectly defined and its developmental
history is perfectly retained – truly the optimal situ-
ation for memorable anatomical “Aha-Erlebnishen”
for students and teachers alike!
Embryology has always been a prerequisite for a
real understanding of gross anatomy and of the tera-
tology that results from development going awry.
Today, however, it is that and much more besides;
contemporary biomedical research requires embry-
ology (or, rather, developmental biology) to play a
central role in its progress – a role with important
societal implications. Assisted reproduction tech-
nologies, for example, are as much applied to
domestic animals as to humans in which vitro fer-
tilization has become a common step ex soma to
bridge one generation of mankind to the next. In
the domestic animals, techniques such as cloning by
somatic cell nuclear transfer have made possible
genetic modifications that offer the prospects of
modifying animals so that they produce valuable
proteins, serve as models of human diseases, or
provide organs for xenotransplantation in the future.
All of these prospects depend upon a thorough
knowledge of embryology and many of them are
contentious. They put the discipline in an ethical
PREFACE
spotlight that, in my view, obliges contemporary
embryologists to participate in scientifically based
societal debates.
For a while, ever more sophisticated assisted
reproduction technologies moved the “cutting edge”
of embryological research out of the body and into
the in-vitro environment. However, the expansion
of this field to encompass embryonic stem cells has
refocused us on the embryo per se; the control of
stem cell differentiation in vitro will depend abso-
lutely on fundamental knowledge of the molecular
regulation of developmental processes in vivo. The
embryo itself has become the key to success – the
circle is closed!
“Cutting edges” are these days fashioned from an
amalgam of conventional embryology, genomics,
transcriptomics and epigenomics applied to the
investigation of the molecular mechanisms of
developmental biology. Information is growing
exponentially, and to combine in-depth molecular
understanding with overall holistic embryology is a
challenge – a challenge that becomes crystal clear
when writing a textbook on the essentials of domes-
tic animal embryology! For years, teaching the
subject has been hampered by the lack of such a
textbook and medical embryology textbooks were
used as a poor compromise. By 2006, when McGaedy
et al published their welcome textbook on veteri-
nary embryology, we had already embarked on the
present book with the goal of making our own
research material, collected over several decades,
palatable and stimulating for students. Working
 Preface

towards this goal has been a great experience for us “Aha-Erlebnis” in the wondrous world of
and we hope that you the reader will find that we embryology!
have accomplished at least part of what we intended.
Future improvements, of course, will depend largely Poul Hyttel
on feedback and I would much appreciate receiving Vidiekjaer
constructive criticism. Valby, Denmark
In the meantime, it is my sincere wish that June 3, 2009
reading this book may lead you into at least one

My painting (2003) of the open horizon of Skagen, Denmark, where I grew up and was ‘imprinted’. I see embryology in the
same light: a vista with infinite potentials that are just waiting to be realized.

