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Embryology
Embryology

Ronald W. Dudek, Ph.D.

Professor
Department of Anatomy and Cell Biology
Brody School of Medicine
East Carolina University
Greenville, North Carolina
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 Preface

The sixth edition of BRS Embryology includes improvements based on suggestions and
comments from the many medical students who have used this book in preparation for
the USMLE Step 1 examination and those students who have reviewed the book. I pay
close attention to these suggestions and comments in order to improve the quality of this
book. The goal of BRS Embryology is to provide an accurate and quick review of important
clinical aspects of embryology for the future physician. In addition, we have added color
to the diagrams. In this regard, I have used the following color scheme. The ectoderm/
neuroectoderm and derivatives are colored blue. The neural crest cells and derivatives are
colored purple. The mesoderm and derivatives are colored red. When multiple mesoder-
mal structures are involved (e.g., reproductive systems), I used light red and dark red. The
endoderm and derivatives are colored yellow.
Many times in the history of science, certain biological concepts become entrenched
and accepted as dogma even though recent evidence comes to light to challenge these con-
cepts. One of these concepts is the process of twinning. Recent evidence calls into question
the standard figures used in textbooks on how the process of twinning occurs. In particular,
it is becoming increasingly difficult to ignore the fact that dizygotic twins are sometimes
monochorionic. Although we by far do not know or attempt to explain exactly how twinning
occurs, it seems that the interesting cell and molecular events involved in twinning occur
in the first few cell divisions during first three or four days after fertilization. You are not a
twin because the inner cell mass splits. The inner cell mass splits because you are a twin.
This evidence warrants a new twinning figure (Figure 6.6) that does not comport with the
standard figures but tries to embrace recent evidence, although many may call it controver-
sial. Progress in our scientific understanding of twinning will never occur if our concept of
the twinning process is overly simplistic and reinforced by standard figures repeated over
and over in textbooks. Some published references that speak to this twinning issue include
Boklage,1,2 Yoon et al.,3 Williams et al.,4 and Hoekstra et al.5
I understand that BRS Embryology is a review book designed for a USMLE Step 1 review
and that you will not be faced with a question regarding this twinning concept, but I know
my readers are sophisticated enough to appreciate the scientific and clinical value of being
challenged to question traditional concepts as “grist for the mill” in discussions with your
colleagues.
I would appreciate receiving your comments and/or suggestions concerning BRS Em-
bryology sixth edition, especially after you have taken the USMLE Step 1 examination. Your
suggestions will find their way into the seventh edition. You may contact me at dudekr@
ecu.edu.
Ronald W. Dudek, PhD

v
vi Preface

REFERENCES
1. Boklage CE. Traces of embryogenesis are the same in monozygotic and dizygotic twins: not
compatible with double ovulation. Hum Reprod. 2009;24(6):1255–1266.
2. Boklage CE. How New Humans Are Made: Cells and Embryos, Twins and Chimeras, Left and
Right, Mind/Self/Soul, Sex, and Schizophrenia. Hackensack, NJ: World Scientific Publishing;
2010.
3. Yoon G, Beischel LS, Johnson JP, et al. Dizygotic twin pregnancy conceived with assisted repro-
ductive technology associated with chromosomal anomaly, imprinting disorder, and monocho-
rionic placentation. J Pediatr. 2005;146:565–567.
4. Williams CA, Wallace MR, Drury KC, et al. Blood lymphocyte chimerism associated with IVF
and monochorionic dizygous twinning: case report. Hum Reprod. 2004;19(12):2816–2821.
5. Hoekstra C, Zhao ZZ, Lambalk CB, et al. Dizygotic twinning. Hum Reprod Update.
2008;14(1):37–47.
 Contents

Preface v

1. PREFERTILIZATION EVENTS 1
I.Sexual Reproduction 1
II.Chromosomes 1
III.Meiosis 2
IV. Oogenesis: Female Gametogenesis 2
V. Spermatogenesis: Male Gametogenesis 4
VI. Clinical Considerations 4
Study Questions for Chapter 1 8
Answers and Explanations 10

2. WEEK 1 OF HUMAN DEVELOPMENT (DAYS 1–7) 12

I.Fertilization 12
II.Cleavage and Blastocyst Formation 13
III.Implantation 14
IV. Clinical Considerations 14
Study Questions for Chapter 2 15
Answers and Explanations 17

3. WEEK 2 OF HUMAN DEVELOPMENT (DAYS 8–14) 18

I.Further Development of the Embryoblast 18
II.Further Development of the Trophoblast 18
III.Development of Extraembryonic Mesoderm 20
IV. Clinical Considerations 20
Study Questions for Chapter 3 22
Answers and Explanations 24

4. EMBRYONIC PERIOD (WEEKS 3–8) 26

I. General Considerations 26
II. Further Development of the Embryoblast 26
vii
viii Contents

III. Vasculogenesis (De Novo Blood Vessel Formation) 28

IV. Hematopoiesis (Blood Cell Formation) 31
V. Clinical Considerations 31
Study Questions for Chapter 4 33
Answers and Explanations 35

5. CARDIOVASCULAR SYSTEM 37
I.Formation of Heart Tube 37
II.Primitive Heart Tube Dilations 37
III.The Aorticopulmonary Septum 39
IV. The Atrial Septum 41
V. The Atrioventricular Septum 43
VI. The Interventricular Septum 45
VII. The Conduction System of the Heart 46
VIII. Coronary Arteries 47
IX. Development of the Arterial System 47
X. Development of the Venous System 49
Study Questions for Chapter 5 50
Answers and Explanations 53

6. PLACENTA AND AMNIOTIC FLUID 55

I. Formation of the Placenta 55
II. Placental Components: Decidua Basalis
and Villous Chorion 55
III.Placental Membrane 58
IV. The Placenta as an Endocrine Organ 59
V. The Umbilical Cord 60
VI. Circulatory System of the Fetus 60
VII. Amniotic Fluid 62
VIII. Twinning 62
IX. Clinical Considerations 65
Study Questions for Chapter 6 68
Answers and Explanations 70

7. NERVOUS SYSTEM 71
I. Overview 71
II. Development of the Neural Tube 71
III. Neural Crest Cells 73
IV. Placodes 75
V. Vesicle Development of the Neural Tube 75
VI. Histogenesis of the Neural Tube 76
VII. Layers of the Early Neural Tube 78
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