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2015v1.0
CONCISE CLINICAL
EMBRYOLOGY
AN INTEGRATED,
CASE-BASED APPROACH
Mark G. Torchia, DipBT, MSc, PhD

Associate Professor, Department of Surgery and Department of
Human Anatomy and Cell Sciences, Max Rady College of Medicine,
Rady Faculty of Health Sciences, University of Manitoba
Vice-Provost (Teaching and Learning), University of Manitoba
Winnipeg, Manitoba, Canada
T.V.N. (Vid) Persaud MD, PhD, DSc, FRCPath (Lond.), FAAA

Professor Emeritus and Former Head, Department of
Human Anatomy and Cell Science
Professor of Pediatrics and Child Health
Associate Professor of Obstetrics, Gynecology,
and Reproductive Sciences
Max Rady College of Medicine, Rady Faculty of Health Sciences,
University of Manitoba, Winnipeg, Manitoba, Canada
Elsevier
1600 John F. Kennedy Blvd.
Ste 1800
Philadelphia, PA 19103-2899
CONCISE CLINICAL EMBRYOLOGY:

AN INTEGRATED, CASE-BASED APPROACH ISBN: 978-0-323-69615-9
Copyright © 2021 by Elsevier, Inc. 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. Details on how to seek permission, further information about the Publisher’s permissions policies
and our arrangements with organizations such as the Copyright Clearance Center and the Copyright Licensing Agency,
can be found at our website: www.elsevier.com/permissions.
This book and the individual contributions contained in it are protected under copyright by the Publisher (other than
as may be noted herein).
Notice
Practitioners and researchers must always rely on their own experience and knowledge in evaluating and using
any information, methods, compounds or experiments described herein. Because of rapid advances in the medical
sciences, in particular, independent verification of diagnoses and drug dosages should be made. To the fullest
extent of the law, no responsibility is assumed by Elsevier, authors, editors or contributors for any injury and/or
damage to persons or property as a matter of products liability, negligence or otherwise, or from any use or
operation of any methods, products, instructions, or ideas contained in the material herein.
Library of Congress Control Number: 2020951836
Content Strategist: Jeremy Bowes

Content Development Specialist: Erika Ninsin
Publishing Services Manager: Deepthi Unni
Project Manager: Srividhya Vidhyashankar
Design Direction: Margaret Reid
Printed in United States of America
Last digit is the print number: 9 8 7 6 5 4 3 2 1

Mark:
For Eddie James Torchia
Our dear grandson and little ray of sunshine.
To my wife, Barbara, our children Erik and Muriel, and their spouses Sarah and Caleb
- thank you for your love and support.
Vid:
For Gisela
My lovely wife and best friend, for her endless support and patience.
Preface
This comprehensive yet concise textbook is designed for ongoing discussion amongst peers, advanced learning and
students in all health fields learning human embryology, knowledge testing.
as well as for the review of human embryology in clinical Each chapter also provides fundamental molecular biol-
practice. The text is copiously illustrated to provide visual ogy considerations. This information is derived from the
cues and resources for better understanding. Accompany- extant literature and is based mainly on experiments with
ing this book, in online format, are 18 exceptional colour animal models including mice because the human cells or
animations, with narrations, which will assist the student tissues required to examine such science are not generally
in learning the various stages of human embryo and fetal available. As such, keep in mind that as knowledge of mo-
development. lecular genetics and biology progresses, the specific genes
A clinical case scenario is provided at the beginning of and their products that are identified may change or be ex-
each chapter. These cases are not straightforward and many panded upon.
of the concepts or considerations will require the reader The section on Clinical Issues in each chapter provides a
to seek information outside the direct field of clinical em- description of the common congenital anomalies and other
bryology; this helps to place the knowledge and details clinical information related to the embryology details con-
of embryology within the larger concept of clinical care. tained in the chapter. Finally, each chapter has a brief refer-
Follow-on scenarios for each case are found at the end of ence list that can be used to find additional details about the
each chapter and provide the reader with an opportunity to clinical cases, molecular biology and clinical embryology.
further expand the ability to problem solve, think broadly Learners wanting to test their knowledge or prepare for
and search for answers beyond this textbook. Answers are examinations will also benefit from the multiple choice
not provided to these cases so that they can be used for questions we have provided through the website.
vi
Acknowledgements
We are indebted to Mr. Jeremy Bowes, Senior Content Project Manager/Health Content Manager for their help-
Strategist, for his invaluable insights and unstinting sup- ful suggestions. Finally, we would like to Dr. Brad Smith,
port in the preparation of Concise Clinical Embryology. University of Michigan, for graciously providing the image
We are particularly grateful to Ms. Erika Ninsin, Content (Carnegie Stage 18 human embryo) which is on the cover
Development Specialist, Ms. Meghan Andress, Content of this book (Imaging performed at the Center for In-Vivo
Development Manager, and Ms. Sri Vidhya Vidhyashankar, Microscopy, Duke University).
vii
Contents
SECTION 1
GENERAL DEVELOPMENT SECTION 2
DEVELOPMENT OF
OF THE EMBRYO AND ORGAN SYSTEMS
FETUS
8 Development of the Cardiovascular,
1 Introduction 2 Haematopoietic and Lymphatic Systems 44
2 Reproductive Organs and Gametogenesis 4 9 Development of the Body Cavities, Diaphragm,

Respiratory System, and Head and Neck 58
3 Fertilisation and Reproductive
Technologies 12 10 Development of the Nervous System, Eyes and
Ears 71
4 Implantation and Week 2 18
11 Development of the Alimentary System 82
5 Weeks 3 to 8—General Organogenesis 23
12 Development of the Urogenital System 89
6 Placentation and Membranes 31
13 Development of Skeletal, Muscular and
7 Fetal and Neonatal Period 37 Integumentary Systems 98
14 Teratogenesis and Birth Defects 107
Multiple Choice Question Answers 112
Index 115
viii
Video Table of Contents
Video 3.1 Fertilisation 12 Video 9.3 Pharyngeal Apparatus 64
Video 3.2 Blastocyst 12 Video 9.4 Face and Palate 66
Video 4.1 Implantation 18 Video 10.1 Nervous System 71
Video 5.1 Gastrulation 23 Video 10.2 Eyes 76
Video 5.2 Folding 28 Video 10.3 Ears 78
Video 8.1 Heart 44 Video 11.1 Gastrointestinal 82
Video 8.2 Vascular 49 Video 12.1 Urinary 89
Video 9.1 Body Cavities 58 Video 12.2 Reproductive System 91
Video 9.2 Respiratory System 61 Video 13.1 Limbs 101
ix
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SECTION 1
GENERAL DEVELOPMENT OF
THE EMBRYO AND FETUS
PART OUTLINE
1 Introduction 4 Implantation and Week 2
2 Reproductive Organs and 5 Weeks 3 to 8—General
Gametogenesis Organogenesis
3 Fertilisation and Reproductive 6 Placentation and Membranes
Technologies 7 Fetal and Neonatal Period
1 Introduction
visual resource to further enhance the textual explanations
Case Scenario and development paths.
A clinical case scenario is provided for each chapter. As
A 26-year-old woman (GW) presents to you, a nurse practitio-
you will discover, the cases are not straightforward, and many
ner at a public health clinic, with severe odynophagia. The his-
tory and physical examination leads you to strongly believe she of the words, concepts or considerations will require the
has a streptococcal pharyngitis; a swab is taken for rapid strep reader to seek information outside the direct field of clinical
test (which is positive) and you prescribe amoxicillin. When you embryology—this helps to situate the knowledge and details
inquire about her obvious pregnancy, GW reports being ap- of embryology within the larger concept of clinical care. The
proximately 5 months pregnant based on when she remem- clinical case in this chapter is a good example. You will need
bered having had her last menstrual period. She said that the to consider, for example, infectious disease, genetics, phar-
father is a 55-year-old companion. GW has been living on the macology and neonatal cardiology and combine that knowl-
street and in shelters for the past year since discharge from edge to answer the question. The follow-on scenarios to the
an inpatient facility for treatment of a crystal methamphetamine original case, found at the end of each chapter, will further
addiction. She vehemently denies use of methamphetamine
expand your need to problem solve, think broadly and search
since that treatment. She has not sought any other medical
care in the interim. GW has had only one previous pregnancy for answers beyond this textbook. Answers are not provided
which resulted in the birth of a son, now 5 years old, and cur- to the cases so that they can be used for ongoing discussion
rently living with the maternal grandparents. Her son was born amongst peers, advanced learning and knowledge testing.
with a bilateral cleft palate. You recommend a fetal ultrasound Each chapter also provides molecular biology consider-
as soon as possible, to which she agrees. ations. This information is based mainly on experiments
Questions for reflection: Why might an ultrasound fetal with animal models including mice because the human cells
assessment be warranted? What concerns might you have or tissues required to examine such science are not gen-
related to the health of GW? What impact, if any, might erally available. As such, keep in mind that as knowledge
these issues have on the health of her fetus, including
of molecular genetics and biology progresses, the specific
risk for anomalies? Is the father’s age or the fact that her
5-year-old son had a cleft palate relevant to the current
genes and their products that are identified may change or
pregnancy? Why? be expanded upon.
The study of embryology is essential for the understanding Molecular Biology Considerations
of both normal anatomy and congenital anomalies. Moreover,
• TFG-ß, BMP, FGF10, MX1, IRF6—most commonly impli-
the practice of obstetrics and neonatal–perinatal medicine
cated pathways for palatine clefting
involves clinical embryology. Although infant mortality rates
have been decreasing steadily in North America for the past
50 years, the 2018 rate in the United States remains at 5.6 The section Clinical Issues in each chapter provides a
per 1000 live births, 4.3 per 1000 in Canada and 11 per 1000 description of the common congenital anomalies and other
in Mexico. Given that congenital anomalies are the second clinical information, related to the embryology details con-
leading cause of infant mortality (behind premature birth), tained in the chapter.
the need to better understand the mechanisms of normal Finally, each chapter provides a brief reference list that
embryo and fetal development and the factors that impact can be used to find additional details about the clinical
this development, leading to congenital anomalies remains cases, molecular biology and details of clinical embryology.
very high. The growing field of molecular biology and the We encourage you to seek additional information during
development of many novel laboratory techniques have led to your studies as the timing of book printing, relative to the
a significant improvement of our knowledge of the temporal constant gain of knowledge and reporting, negates the pos-
and regional expression of genes and their products to con- sibility of including the very most recent literature, although
trol such processes as morphogenesis. the authors have tried their utmost to provide citations that
are as current as possible.
USING THIS TEXTBOOK
OTHER IMPORTANT INFORMATION
This textbook is designed to offer a concise knowledge base
for the study or review of clinical embryology. The accompa- Throughout this textbook, the specified age of embryos
nying illustrations (drawing and medical imaging) provide a and fetuses as it relates to specific structures and other
2
CHAPTER 1 — Introduction 3
developments, has been quoted as fertilisation age—length • The primary palate, with each other, and the nasal sep-
of time from the date of fertilisation. tum (secondary palate cleft).
In the clinical context, gestational age is indicated as the
Some clefts appear as part of single mutant gene or chro-
time from the date of the start of the last menstrual period
mosomal syndromes or following the effects of teratogenic
(LMP). Given that ovulation (and shortly thereafter, fertili-
substances.
sation) occurs typically around 14 days after the start of the
menstrual period, gestational age LMP is approximately
2 weeks or 14 days greater than fertilisation age.
It is important to specifically describe the method used Case Outcome
for indicating ‘gestational age’, so that confusion does not
arise, especially when ordering or interpreting ultrasound Fetal ultrasound showed a male fetus of approximately 22
images or comparing between times within a patient history. weeks of age (based on femur length, biparietal diameter, head
Because the Federative International Committee on circumference and abdominal circumference), which would ap-
Anatomical Terminology does not recommend the use of proximately align with the predicted age based on the patient’s
eponyms, for the most part, this book follows suit (there last menstrual period. The ultrasound also detected an isolated
are few exceptions to this when the clinical eponym is most membranous ventricular septal defect (VSD). The remainder of
the examination was normal. Sixteen weeks later, GW had a
commonly used).
vaginal delivery. The neonate had good Apgar scores (7/8 at
There will be a number of terms in this book that may 1/5 minutes). The birth weight was at the 4th percentile. Other-
not be familiar to the reader, not limited to just those of wise the infant appeared normal.
embryology. It is recommended that the reader search for Additional reflection: What is the error rate for estimating
those definitions from a reliable source of such medical delivery dates from a single ultrasound examination at 22
information. weeks? Was it likely that the ultrasound was in error or that
Anatomical position and direction terms are used GW delivered early or both or neither? Why? What is the
throughout this book. In adults, the terms anterior and pos- likely cause of the VSD? How common are these anoma-
terior are used to describe the front and back of the body lies and what treatment is required and when, if any? What
or limbs or relative positions of one structure to another. In might be the causes for the baby to be born at such a low
percentile birth weight? What other concerns might you
the fetus or embryo, the terms ventral and dorsal are used,
have regarding the health of the neonate or GW?
respectively. In addition, the terms caudal or rostral are used
to denote a relationship to the head, whereas caudal is used
to denote relationship to the caudal eminence or tail.
CLINICAL ISSUES
BIBLIOGRAPHY
CLEFT PALATE Methods for estimating the due date. Committee Opinion No. 700.
American College of Obstetricians and Gynecologists. Obstet Gyne-
Palate clefts arise from failure of the lateral palatine process col 2017;129:e150–154.
Deshpande AS, Goudy SL. Cellular and molecular mechanisms of
to fuse with: cleft palate development. Laryngoscope Investig Otolaryngol
• The primary palate (anterior palate cleft) 2019;4(1):160–4.
• Each other and the nasal septum (posterior palate cleft)
2 Reproductive Organs and
Gametogenesis
tissues of the uterus and vagina (endometrium and mucosa,