x

Over the last 20 years, modern life science research
efforts have rapidly advanced our knowledge of the
normal and abnormal processes of domestic animal
development. As our depth of understanding of the
cellular and molecular mechanisms has grown, so
too has the recognition of the potential for, and
successful application of, this knowledge to enhance
animal-based food and fiber production. It is during
embryo and fetal development, from the formation
of competent gametes to parturition, that powerful
advancements in molecular genetic manipulation
and assisted reproductive technologies are employed,
and these efforts have had profound impact on
animal production worldwide. As a result of these
advances, there remains an unmet need for a con-
temporaneous text of domestic animal development
to support education and training of today’s veteri-
narians, animal scientists and developmental biolo-
gists. Essentials of Domestic Animal Embryology fulfills
this need by providing the student, the instructor
and the veterinary practitioner with an in depth
presentation of the elaborate, chronological proc-
esses that culminate in formation of functional
embryonic structures from the development of
gametes through the peri-partum period. As our
understanding of the precisely orchestrated proc-
esses of animal development advances, and animal
genomes are further unveiled and analyzed, the
importance of animal development becomes central
to understanding and enhancing animal growth,
sustaining health and determining the underlying
causes of disease.
Although there continues to be available a
number of quality texts of human embryology (for
example Langman’s Medical Embryology), focus on a
single species remains a serious limitation for vet-
erinary and animal science audiences and prevents
a thorough view of the wide and distinct variations
that exist among domestic animal species, with
particular reference to processes of blastogenesis,
implantation and placentation. Certainly, Bradley
FOREWORD
Patton’s 1927 benchmark publication in English,
Embryology of the Pig, provided a wonderfully illus-
trated, descriptive account of this often utilized
example of mammalian embryonic development.
A concise descriptive publication of development
in the pig, The Embryonic Pig: A Chronological
Account, was later published by A.W. Marrable in
1971. In 1984, Drew Noden and Alexander de
Lahunta, similarly recognizing the lack of an ade-
quate text on domestic species embryology that
would be useful for veterinary students, published
The Embryology of Domestic Animals: Developmental
Mechanisms and Malformations. An important con-
tribution to veterinary curricula for many years to
follow, this book presented traditional system-by-
system descriptive material on the developmental
anatomy of domestic species including birds, and
the authors also included many relevant experimen-
tal and clinical case references throughout the book.
Unfortunately, a revised edition was not forthcom-
ing, and is not currently available. More recently,
the finely detailed German text, Lehrbuch der
Embryologie der Haustiere (1991), was published by
Imogen Russe and Fred Sinowatz (current co-
author). Excellent illustrations and micrographs
characterize this comprehensive embryology refer-
ence text of domestic species. Printed only in
German, broad international adoption has been
limited. In 2006, McGaedy and colleagues pub-
lished Veterinary Embryology, a text targeting the
particular needs of the veterinary student. Accord-
ingly, the publication of Essentials of Domestic
Animal Embryology is particularly timely as it fills a
resource void for those students keen to study and
understand the fundamental processes of animal
development, be they students, instructors, research
scientists or veterinary practitioners.
On reflection, this book project was conceived
following from a discussion Poul and I had during
a scientific meeting in 2000. As we shared and com-
pared our experiences teaching animal embryology
 Foreword
and anatomy to veterinary students, there was
mutual recognition that a modern text in domestic
animal development was sorely needed to contrib-
ute to the essential academic underpinnings for 21st
century veterinary and animal science curricula. To
appeal to a more global audience, contributions
were solicited from established co-authors recog-
nized internationally as experts in the field. Their
chapters have been seamlessly woven into a very
readable and informative text book. Of particular
note are the inclusion of Chapters 1, 2, 20 and 21,
each of which provides a distinguishing topic per-
spective, and include a historical account of the
study of animal embryology, a discussion of current
cell and molecular-based regulatory mechanisms
that govern key developmental processes, compara-
tive embryology of the chicken and mouse, and a
succinct summary of the advent, development and
broad application assisted reproductive technolo-
gies (ARTs) to enhance production of domestic
animals, respectively.
Lead author, professor Poul Hyttel is recognized
internationally as a distinguished research scientist,
and a passionate educator and student mentor of
animal reproduction and cell biology at the Royal
Veterinary and Agricultural University, and most
recently the University of Copenhagen. He has
directed both the veterinary anatomy and histology/
cell biology courses for many years in Copenhagen,
xii
and has received formal awards and recognition, as
well as continuous accolades from students for his
passion and dedicated commitment to teaching.
Under his direction, Poul has assembled a distin-
guished international group of contributing co-
authors including professor Fred Sinowatz (Munich)
and Dr. Morten Vejlsted (Copenhagen), among
others. Keith Betteridge, distinguished professor of
animal reproduction at the University of Guelph,
has provided thorough editorial review of the text,
This first edition presents a logical, contemporane-
ous and comprehensive view of the current state of
knowledge in the field. The format provides succinct
text information that is well supported by quality
illustrations, photographs and micrographs. I
believe that the student, instructor and practitioner
alike will embrace this text as it will prove to be an
invaluable resource to further both their education
and their knowledge in the field of domestic animal
embryology.
May 2009      Worcester, Massachusetts, USA
Eric W. Overström, Ph.D.
Professor and Head
Department of Biology & Biotechnology
Director, Life Sciences & Bioengineering Center
Worcester Polytechnic Institute
Worcester, Massachusetts

Writing this embryology textbook set the authors on
a long and winding road. The project was initially
proposed by Eric Overström in 2000. At that time
Eric taught a course on developmental biology at
Tufts University, Boston, US, and held a Fulbright
stipend allowing regular trips to Copenhagen,
Denmark. I am indebted to Eric for his enthusiasm
in initiating the book project which resulted in
many happy hours together in Copenhagen.
I feel enormously privileged to have been able to
undertake this book project with cutting edge scien-
tists who share my passion for scholarly university
life. My co-authors, Fred Sinowatz and Morten Vejl-
sted, have both made extraordinary efforts in writing
their many chapters. In particular, Fred’s astonish-
ing embryological breadth, ranging from the molec-
ular to the gross anatomical levels, has been
indispensable to the setting of our goals, and I am
truly grateful that we were allowed to use high
quality drawings from the previous embryology
textbook that he produced with Imogen Rüsse and
published in German. Other highly qualified con-
tributors, Gry Boe-Hansen, Henrik Callesen, Ernst-
Martin Füchtbauer, Vanessa Hall, Palle Serup and
Gábor Vajta, have each brought their particular
expertise to bear on other chapters to ensure that the
text is as up-to-date as possible. I am greatly indebted
to all these colleagues who have so generously
shared their time and ideas with me.
As non-native English speakers the authors are
extremely grateful for the willingness of Keith Bet-
teridge, one of the pioneers of embryo transfer, to
ACKNOWLEDGEMENTS
undertake a thorough linguistic edit of the com-
pleted text. Due to the breadth of Keith’s scientific
view, the linguistic affair developed into an inspir-
ing dialogue about many conceptual subjects of
embryology. It has been a pleasure to learn from the
extreme precision with which Keith has tackled each
step in the process.
Several qualified persons have devoted time and
given extremely valuable comments to the text.
I would like to thank Marie Louise Grøndahl, Vibeke
Dantzer and Kjeld Christensen for their highly
appreciated efforts.
Images have been an important issue in the pro-
duction of the book. I would like to thank Jytte
Nielsen and Hanne Marie Moelbak Holm for their
skilled contribution to the preparation of thousands
of sections for light and electron microscopy over
the years as well as for digital processing of the
micrographs.
Finally, I would like to thank Danish Pig
Production for a very fruitful collaboration enabling
the collection of thousands of porcine embryos
over the years. The data generated from these
resources have contributed significantly to the book
and many of the photographs are taken from these
embryos.
Poul Hyttel
Vidiekjaer
Valby
Denmark
June 3, 2009
This page intentionally left blank
 CHAPTER
Poul Hyttel and Gábor Vajta
History of embryology
Embryology, the study of development from fertili­
zation to birth, has always intrigued philosophers
and scientists. Universal fascination with the way in
which ‘life’ unfolds has led to spirited discussion of
the complexities of the process amongst embryo­
logists over the years. In 1899, Ernst Haeckel
(1834–1919) considered that ‘Ontogeny is a short
recapitulation of phylogeny’. In other words, ontog-
eny (the development of an organism from the fer­
tilized egg to its mature form) reflects, in a matter
of days or months, the origin and evolution of a
species (phylogeny), a continuing process that is
measured in millions of years. Although this is not
entirely true, one has only to look at a 19-day-old
sheep embryo to understand Haeckel’s viewpoint;
the gill-like pharyngeal arches and the somites, for
example (Fig. 1-1) are common to embryos of all
chordates. Although trained as a physician, Haeckel
abandoned his practice after reading The Origin of
Species by Charles Robert Darwin (1809–1882)
published in 1859 (and available at that time for
fifteen shillings!). Always suspicious of teleological
and mystical explanations of life, Haeckel used
Darwin’s theories as ammunition for attacking
entrenched religious dogma on the one hand and
elaborating his own views on the other. However,
in projecting phylogeny into ontogeny, Haeckel
made one mistake: his mechanism of change
required that formation of new characters, diagnos­
tic of new species, occur through their addition to a
basic developmental scheme. For example, because