Case Scenario respectively) increase in thickness. It is not until approxi-
mately 30 months later that menstruation begins, although
A 24-year-old woman (KR) presents at her new family physician
the regularity of menstruation may be variable for a number
with difficulty conceiving. She and her husband have been try-
ing to have a child for almost 5 years. Her husband recently had
of months as anovulatory cycles are common. In general,
his sperm count and morphology tested, and this has proven the age at which puberty begins in females has been decreas-
to be normal. KR is now seeking additional advice and investi- ing since the mid-1940s; the reasons for this are not known,
gation for herself. KR describes her menstrual cycle as varying but may be related to the increase in child obesity or other
in length, and occasionally she has missed her period entirely. environmental factors.
Otherwise, she has been healthy. KR also has severe acne. It In males, the enlargement of the testicles and develop-
had been previously controlled after a course of antibiotics and ment of pubic hair are the early signs of the onset of pu-
topical gel treatment when she was 21 years old, but the acne berty, typically occurring at approximately 10 years of age
has now returned. KR mentioned that she has had acne since (SMR 2). The testes and the penis continue to enlarge until
she was 13 years old. Recently, she began to notice more dark
late adolescence under the influence of both LH and tes-
hair growth on her chin and areolas, and that her leg hair has
had a noticeable regrowth after shaving. Her body mass index
tosterone secretion as do the prostate and seminal vesicles.
(BMI) is 20.6. Sperm appear approximately 3 to 4 years following the onset
Questions for reflection: Are there other fertility consid- of puberty. Although males undergo some degree of breast
erations for KR’s husband beyond the semen analysis that enlargement, gynaecomastia, during puberty, this tends to
might be investigated? What might be the connection, if resolve spontaneously in later stages of adolescence.
any, between KR’s recurrent acne and potential fertility
concerns? What further testing or consultation might be
appropriate for KR?
MALE REPRODUCTIVE ORGANS
The penis (Fig. 2.1) acts as the conduit for both urine and
ejaculate to exit the body. It consists of the glans or head,
PUBERTY which in uncircumcised men is covered by the prepuce or
foreskin. The urethral opening is found at the tip of glans
The reproductive organs (or primary sex characteristics) penis which forms from the expanded distal end of the cor-
develop in utero. Maturation of the reproductive organs pus spongiosum. The vascular corpus cavernous surrounds
and the appearance of secondary sex characteristics (such the corpus spongiosum, which when expanded by blood,
as breast growth, presence of axillary and public hair) oc- provide the erectile function of the penis. The erectile tissue
cur after puberty—the transitional process from childhood of the corpus spongiosum supports the urethra and main-
to adulthood. The exact biological trigger that starts the tains its patency during an erection.
process of puberty is unclear; however the initiation of go- The testes are the oval-shaped, sperm- and testosterone-
nadotropin-releasing hormone (GnRH) pulsing leads to the producing organs found within the scrotum. The testes are
secretion of luteinising hormone (LH) and follicle-stimulat- covered with a thick fibrous capsule, the tunica albuginea,
ing hormone (FSH) by the pituitary. LH and FSH, in turn, and contain a series of coiled seminiferous tubules within
stimulate the secretion of androgens and oestrogens from which sperm development occurs. The seminiferous tu-
the gonads (the hypothalamic–pituitary–gonadal axis). The bules are connected to the tubuli recti. The rete testes are
Tanner scale or sexual maturity rating (SMR; 1 = preadoles- connected to the epididymis. The duct of the epididymis
cence to 5 = sexual maturity) is used as a framework on (ductus deferens) passes from the epididymis through the
which to objectively classify the development of secondary inguinal canal into the pelvic cavity. The ductus deferens
sexual characteristics. traverses the prostate gland where it joins the urethra. The
In females, the appearance of breast buds is the start of prostate gland secretes prostatic fluid into the semen, which
SMR 2, the first indication of the onset of puberty, and typi- supports transportation and nutrition of the sperm. Paired
cally occurs between the ages of 8 and 12 years. Simultane- seminal vesicles and the bulbourethral glands provide ad-
ously, the labia, uterus and ovaries increase in size, and the ditional secretion to the semen.
4
CHAPTER 2 — Reproductive Organs and Gametogenesis 5
Fig. 2.1 Sagittal section of the male pelvic region. (From Moore KL, Persaud TVN, & Torchia MG. The Developing Human: Clinically Oriented Em-
bryology. 10th ed. Philadelphia: Elsevier; 2015.)
(internal), myometrium (middle muscular) and perime-

FEMALE REPRODUCTIVE ORGANS trium (external). The endometrium can be further distin-
guished into the compact, spongy and basal layers, and var-
The vagina (Fig. 2.2) is a fibromuscular organ that extends ies in thickness according to stages of the menstrual cycle.
from the external genitalia (vulvar structures) to the cervix The uterine tubes are continuous with the uterine horns
of the uterus. The opening of the vagina is situated posterior found at the superior end of the uterus, the fundus. The
to the opening of the urethra and is covered by the labia uterine tubes are approximately 10-cm long, and consist
minora. The uterus is a thick-walled muscular organ con- of four parts: infundibulum, ampulla, isthmus and uterine
sisting of the body (upper two-thirds) and the cervix (lower part. The tubes are lined with cilia that help to propel the
one-third). The cervix is cylindrical with constricted open- ovum and sperm, first to the site of fertilisation (ampulla)
ing at both ends, the internal and external os. The body of and then to assist in moving the cleaving zygote to the uterus
the uterus is comprised of three tissue layers, endometrium for implantation.
Fig. 2.2 Sagittal section of the female pelvic region. (From Moore KL, Persaud TVN, & Torchia, MG. Before We Are Born: Essentials of Embryology
and Birth Defects. 9th ed. Philadelphia: Elsevier; 2016.)
6 SECTION 1 — General Development of the Embryo and Fetus
The ovaries are oval-shaped glands adjacent to the uterus

and the uterine tube infundibulum, with its finger-like fim- GAMETOGENESIS
briae. The ovaries produce the oocytes, as well as hormones
(oestrogen and progesterone) that regulate the process of Gametogenesis (oogenesis and spermatogenesis) (Fig. 2.3)
sexual development, menstruation and pregnancy. is the process that produces oocytes and sperms from bipo-
The external female genitalia consist of the labia minora, tential primordial germ cells and prepares these gametes for
labia majora and the clitoris. fertilisation. The sperm and oocyte are highly specialised
Fig. 2.3 Simplified diagram showing normal gametogenesis. (From Moore KL, Persaud TVN, & Torchia MG. The Developing Human: Clinically
Oriented Embryology. 10th ed. Philadelphia: Elsevier; 2015.)
sex cells, each of which contains the haploid number of containing several enzymes and other factors which, when
chromosomes that are present in somatic cells. The number released, facilitate dispersion of the follicular cells of the co-
of chromosomes is reduced during meiosis, a special type of rona radiata and sperm penetration of the zona pellucida
cell division that occurs only during gametogenesis. Meiosis during fertilisation. The tail has four segments: the connect-
involves two meiotic cell divisions resulting in diploid germ ing, middle, principal and end pieces, and it provides the
cells giving rise to haploid gametes. motility of the sperm for transport to the site of fertilisation.
The first meiotic division is a reduction division because The axoneme is the motility machinery of the sperm and is
the chromosome number is reduced to haploid by pairing comprised of cytoskeleton and dyneins (ATPase molecular
of homologous chromosomes in prophase and their seg- motors). The helically arranged mitochondria in the middle
regation at anaphase with one representative of each pair piece provide the energy required for motility. Sperm travel
randomly going to each pole of the meiotic spindle. At this at approximately 3 mm/min.
stage, the chromosomes are double-chromatid chromo- Spermatogonia (primordial male germ cells) are dor-
somes. (The X and Y chromosomes are not homologues, mant in the seminiferous tubules of the testes during the
but they have homologous segments at the tips of their fetal and postnatal periods. At puberty, spermatogenesis
short arms and pair in these regions only.) This disjunction begins, a 2-month highly complex process that transforms
of paired homologous chromosomes is the physical basis of spermatogonia into mature sperms. More than one dozen
segregation, the separation of allelic genes during meiosis. different subtypes of male germ cells have been identified.
The second meiotic division does not have an interphase, There are also a number of cells and factors within the tes-
but each double-chromatid chromosome divides, and each tes involved in sperm development. Peritubular myoid cells
half, or chromatid, is drawn to a different pole. Thus the are found surrounding and supporting the seminiferous tu-
haploid number of chromosomes remains, and each daugh- bules and are thought to regulate Sertoli cells, assist in man-
ter cell has one representative of each chromosome pair aging the blood–testis barrier (an important controller of
(now a single-chromatid chromosome). The process of mei- the germ cell microenvironment) and push testicular fluid
osis provides constancy of the chromosome number from with sperm towards the rete testis. Leydig cells (LCs) are
generation to generation, allows random assortment of ma- found clustered near seminiferous tubules and the adjacent
ternal and paternal chromosomes between the gametes and blood vessels. LCs produce testosterone, which is released
relocates segments of maternal and paternal chromosomes into the systemic circulation. LCs ensure a much higher lo-
by crossing over of chromosome segments, which produces cal concentration of testosterone, which is required for nor-
a recombination of genetic material. mal sperm production. LCs also produce oestradiol from
testosterone, which appears to be required for successful
spermatogenesis. Sertoli cells (SCs) make up approximately
SPERM CHARACTERISTICS AND 20% of the epithelial cells of the seminiferous tubules. The
DEVELOPMENT role of the SCs is complex and broad. Their unique struc-
ture allows each SC to shepherd up to 50 germ cells during
Sperms are highly differentiated, actively motile cells con- differentiation; this is accomplished by sophisticated cyto-
sisting of a head and a tail (Fig. 2.4) and approximately skeletal elements. SCs produce anti-Müllerian hormone,
4 µm in length. The head forms most of the bulk of the critical to the normal embryological develop of male and
sperm and contains the nucleus. The anterior two-thirds of female reproductive organs. SCs also act as macrophages,
the head is covered by the acrosome, a saccular organelle and produce inhibin B (regulating FSH production) and
androgen-binding protein.
The male germ cells are arranged in the seminiferous
tubules in a specific manner with least-mature cells in the
basal compartment and more-mature cells found adjacent
to the lumen.
The earliest germ cells in the testes (gonocytes) remain
in G0 phase of the cell cycle until after birth. In the first
few neonatal months, they are transformed into inactive
spermatogonia which begin to undergo rapid mitosis at ap-
proximately 6 years of age. Later, at puberty, the spermato-
gonia undergo the process of spermatogenesis. Briefly,
spermatogonia first develop into primary spermatocytes,
the largest germ cells in the seminiferous tubules of the
testes. Each primary spermatocyte subsequently undergoes
the first meiotic division to form two haploid secondary
spermatocytes. These secondary spermatocytes undergo
the second meiotic division and form four haploid sper-
matids. The spermatids are gradually transformed into
four mature sperms by a process known as spermiogenesis.
When spermiogenesis is complete, the sperms enter the lu-
Fig. 2.4 Main parts of the human sperm. (From Moore KL, Persaud mina of the seminiferous tubules. Sperms are transported
TVN, & Torchia MG. The Developing Human: Clinically Oriented Embryol- passively from the seminiferous tubules to the epididymis,
ogy. 10th ed. Philadelphia: Elsevier; 2015; Fig. 2.5.) where they are stored.
progresses only to metaphase. If a sperm penetrates the sec-

OOCYTE CHARACTERISTICS AND ondary oocyte at fertilisation, the second meiotic division
DEVELOPMENT is completed, and most cytoplasm is again retained by one
cell, the fertilised oocyte. The second polar body is formed
The mature (secondary) oocyte (Fig. 2.5) is an immotile cell and will degenerate.
with a diameter of approximately 100 µm, making it one of There are approximately 2 million primary oocytes in the
the largest cells in the female human body, and just visible ovaries of a neonate, but most of them regress during child-
to the unaided eye. It typically contains a transparent mod- hood so that by adolescence no more than about 40,000
erately granular cytoplasm with refractile structures such as primary oocytes remain. Of these, only approximately 400
lipid, lipofuscin bodies, and autophagic vacuoles. A single become secondary oocytes and are expelled at ovulation
polar body is also associated with the secondary oocyte (see during the reproductive period. Very few of these oocytes, if
later). any, are fertilised. The long duration of the first meiotic divi-
Oogenesis transforms oogonia (primordial female germ sion (up to 45 years) may account in part for the relatively
cells) into mature oocytes. All oogonia develop prenatally, high frequency of meiotic errors that occur with increasing
and the process of oogenesis ceases following menopause. maternal age.
In early fetal life, oogonia proliferate by mitosis and enlarge
to form primary oocytes, each surrounded by a single layer
of flattened, connective tissue follicular cells. The primary
oocyte enclosed by this layer of follicular cells constitutes
FEMALE REPRODUCTIVE CYCLE
a primordial follicle. As the primary oocyte enlarges dur-
The female reproductive cycle (Fig. 2.6) is highly complex
ing puberty, the follicular epithelial cells become columnar
and involves activities of the hypothalamus, pituitary gland,
shaped and the oocyte becomes covered with the glycopro-
ovaries, uterus, uterine tubes, vagina and mammary glands,
teinaceous zona pellucida. Primary oocytes begin the first
all towards preparation of the reproductive system for preg-
meiotic divisions before birth, but completion of prophase
nancy.
does not occur until puberty. The follicular cells surround-
GnRH secreted by the hypothalamus stimulates the an-
ing the primary oocytes secrete a substance, oocyte matura-
terior lobe of the pituitary gland to release FSH, which
tion inhibitor, which keeps the meiotic process of the oo-
stimulates the development of ovarian follicles and the pro-
cyte arrested. With puberty, the ovarian follicle (typically
duction of oestrogen by the follicular cells, and LH, which
only one) matures each month. As a follicle matures, the
triggers ovulation and stimulates follicular cells and corpus
primary oocyte increases in size and shortly before ovula-
luteum to produce progesterone and causes growth of the
tion it completes the first meiotic division to give rise to a
follicles and endometrium.
secondary oocyte and the first polar body. The secondary
As the primary follicle increases in size, the adjacent con-
oocyte receives almost all the cytoplasm; the polar body is
nective tissue organises into a capsule, the theca folliculi.
destined for degeneration. At ovulation, the nucleus of the
This theca soon differentiates an internal vascular and glan-
secondary oocyte begins the second meiotic division, but it
dular layer (theca interna) and a capsule-like layer (theca
externa). The follicular cells produce a stratified layer
around the oocyte. Fluid-filled spaces appear around the
follicular cells, which coalesce to form the antrum, contain-
ing follicular fluid at this stage; the ovarian follicle is then
called a secondary follicle. The primary oocyte is pushed
to one side of the follicle. At approximately the midpoint
of the cycle, FHS and LH stimulation cause rapid follicle
growth leading to the formation of a small avascular spot,
follicular stigma, on the surface of the ovary. Rupture of the
stigma and expulsion of the secondary oocyte (ovulation)
occurs 12 to 24 hours after this surge of LH production. The
expelled secondary oocyte is surrounded by the zona pel-
lucida and one or more layers of radially arranged follicular
cells (corona radiata). Shortly after ovulation, the walls of
the ovarian follicle and theca folliculi collapse and develop
into the corpus luteum. The corpus luteum secretes proges-
terone and some oestrogen, causing the endometrial glands
to secrete and prepare the endometrium for implantation
of the blastocyst. If the oocyte is fertilised, the corpus lu-
teum enlarges to form a corpus luteum of pregnancy and
increases its hormone production. The corpus luteum of
pregnancy remains functionally active throughout about
the first 20 weeks of pregnancy. By this time, the placenta
has assumed the production of oestrogen and progesterone
Fig. 2.5 Photomicrograph of a human oocyte. (From Zhang P, Zuc- necessary for the maintenance of pregnancy. If the oocyte is
chelli M, Bruce S, et al. Transcriptome profiling of human pre-implan- not fertilised, the corpus luteum involutes and degenerates
tation development. PLoS One 2009; 4(11): e7844. With permission.) 10 to 12 days after ovulation.
Fig. 2.6 Schematic drawing of the ovarian and menstrual cycles. (From Moore KL, Persaud TVN, & Torchia MG. The Developing Human: Clinically
Oriented Embryology. 10th ed. Philadelphia: Elsevier; 2015.)
MENSTRUAL CYCLE by the corpus luteum stimulates the glandular epithelium

to secrete a glycogen-rich material. The glands become
Changes in the oestrogen and progesterone levels cause cy- wide, tortuous and saccular, and the endometrium thickens
clic changes in the structure of the female reproductive tract, because of the influence of progesterone and oestrogen
notably the uterine endometrium (Fig. 2.6). The menstrual from the corpus luteum. As the spiral arteries grow into the
cycle is a continuous process lasting on average 28 days, with superficial compact layer, they become increasingly coiled.
each phase gradually passing into the next. Day 1 is desig- The venous network becomes complex, and large lacunae
nated as the day menstrual flow begins. In the menstrual (venous spaces) develop in the endometrium. If fertilisation
phase (4–5 days), the functional layer of the endometrium does not occur the corpus luteum degenerates, oestrogen
is sloughed off as the menses (blood discharged from the va- and progesterone levels fall, and the secretory endome-
gina combined with small pieces of endometrial tissue). After trium enters an ischemic phase. The spiral arteries con-
menstruation, the remaining endometrium is thin. strict, glandular secretion stops, interstitial fluid is reduced,
During the proliferative phase (9 days), the ovarian fol- endometrium shrinks and venous stasis occurs. This leads
licles grow and the uterine surface epithelium reforms to patchy ischaemic necrosis of the functional layer of the
and covers the endometrium. The uterine glands increase endometrium. Rupture of damaged vessel walls allows blood
in number and length and endometrial spiral arteries to leak into the surrounding connective tissue, resulting in
elongate. The luteal (secretory) phase lasts approximately bleeding (typical loss of 20–80 mL).
13 days and coincides with the formation, functioning and If fertilisation occurs, the zygote undergoes cleavage and
growth of the corpus luteum. The progesterone produced blastogenesis, and the blastocyst begins to implant in the
steps!
endometrium (sixth day of the luteal phase). The embryo implantation. No hormone-based contraceptive is avail-
syncytiotrophoblast produces chorionic gonadotropin, able for men. Barrier methods, including condoms (male
which keeps the corpora luteum secreting oestrogens and or female) and contraceptive diaphragms, prevent sperm
progesterone; the luteal phase continues and menstruation from entering the vagina or uterus, respectively. Sterilisation
does not occur. (implant, vasectomy or tubal ligation) are permanent forms
of birth control.
Molecular Biology Considerations NONDISJUNCTION

• PI3K/PTEN and TSC/mTOR pathways—activation of pri- Nondisjunction is an error in cell division in which there is
mordial follicles failure of a chromosomal pair (autosome or sex chromo-
• GDF9 and BMP15—development of secondary and pre- some) to separate during mitosis or meiosis, resulting in
ovulatory follicle numeric aberrations of chromosomes. Nondisjunction may
• cAMP meiotic arrest occur during maternal or paternal gametogenesis, resulting
• MAPK3/1—ovulation control in some gametes having 24 chromosomes while others have
• bFGF—maintenance of blood–testis barrier only 22. If these gametes should become fertilised with a
• TGF-a/b and GNDF—maintenance of spermatogenesis normal gamete, a zygote with either trisomy (three copies of
microenvironment
a chromosome) or monosomy (one copy of a chromosome)
• PModS—regulates Sertoli cell function
• HOX—shaping of sperm head results. Clinical conditions resulting from such nondis-
junction include trisomy 21 (Down syndrome), trisomy 18
(Edwards syndrome), XXY trisomy (Klinefelter syndrome)
and monosomy X (Turner syndrome).
CLINICAL ISSUES Case Outcome
FERTILITY KR was sent for blood tests as well as endocrine and gyn-
aecological consultations. Higher than normal levels of andro-
In 85% to 90% of cases, heterosexual couples are able to gens were detected in her blood; there were no other abnormal
achieve pregnancy through sexual intercourse. In the re- findings. A pelvic examination was normal. A pelvic ultrasound
maining couples, fertility issues for both the male and fe- demonstrated multiple cysts on her ovaries (see Fig. 2.7). She
male require investigation; these male/female concerns was diagnosed with polycystic ovarian syndrome (PCOS) and
returned to her family physician for discussion of treatment op-
often coexist. In men, the most common causes of reduced
tions and follow-up.
fertility are a blockage of sperm delivery, altered sperm mor- Additional reflection: Did KR present with the typi-
phology, motility and function and reduced sperm numbers. cal signs for PCOS? Given KR’s desire for children, how
Previous infection, retrograde ejaculation, prior trauma and might this be a consideration for her long-term treatment
tumours are examples of causes of blocked semen flow. Ab- of PCOS? What is the likelihood of KR conceiving a child,
normal sperm morphology includes large or double heads should her husband’s fertility prove to be normal? What
and bent or double tails; causes include genetic disorders, might be some psychological implications of PCOS?
exposure to environmental toxins or high testicular tem-
peratures. Men with fewer than 10 million sperms per milli-
litre of semen are less likely to be fertile, especially when the
specimen contains immotile and abnormal sperms. Envi-
ronmental factors (drug or alcohol abuse, exposure to envi-
ronmental toxins), medication and hormone imbalance are
only a few of the reasons that low sperm counts may occur.
In women, the most common causes of infertility are
blockage of oocyte transportation attributed to tubal scar-
ring or endometriosis, reduced production of oocytes be-
cause of increased age, and hormonal imbalances such as
from polycystic ovarian syndrome (PCOS) and obesity.
CONTRACEPTION
The use of hormonal methods of female contraception can
result in some or all of thickened cervical mucus, alteration
of the endometrium, prevention of ovulation or blockage
of sperm. These contraceptives include progestin-only pills,
combined oestrogen–progestin pills, emergency contracep- Fig. 2.7 Pelvic ultrasound demonstrating cystic structures on
tive pills, vaginal rings, contraceptive patches and inject- the oval in polycystic ovarium syndrome. (From Karakas SE. New
able long-acting medications including drug-integrated biomarkers for diagnosis and management of polycystic ovary syn-
implants. Intrauterine devices may contain either hormones drome. Clin Chim Acta 2017; 471: 248–253.With permission.)
or copper and prevent sperm from reaching the ovum or
QUESTIONS
1. Which of the following types of germ cell does not un- c. disturbances in spermiogenesis
dergo cell division? d. disturbances in mitosis
a. spermatogonia e. abnormal spermatogonia
b. primary oocytes
c. spermatids BIBLIOGRAPHY
d. secondary spermatocytes Datta J, Palmer MJ, Tanton C, et al. Prevalence of infertility and help seek-
e. oogonia ing among 15,000 women and men. Hum Reprod 2016;31:2108–18.
Neto FTL, Bach PV, Najari BB, Li PS, Goldstein M. Spermatogenesis in
2. An infant is diagnosed as having 47 chromosomes instead humans and its affecting factors. Sem Cell Dev Biol 2016;59:10–26.
Pasquali R. Contemporary approaches to the management of polycystic
of 46. This abnormal condition (trisomy) results from:
ovary syndrome. Ther Adv Endocrinol Metb 2018;9(4):123–34.
a. gene mutation
b. nondisjunction
3 Fertilisation and Reproductive
Technologies
genital tract. Oocytes are usually fertilised within 12 hours

Case Scenario of ovulation, and it appears that they cannot be fertilised
after 24 hours (see Video 3.1).
A 49-year-old woman (PG) presents to her family physician
The usual site of fertilisation is in the ampulla of the uter-
reporting a positive home pregnancy test and claims to be
approximately 2-months pregnant based on timing of her last ine tube. If the oocyte is not fertilised, it slowly passes along
normal menstrual period. She is concerned because this is her the tube to the body of the uterus, where it degenerates and
first pregnancy; she and her partner have not used birth control is resorbed.
for the past three months because her menstrual periods have Fertilisation is a sequence of coordinated events (Fig. 3.2),
been very irregular for the past year and she believed that she beginning with the passage of a sperm through the corona
was ‘in menopause and not infertile’. radiata. Hyaluronidase released from the sperm acrosome,
A second pregnancy test was ordered which was positive. tubal mucosa enzymes and sperm motion appear to cause
An ultrasound (endovaginal sonogram) showed a live embryo dispersal of the follicular cells of the corona radiata. Passage
with crown–rump length of approximately 9 mm, aged between of a sperm through the zona pellucida is the next phase and
35 and 38 days.
also results from the action of enzymes released from the
Given PG’s age, she was counselled regarding the options
for (or no) prenatal screening for fetal aneuploidies. acrosome, including acrosin, esterase and neuraminidase.
Questions for reflection: Why is PG’s age a concern and Once a sperm penetrates the zona pellucida, a change in the
how may age impact normal gametogenesis. What types properties of the zona pellucida (zona reaction) occurs that
of prenatal screening are available and at what gestational makes it impermeable to other sperms. The zona reaction is
age? Which of the tests is considered diagnostic? What believed to result from the action of lysosomal enzymes re-
are the most common aneuploidies? leased by cortical granules near the plasma membrane of the
oocyte. The contents of these granules also cause changes in
the plasma membrane that make it impermeable to other
sperms. Fusion and localised breakdown of cell membranes
of the oocyte and sperm occurs next, resulting in the head
FERTILISATION and tail of the sperm entering the cytoplasm of the oocyte
(the cell membrane and mitochondria of the sperm remain
During ovulation, the fimbriated end of the uterine tube be- behind). Penetration of the oocyte by a sperm activates the
comes closely applied to the surface of the ovary. The sweep- oocyte into completing the second meiotic division and
ing action of the tube and of fluid currents produced by the forming a mature oocyte and a second polar body. The ma-
ciliated mucosal cells of the fimbriae, causes the extruded ternal chromosomes decondense and the nucleus of the ma-
oocyte to enter the infundibulum of the uterine tube. The ture oocyte becomes the female pronucleus.
oocyte then passes into the ampulla of the tube, mainly as The nucleus of the sperm enlarges to form the male pro-
the result of tube peristalsis. During sexual intercourse and nucleus and the tail of the sperm degenerates. Both pronu-
ejaculation, sperms are rapidly transported from the epidid- clei duplicate their DNA and the oocyte becomes an ootid.
ymis to the urethra by peristaltic contractions of the thick When the pronuclei fuse into a single diploid aggregation
muscular coat of the ductus deferens. Between 200 and 600 of chromosomes, the ootid becomes a zygote. The chromo-
million sperms are deposited in the vagina, around the ex- somes in the zygote become arranged on a cleavage spin-
ternal os and the fornix, and then some pass through the dle in preparation for cleavage of the zygote. The zygote is
cervical canal. The cervical mucus increases in amount and genetically unique.
becomes less viscid during ovulation, making it more favour-
able for sperm passage. Approximately 200 sperms reach
the ampulla of the uterine tube; the remainder degenerate
and are absorbed in the female genital tract. Sperms must CLEAVAGE OF THE ZYGOTE AND
undergo capacitation, lasting approximately 7 hours, before FORMATION OF THE BLASTOCYST
they are able to fertilise the oocyte. During this process, a
glycoprotein coat and seminal proteins are removed from Cleavage occurs approximately 30 hours after fertilisation
the surface of the sperm acrosome and the membrane com- as the zygote, within the zona pellucida, passes along the
ponents of the sperms are extensively altered. Sperms are uterine tube towards the uterus. Cleavage consists of re-
usually capacitated while they are in the uterus or uterine peated mitotic divisions of the zygote, resulting in a rapid
tubes by substances secreted by these parts of the female increase in the number of cells (blastomeres) and decrease
12
CHAPTER 3 — Fertilisation and Reproductive Technologies 13
Fig. 3.1 Timeline of development related to fertilisation.
Fig. 3.2 Events taking place in fertilisation. (From Moore KL, Persaud TVN, & Torchia MG The Developing Human: Clinically Oriented Embryology.
10th ed. Philadelphia: Elsevier; 2015.)
in the size of subsequent blastomeres with each successive After the blastocyst has floated in the uterine secretions
cleavage division. After the nine-cell stage, the blastomeres for approximately 2 days, shedding of the zona pellucida
undergo compaction, changing their shape and tightly occurs, permitting the blastocyst to increase rapidly in size.
aligning themselves against each other to form a compact While in the uterus, the embryo derives nourishment from
ball of cells. Compaction changes the cell cytoskeleton, per- secretions of the uterine glands. At approximately 6 days, the
mitting greater cell-to-cell interaction. Polarisation of the blastocyst (usually adjacent to the embryonic pole) attaches
blastomeres into apical and basolateral domains also takes to the endometrial epithelium. The trophoblast proliferates
place. Compaction is necessary for segregation of the inter- rapidly and differentiates into two layers—an inner layer
nal cells that will form the embryoblast (inner cell mass) of of cytotrophoblast that is mitotically active and forms new
the blastocyst from surrounding cells that form the tropho- mononuclear cells that migrate into the increasing mass of
blast (Fig. 3.3). At the 12- to 32-blastomeres stage, the devel- syncytiotrophoblast, and an outer layer of syncytiotropho-
oping embryo is called a morula. Shortly after the morula blast (multinucleated protoplasmic mass) (Fig. 3.4). The
enters the uterus (approximately 4 days postfertilisation), syncytiotrophoblast begins to invade the uterine connective
the fluid-filled blastocystic cavity appears inside the morula tissue so that the blastocyst can now derive its nourishment
separating the blastomeres into the trophoblast (thin outer from the eroded maternal tissues. Endometrial cells assist
cell layer giving rise to the embryonic part of the placenta) to control the depth of penetration of the syncytiotropho-
and the embryoblast (centrally located blastomeres which blast. At approximately 7 days, a layer of cells, the hypoblast
form the embryo). Early pregnancy factor (EPF), an immu- (primary endoderm), appears on the surface of the embryo-
nosuppressant protein, is secreted by the trophoblastic cells blast facing the blastocystic cavity. Comparative embryologi-
and aids in the prevention of early maternal immune attack cal data suggest that the hypoblast arises by delamination of
of the embryo (see Video 3.2). blastomeres from the embryoblast.
Fig. 3.3 Stages of development during the first week. (A) Ovulated oocyte; (B) fertilisation; (C) pronuclei formation; (D) first cleavage spindle; (E–G)
cleavage of zygote; (H) morula; (I) blastocyst. (From Mitchell B, Sharma R. Embryology: An Illustrated Colour Text. 2nd ed. London: Elsevier; 2009.)
many women who are sterile to have children. Since 1978,