most metazoans pass through a developmental
stage called a gastrula (a ball of cells with an infold­
ing that later forms the gut) Haeckel thought that at
one time an organism called a ‘gastraea’ must have
1
existed, looking much like the gastrula stage of
ontogeny. This hypothesized ancestral metazoan, he
thought, gave rise to all multi-celled animals. Such
a ‘single straight line’ conception of phylogeny, with
all creatures standing on the shoulders of predeces­
sors in one trajectory, has now been abandoned;
phylogeny divides into a multitude of lines.
Many of the ancient Greek philosophers were
interested in embryology. According to Democritus
(ca. 455–370 BC), the sex of an individual is deter­
mined by the origin of the sperm: males arising
from the right testicle (of course!) and females from
the left. This hypothesis was somewhat modified by
Pythagoras, Hippocrates and Galen. However,
gender bias was always evident, for science was the
privilege of men; philosophers mostly positioned
females between man and animals, so males were
supposed to originate from the stronger sperm of
the right testicle.
The first real embryologist that we know about
was the Greek philosopher Aristotle (384–322 BC).
In The Generation of Animals (ca. 350 BC) he
described the different ways that animals are born:
from eggs (oviparity, as in birds, frogs and most
invertebrates), by live birth (viviparity, as in placen­
tal animals and some fish) or by production of an
egg that hatches inside the body (ovoviviparity,
which occurs in certain reptiles and sharks). It was
Aristotle who also noted the two major patterns of
cell division in early development: holoblastic
cleavage in which the entire egg is divided into pro­
gressively smaller cells (as in frogs and mammals)
and the meroblastic pattern in which only that
part of the egg destined to become the embryo
proper divides, with the remainder serving nutritive
 Essentials of Domestic Animal Embryology
purposes (as in birds). The fetal membranes and the
umbilical cord in cattle were described by Aristotle
and he recognized their importance for fetal nutri­
tion. By sequential studies of fertilized chick eggs,
Aristotle made the very important observation that
the embryo develops its organ systems gradually –
they are not preformed. This concept of de novo
formation of embryonic structures, which is referred
to as epigenesis, remained enormously controver­
sial for more than 2000 years before becoming fully
accepted; Aristotle was way ahead of his time. The
enigma of sexual reproduction also intrigued Aris­
totle. He realized that both sexes are needed for
conception but felt that the male’s semen did not
contribute to conception physically, but by provid­
ing an unknown form-giving force that interacts
with menstrual blood in the womb of the female to
materialize as an embryo. In this case Aristotle was
wrong. Ironically, in contrast to the non-acceptance
of his correct views on epigenesis, his error about
conception prevailed for about 2000 years, and it
took a battle of almost a hundred years to correct it!
During those 2000 years, the science of embryo­
logy developed very slowly along descriptive ana­
2
Fig. 1-1: Sheep embryo at Day 19 of
development.
tomical lines. One of the first anatomical descriptions
of the pregnant uterus of the pig was by Kopho
(years of birth and death unknown) in the early 13th
century in his work Anatomio Porci. Kopho worked
at the famous medical school of Salerno using pigs
as models because human cadavers could not be
dissected for religious reasons. During the early
Renaissance, the famous and incomparably artistic
anatomical studies of Leonardo da Vinci (1452–
1519) included investigation of the pregnant uterus
of a cow. His drawings of a pregnant bicornuate
uterus and of the fetus and fetal membranes released
from the uterus are reproduced in Fig. 1-2. In an
accompanying drawing, Leonardo depicted a human
uterus cut open to ‘reveal’, not a human placenta
but a multiplex, villous, ruminant placenta (see
Chapter 9)! Clearly it had not been possible for him
to actually study a human pregnancy. The structures
are described in the characteristic mirrored writing
of Leonardo, who worked mostly in Florence.
The Renaissance, from the 13th to the 15th century,
was a great time for anatomical studies and, happily,
coincided with the invention of book printing by
Johann Gutenberg. Thus, the first major publication
 History of embryology