when Robert G. Edwards and Patrick Steptoe pioneered IVF,
several million children have been born following an IVF pro-
cedure. The steps involved during IVF and embryo transfer
are briefly noted. Beginning on day 1 of the menstrual cycle,
ovarian follicles are stimulated to grow and mature (super-
ovulation), typically by the administration of a drug that in-
creases follicle-stimulating hormone (FSH) and/or luteinis-
ing hormone (LH) secretion by the pituitary. At the optimal
time (often determined by ultrasound), another medication
(synthetic human chorionic gonadotropin [hCG]) is given to
trigger ovulation. Using an ultrasonically guided, minimally
invasive procedure, several mature oocytes (typically 8–15)
are aspirated from mature ovarian follicles. The oocytes are
then placed in a Petri dish containing a special culture me-
Fig. 3.4 A 7-day blastocyst beginning to implant. (From Mitchell B, dium and capacitated sperms. Fertilisation of the oocytes and
Sharma R. Embryology: An Illustrated Colour Text. 2nd ed. London: cleavage of the zygotes are monitored by microscope for 3
Elsevier; 2009.) to 5 days. Depending on the mother’s age, one to three of
the resulting embryos (four-cell to eight-cell stage, or early
blastocysts) are transferred by introducing a catheter through
the vagina and cervical canal into the uterus. Any remaining
Molecular Biology Considerations embryos are frozen for later use. Approximately 2 weeks later,
a pregnancy test is performed.
• SPAM1, HYAL5, ACE3, ADAMS1–3—gamete fusion
• Hippo—segregation of embryoblast from trophoblast CRYOPRESERVATION OF EMBRYOS
• TGF-b—proliferation and differentiation of the trophoblast
Early embryos resulting from IVF can be preserved for long
periods by freezing them in liquid nitrogen with a cryopro-
tectant (e.g., glycerol or dimethyl sulfoxide). Successful
transfer of four- to eight-cell embryos and blastocysts to the
CLINICAL ISSUES uterus after thawing is now a common practice. The longest
period of sperm cryopreservation that resulted in a live birth
ASSISTED REPRODUCTIVE TECHNOLOGIES was reported to be 21 years.
IN VITRO FERTILISATION AND EMBRYO TRANSFER INTRACYTOPLASMIC SPERM INJECTION

In vitro fertilisation (IVF) of oocytes and transfer of cleav- A sperm can be injected directly into the cytoplasm of a ma-
ing zygotes into the uterus have provided an opportunity for ture oocyte. This technique has been successfully used for
the treatment of couples in whom typical IVF has failed, or

in cases where there are too few sperms available.
ASSISTED IN VIVO FERTILISATION

Gamete intrafallopian (intratubal) transfer enables fertilisa-
tion to occur in the uterine tube. It involves superovulation
(similar to that used for IVF), oocyte retrieval, sperm col-
lection and laparoscopic placement of several oocytes and
sperms into the uterine tubes. Using this technique, fertilisa-
tion occurs in the ampulla, its usual location.
SURROGATE MOTHERS
Some women produce mature oocytes but are unable to
become pregnant, for example, a woman who has had a
hysterectomy. In these cases, IVF may be performed, and
the embryos transferred to another woman’s uterus for fetal
development and delivery.
Fig. 3.5 Three-dimensional ultrasound image of a fetus with trisomy
21, showing characteristic features including a protruding tongue
PREGNANCY TESTING (macroglossia).
Most pregnancy tests are based on the detection or measure-

ment of human chorionic gonadotropin (hCG) produced nose and eyes, holoprosencephaly, polydactyly, skin defects;
by the syncytiotrophoblast. hCG can be measured in urine trisomy 18—includes clenched hands, prominent occiput,
or in blood. hCG rapidly increases in concentration from short sternum, small pelvis, cryptorchidism) and have ma-
the time of early implantation (approximately day 6–12). jor neurodevelopmental disabilities. Infants with these life-
The blood testing performed by clinical laboratories uses limiting disorders (trisomy 18 and 23) have a 1-year survival
a more sensitive assay, and it also measures hCG concentra- rate of approximately 6% to 12%. Infants with trisomy 21
tion which can be followed over time depending on clinical (Fig. 3.5) have abnormalities that include cognitive defi-
needs. There are a number of biological and pharmacologi- ciency, hypotonia, bradycephaly, upward slanting palpebral
cal factors that can produce false positive and negative hCG fissures, protruding tongue, small ears, heart defects such as
test results including heterotrophic antibodies, rheumatoid endocardial defects, ventricular septal defect or atrial septal
factors and ectopic pregnancies. defect, and a transverse palmar flexion crease. More than
Transvaginal ultrasonography can detect the gestational one-half of trisomic embryos spontaneously abort early. Tri-
sac at approximately 2.5 to 3 weeks following conception. somy of the autosomes occurs with increasing frequency
as maternal age increases. For example, trisomy 21 occurs
ANEUPLOIDY once in approximately 1400 births among mothers between
the ages of 20 and 24 years but once in approximately 25
Aneuploidy, usually resulting from nondisjunction, is any births among mothers 45 years and older. Because of the
deviation from the diploid number of 46 chromosomes and current trend of increasing maternal age, it has been esti-
is the most common (3%–4% of pregnancies) and clinically mated that children born to women older than 34 years will
significant numeric chromosomal abnormalities. account for 39% of infants with trisomy 21.
MONOSOMY X TRISOMY OF SEX CHROMOSOMES
The incidence of Turner syndrome (45,X) is about 1:8000 These disorders are not usually detected until puberty be-
live births. Only 1% of monosomy X female embryos survive, cause there are no characteristic physical findings in in-
with 45,X being the most common abnormality detected in fants or children. These disorders include XYY syndrome
all spontaneous abortions. When it is possible to trace, it is (1:1000; tall stature, cognitive disabilities, severe acne, au-
the paternal X chromosome that is missing in approximately tism spectrum disorder, normal fertility), XXX (1:1000;
75% of cases. In some cases, mosaicism occurs (XX/X and normal puberty, normal fertility, some degree of cognitive
XY/X mosaics) and in these cases there is a lesser degree deficiency can occur); and XXY syndrome (Klinefelter syn-
of abnormalities. The abnormalities typically seen with 45,X drome) (1:500; most common cause of hypogonadism and
include small stature, ovarian dysgenesis, broad chest with infertility, gynaecomastia, inadequate virilisation, long limbs
wide-spaced nipples, congenital lymphedema and a short and possible developmental delay).
and/or webbed neck.
AUTOSOMAL TRISOMY MOSAICISM

Trisomy is the most common aneuploidy. Trisomy of auto- A person with at least two cell lines with two or more geno-
somes is mainly associated with three syndromes: trisomy 18 types is considered a mosaic. The autosomes or sex chromo-
(Edwards syndrome, 1.3:10,000 live births), trisomy 13 (Pa- somes may be involved. The defects usually are less serious
tau syndrome, 0.8:10,000) and trisomy 21 (Down syndrome, than in persons with monosomy or trisomy. For instance,
12:10,000). Infants with trisomy 13 and trisomy 18 are se- the features of Turner syndrome are not as evident in
verely malformed (trisomy 13—includes defects of the lip, 45,X/46,XX mosaic females as in the usual 45,X females.
Although mosaicism usually results from nondisjunction, it genotype of the embryo and allow selection of a chromo-
can also occur through the loss of a chromosome by ana- somally healthy embryo for transfer. Preimplantation genet-
phase lagging; chromosomes separate normally, but one of ic diagnosis can be carried out 3 to 5 days after IVF of the
them is delayed in its migration and is eventually lost. oocyte. One or two cells (blastomeres) are removed from
the embryo and these cells are then analysed before transfer
into the uterus. The sex of the embryo can also be deter-
MULTIPLE GESTATIONS mined from one blastomere taken from a six- to eight-cell
dividing zygote, and analysed by polymerase chain reaction
In North America, twins naturally occur approximately once
and fluorescence in situ hybridisation techniques. This pro-
in every 85 pregnancies, triplets approximately once in 902
cedure has been used to detect female embryos during IVF
pregnancies, quadruplets once in 903 pregnancies and quin-
in cases in which a male embryo would be at risk of a serious
tuplets approximately once in every 904 pregnancies. Twins
X-linked disorder. The polar body may also be tested for dis-
that originate from two zygotes are dizygotic (DZ) twins
eases where the mother is the carrier (Fig. 2.15A).
whereas twins that originate from one zygote are monozy-
gotic (MZ) twins. Two-thirds of twins are DZ, with marked
racial differences whereas the incidence of MZ twinning is
approximately the same in all populations. DZ twins may be
of the same sex or different sexes and are no more alike Case Outcome
genetically than brothers or sisters born at different times.
Patient PG opted for noninvasive prenatal testing (NIPT)
The fetal membranes and placentas vary according to the
through cell-free DNA (cfDNA) screening. This testing was con-
origin of the twins. DZ twins always have two amnions and ducted approximately 3 weeks after her previous visit (8 weeks
two chorions, but the chorions and placentas may be fused. post conception—10 weeks gestational age). The test results
Anastomoses between blood vessels of fused placentas of demonstrated a high risk for trisomy 21 (Down syndrome).
DZ twins may result in erythrocyte mosaicism. MZ twins are A diagnostic chorionic villus sampling (CVS) was then per-
genetically identical; physical differences between MZ twins formed that confirmed the diagnosis of trisomy 21. PG opted to
are caused by environmental differences, chance variation continue the pregnancy. A later fetal ultrasound demonstrated
and uneven X-chromosome activation. MZ twinning usually enhanced nuchal translucency (Fig. 3.6), but no cardiovascu-
results from division of the embryoblast into two embryonic lar anomalies. The remainder of the pregnancy was uneventful,
primordia, with each embryo in its own amniotic sac but and PG delivered a baby girl at 38 weeks.
Additional reflection: What is the difference between
sharing the same chorionic sac and placenta (monochori-
a screening test and a diagnostic test? How is CVS per-
onic–diamniotic twin). Uncommonly, early separation of formed, at what gestational age, and with what possible
embryonic blastomeres (e.g., during the two-cell to eight- risks to fetus and the mother? What is a nuchal translu-
cell stages) results in MZ twins with two amnions, two cho- cency, and why was there a concern about cardiovascular
rions and two placentas that may or may not be fused. Twin anomalies?
transfusion syndrome occurs in as many as 10% to 15% of
monochorionic–diamniotic MZ twins. There is shunting of
arterial blood from one twin through unidirectional um-
bilical–placental arteriovenous anastomoses into the venous
circulation of the other twin. The donor twin is small, pale
and anaemic whereas the recipient twin is large and has
polycythaemia. In lethal cases, death results from anaemia
in the donor twin and congestive heart failure in the recipi-
ent twin. Late division of early embryonic cells, such as divi-
sion of the embryonic disc during the second week, results
in MZ twins that are in one amniotic sac and one chorionic
sac. A monochorionic–monoamniotic twin placenta is as-
sociated with fetal mortality rates that are higher by up to
10%, with the cause being cord entanglement. Because ul-
trasonographic studies are a common part of prenatal care,
it is known that early death and resorption of one member
of a twin pair is common. Triplets may be derived from one
zygote and be identical, two zygotes and consist of identical
twins and a singleton or three zygotes and be of the same sex
or of different sexes. The determination of twin zygosity is
done by molecular diagnosis.
PREIMPLANTATION GENETICS Fig. 3.6 Ultrasound of a fetus demonstrating an enhanced

nuchal translucency (arrow).
In couples with inherited genetic disorders and using
IVF, preimplantation genetic diagnosis can determine the
QUESTIONS
1. How many sperms would probably be deposited by a nor- c. neck

mal young adult male in the vagina during intercourse: d. main piece
a. 300,000 e. head
b. 3 million
c. 30 million 4. Morphologically abnormal sperm may cause:
d. 300 million a. monosomy
e. 3 billion b. congenital anomalies
c. trisomy
2. The secondary oocyte completes the second meiotic divi- d. abnormal embryos
sion: e. infertility
a. before ovulation
b. during ovulation BIBLIOGRAPHY
c. at fertilisation Jelin AC, Sagasser KG, Wilkins L. Prenatal genetic testing options. Pedi-
d. before birth atr Clin North Am 2019;66:281–93.
e. at puberty Bamberg C, Hecher K. Update on twin-to-twin transfusion syndrome.
Best Pract Res Clin Obstet Gynaecol 2019;58:55–65.
Katz DJ, Teloken P, Shoshany O. Male infertility – The other side of the
3. The sperm penetrates the zona pellucida, partially as-
equation. Aust Fam Physician 2017;46:641–6.
sisted by enzymes that are released from which portion
of the sperm:
a. middle piece
b. acrosome
4 Implantation and Week 2
endometrium. The window of implantation is relatively

Case Scenario brief, 2 to 3 days, and occurs during a restricted time pe-
riod, 6 to 10 days after ovulation and fertilisation—the mid-
A 32-year-old woman, having had two previous successful
pregnancies, presented to her family physician with unusually
secretory stage of the menstrual cycle. At this time, and
heavy menstruation. Her last normal menstrual period was ex- under the influence of progesterone and oestrogen, the
actly 2 months ago. She previously had a very regular cycle and cellular features of the endometrium are altered; pinopods
normal menstrual flow. She did not complain of any abdominal (microvilli) form on the epithelial cells, cellular vacuoles
pain. She has been on oral contraception for the past 3 years move towards the apical end of cells, the uterine connec-
and felt that she had been compliant. tive tissue is more oedematous, and some of the connec-
On physical examination, there was mild tenderness in her tive tissue cells accumulate large quantities of glycogen and
lower abdomen. Her blood pressure and heart rate were nor- lipids and expand in size, becoming the decidual cells (see
mal. Her haemoglobin level and white cell count were normal. Video 4.1).
Vaginal examination did not reveal any causes of bleeding al-
The syncytiotrophoblast and the blastocyst slowly implant
though her posterior fornix was tender on digital examination.
A pregnancy test was positive and an ultrasound examination
in the endometrium. Syncytiotrophoblastic cells displace
in the office revealed an empty uterine cavity and fluid present endometrial cells at the implantation site, reach the base-
in the rectouterine pouch (Fig. 4.1). ment membrane and then extend invadopodia between
She was sent immediately to the nearby hospital for further the cells, leading to degradation of the extracellular matrix.
care and treatment. This invasion allows the syncytiotrophoblast to reach the
Questions for reflection: How could the pregnancy test be vascular connective tissues and blood vessels. The syncy-
positive yet the woman is menstruating? Similarly, how could tiotrophoblast engulfs decidual cells to provide embryonic
she be pregnant yet the uterine cavity is empty? What kind nutrition. Some endometrial cells undergo apoptosis which
of fluid might be in the rectouterine pouch? What is your di- facilitates the invasion. Endometrial cell signalling also
agnosis and why? What further testing might be important?
helps to modulate the depth of penetration of the syncy-
tiotrophoblast. The blastocyst is completely embedded by
day 8 to 10. Initially the entry location is covered with fibrin
after which endometrial cells proliferate to cover the im-
plantation site.
Individual lacunae (Fig. 4.2) soon appear in the syn-
cytiotrophoblast. These become filled with a mixture of
maternal blood from the ruptured endometrial capillar-
ies and cellular debris of eroded uterine glands, provid-
ing nutritive material to the embryo. Communication of
the eroded endometrial capillaries with the lacunae in the
syncytiotrophoblast establishes the earliest uteroplacen-
tal circulation. Oxygenated blood passes into the lacunae
from the spiral endometrial arteries and poorly oxygenat-
ed blood is removed from them through the endometrial
veins.
In a 12-day embryo, adjacent syncytiotrophoblastic lacu-
nae have fused to form lacunar networks, the primordia
Fig. 4.1 Ultrasound image as per case.
of the intervillous spaces of the placenta. The endometrial
capillaries around the implanted embryo become congested
and dilated to form maternal sinusoids, thin-walled termi-
nal vessels that are larger than ordinary capillaries. The syn-
IMPLANTATION cytiotrophoblast erodes the sinusoids, and maternal blood
flows freely into the lacunar networks. The trophoblast
The mechanisms of implantation involve synchronisa- absorbs nutritive fluid from the lacunar networks, which is
tion between the invading blastocyst and a prepared transferred to the embryo.
18
CHAPTER 4 — Implantation and Week 2 19
Fig. 4.2 Timeline of development related to implantation and week 2 of development.
BILAMINAR EMBRYO
As implantation of the blastocyst progresses, the primordi-