A B

Fig. 1-2: Drawings of Leonardo da Vinci. A: Top: A pregnant bicornuate uterus of the cow. Bottom: Fetus and fetal membranes showing the cotyledons of
the placenta (see Chapter 9). B: Opened simplex uterus of human showing the fetus with the umbilical cord. Note that the placenta is of the ruminant
multiplex type with several placentomes drawn on the cut wall of the uterus and, at the right top, a single cotyledon drawn at a higher magnification
displaying its villous surface.
1

3
 Essentials of Domestic Animal Embryology
on comparative embryology was De Formato Foetu
in 1600 by the Italian anatomist Hieronymus
Fabricius of Acquapendente (1533–1619). Fabri­
cius described and illustrated the gross anatomy of
embryos and their membranes in that book, but was
not actually the first to do so; another Italian anato­
mist, Bartolomeo Eustachius (1514–1574), had
previously published illustrations of dog and sheep
embryos in 1552. We now recognize the names of
Fabricius, in the term bursa Fabricii (the immuno­
logically competent portion of the bird gut), and of
Eustachius in the Eustachian tube.
The work of Eustachius, Fabricius and others gave
insight into how organs develop from their imma­
ture to mature forms, but left unanswered the basic
enigma of how and where the mammalian embryo
originates. However, the development of the micro­
scope by Zacharias Janssen, a Dutch eyeglass maker,
in 1590 ushered in a new era of embryological
science to tackle that 2000-year-old question. The
Dutch dominance in the optical field at that time
may not be just a coincidence; the naval ambitions
of their new empire required excellent telescopes
and lens systems. Janssen’s microscope in its origi­
nal form, however, was not really appropriate for
cell and tissue research; it was approximately 2
metres long, achieved only 10 to 20 times magnifi­
cation, and its principal use was to attract an audi­
ence at country fairs! In 1672, the Italian medical
doctor Marcello Malpighi (1628–1694) published
the first microscopic account of chick development,
identifying the neural groove, the somites, and cir­
culation of blood in the arteries and veins to and
from the yolk. Malpighi also observed that even the
unincubated chick egg is considerably structured,
leading him to think that a preformed version of the
chicken resided in the egg. Later (in 1722), the
French ophthalmologist Antoine Maître-Jan (1650–
1730) pointed out that although the egg examined
by Malpighi was technically ‘unincubated’, it had
been left sitting in the Bolognese sun in August and
so was certainly not ‘unheated’. Nevertheless, Mal­
pighi’s notion of a preformed chicken initiated one
of the great debates in embryology that was to last
throughout the 17th and 18th centuries. The question
was: are the organs of the embryo formed de novo
4
(epigenesis), or they are already present in a mini­
ature form in the egg (or the sperm when this cell
was discovered), a concept referred to as preforma-
tion. We will return to this debate in a moment.
The new microscopic techniques also prompted
a vigorous search for the mammalian gametes. The
chicken egg and its initial transformation into a
chick were obvious, as Aristotle had described, but
what mediated the formation of the embryo in
mammals? Where was the mammalian egg to be
found?
One of the earliest and most influential names in
the fascinating story of the discovery of the mam­
malian egg was that of William Harvey (1578–
1657), personal physician to the English kings
James I and Charles I, and famous for his descrip­
tion of the circulation of the blood. In 1651, Harvey
published De Generatione Animalium (Disputations
touching the Generation of Animals) with a famous
frontispiece showing Zeus freeing all creation from
an egg bearing the inscription Ex ovo omnia (All
things come from the egg). However, it should be
realized that, far from advancing 17th century knowl­
edge of reproduction and embryology, Harvey’s
observations in some ways impeded progress. From
having studied with Fabricius, Harvey was imbued
with Aristotle’s view that the semen provided a force
that interacted with the menstrual blood to materi­
alize as an embryo. Harvey set out to understand
this process by looking for the earliest products of
conception in female deer killed during the breed­
ing season in the course of King Charles I’s hunts in
his Royal forests and parks over a 12-year period. In
the red and fallow deer that he studied, the male’s
rut begins in mid-September and so Harvey dis­
sected uteri throughout the months of September to
December. Believing, wrongly, that copulation coin­
cides with the onset of the rut, Harvey was mystified
to find nothing that he recognized as an embryo
until mid-November, some two months later. This
forced him to the erroneous, but entirely logical
conclusion that ‘nothing after coition is to be found
in [the] uterus for many days together’. When he
did find a conceptus, that, for Harvey, was the egg:
‘Aristotle’s definition of an egg applies to it, namely,
an egg is that out of a part of which an animal is