um of the amniotic cavity appears in the embryoblast. Am-
niogenic cells, amnioblasts, separate from the epiblast and
form the amnion, which encloses the amniotic cavity. Con-
currently, morphological changes occur in the embryoblast
resulting in a flat, almost circular embryonic disc, which is
bilaminar (Fig. 4.3), consisting of the:
• epiblast—the thicker layer of high columnar cells related
to the amniotic cavity; and
• hypoblast—the thinner layer of small cuboidal cells adja-
cent to the exocoelomic cavity.
The pluripotent epiblast forms the floor of the amniotic
cavity and is continuous peripherally with the amnion. The
Fig. 4.3 Implanted blastocyst at 12 days. (Modified from Mitchell B,
hypoblast forms the roof of the exocoelomic cavity and is con- Sharma R. Embryology: An Illustrated Colour Text. 2nd ed. London:
tinuous with the thin exocoelomic membrane (Fig. 4.3). The Elsevier; 2009.)
exocoelomic membrane, together with the hypoblast, lines
the primary umbilical vesicle. The bilaminar embryonic disc
now lies between the amniotic cavity and umbilical vesicle. umbilical vesicle forms. This smaller vesicle is formed by ex-
Cells from the umbilical vesicle endoderm form a layer of traembryonic endodermal cells that migrate from the hypo-
connective tissue, the extraembryonic mesoderm which sur- blast inside the primary umbilical vesicle.
rounds the amnion and umbilical vesicle. The extraembry-
onic mesoderm increases and isolated coelomic spaces ap- DEVELOPMENT OF CHORIONIC SAC
pear within it which rapidly fuse to form the extraembryonic
coelom surrounding the amnion and umbilical vesicle (ex- The end of the second week is characterised by the appear-
cept where they are attached to the chorion by the connect- ance of primary chorionic villi (Fig. 4.4), that form columns
ing stalk). As the extraembryonic coelom forms, the primary with syncytial coverings. These cellular extensions grow into
umbilical vesicle decreases in size and a smaller secondary the syncytiotrophoblast forming primary chorionic villi, the
steps!
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different content
out into the kitchen, and began acting as though in pursuit of a
mouse. He and puss had often engaged in such diverting business.
This was more than the latter could stand. She was down from her
cozy bed in an instant, and was soon by the side of the dog. But as
soon as puss, all ablaze with excitement, had her head in a corner
and was straining her eyes to get a glimpse of the supposed mouse,
the dog ran to the sofa at full speed, jumped on the cushion, curled
himself round, and was happy. Poor puss, perceiving that the dog
had left her, was not slow to discern that she had been imposed
upon by the latter, and that it was only a trick that had been played
upon her by her shrewd companion, that he might get possession of
the soft spot upon the sofa. She, however, bore it good-naturedly
and decorously, and was ever afterward on the alert for these little
tricks of her canine friend.
Birds can be as capable of cheating, not only each other, but other
animals. A crow, belonging to John Smedley, a resident of Lima, Pa.,
was an adept in the business. When dinner was preparing, he would
fly around the corner of the house, set up a terrific cawing as
though in great distress, and when the mistress of the house, with
whom he was a great favorite, would come out on a tour of
investigation, the rascally bird would elude her and manage to steal
round to the table in the opposite direction and seize what food
suited him the best, which he would carry to the top of the house,
where he would eat it at his leisure. No persuasions would induce
him to come down, for he knew that such action was a breach of the
peace, and he was fearful of the punishment, that of confinement to
a cage, which would follow. When, however, he felt assured that his
mistress had forgiven the wrong-doing, he would fly down to the
porch, and do his utmost to convince her that he was a well-
meaning bird, and that he was thoroughly ashamed of his actions.
But there was one member of the family that utterly detested the
bird. It was the dog Rover. Many a trick had the bird practised upon
the latter, especially at meal time. Poor Rover was not allowed to eat
in peace. When he would be wholly absorbed in his dinner, the crow
would approach him in the rear, give him a severe twirl of the tail,
and then in a twinkling fly to one side, looking the very picture of
innocence. But ere the dog had recovered his self-possession and
was ready to resume his feeding again, the bird had captured the
daintiest morsel, and was off to the tree-top. Discomfited and
outwitted, the dog would rush to the base of the tree, bark his
growls of anger and defiance, while the crow would look quizzically
down from above, and chuckle with delight.
Many of my readers may, perhaps, remember the story of the two
dogs that used to hunt the hare in concert, the one starting the hare
and driving it toward the spot where his accomplice lay concealed. I
recall an instance where a somewhat similar arrangement was
made, only the two contracting parties, instead of being two dogs,
were a dog and a hawk, the latter making use of his wings in driving
the prey out of the copse into the open ground. Innumerous
examples of such alliances are known, and in all of them there is
manifest the curious fact that two animals can arrange a mode of
cheating a third. One of the principal stratagems used in war, that is
the ambuscade, whereby the enemy is induced to believe that
danger is imminent in one direction, when it really lies in the
opposite and unsuspecting direction, is employed. No one would
admit that a general who contrived to draw the enemy into an
ambuscade acted by instinct. The act would be construed as proof of
the possession of reasoning powers surpassing those of the
adversary. And if this be the case with the man, why not with the
dog, or with the raven or hawk, when the deception is carried out by
precisely the same line of reasoning?
Beasts possess, in common with man, the sense of Humor. This is
developed in many ways. Generally it assumes the phase of teasing
or annoying others, and thus deriving pleasure or amusement from
their discomfort. Sometimes, both with man and beast, it takes the
form of bodily torture, the struggles of the victim being highly
amusing to the torturer. Civilized man has now learned to regard the
infliction of pain upon a fellow as anything but an amusement, and
would rather suffer the agony than inflict it upon another. But with
the savage it is otherwise, for there is no entertainment so
fascinating as the infliction of bodily pain upon a human being.
Among our Indian tribes, torture is a solemn usage of war, which
every warrior expects for himself if captured, and which he is certain
to inflict upon any prisoner whom he may happen to take. The
tortures which he inflicts are absolutely fiendish, and yet a whole
tribe will assemble around the stake, and gloat upon the agonies
which are being borne by a fellow-creature. Similarly the African
savage inflicts the most excruciating sufferings upon the man or
woman accused of witchcraft, employing means too horrible to be
mentioned. But in all these cases the cruelty seems to be in a great
measure owing to obtuseness of perception. Yet the savage who
binds his victim to a stake, and perforates the sensitive parts of his
body with burning pine-splinters, behaves very much like a child who
amuses itself by catching flies, pulling off their wings and legs, and
watching their unavailing efforts to escape.
Many years ago cockchafers were publicly sold in Paris for children
to torture to death. The amusement consisted in running a hooked
pin through the insect’s tail, fastening a thread thereto, and
watching the poor creature spin in the air. After the poor beetle was
too enfeebled to expand its wings, it was slowly dismembered, the
child being greatly amused at its endeavors to crawl, as leg after leg
was pulled from the body. A similar custom, though in a more cruel
form, prevails in Italy, the creatures which are tortured by way of
sport being more capable of feeling pain than are insects. Birds are
employed in this country for the amusement of children. A string is
tied to the leg of the bird, and the unfortunate creature, after its
powers of flight are exhausted, is generally plucked alive and
dismembered. The idea of cruelty does not seem to enter at all in
these practices, but they are done from the sheer incapacity of
understanding that a bird or a beast can be a fellow-creature.
Italians are notorious for their cruel treatment of animals, and if
remonstrated with become very much astonished and reply, “Non è
Cristiano,” that is to say, “It is not a Christian.” Englishmen have little
to boast of on this score. Bear-baiting was abolished by the Puritans,
not because it gave pain to the bear, but because it gave pleasure to
the spectators. Even at the present day, both in England and in this
country, there is a latent hankering after similar scenes, and dog-
fighting, rat-killing and cock-fighting, even though they are now
contrary to law, are still practised in secret. Similarly the sense of
humor is developed in the lower animals by causing pain or
annoyance to some other creature, and the animal acts in precisely
the same manner as a savage or a child.
Sparrows, as might be expected from their character, will gratify
their feelings of aversion by banding together for the purpose of
mobbing some creature to which they have an objection. In
Hardwicke’s Science Gossip for December, 1872, there is a short
account of a number of sparrows mobbing a cat. Evidently the cat
had intended making a meal on one of the birds, but was greatly
mistaken, for the sparrows dashed upon him so fiercely, that he
soon turned tail and ran into the house, one of the sparrows actually
pursuing him into the house. The poor cat ran up-stairs, and was
found crouching in terror under one of the beds. This happened in
London, where the sparrows are less numerous now than they used
to be.
No bird of my knowledge possesses a larger amount of humor
than the crow. I have known him to feign an attack upon a distant
part of a field of newly-sprouted corn, which was being guarded by a
farmer with his gun. When the latter would be drawn to that part of
the field where the attack was to be made, the sagacious bird would
manage to outwit him, slip around to the other side, drop down into
the field and obtain a few tender sprouts before the farmer hardly
knew what was going on. But he was always up and away at the
opportune moment, and, perched upon a fence-rail, beyond the
range of the gun, would enjoy one of his rollicking cawing laughs at
the farmer’s expense. Crows that are tame have the sense of humor
more keenly developed than their wild brethren of the fields and the
woods. I once knew a tame crow that took great pleasure in
annoying a dog that lived in the same family. Carlo, as the dog was
called, was never so contented as when allowed to sleep the hours
of the morning away, after a night’s carousal, in a quiet, sunny spot
in the backyard. When the dog had become fast wrapped in the
arms of the god of slumber, the crow would steal to his side, give his
ear a sharp pull, and when the dog would awake and look around
the crow would be busy in gleaning, the most unconcerned creature
in the whole yard. Again and again would she annoy the poor
animal, and always with the same evident sense of delight, which I
could always read in the mischievous twinkle that lurked in her eyes,
till the dog, bewildered and unable to account for such mysterious
actions, would silently skulk away to other parts, where he hoped to
be free from all intrusion. Even the mistress of the house was not
exempt from her annoyance. She would carry off everything she
could lay hold of, and always hid them away in one place, that is, in
a large crevice on the top of the house between the peak of the roof
and the chimney. One day the mistress’s spectacles disappeared.
Search was instituted everywhere, but without effect. None knew
better than the bird what the trouble was. While the search was
going on, she busied herself in looking around, and seemed as
desirous of finding the missing glasses as any member of the
household. The look which the bird gave showed that she enjoyed
the situation of affairs immensely, and considered it a fine joke that
she had played upon her mistress. After a few days the lost
spectacles were restored to their accustomed place, but no one ever
positively knew how they came thither.
Domestic birds, as a rule, are remarkable for the generosity which
the master-bird shows to his inferiors. He will scratch the ground,
unearth some food, and then, instead of eating it himself, will call
some of his favorites, and give them the delicacy for which he
labored. But I have met with a few cases where the cock scratched
as usual, called his wives, and, when they had gathered round him,
ate the morsel himself. It was but a practical joke that he had
perpetrated upon them, and that they felt it as such their looks only
too strongly testified. There was a relish of delight in it for the cock,
for the cackle, which he immediately gave, assured me of this fact
as much as the laugh of a man could have done who had played
such a joke upon one of his fellows.
Parrots are much given to practical joking, after the ways of
mankind. A parrot, belonging to an aunt, had a bad habit of
whistling for a dog, and then enjoying the animal’s bewilderment
and discomfiture. She would call the cat, as her mistress was
accustomed to do, and when puss would come, expecting some
dainty article of food, she would call out in her severest tone, “Be
off, you hussy!” and the cat would make all possible speed for a
place of security, greatly to the amusement of the parrot from her
perch in the cage. There have been known parrots that would play
practical jokes upon human beings, but dogs and cats seem to be
the principal victims of the parrot’s sense of humor.
Animals not only show their playfulness in such tricks as have
been mentioned, but many of them are able to appreciate and take
part in the games played by children. When I was a boy I knew a
dog, a species of greyhound, which was an accomplished player at
the well-known game called tag, or touch. Quite as much
enthusiasm was displayed by the animal as by any of the human
players. He would dart away from the boy who happened to be
“touch” with an anxiety that almost appeared terror. It was an
impossibility to touch the clever canine player; but he was a
generous creature, with a strong sense of justice, and so, when he
thought that his turn ought to come, he would stand still and wait
quietly to be touched. His manner of touching his play-fellows was
always by grasping the end of their trousers with his teeth, and as it
was impossible for the boy to stop when so seized in full course, the
dog was often jerked along the ground for some little distance.
Hide-and-seek is a game which is often learned and enjoyed by
many animals. I have often been an interested spectator of the play
in which two dogs were the participants. It was as exciting as such a
diversion could possibly be between two children. For an hour at a
time I have watched the fun, and the players seemed not to abate
the least jot or tittle from their ardor and enthusiasm. They were
apparently as fresh then as at the beginning. In due time the game
ceased as if by mutual consent, but the animals did not seek some
cool, quiet spot for comfort and rest, but started off to the woods for
some further diversion, from which their voices were soon heard,
telling that they were in pursuit of a rabbit or the ignoble ground-
hog.
We have far from exhausted the list of examples at hand to show
that the lower animals possess a sense of humor. But what use, it
may be asked, can the capacity of humor subserve in the next
world? Much the same, I presume, that it subserves in this. There
are some in this world in whom the sense of humor is absolutely
wanting. Estimable as they may be in character, they are just solemn
prigs, and I should be very sorry to resemble them in the world,
whither, it is hoped, all life tendeth.
Pride, Jealousy, Anger, Revenge and Tyranny, while not very
pleasing characteristics, belong, as such, to the immaterial, and not
to the material, part of man. That the lower animals possess these
qualities will be seen from what follows. Hence the inference to be
drawn from that fact must be quite obvious.
Taking these characteristics in order, Pride, or Self-esteem, is
developed as fully in many animals as in the proudest of the human
race. Most conspicuously is this shown in animals which herd
together. There is always one leader at the head, who will not permit
any movement to be made without his order, and who resents the
least interference with his authority. This is particularly the case with
the deer, the horse and the ox. Even when these animals are
domesticated, and the habits of their feral life have materially
changed, the feeling of pride exists to the fullest extent.
Whoever has carefully watched and studied the inhabitants of a
farm-yard cannot fail to have observed that the cows have their laws
of precedence and etiquette as clearly defined as those of any
European Court. Every cow knows her own place and keeps it. She
will never condescend to take a lower, nor would she be allowed to
assume a higher. A new-comer in a farm-yard has about as much
chance of approaching the rack at feeding-time as a new boy at
school has of getting near the fire on a cold winter day. But as the
young calf increases in growth, and is nearing maturity, she is
allowed to mingle with her companions on tolerably equal terms.
Should, however, a younger animal than herself be admitted, it is
amusing to see with what gratification she bullies the new-comer,
and how much higher she ranks in her own estimation when she
finds she is no longer the junior.
But should the fates be propitious, and she should arrive at the
dignity of being senior cow, she never fails to assert that dignity on
every occasion. When the cattle are taken out of the yard to their
pasture in the morning, and when they are returned to it in the
evening, she will not allow any except herself to take the lead. An
instance is recorded where the man in charge of a herd of cows
would not permit the “ganger,” as the head cow is often called, to go
out first. The result was that she refused to go out at all. Therefore,
to get her to go out of the yard, the man had to drive all the other
cows back again, so that she might take her proper place at the
head of the herd.
Few people know much about the real disposition of the mule.
Judging from popular ideas respecting the animal, one would think
that it had no pride in its composition. It is in reality a very proud
animal, and fond of good society. One of his most striking
characteristics is his aversion to the ass, and the pride which he
takes in his relation to the horse. An ass would be hardly safe in a
drove of mules, for he would, in all probability, be kicked and lamed
by his proud relatives; whereas a horse, on the contrary, takes a
distinguished position, the mules not only crowding around him and
following his movements, but exhibiting a violent jealousy, each
striving to get the nearest to their distinguished relative.
EXHIBITION OF GRANDEUR.
Male Peacock in Presence of Some Barn-Yard Fowls.
We have seen the pride of rank and love of precedence in cows,
and the pride of ancestry in mules. There is, however, a pride that
takes the form of sensitiveness to ridicule. Nothing is so galling to a
proud man as to find himself the object of ridicule. The same trait of
character is to be found in many animals, and especially in those
that have been domesticated, for it is in these that we have the
most opportunities for observation. All high-bred dogs are
exceedingly sensitive to ridicule. We knew of a cat that was quite
conscious if spoken of in a disparaging manner, and testified his
disapprobation by arching his tail, holding himself very stiff indeed,
and marching slowly out of the room.
There is, however, another form of pride which is often to be seen
among the lower animals, but more especially among birds notable
for gaudy or abundant plumage. This is the pride which manifests
itself in personal appearance. Vanity is the name which is currently
applied to this form of pride. Those who have observed a peacock in
all the glory of his starry train will recognize the intense pride he
feels at his own splendor. This display of his magnificent train is not
for the purpose of attracting the homage of his plainly-attired mates
solely, but seems to be intended to evoke the admiration of human
beings as well. Not even the homage of birds, whom he regards his
inferiors, is to be despised.
We have seen him, with his train fully spread, displaying his
grandeur around a dozen or more barn-yard fowls, and apparently
as satisfied with the effect he produced, as he stalked majestically
among them, as if he had been surrounded by his own kith and kin.
Then there is the turkey. No movements are more grotesque than
his. See him as he struts about in his nuptial plumage, and yet no
bird, notwithstanding the ludicrousness of his behavior, surveys
himself with greater complacency. The whidah-bird, or widow-bird,
as it is often called, exhibits this trait of character in its highest
development. He is wonderfully proud of his beautiful tail, and, as
long as he wears it, loses no opportunity of displaying it to every
person who visits his cage. But when the moulting season has
arrived, and he has taken on the plain, tailless attire of his mate, a
change as great has come over his manner, and, instead of
exhibiting himself in all his pride and glory, he mopes listlessly and
stupidly about, and seemingly ashamed of his mean condition. In all
these instances the character of pride in personal appearance is as
strongly developed as it is possible for it to be in any human being.
That peculiar uneasiness, which arises from the fear that a rival
will dispossess us of the affection of one whom we love, or the
suspicion that he has already done so, is termed jealousy. There are
two forms of this passion, one connected with the love of some
other being, and the other dependent on the love of self. But it is
the former, whose definition begins the present paragraph, with
which we shall exclusively deal. It is evident from the meaning of
jealousy, as given above, that the power of reasoning is implied, and
that any creature by which it is manifested must be able to deduce a
conclusion from premises. No matter if the conclusion drawn by the
animal be wrong, the process, however incorrect it may be, is, it
cannot be denied, still one of reasoning. All who have possessed pet
animals must be familiar with the exceeding jealousy displayed by
most of them. Most strongly is this feeling manifested when an
animal has been the only pet and another is introduced into the
house. Where there are two or more dogs in the same family, one is
often amused at the boundless jealousy displayed toward each other
while engaged in the service of the master, although at other times
they were on the most excellent terms. Bus is the name of a favorite
dog belonging to a friend. No more affectionate dog ever lived.
Beagle was his companion. When they were by themselves, life was
a round of frolics and rambles. No matter how rough and exciting
their plays were, they never got cross, but endured everything with
patience and forgiveness of spirit. Beagle was a clever animal, and
very fond of the chase. Many a ground-hog would he dislodge from
its burrow and fight to the death, while Bus would look on with
wonder and admiration. But let the slightest attention be shown by
the master to Beagle, and Bus’s jealousy and anger became
unbounded. He would fly at his friend in the most infuriated manner,
rending him with tooth and claw, while Beagle would quietly slip
around the corner of the house to get out of the reach of his
companion’s temper. Beagle, being a large and powerful dog, had in
him the ability to give Bus a very sound whipping, but he was too
noble and magnanimous a creature to take advantage of one
younger and smaller than himself. He would always allow Bus to
have his own way, knowing that the passion which was lacerating
the bosom of his young companion and playmate would soon spend
itself, and the latter, ashamed and abashed, would be soon seeking
forgiveness and reconciliation.
Even in such rarely tamed animals as the common mouse the
feeling of jealousy has been known to be so intense as to lead to
murder. A young lady, one of Rev. J. G. Wood’s correspondents, had
succeeded in taming a common brown mouse so completely that it
would eat out of her hand and suffer itself to be taken off the floor.
She had also a tame white mouse in a cage. One morning when she
went to feed the white mouse, as was her usual custom, she found
it lying dead on the bottom of the cage, and beside it was its
murderer, the brown mouse. The cage being opened, the latter
made its escape, as though fearful of the consequences that might
ensue, but how it had managed to gain admission was always a
mystery.
Instances are on record where the jealousy of a rival has been
restrained for long years through fear, and has ultimately broken out
when the cause of the fear has been removed. A case of the kind
came under our notice some few years ago. There were two cocks,
belonging to different breeds, whom fate had placed as denizens of
the same family. One was a magnificent dunghill cock, and the other
a Malay, a cowardly caitiff, that was kept in fear and subjection by
the former. In the course of events the dunghill cock suddenly died.
His rival, coming by chance on his dead body, and perceiving that
the time had come to wreak out the mixture of hatred and revenge
that had lain smouldering in his bosom for years, instantly sprang
upon it, kicked, spurred and trampled upon the lifeless bird, and,
standing upon the corpse, flapped his wings in triumph, as it were,
and crowed himself hoarse with the most disgusting energy. He
immediately took possession of the harem, but he was far from
being the noble, generous and unselfish creature that his
predecessor had been. Again, comparing man with beast, it is at
once apparent that the bird in this instance acted exactly as a
savage does when his enemy has fallen, for the savage not only
exults over the dead body of an enemy, especially if the latter has
been very formidable in life, but also mutilates in futile and silly
revenge the form which he feared when alive.
Tyranny, or the oppression of the weak by the strong, is another
of the many traits of character common to man and the lower
animals. But whether or not that strength belongs to the body or the
mind, it is tyranny all the same. Taken in its most obvious form, it
not only manifests itself in many of the animals in the oppression of
the weak by the strong, but also in the killing and the eating of the
same, even though they be of the same species. Human cannibals
act in just the same manner, eating their enemies after they have
killed them. There is hardly an animal in which the milder forms of
tyranny may not be found. Insects, especially, manifest it in a light
manner when they drive away their fellows from some morsel of
food which they desire to keep to themselves. Among gregarious
animals, the herd or flock is always under the command of an
individual who has fought his way to the front, and who will rule
with imperious sway until he has become old and in turn has been
supplanted by a younger and more vigorous rival. In the poultry-
yards the same form of tyranny is manifest, one cock invariably
assuming the leadership, no matter how many may be the number
of birds.
There is a curious analogy between these birds and human
beings, especially those of the East, whether at the present day or in
more ancient times. Many petty chieftains are found in Eastern
countries, but there is always to be met with one among them who
is more mighty than the rest, and who holds his place by superior
force, either of intellect or military power. Challenged by one of the
inferior chiefs and victorious, he retains his post, but if vanquished,
his conqueror takes his place, his property and his wives. But curious
to relate, with men as with birds, the members of the harem seem
to trouble themselves very little, if any, about the change of master.
The Scriptures are full of allusions to the invariable custom that the
conqueror takes the possession of the harem belonging to the
vanquished. David did so with regard to the women of Saul’s
household, and when Nabal died, who had defied the authority of
David, so the latter, as a matter of course, took possession of his
wife, together with the rest of his property. And when Absalom
rebelled against David, he publicly took possession of his father’s
harem, which was a sign that he had assumed the kingdom.
Where a number of creatures are confined in the same place, a
very curious sort of tyranny is sometimes manifested. Mandarin
ducks, according to Mr. Bennett, when confined to an aviary, show a
very querulous disposition at feeding-time. The males of one and the
same kind of a different species endeavor to grasp all the
nourishment for themselves, unmindful of the wants of others, and
will not even permit their companions to perform their ablutions
without molestation, although they may themselves have completed
what they required. Often the mandarin ducks have been observed
to excite the drakes to assail other males or females of the same
species, and other kinds of birds in the aviary, against whom the
ladies, from some cause or other, have taken a dislike. One pair of
these ducks are always to be noticed that exercise a tyranny over
the others, not allowing them to wash, eat or drink, unless at their
pleasure and approval.
But, of all tyrants, none can be compared to a spoiled dog, who is
even worse than a spoiled child. Obedience is a stranger to his
nature. Does his master want him to go out for a walk, and he
prefers to stay at home, he stays at home, and his master is
compelled to go out without him. But if he wants to go for a walk,
he makes his master go with him, and even to take the direction he
prefers. Duchie is the name of a Skye terrier whose history is given
in a work on the latter breed of dogs by Dr. J. Brown. So completely
had this little animal domineered over her mistress, that the latter
could not even choose her own dinner, but was obliged to have
whatever the dog preferred. It is related that for a half of a winter’s
night she was kept out of bed, because Duchie had got into the
middle and refused to move. Certainly, no better example of tyranny
could be adduced.
That so-called brutes possess, in common with ourselves, a
Conscience, that is, a sense of Moral Responsibility, and a capability
of distinguishing between right and wrong, may seem a very strange
assertion to be made, especially to those who have never studied
the ways of the lower animals. Animals which are placed under the
rule of man, and those, like the dog, which belong to his household
and are made his companions more particularly, would naturally be
expected to show the strongest development of the principle.
Conscience, in their dealings with man, constitutes their religion, and
they often exercise it in a way which would put many a human being
to the blush. This feeling it is that induces the dog to make himself
the guardian of his master’s property, and often to defend that
property at the risk of his life. However hungry may be the dog that
is placed in charge of his master’s dinner, nothing would, as a rule,
tempt him to touch a morsel of the food, for he would rather die of
starvation than eat the food which belongs to his master. Often have
we seen field-laborers at work at one end of a large field, while their
coats and their dinner were at the other end, guarded by a dog. Not
the least uneasiness did they seem to manifest about the safety of
their property, for well they knew that the faithful animal would
never allow any one to touch either the clothes or the provisions.
There could hardly be a stronger instance of moral responsibility
than the one which I shall now relate, which is substantially the
same as appears in Wood’s “Man and Beasts Here and Hereafter.”
Living in an unprotected part of Scotland was a poor woman, who
unexpectedly became possessed of a large sum of money. She
would have taken it to the bank, could she have left the house, but
lack of bodily health prevented her from so doing. At last she asked
the advice of a butcher of her acquaintance, telling him that she was
afraid to live in the house with so much money about her. “Never
fear,” said the butcher, “I will leave my dog with you, and I’ll warrant
you that no one will dare to enter your house.” Towards the close of
the day the dog was brought, and chained up close to the place
where the money was deposited. That very night a robber made his
way into the house and was proceeding to carry off the money,
when he was seized by the dog, who held him a prisoner until
assistance arrived. The thief turned out to be the butcher himself,
who thought he had made sure of the money, but he had not
considered that his dog was a better moralist than himself, for who
would, rather than betray a defenceless woman, take her part
against his own master. Kindly pardoned by the woman, the
intending robber made his way home, and it is to be hoped that for
the future he learned a lesson from his own dog and amended the
evil of his ways.
Not only does the dog guard the property which is intrusted to its
charge, but frequently goes a little further and assumes a charge on
its own account. When the writer was a boy living in the country,
where much of the spring and summer of the year was spent in
working upon a farm, he became on very excellent terms with a little
bull-terrier, named Tip, that belonged to a certain farmer by whom
he was employed. Upon my first introduction to Tip, I felt a sort of
aversion towards him. This grew out of the mysterious actions of the
animal. He was always around when I was busy at work and seemed
to be eying me in a suspicious sort of manner, which at times made
me feel very unpleasant. After the lapse of a few days I discovered
that I was not so closely watched as before, and that I was treated
by him as he was accustomed to treat the other members of the
family. Upon inquiry I learned that he always acted in this way
toward people whom he did not know intimately, and that, after a
time, he had confidence in their honesty and left them alone. While
in many instances Tip was entirely wrong in his surmises, yet cases
are recalled where the dog was right and acted in a manner that
would have been creditable to a human being. One of the men
employed upon the place, presuming upon the friendship of the dog,
sought to carry away under cover of darkness something belonging
to the farmer, but he was immediately beset by the animal, who was
an eye-witness of the proceeding, and compelled to desist from the
intended theft. From that time the man was under the closest
surveillance by the dog. Unable to effect a reconciliation, and chafing
under the look of suspicion with which he was always greeted, the
man soon took his departure, much to the delight and satisfaction of
the faithful canine, and was never afterwards seen.
Quite a common form of conscience among the lower animals is
that which may be defined as a recognition of having done wrong,
and acknowledgment that punishment is deserved. Animals have in
their way very pronounced ideas as to right and wrong. When they
have committed an act which they know will offend their master,
they display as keen a conscience as any human being self-convicted
of sin could exhibit. In many instances, the offence in not merely
acknowledged, but the creature remains miserable until forgiveness
has been granted. This condition of mind, if manifested by man, is
called Penitence, and, assuredly, it cannot be known by any other
name when manifested by animals that are lower down in the scale
of life. My little dog Frisky, about whom mention has already been
made, affords a very fine illustration of this phase of conscience.
Whenever he did wrong, the severest punishment that could be
meted out to him was to ignore his presence and decline his offered
paw. For hours the poor fellow would moan and cry, and even refuse
food, when he thought I was angry with him. But a word or a look of
forgiveness was sufficient to change his sadness into joy. A shaking
of hands, so to speak, would then follow, and master and dog would
be good friends again. No love could be more intense than his, and
this was especially shown when I would return from a short
absence, when the little fellow would almost overwhelm me by his
affectionate caresses.
No loftier characteristic adorns humanity than Love. But how far it
is shared by the lower animals it is now our purpose to inquire. That
there are many phases of development cannot be doubted.
Sympathy, or that capacity of feeling for the sufferings of another, is
the first phase. Many, and perhaps all, living creatures possess the
capacity of sympathy. In the majority of cases it is not restricted to
their own species, but is extended to those beings which appear to
have very little in common with each other. Ordinarily, however, it is
exhibited between animals of the same species, and it is often seen
in the dog, as, for example, where a dog, having been cured of an
injury, has been observed to take a fellow-sufferer to his benefactor.
Such sympathy, it need hardly be remarked, could not be carried out
unless the animals possessed a language adequately defined to
enable them to transmit ideas from one to the other. Cats are often
kind to each other, sympathizing under difficulties, and helping their
friends who require assistance. A cat, belonging to a friend, has
been known, when oppressed with the cares of a family, to employ a
half-grown kitten to take charge of the young while she went for a
ramble. Between the cat and the dog an enmity exists that is
hereditary, and yet, when in good hands, they are sure to become
very loving friends, and even to show considerable sympathy
towards each other. Such an exhibition of good feeling was observed
by the writer a few years ago. The dog, a large black Newfoundland,
had contracted a warm and devoted friendship for a gray cat that
was an inmate of the same family. When the cat was assailed by one
of her kind, or by a strange dog, the Newfoundland would pick her
up in his mouth and carry her to the house out of reach of danger,
the cat maintaining all the while the most perfect serenity of
composure, knowing that she was in the care of one who meant her
no ill. When the same cat would become sick, the Newfoundland
would lie down by her side, caress her with his tongue, and show in
every way possible that he was sorry that she was sick.
Many examples are recorded of birds feeling sympathy with the
lost or deserted young of other species, and that have taken upon
themselves the task of feeding the starving children. A pair of robins
had constructed a nest near to the writer’s home in the country,
where in due season a family of four children was raised. Disaster
soon came to the little ones, for both parents were slain by some
wicked boys of the neighborhood. There dwelt in the same locality a
pair of bluebirds, but between the two families there had never been
apparent the least interchange of friendship. Each family kept to
itself, and attended to its own business. But when the cry of the
young robins in their piteous demands for food rent the air, the
bluebirds came over to their home to discover what the trouble was.
They were not slow to perceive the sad state of things. Their
sympathies were at once aroused, and their energies soon bent in
the direction of relieving the sufferings of the little orphaned robins.
For the next two weeks they had all they could do in providing meat
for their own and the robins’ young.
FOUR ORPHANED ROBINS.
Kind-Hearted Bluebirds Assuming the Role of Parents.
While capable of showing sympathy for near as well as distant kin,
the lower animals have also the capacity to sympathize with human
beings in distress. Cats occasionally manifest a sympathy for
suffering humanity. As for sympathy displayed by dogs, there is no
need to cite examples. No human being, I am safe in saying, was
ever free from troubles of some kind, and I am equally sure that no
one who had a companionable dog felt that he was without
sympathy. Full well does the dog know when his master is suffering
pain or sorrow, and his nose pushed into his master’s hand, or laid
affectionately upon his knee, is a sign of sympathy worth
possessing, even though it exists only in the heart of a dog. From
that moment there has been established a bond between the soul of
the master and the dog, and certainly no one can believe that the
bond can ever be severed by the death of the material body,
whether of the man or the animal.
That Friendship, which is another branch of love, exists among
animals, is a well-known fact. But it is among the domesticated
animals that it most frequently exhibits itself. Horses, as every one
knows, which have been accustomed to draw the same carriage are
usually sure to be great friends, and if one be exchanged the other
becomes quite miserable for want of his companion and seems
unable to throw any spirit into his work. Dogs, too, are very apt to
strike up friendships with each other. Among animals it is not
confined to one species, but is occasionally found to exhibit itself in
those which might be supposed to be peculiarly incongruous in their
nature. That cows and sheep live, as a rule, on good terms with
each other in the same pasture is a familiar experience, though
sometimes the former are a little prone to domineer over the latter.
But a very strong affection sometimes exists between animals so
different, and when once they have accustomed themselves to each
other’s society neither can be happy without the other. The goat and
the horse frequently become friends, and a peculiarly vicious horse
has been known to allow a goat to take undue liberties with him
without the least manifestation of resentment. In many places the
stable-cat is quite an institution. Its usual place of repose is upon
the back of the horse, and the latter has been known to grow very
uneasy if left for any length of time without the companionship of
his little friend. A very singular instance of friendship occurred at the
rural home of a near relative. He had a fine mastiff which had taken
a fancy to a brood of young chickens, and which acted as their
protector. They were not at all unwilling to accept him in this
capacity, as they followed him about just as though he had been
their mother. Quite an interesting sight it was to watch the dog and
the chickens as they would take their siesta. The dog used to lie on
his side, and the chickens would nestle all about him, though one
chicken in particular would invariably scramble upon the dog’s head,
and another just over his eye, but both parties appeared equally
satisfied with this remarkable arrangement.
Already have we referred to the intense yearning which is felt by
many of the lower animals for human society. This yearning is
indeed but the aspiration of the lower spirit developed by contact
with the higher in domesticated animals or those which are in
perpetual contact with man. This feeling is a matter of no great
surprise. But that it should be exhibited in feral animals and birds,
and even in insects, is a fact well worth considering, as it furnishes a
clew to some of the many problems of life which are as yet
unsolved. That power of attraction exercised by the spirit of man
upon that of the lower creation is well exemplified in many wild
animals, who are known to forsake the society of their own kind for
the companionship of the being whom they feel to be higher than
themselves.
Perhaps one of the wariest of wild animals is the squirrel. He is
horribly afraid of human beings, and if a man, woman or child come
to the windward of him, the little animal is sure to scamper off at his
fleetest pace, scuttle up the nearest tree, and conceal himself behind
some branch. Yet, wild as he may be, he is peculiarly susceptible to
the influence of the human spirit, and for the sake of human society
will utterly abandon that of his own kind. I once knew a pet gray
squirrel by the name of Charley. He had been taken from the nest
when very young. His home for awhile was one of those whirl-about
cages. Charley did not like his cage, but preferred to be outside in
the unrestrained enjoyment of the dictates of his own free will. So it
was difficult to keep him behind the bars. When awake he loved to
follow his own devices; but when tired he usually slept on a soft
cushion on the sofa, or found his way into some bed-room where he
would nestle under a pillow. Nothing was more to his satisfaction
and pleasure than a share of the bed of his mistress, but he was
always a troublesome nest-fellow. Charley had, as must be obvious,
perfect freedom. He was allowed to go as he pleased. There was no
coercion in his case. Had he wished to escape, there was nothing to
prevent, and nothing bound him to his mistress but an “ever-
lengthening chain” of love and aspirations which none but a human
being could satisfy. The sparrow, one of the most independent and
self-reliant of birds, has been known to abandon its kind for the sake
of human beings. Wood cites a case of a bird of this species that had
been rescued from some boys who had been robbing the nest. The
bird was brought home, but was never confined in a cage, but was
permitted to fly freely about the house. As there was a cat about the
house, she had to be closely watched lest she might do the bird
some injury. On Sundays, when the family went to church and no
one remained to keep an eye on the cat, the sparrow was turned
into the garden, where it flew about until the family’s return. The
opening of the dining-room window by its mistress, and the display
of her ungloved hands, was the signal for its entry. But if the
mistress stood by the window with her gloves on, then the bird
showed not the slightest disposition to enter.
Such is the intensity of the love which the lower animals
sometimes entertain toward man that they have been known to
grieve themselves to death on account of his loss. A dog by the
name of Prince, who lived in the family where the writer spent a few
weeks of a summer, is a case in point. He had a good master, and
one to whom he was strongly attached. The year before the master
sickened and died, and Prince felt the loss so keenly that he refused
to take any food, and even to notice the surviving members of the
family. He was pitiable to behold. Life had lost all attractions to him,
and he showed that he was slowly but surely grieving his life away.
Some few weeks after the writer’s departure, the poor animal
breathed his last, and his spirit, it is to be hoped, went to join that of
his master, while his ashes became mingled with the dust of the
earth as his master’s had been.
What a wonderful power do some animals have of returning to
their beloved master, even though they have been conveyed to a
considerable distance. This is especially true of the dog. So many
examples of such feats are on record that I refrain from mentioning
them, but will give but a single example. Rover, a pet greyhound
that belonged to the writer, had become such an annoyance to the
neighborhood where he lived, that the master determined to provide
him a home in the country some fifty miles away. He was conveyed
to his destination in a covered wagon, and after his new master had
reached home, the poor animal was placed in a stable for several
days, where he was daily visited and fed, and every effort possible
made to attach him to the place and family. On the fourth day of his
arrival he was given his freedom. With a long, loud wail he saluted
the neighborhood, and the next moment was off at full speed across
the country, all efforts to stop him being unavailing. In less than a
week from his leaving he was at home again, hungry and jaded out
with fatigue and travel, but not too tired nor too hungry to express
the great joy he felt for the old master. How he ever accomplished
the journey, and what vicissitudes and difficulties he encountered on
the way, no one will ever know. After this I had not the heart to
send him away again, but put up with his capers and tricks as best I
could, and when complaints were preferred against him endeavored
to excuse them as a parent is prone to do in the case of a spoiled
and wayward child. But a day arrived when Rover to me was no
more. What had become of him I was never able to discover, but I
always blamed a near-by neighbor, a man who had neither love nor
charity in his soul, for his sudden disappearance.
That cats are selfish animals, attaching themselves to localities
and not to individuals, I do not believe. This idea has, perhaps,
some ground of truth, for the nature of a cat is not so easy to
understand as that of a dog. But when a cat is not understood, it is
very probable that she cares less for the inhabitants of the house
than for the house itself. Frequent instances are known by the writer
where cats have been in the habit of moving about with their
owners, and have been as much unconcerned as dogs would have
been. True they have, like women, a curious and prying disposition.
I have seen them in new and strange quarters go sniffing about
every room of a house, and at last settle down in some cozy,
comfortable place, well satisfied with their tour of investigation.
Where the house fell short of their expectations, if they have been
cats that have received due consideration from their mistresses or
masters, they have tried to live down their objections and to learn to
be happy and contented with their lot. Only cats that have not been
much thought of are inclined to show their disapproval to changes of
residence which they deemed unsuitable by refusing to stay with
their masters. Blackie, a favorite cat of ours, never seemed to care
where her home was, so long as her friends were there to pet,
caress and pamper her with choice dainties.
All animals, so far as can be learned, have not only a capacity for
the society of man, but an absolute yearning for it. This feeling may
be in abeyance, from not having received any development at the
hands of man, but it nevertheless exists, and only awaits to be
educed by some one capable of appreciating the character of the
animal. Tigers, as is well known, are not generally considered the
friends of mankind, and yet the Indian fakirs will travel over the
country with tame tigers, which they simply lead about with a slight
string, and which will permit small children to caress them with their
hands without evincing the least disposition to hurt them.
When we survey the examples of love displayed by animals
towards human beings, which we have just detailed, and recall the
hundreds that we know and have read about, is it possible to believe
that such love can perish? We apprehend not. Unselfish love as this,
which survives ingratitude and ill-treatment, belongs to the spirit and
not to the body, and all beings capable of feeling such love must
possess immortal spirits. All may not have an opportunity of
manifesting it, but all possess the capacity and would, were the
conditions favorable, manifest it openly.
Few animals, as may easily be imagined, manifest Conjugal Love.
Most species have no particular mates, but merely meet by chance,
and seemingly never trouble themselves about each other again. No
real conjugal love, therefore, can exist, and it is rather curious that
in such animals a durable friendship is frequently formed between
two individuals of the same sex. But when we come to polygamous
animals, such as the stag among mammals and the domestic poultry
among birds, we meet with a decided advance towards conjugal
love, although as in the case of polygamous man, that love must
necessarily be of an inferior character. There is seen, at all events, a
sense of appropriation on either side. Take the example of the barn-
yard fowl, as has already been mentioned in that part of the chapter
which deals with jealousy, where it is shown that the proprietor of
the harem resents any attempt on the part of another male to
infringe on his privileges.
This brings us to the consideration of birds, where the many are
mated for the nesting-season, but subsequently do not seem to care
more for each other than they do for their broods of children. If one
of the pair be killed at the nesting-time the survivor, after a brief
lamentation, consoles itself in a few hours or days with another
partner, for there really appears to be a supply of spare partners of
both sexes always at hand. And now we come to those creatures
which are mated for life, and often we find among them a conjugal
love as strong and as sincere as among monogamous mankind.
Prominent among them are the eagle, the raven and the dove. And
while we praise the turtle-dove for its conjugal fidelity, and credit it
with the possession of all that is sweet, and good, and gentle, how
remarkable is it that we forget to accredit with the same virtue the
eagle and the raven, that are the types of all that is violent, and
dark, and cunning. There are many examples in existence of the
conjugal love among such birds, but they are so well known that
reference to them is unnecessary. The case of the mandarin duck,
already narrated, affords a strong instance of conjugal love wherein
the lady was faithful and the husband avenged himself on the
destruction of his domestic peace.
MATED FOR LIFE.
Conjugal Fidelity Shown by a Pair of Doves.
So numerous as are the instances of love shown by parents

among the lower animals towards their offspring, yet it is a very
singular fact that few, if any, trustworthy accounts of Filial Love, or
the love of children toward their parents, are to be found. But we
must look to man if we would understand the lower animals. Even
human nature must attain a high state of development before filial
love can find any place in the affections. In savages it barely exists
at all, and certainly does not survive into mature years. It is the
glory of the North American Indian boy, at as early an age as
possible, to despise his mother and defy his father. And the women
are just as bad as the men. Rejoicing in the pride of youth and
strength, they utterly despise the elder and feeble women, even
though they be their own mothers, and will tear from their hands
the food they are about to eat, on the plea that old women are of no
use, and that the food would be much better employed in giving
nourishment to the young and strong. The Fijians have not the least
scruple in burying a father alive when he becomes infirm, and assist
in strangling a mother that she may keep him company in the land
of spirits. Both the Bosjesmen of South Africa and the Australian
seem to have not the least idea that any duty is owing to a parent
from a child, nor have they much notion of duty from a parent
toward the child. If the father be angry with any one for any reason,
he has a way of relieving his feelings by driving his spear through
the body of his wife or child, whichever one of the two happens to
be the nearer. Even the mother treats her child with less
consideration than a cow does her calf, and leaves the little creature
to shift for itself at an age when the children of civilized parents are
hardly thought fit to be left alone for a few minutes. This being the
case with parental love, it may be readily imagined that filial
affection can have not the slightest chance for development, and it
is very much to be questioned whether in the savage it can really be
said to exist at all in the sense understood by enlightened peoples.
Therefore, as in the lower human races, we find that filial love either
is very trifling, or is absolutely non-existent, need we wonder that in
the lower animals such few, if any, indications of its presence should
be found?
Now, as to the subject of Parental Love, and the various ways in
which it manifests itself. There are many writers who claim that
parental love in the lower animals is not identical with that of man.
They affirm that it is only a blind instinct, and, in order to mark more
strongly the distinction between man and beast, call the parental
love of the latter by the name of storgë. Speaking for myself, I must
declare that I am unable to perceive any distinction between the
two, save that in civilized man the parental love is better regulated
than among the lower animals. But, as has been seen, it is not
regulated at all among the uncivilized races, and, in truth, many of
the beasts are far better parents than most savages. Nor can I
understand why the word storgë should be applied to parental love
among the lower animals and not to the same feeling in man.
Among Greek writers the word, together with the verb from which it
is derived, is applied to the love between human parents and
children. It is so applied by Plato, and in the same sense by
Sophocles and others. One argument adduced by those who deny
the identity of the feeling in both cases is that parental love endures
throughout life in man, while it expires with the adolescence of the
young in the lower animals. This is doubtless true, as a rule, with
civilized man, but in the case of the savage, as has previously been
shown, it does not last longer than that of a bird, a cat or a dog,
taking into consideration the relative duration of life. And the reason
is identical in both cases. Were this love to exist through life in the
savage, the beast or the bird, the race would become extinct, for
neither race is able to support its children longer than their time of
helplessness. The beast and the bird cannot, and the savage will
not, provide for the future. It is therefore evident that if the young
had to depend upon their parents for subsistence, they would soon
perish from lack of food. Exceptions there are to this general rule,
and always, as far as can be determined, in the case of
domesticated animals whose means of subsistence are already
insured.
Several of such cases have come to my notice. I shall instance but
one. A friend of mine has two terriers, a mother and a daughter. The
strongest bond of love and fellowship unites them. They always sit
close together, and the mother playfully pinches her daughter all
over. Should they by chance become separated, even for a very
short time, the daughter comes up wagging her tail, and then licks
her mother’s nose and mouth. When hunting together, they always
act in concert, each one taking a hole, and one keeping watch while
the other scrapes away the earth. The meaning of each other’s
whine or bark is perfectly understood, and no two persons could
understand their own language better than do these dogs theirs, nor
be more comprehensible to each other.
Self-abnegation is perhaps one of the most beautiful
characteristics which parental love can give. This is particularly
shown when the young are in danger. A human mother in charge of
her child will defy a danger before which she would shrink if alone,
and in its defence would dare deeds of which most strong men
would be incapable, for during the time her selfhood is extinguished,
and her being is sunk into that of her child. Such abnegation
becomes a true mother, for if she would not consent to do and dare
for the sake of her offspring, she would degrade herself below the
beasts and the birds, who hesitate not in performing that duty to
their children, though savants do declare that they possess only
storgë, whatever they may mean by it, and not parental love.
EVIDENCE OF CONJUGAL AFFECTION.
Male Humming-Bird Feeding His Partner, and Ready to Act in Her Defence.
Everyone who has paid even a passing attention to the habits of