begotten and the remainder is the food for that
which is begotten’.
Three factors of veterinary interest had led this
brilliant man astray. First, he did not appreciate that
the females did not come into oestrus until early
October and so his estimates of breeding dates were
wrong. Second, he dismissed the ovaries (‘female
testicles’) as making no contribution to conception
because they failed to swell up as the testes of males
do during rut. Third, expecting to find an egg-shaped
conceptus, he failed to recognize the ‘purulent
matter … friable … and inclining to yellow’, which
he observed much earlier after mating and describes
quite vividly, as being the filamentous blastocyst so
characteristic of ruminants. Had Harvey used a lens,
or conducted his studies on species (like rabbits or
horses) with spherical early conceptuses, discovery
of the real egg might have been advanced
considerably!
It is easy to be critical with hindsight of course,
and we should remember that Harvey made impor­
tant contributions to embryology: his descriptions
of early development were impeccable; he was the
first to observe the blastoderm of the chick embryo
(the small region of the egg containing the yolk-free
cytoplasm that gives rise to the embryo proper) and
to indicate that blood islands form before the heart
does; and he was aware of the gradual development
of the embryo, subscribing to the school of epi­
genesis as did Aristotle.
The observations by Harvey and the search for the
mammalian egg were extended by Regnier de Graaf
(1641–1673) who performed detailed studies of the
female reproductive organs, especially the ovary. De
Graaf, like his friend Leeuwenhoek, worked in Delft.
From a comparison of mammalian ovaries with
those of chicken, de Graaf considered mammalian
antral ovarian follicles to be the eggs; an assumption
he confirmed by tasting! His contribution to science
was later acknowledged by the German medical
doctor Theodor Ludwig Wilhelm Bischoff (1807–
1882) who introduced the nomenclature ‘Graafian
follicle’. De Graaf also noted some connection
between follicular maturation and the development
of oocytes but, without an appropriate microscope,
he could not substantiate this and his observations
History of embryology 1
were for a long time forgotten. Had he lived a little
longer (he died tragically early, at the age of 32), the
discovery of the mammalian egg could probably
have avoided a delay of about 150 years!
As it was, the first scientist to actually see the
mammalian egg (which everyone believed to exist,
but no one had seen) was the Estonian medical
doctor Karl Ernst von Baer (1792–1876). He
opened the ‘Graafian egg’, as the follicle was known
at that time, and saw with his naked eye a small
yellow point which he released and examined under
the microscope (Baer, 1827). There, upon a first
glance, Baer was stunned and could hardly believe
that he had found what so many famous scientists
including Harvey, de Graaf, Purkinje and others had
failed to find. He was so overcome that he had to
work up courage to look into the microscope a
second time. The mammalian egg had been
identified.
How about the spermatozoa? Anton van Leeu-
wenhoek (1632–1723), a Dutch tradesman and
scientist from Delft, was the first to report having
seen moving spermatozoa. He constructed a single
lens microscope that magnified up to about 300
times. Technically, this microscope was an amazing
achievement – no bigger than a small postage stamp
and resembling a primitive micromanipulator. It
was used close to the eye like a magnifying glass.
Using this, Leeuwenhoek drew spermatozoa from
different species. Initially, Leeuwenhoek was reluc­
tant to study sperm and he questioned the propriety
of writing about semen and intercourse. When he
first focused his microscope on semen, Leeuwen­
hoek discovered what he then took to be globules.
However, he so disliked the prospect of having to
discuss his findings that he quickly turned to other
matters. Three or four years later, however, in 1677,
a student from the medical school at Leiden brought
him a specimen of semen in which he had found
small animals with tails, which Leeuwenhoek now
observed as well. Consequently, Leeuwenhoek
resumed his own observations and, in his own
semen (acquired, he stressed, not by sinfully defiling
himself but as a natural consequence of conjugal
coitus), observed a multitude of ‘animalcules,’ less
than a millionth the size of a coarse grain of sand
5
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certain feeling between them. But now each of them had discerned
something in the other that transcended anything that they had
previously known.
In short, I saw a man’s affection for another man shining from
each pair of eyes.
The rest of the gang, knowing the entire situation, chimed in with
a lot of Air Service kidding, about that last ride, as for instance,
Sleepy Spears’ remark:
“Any hop is foolish, and a last one is suicide. You’ve made it sure
death by letting Penoch fly you.”
“In fact,” Tex MacDowell chimed in, in his soft southern drawl,
“I’ve had a shovel all ready to pick Penoch up with, for a long time.
I’d pick a hang-over in preference to Penoch any time.”
I guess nobody outside of the men who fly understand what air
kidding is. Probably I don’t myself. But in my dumb way I think that
it’s like a kid’s whistling when he passes a graveyard, or, perhaps,
laughing at the worst that could happen, so that, when it does
happen, it won’t mean anything.