birds must have noticed the vigilance a pair of catbirds exercise over
their nest when containing young birds. Neither parent, when the
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For technical assistance:

Mark G. Torchia, DipBT, MSc, PhD

T.V.N. (Vid) Persaud MD, PhD, DSc, FRCPath (Lond.), FAAA

CONCISE CLINICAL EMBRYOLOGY:

Copyright © 2021 by Elsevier, Inc. All rights reserved.

Library of Congress Control Number: 2020951836

Content Strategist: Jeremy Bowes

Printed in United States of America

Last digit is the print number: 9 8 7 6 5 4 3 2 1

2 Reproductive Organs and Gametogenesis 4 9 Development of the Body Cavities, Diaphragm,

14 Teratogenesis and Birth Defects 107

Multiple Choice Question Answers 112

Video 3.1 Fertilisation 12 Video 9.3 Pharyngeal Apparatus 64

Video 3.2 Blastocyst 12 Video 9.4 Face and Palate 66

Video 4.1 Implantation 18 Video 10.1 Nervous System 71

Video 5.1 Gastrulation 23 Video 10.2 Eyes 76

Video 5.2 Folding 28 Video 10.3 Ears 78

Video 8.1 Heart 44 Video 11.1 Gastrointestinal 82

Video 8.2 Vascular 49 Video 12.1 Urinary 89

Video 9.1 Body Cavities 58 Video 12.2 Reproductive System 91

Video 9.2 Respiratory System 61 Video 13.1 Limbs 101

tissues of the uterus and vagina (endometrium and mucosa,

(internal), myometrium (middle muscular) and perime-

The ovaries are oval-shaped glands adjacent to the uterus

progresses only to metaphase. If a sperm penetrates the sec-

MENSTRUAL CYCLE by the corpus luteum stimulates the glandular epithelium

Molecular Biology Considerations NONDISJUNCTION

CLINICAL ISSUES Case Outcome

genital tract. Oocytes are usually fertilised within 12 hours

Fig. 3.1 Timeline of development related to fertilisation.

many women who are sterile to have children. Since 1978,

IN VITRO FERTILISATION AND EMBRYO TRANSFER INTRACYTOPLASMIC SPERM INJECTION

the treatment of couples in whom typical IVF has failed, or

ASSISTED IN VIVO FERTILISATION

Most pregnancy tests are based on the detection or measure-

AUTOSOMAL TRISOMY MOSAICISM

PREIMPLANTATION GENETICS Fig. 3.6 Ultrasound of a fetus demonstrating an enhanced

1. How many sperms would probably be deposited by a nor- c. neck

endometrium. The window of implantation is relatively

Fig. 4.2 Timeline of development related to implantation and week 2 of development.

As implantation of the blastocyst progresses, the primordi-

So numerous as are the instances of love shown by parents

Everyone who has paid even a passing attention to the habits of

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