When lunch was over, and we were all drifting out of the mess-hall, I
suddenly realized that I would like to share, a little bit, those last
hours. I’d been so close to the thing that I wanted to hang around the
outskirts of it, until Kennedy left. In other words, I was sentimental,
and I thought quite a lot of Penoch, at that. So I said casually—“I
think I’ll take a little private hop for myself when you do—a sort of
chaser for the poker game, eh?”
Kennedy, I think, bewildered as he was at the world that had been
opened to him so recently, appreciated the impulse behind my
suggestion.
“Sort of be my guard of honor, eh?” he said. But those cold eyes
were soft. “In fact, we’ll be glad to have company, won’t we
Peewee?”
And so it happened that a half-hour later the three of us were on
the line. Our two ships were being warmed up, and the mechanics,
satisfied, had brought them down to idling.
 “I’ll sit in the back seat, big boy,” Penoch told Kennedy, “but don’t
think that I won’t take the stick away from you any time!”
You know, of course, that De Havilands are dual-control ships,
but all the instruments are in the front cockpit, and that, in a manner
of speaking, is the driver’s seat.
I got into my own plane and, as I taxied out for the take-off, I
couldn’t exactly analyze my reason for being there. I guess it was a
sort of vague tribute to Kennedy, and yet I had a funny feeling that
something might happen—so much so that as I turned the motor full
on and pushed the stick forward I was so absent-minded that I nearly
broke the propeller, because I had the nose of the plane down so far.
Kennedy and Penoch had taken off first, and I just followed them
as they circled the field for altitude. When we got to the tremendous
height of fifteen hundred feet, Kennedy, who was doing the piloting,
started due west for Laredo.
I’ll swear that we were not a thousand yards from the airdrome
when it happened. I was flying possibly a hundred yards back of
them, and almost the same distance to the right. All of a sudden I
saw their ship go into a dive.
That meant something. When Kennedy started turning back
toward the field I knew that the motor had cut out. Then, as I noticed
the propeller, I knew that the motor had not only cut out, but had cut
dead. The stick was revolving slower and slower. There was no
motor power behind it.
The next second their ship was blanketed in fire. The motor was a
mass of blue flame. Kennedy had not cocked the ship up into a side-
slip soon enough. That blows the flames upward, away from the
pilot. The left wing was afire before he started slipping, and that
second of backward draught, because they were in a dive when the
fire started, had caused the fuselage to catch fire in a dozen places.
My body and brain were numb. Even so, I subconsciously knew
what had happened. The gas line had broken, and the gas, sprayed
over the hot motor, had ignited.
They had gone into the slip—too late—before I started toward
them. I was diving my ship, motor full on. There was nothing I could
do—Penoch and Kennedy burned to death before my eyes—I was
just getting nearer for no reason.
I suppose that it registered on me at the time, because I
remember it so vividly now. Penoch told me what was said. Anyway,
I saw Kennedy, bearing the brunt of the fire in the front cockpit, turn
and gesture. He was talking. I could see through the smoke and fire
his lips moving. Penoch told me that what he said was:
“Get out on the wing! I’m not leaving the ship— Let me jump! If
Slim sees you, he’ll get close. God! I can’t last long! It’ll just be two
instead of one—”
What I saw was Penoch getting out of the back cockpit, hanging
by his hands from the cowling. He hauled himself along the side of
the ship, his feet dangling over space. His head was turned
backward, to protect his eyes, and his clothing was charring,
because he was out where the air-stream could reach him, and no
flame could get a real start. But Kennedy, in the cockpit—
I was going through the most horrible nightmare that can be
conceived. And yet, I instinctively sensed the possibility of saving
Penoch, as he reached the right wing and started crawling along it.
Then Ralph brought the burning ship level; that blew the fire right
back on him. I was close, and for that horrible minute I guess I
ceased to think of my own safety. I knew that there was one
desperate chance to save one of the two, and Penoch, of course,
was the one, because Kennedy had willed it so.
Penoch was out of the fire now, at the edge of the right wing. He
was hanging from the edge of it by his hands as I flew my ship up
into position, my left wing underneath the other’s right one. And
Kennedy—I don’t know by what transcendent power he was able to
do it, as he burned to death—kept his ship level. Penoch dropped—
his only chance for life—and he landed on my left wing. He grabbed
the cabane strut, the little metal horn at the edge of the wing to which
the control wires for the ailerons are attached, and passed out.
An instant later, Kennedy, a human bonfire, leaped from his
burning ship. He fell out—blessed surcease from pain. And the ship,
like a flaming coffin, seemed to follow his body down.
 Penoch eventually got back into the rear cockpit, of course, and
we’re both here to tell the tale.
Sometime I hope that a burning ship will cease to trace a crimson
path across my dreams. Probably it won’t.
Anyway, if Ralph is a spook in some spiritual village, teaching the
Twelve Apostles how to play poker, I hope he has time to tip his halo
in acknowledgment of the salute of Slim Evans to a crook and a
hero, a scoundrel and a man.

THE END

Transcriber’s Note: This story appeared in the November 1, 1927

issue of Adventure magazine.
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